Search

The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 are recorded from southern Africa for the first time, with the description of five new species: Leptodrassex murphyi sp. nov. (♂ ♀) from Mozambique and South Africa, and L. capensis sp. nov. (♀) from South Africa; Leptopilos butleri sp. nov. (♂ ♀) and L. vasivulva sp. nov. (♂ ♀) from Botswana, South Africa and Zimbabwe, and L. digitus sp. nov. (♂ ♀) from South Africa. Further, the new genus Afrodrassex gen. nov. is described, with the type species A. balrog sp. nov. (♂ ♀) from South Africa and Angola, and A. catharinae sp. nov. (♂ ♀) from South Africa described therein. Details of the somatic and genitalic morphology of all three genera are examined by scanning electron microscopy, and revised descriptions of Leptodrassex and Leptopilos are presented.

Bunyoronius Bonaldo, Ramírez & Haddad gen. nov. Type species: Bunyoronius femoralis sp. nov. Etymology. The generic name, masculine, is a contraction of the words Bunyoro and Carteronius, honoring the Bunyoro people, a pre-colonial Kingdom on the territory of modern Uganda, including the Budongo Forest, type locality of the type species. Diagnosis. Members of the genus Bunyoronius gen. nov. share with those of Carteronius the trilobulated RTA, with an apical spur inserted in the base of the ventral lobe, sperm duct with a single ventral fold, and the basally widened, curved embolus surrounding tegular margins. They can be readily recognized by large apical retrolateral femoral apophysis, ventral fold of sperm duct oriented retro-dorsally; median apophysis absent, pit vestigial; embolus inserted basally, with wide bifid tip (Figs 22A–C, 23A–C, F–I). Females can be recognized by the epigynal plate divided by two lateral, longitudinal ridges, forming a median atrium, and by the copulatory openings oriented anteriorly (Fig. 22D, E). Description. Medium-sized spiders, 6.03 7.70 mm in length. Carapace reddish-brown, surface finely granulate, with few long hairs; very broad, sub-oval, as long as wide, cephalic region weakly demarcated posteriorly, swollen antero-laterally, higher than thoracic region; thoracic region abruptly depressed, posteriorly rounded, thoracic fovea present (Fig. 20B). Clypeus low, height nearly one AME diameter. AER straight in frontal view, ALE oblique, eyes equidistant; PER slightly procurved in dorsal view, slightly wider than AER, eyes equidistant; AME largest, approximately two times ALE diameter, remaining eyes sub-equal in width, medians and PLE circular; ALE suboval. Chilum present, entire, with large median tubercle in both males and females. Chelicerae nearly as long as half the length of carapace, frontal surface granulate, slightly geniculated in both sexes, unmodified in males; basal boss evident, promargin with three teeth, retromargin with two teeth (Fig. 21A). Endites convergent, promargin slightly protruded anteriorly, retromargin slightly excavated; labium sub-squared, as long as wide, slightly longer than half endite length, proximal lateral constrictions shallow. Sternum shield-shaped, slightly longer than wide; surface covered by small hair-bearing tubercles, precoxal and intercoxal sclerites present, margins well defined, especially antero-laterally. Retrocoxal hymen present (Fig. 21B, C). Leg formula: I.II.V.III. Legs long, I–II sturdier; femur I (Fig. 21D) with one dorsal spine, tibia I (Fig. 21F) with seven pairs of ventral spines, metatarsus I with two pairs of ventral spines. Tarsus I with dorsal cluster of trichobothria; tarsal organ sub-apical (Fig. 21E). Abdomen oval, dorsal and ventral scuta absent. Spinnerets not surveyed with SEM; female (under light microscopy): PMS with three cylindrical gland spigots; PLS with two cylindrical gland spigots., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 367-368, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Carteronius gentilis (Simon, 1909) comb. nov. Figs 18, 19; Map 1 Procopius gentilis Simon, 1909: 382 (♂ holotype from “ Fernando Poo: Basilé”, leg. L. Fea, MNHN 22.254 —examined). Note. The male holotype, from Bioko Island, Equatorial Guinea, is here tentatively associated with a female from continental Africa (Cameroon). Bioko is a continental-shelf island that supports relatively low levels of endemism of angiosperms, bats, birds, reptiles, and amphibians when compared to oceanic islands of the Gulf of Guinea (Príncipe, São Tomé and Annobón) (Jones 2004). This island was separated from mainland Africa by the rise in sea level at the end of the last glacial period, approximately 10 000 years ago (Schabetsberger et al. 2004; PérezPérez & Yu 2021). Both the Bioko and Cameroon specimens share a strikingly similar dorsal abdominal pattern (Fig. 18A, F), as well as similarities in leg spination. Nevertheless, this association must be tested when additional samples come to light. From a strictly nomenclatural point of view, proposing this doubtful association is preferable to making available a possibly unnecessary specific name. Diagnosis. Males of C. gentilis comb. nov. differ from all other Carteronius species in having a bifid dorsal lobe on the RTA (Fig. 19A, B). Females resemble those of C. myene sp. nov. by the slightly curved transverse ridge (Figs 14A, 19C), but differ by the relatively small posterior sector and the extremely narrow copulatory duct in relation to the spermathecae (Fig. 19C, D). Description. Male (holotype). Measurements: Total length 8.03, CL 3.82, CW 3.57, AL 3.95, AW 2.80, SL 1.93, SW 1.81. Eye diameters and interdistances: AME 0.32, ALE 0.21, PLE 0.24, PME 0.26, AME-AME 0.21, AME-ALE 0.24, ALE-ALE 1.66, PME-PME 0.85, PME-PLE 0.52, PLE-PLE 2.07. Length of leg segments: I 4.13+ 1.74+4.05+3.10+1.22=14.24; II 4.02+1.70+3.95+3.20+1.26=14.13; III 2.61+1.26+2.18+2.15+1.00=9.20; IV 3.36+ 1.29+2.82+3.12+1.10=11.69. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-1 pl 0-1-0, II do 0-1-0, III do 0-1-2, IV do 0-1-2; tibiae: I ve 2-2-2-2-2-2, II ve 1 p -1 r -2-2-2-2, III ve 0-2-2-0, IV ve 0-0-0-1 r; metatarsi: I ve 0-2-2-0, II ve 0-2-2-0, III pl 0-1-1 rl 0-1-1 ve 2-2-1, IV pl 0-1-1 rl 0-1-1 ve 2-2-0. Coloration: carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs dark yellow, with femora I and II reddish-brown. Abdomen dorsally with yellowish area anteriorly and five transversal follicular bands in posterior half (Fig. 18A). Palp: RTA with apical spur approximately same size as ventral lobe, medial lobe short and pointed; ventral lobe sub-squared and excavated; sperm duct with long loop, retrolateral apical tegular process present, blunt; embolus long and thin, without projections (Figs 19A, B). Female (MRAC 162.128). Measurements: Total length 7.92, CL 3.26, CW 2.78, AL 4.62, AW 3.26, SL 1.57, SW 1.60. Eyes diameters and interdistances: AME 0.22, ALE 0.18, PLE 0.19, PME 0.19, AME-AME 0.63, AME-ALE 0.20, ALE-ALE 1.35, PME-PME 0.32, PME-PLE 0.38, PLE-PLE 1.60. Length of leg segments I 3.13+1.33+3.01+2.30+1.10=10.87; II 3.05+1.30+2.90+2.29+1.09=10.63; III 2.23+1.00+1.89+1.73+0.87=7.72; IV 2.74+1.06+2.30+2.40+0.93=9.43. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two teeth, inner tooth larger. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-2, IV do 0-1-2; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve 1 p -1 r -2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 p- 1 p -0-0; metatarsi: I ve 0-2- 2-0, II ve 0-2-2-0, III pl 1-0-0-1 rl 0-0-1 ve 2-2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 p- 1 p -1 r -1 r. Coloration: carapace and chelicerae dark reddish brown. Endites, labium and sternum orange red. Legs whitish with femora I and II reddish. Abdomen as in male (Fig. 18F). Epigynum: CDv long, straight, narrow; ST2 tappering, anteriorly located, gland ducts conspicuous, smaller than ST1; CDd large, S-shaped (Fig. 19D). Other material examined: CAMEROON: Mount Cameroun, near Buea, 04°12’N, 09°11’E, 1400 m.a.s.l., II-III.1981, Bosmans & Van Stalle leg. (montane forest with arable fields, V.H.F), 1♀ (MRAC 162.128). Distribution. Recorded from the island of Bioko (Equatorial Guinea) and Cameroon (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 364-367, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035, {"references":["Simon, E. (1909) Arachnides recueillis par L. Fea sur la cote occidentale d'Afrique. 2 e partie. Annali del Museo Civico di Storia Naturale di Genova, 44, 335 - 449.","Jones, P. J. (2004) Biodiversity in the Gulf of Guinea: an overview. Biodiversity & Conservation, 3, 772 - 784. https: // doi. org / 10.1007 / BF 00129657","Schabetsberger, R., Drozdowski, G., Drozdowski, I., Jersabek, C. D. & Rott, E. (2004) Limmnological aspects of two tropical crater lakes (Lago Biao and Lago Loreto) on the island of Bioko (Equatorial Guinea). Hydrobiologia, 524, 79 - 90. https: // doi. org / 10.1023 / B: HYDR. 0000036121.07007. e 8"]}

Carteronius myene Bonaldo & Labarque sp. nov. Figs 13A, B; 14A, B; Map 1 Type. ♀ holotype from Parc National de Moukalaba-Doudou, Département de Ndougou, Province de Ogooué Maritime (02°35’S, 10°14’E), GABON, III–IV.2003, O.S.G. Olivier & M. Burger leg. (forêt marecaguese) (MRAC 220.827). Etymology. The specific name is a noun in apposition referring to the Myene people, which settled fishing communities along the coast of Gabon. Diagnosis. Females of Carteronius myene sp. nov. resemble those of C. gentilis comb. nov. by the slightly curved epigynal transversal ridge (Figs 14A, 19C), but can be recognized by the posterior sector relatively larger; spermathecae barely visible by transparency in anterior sector (Fig. 14A). Description. Male. unknown. Female. (MRAC 220.827). Measurements: Total length 11.29, CL 5.29, CW 4.85, AL 6.33, AW 4.59, SL 2.64, SW 2.30. Eye diameters and interdistances: AME 0.42, ALE 0.34, PLE 0.29, PME 0.32, AME-AME 0.32, AME-ALE 0.37, ALE-ALE 2.32, PME-PME 0.39, PME-PLE 0.62, PLE-PLE 2.71. Length of leg segments: I 4.68+2.28+4.48+3.46+1.34=16.24; II 4.96+2.29+4.27+3.58+1.41=16.58; III 3.59+1.71+2.83+2. 81+1.00=11.29; IV 4.03+1.69+3.43+3.44+1.15=13.74. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, proximal tooth largest; Leg spination: femora: II do 0-1-0, III do 0-1-0 ve 0-1-0, IV do 1-0-0; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve 2-2, IV ve 1-0-1; metatarsi: I ve 2-2-2-2, II ve 1 p -1 p -0-1 r -2, III pl 0-0-0-1 rl 0-0-0-1 ve 2-2-1, IV pl 2-0-1-0 ve 0-1-0 -0. Coloration: Carapace and chelicerae dark reddish-brown. Endites, labium and sternum dark reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen dark gray, with several small white spots, two pairs of small white spots in middle dorsally, and three faint chevrons posteriorly. Venter gray, with irregular white spots forming two transversal lines (Fig. 13A). Epigynum: CDv long, gently arched, ST2 tapering, anteriorly located, gland ducts present, approximately same size as ST1; CDd folded ventrally (Fig. 14B). Other material examined. None. Distribution. Only known from Gabon (Map 1).

Carteronius simoni Bonaldo & Shimano sp. nov. Fig. 17; Map. 1 Holotype. ♀ from GABON: leg. Mocquery (No further data) (MNHN-17.436). Etymology. The specific name is a patronym honoring French arachnologist Eugène Simon, who described Carteronius. Diagnosis. Females of C. simoni sp. nov. are similar to those of C. lumumba sp. nov. by the inconspicuous lateral plates of the posterior epigynal sector (Figs 16D, 17C) but can be recognized by the wider, divergent copulatory ducts (Fig. 17C). Description. Male. unknown. Female. (MNHN-17.436) Measurements: Total length 9.15, CL 4.14, CW 3.55, AL 4.99, AW 4.03, SL 1.68, SW 1.82. Eye diameters and interdistances:AME 0.35, ALE 0.24, PLE 0.25, PME 0.30, AME-AME 0.30, AME-ALE 0.40, ALE-ALE 2.15, PME-PME 0.56, PME-PLE 0.61, PLE-PLE 2.60. Length of leg segments: I 2.80+1.47+2.58+2.10+1.09=10.04; II 2.95+1.40+2.48+2.12+0.96=9.91; III 2.35+1.09+1.77+1.76+0.78 =7.75; IV 2.80+1.14+2.43+2.27+0.87=9.51. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two subequal teeth. Leg spination: tibiae: I ve 2-2-2-2-2, II ve 2-2-2-2, IV ve 0-0-1 p; metatarsi: I ve 2-2, II ve 2-2, III ve 2-1 p -2, IV pl 0-0-0-1 rl 0-1-0-1 ve 1 p -1 r - 1 p. Coloration: carapace, chelicerae, endites, labium and sternum reddish. All leg segments yellowish-red. Abdomen pale gray dorsally, with indistinct white spots (Fig. 17A). Epigynum: CDv long, straight, ST2 a small globe, anteriorly located, gland ducts inconspicuous, smaller than ST1; CDd large, S-shaped (Fig. 17D). Other material examined. None. Distribution. Only known from Gabon (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on page 363, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Carteronius arboreus Bonaldo & Haddad sp. nov. Figs 7, 8; Map 1 Types. ♂ holotype from Bas Congo, Mayombe, Luki Forest Reserve (05°37’S, 13°05’E), 28.IX.2007, DEMOCRATIC REPUBLIC OF THE CONGO, D. de Bakker & J. P. Michiels leg. (sieving along trail in primary rainforest) (MRAC 223.475). ♀ paratype, same locality and collectors, 10.XI.2006 (Fog 3, primary rainforest) (MRAC 220.925). Etymology. The specific name is a Latin adjective meaning arboreal, in reference to the fact that most of the known specimens were collected by canopy fogging. Diagnosis. Males of Carteronius arboreus sp. nov. are similar to those of C. ashanti sp. nov. by the presence of a sub-apical embolar process (Figs 8A, 10A), but differ by the dorsal lobe of the RTA, which is broad and retrolaterally oriented, and the rounded medial lobe, which is small in relation to the dorsal lobe, and share the same base (Fig. 8B). In C. ashanti sp. nov., the dorsal lobe is spoon-shaped and the medial lobe longer and fang-shaped, with its own base. Females resemble those of C. sudanus comb. nov. in the strongly recurved epigynal transversal ridge (Figs 4C, 8C), but differ by the lateral plates of the epigynal posterior sector being strongly sclerotized (Fig. 8C). Description. Male. (MRAC 223475). Measurements: Total length 6.03, CL 3.16, CW 2.71, AL 2.95, AW 2.24, SL 1.39, SW 1.49. Eye diameters and interdistances: AME 0.28, ALE 0.17, PLE 0.18, PME 0.16, AME-AME 0.65, AME-ALE 0.16, ALE-ALE 1.34, PME-PME 0.33, PME-PLE 0.37, PLE-PLE 1.57. Length of leg segments: I 3.08+1.17+2.67+2.29+1.22=10.43; II 3.10+1.24+3.10+2.24+1.27=10.95; III 2.07+0.93+1.61+1.65+0.96=7.22; IV 2.55+0.94+2.11+2.25+1.05=8.90. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-0-1-0, II do 0-1-0, III do 0-1-0, IV do 0-1-0; tibiae: I ve 2-2-2-2-2, II ve 1 p -1 r -1 p -2-2-2, III ve 1 r -1 p -2, IV rl 0-1-0-1 ve 1 p -0-1 p; metatarsi: I ve 2-2, II ve 2-2, III pl 0-1-0-1 rl 0-1-0-1 ve 2-2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 r -1 p -1 r -0. Coloration: carapace and chelicerae reddish-brown. Endites, labium and sternum reddish-brown. Legs I and II: coxae and trochanters reddish-brown; femora reddish-brown, yellowish distally; tibiae, metatarsi and tarsi yellowish. Legs III and IV yellowish. Abdomen dark gray dorsally, with two well defined white bands; white ventrally (Fig. 7A). Palp: RTA with apical spur short, curved and pointed, dorsal lobe with apical edges bent ventrally, ventral lobe rounded and excavated. Tegulum with short retrolateral apical tegular process, spermatic duct with long loop (Fig. 8A, B). Female. (MRAC 220925). Measurements: Total length 10.58, CL 4.10, CW 3.86, AL 6.37, AW 4.91, SL 1.97, SW 1.93. Eye diameters and interdistances: AME 0.31, ALE 0.21, PLE 0.21, PME 0.18, AME-AME 0.25, AME-ALE 0.29, ALE-ALE 1.75, PME-PME 0.45, PME-PLE 0.49, PLE-PLE 2.13. Length of leg segments: I 3.87+1.81+3.70+2.88+1.28=13.54; II 3.70+1.73+3.66+2.99+1.31=13.39; III 2.75+1.32+2.17+2.08+0.91=9.23; IV 3.25+1.31+2.76+2.84+0.98=11.14. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-0-1 p, IV do 0-1-0-1 p; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-1 r -1 p -1 r -1 p -1 r, III ve 1 p -1 r -2, IV rl 0-1-0-1 ve 1p-0-1 p; metatarsi: I ve 2-0-2-0, II ve 2-2-2, III pl 0-1-0-1 rl 0-0-0-1 ve 2-2-1, IV pl 0-1-0-1 rl 1-0-1 ve 1 p -1 r -1 p - 0-0. Coloration: Carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen gray, dorsum with scattered small white spots, denser in middle, forming irregular white longitudinal band; posteriorly with large triangular white spot. Ventrally gray with two irregular lateroventral white bands (Fig. 7C). Epigynum: CDv slight folded posteriorly, ST2 globose, anteriorly located, larger than ST1, CDd almost straight (Fig. 8D). Other material examined. DEMOCRATIC REPUBLIC OF THE CONGO: Bas Congo, Mayombe, Luki Forest Reserve (05°37’S, 13°05’E), 18.IX.2007, D. de Bakker & J.P. Michiels leg. (Fog 5, old secondary forest), 1♀ (MRAC); same data but 22.IX.2007, 1♀ (MRAC); same data but 30.IX.2007, 1♂ (MRAC). Distribution. Only known from the Democratic Republic of the Congo (Map 1).

Carteronius ashanti Bonaldo & Silva-Junior sp. nov. Figs 9 – 12; Map 1 Types. ♂ holotype from Kakum National Park, Kakum Forest (05°20’N, 01°23’W), GHANA, 17.XI.2005, R. Jocqué, D. de Bakker & L. Baert leg. (secondary forest, fogging), (MRAC 218261). ♂ paratype, same data (MRAC). Etymology. The specific name is a noun in apposition referring to the Ashanti people, a major ethnic group of modern Ghana. Diagnosis. Males of Carteronius ashanti sp. nov. resemble those of C. arboreus sp. nov. by the presence of a sub-apical embolar process (Figs 8A, 10A), but can be recognized by the long, narrow, upwardly oriented dorsal lobe of the RTA, by the prominent retrolateral apical tegular process and by the long, pointed apical spur in relation to the dorsal lobe (Figs 10A, B, 12A, B, D, F). Description. Male. (MRAC 218.261). Measurements: Total length 7.42, CL 3.60, CW 3.29, AL 3.80, AW 2.11, SL 1.61, SW 1.56. Eye diameters and interdistances: AME 0.23, ALE 0.16, PLE 0.19, PME 0.18, AME-AME 0.24, AME-ALE 0.21, ALE-ALE 1.54, PME-PME 0.45, PME-PLE 0.44, PLE-PLE 1.85. Length of leg segments: I 3.39+1.51+3.47+2.89+1.18=12.37; II 3.57+1.54+3.41+2.96+1.30=12.78; III 2.34+1.04+1.88+1.96+0.83=8.05; IV 2.82+1.07+2.47+2.47+0.91=9.74. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-0, IV do 0-1-0; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-2-2-2, III ve p 1- r 1-2, IV pl 1-0-1 rl 0-1-1; metatarsi: I ve 2-2, II ve 2-2, III pl 0-1-0 vl 1-0-0 ve 2-2, IV pl 0-1-0 rl 0-1-0 ve 2-2. Coloration: carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen gray, dorsum with scattered small white spots, denser in middle, forming large irregular white longitudinal bands. Venter gray, with irregular white spots (Fig. 9A). Palp: RTA with ventral lobe sub-rectangular, without excavations, medial lobe narrow, as long as dorsal lobe. Sperm duct with loop restricted to middle of tegulum. Embolus with small subapical embolar process (Figs 10A, B, 12A–I). Female. unknown. Other material examined. None. Distribution. Only known from Ghana (Map 1).

Carteronius ashanti Bonaldo & Silva-Junior sp. nov. Figs 9 – 12; Map 1 Types. ♂ holotype from Kakum National Park, Kakum Forest (05°20’N, 01°23’W), GHANA, 17.XI.2005, R. Jocqué, D. de Bakker & L. Baert leg. (secondary forest, fogging), (MRAC 218261). ♂ paratype, same data (MRAC). Etymology. The specific name is a noun in apposition referring to the Ashanti people, a major ethnic group of modern Ghana. Diagnosis. Males of Carteronius ashanti sp. nov. resemble those of C. arboreus sp. nov. by the presence of a sub-apical embolar process (Figs 8A, 10A), but can be recognized by the long, narrow, upwardly oriented dorsal lobe of the RTA, by the prominent retrolateral apical tegular process and by the long, pointed apical spur in relation to the dorsal lobe (Figs 10A, B, 12A, B, D, F). Description. Male. (MRAC 218.261). Measurements: Total length 7.42, CL 3.60, CW 3.29, AL 3.80, AW 2.11, SL 1.61, SW 1.56. Eye diameters and interdistances: AME 0.23, ALE 0.16, PLE 0.19, PME 0.18, AME-AME 0.24, AME-ALE 0.21, ALE-ALE 1.54, PME-PME 0.45, PME-PLE 0.44, PLE-PLE 1.85. Length of leg segments: I 3.39+1.51+3.47+2.89+1.18=12.37; II 3.57+1.54+3.41+2.96+1.30=12.78; III 2.34+1.04+1.88+1.96+0.83=8.05; IV 2.82+1.07+2.47+2.47+0.91=9.74. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-0, IV do 0-1-0; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-2-2-2, III ve p 1- r 1-2, IV pl 1-0-1 rl 0-1-1; metatarsi: I ve 2-2, II ve 2-2, III pl 0-1-0 vl 1-0-0 ve 2-2, IV pl 0-1-0 rl 0-1-0 ve 2-2. Coloration: carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen gray, dorsum with scattered small white spots, denser in middle, forming large irregular white longitudinal bands. Venter gray, with irregular white spots (Fig. 9A). Palp: RTA with ventral lobe sub-rectangular, without excavations, medial lobe narrow, as long as dorsal lobe. Sperm duct with loop restricted to middle of tegulum. Embolus with small subapical embolar process (Figs 10A, B, 12A–I). Female. unknown. Other material examined. None. Distribution. Only known from Ghana (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 356-358, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Eb.Bonaldo, Ramírez, Martín J., Om.Labarque, Shimano, Yulie, Silva-Junior, Cláudio J., Haddad, Charles R. (2022): Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group. Zootaxa 5205 (4): 343-373, DOI: https://doi.org/10.11646/zootaxa.5205.4.3

Carteronius simoni Bonaldo & Shimano sp. nov. Fig. 17; Map. 1 Holotype. ♀ from GABON: leg. Mocquery (No further data) (MNHN-17.436). Etymology. The specific name is a patronym honoring French arachnologist Eugène Simon, who described Carteronius. Diagnosis. Females of C. simoni sp. nov. are similar to those of C. lumumba sp. nov. by the inconspicuous lateral plates of the posterior epigynal sector (Figs 16D, 17C) but can be recognized by the wider, divergent copulatory ducts (Fig. 17C). Description. Male. unknown. Female. (MNHN-17.436) Measurements: Total length 9.15, CL 4.14, CW 3.55, AL 4.99, AW 4.03, SL 1.68, SW 1.82. Eye diameters and interdistances:AME 0.35, ALE 0.24, PLE 0.25, PME 0.30, AME-AME 0.30, AME-ALE 0.40, ALE-ALE 2.15, PME-PME 0.56, PME-PLE 0.61, PLE-PLE 2.60. Length of leg segments: I 2.80+1.47+2.58+2.10+1.09=10.04; II 2.95+1.40+2.48+2.12+0.96=9.91; III 2.35+1.09+1.77+1.76+0.78 =7.75; IV 2.80+1.14+2.43+2.27+0.87=9.51. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two subequal teeth. Leg spination: tibiae: I ve 2-2-2-2-2, II ve 2-2-2-2, IV ve 0-0-1 p; metatarsi: I ve 2-2, II ve 2-2, III ve 2-1 p -2, IV pl 0-0-0-1 rl 0-1-0-1 ve 1 p -1 r - 1 p. Coloration: carapace, chelicerae, endites, labium and sternum reddish. All leg segments yellowish-red. Abdomen pale gray dorsally, with indistinct white spots (Fig. 17A). Epigynum: CDv long, straight, ST2 a small globe, anteriorly located, gland ducts inconspicuous, smaller than ST1; CDd large, S-shaped (Fig. 17D). Other material examined. None. Distribution. Only known from Gabon (Map 1).

Carteronius sudanus (Karsch, 1880) comb. nov. Figs 2 – 6; Map 1 Mandane sudana Karsch, 1880: 377 (♂ holotype from Adafoah, Volta River Basin, Ghana, Ungar leg., ZMB 2143, examined). Carteronius helluo Simon, 1896: 400 (♂ holotype from Freetown, Sierra Leone, MNHN 10611, accidentally switched with the male holotype of Carteronius scriptus Simon, 1896, from Diego Soares, Madagascar, MNHN 14.625). syn. nov. Mandaneta sudana; Strand 1932: 140; Haddad & Bosselaers 2010: 7, figs 16–19; Ramírez 2014: 365, figs 20A, 69D. Medmassa laurenti Lessert, 1946: 211, fig. 10 (♂ lectotype and ♀ paralectotype from Eala, the Democratic Republic of the Congo, leg. H. J. Bredo, MRAC 12419-12420, one palp of the ♂ lectotype in Museum of Natural History, Geneva, not re-examined). Synonymized with Mandaneta sudana by Haddad & Bosselaers 2010: 7. Diagnosis. Males of C. sudanus comb. nov. resemble those of C. lumumba sp. nov. by the absence of a retrolateral apical tegular process and by the bifid embolus tip (Figs 4A, 16B), but differ by the chelicerae with a frontal anterior excavation (Fig. 2A), the wider proximal half of embolus, and by the short distal loop of the spermatic duct, which is restricted to the middle of the bulb (Fig. 4A). Females resemble those of C. arboreus sp. nov. by the strongly recurved epigynal transversal ridge (Figs 4C, 8C), but differ by the lateral plates of the posterior sector not being sclerotized (Fig. 4C). Description. Male. (MRAC MT 207.386). Measurements: Total length 9.21, CL 4.52, CW 4.00, AL 4.69, AW 2.72, SL 2.20, SW 1.94. Eye diameters and interdistances: AME 0.30, ALE 0.19, PLE 0.18, PME 0.23, AME-AME 0.32, AME-ALE 0.36, ALE-ALE 1.62, PME-PME 0.52, PME-PLE 0.62, PLE-PLE 2.18. Length of leg segments: I 4.36+1.86+4.44+3.52+1.62=15.80; II 4.52+2.04+4.56+3.60+1.58=16.30; III 3.16+1.36+2.84+2.36+1.18=10.90; IV 3.68+1.48+3.52+3.52+1.22=13.42. Chelicerae: promargin with three spaced teeth, median tooth largest, distal tooth smallest; retromargin with two teeth, subequal in size. Leg spination: femora: I do 0-0-1-0 pl 0-0-1, II do 0- 1-0, III - IV do 0-1-0-1; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve p 1- r 1-2, IV pl 1-0-1-1 rl 0-1-1; metatarsi: I-II ve 2-2, III pl 1-1-0 lv 0-1-0 ve 2-2, IV pl 0-1-0 lv 0-1-0 ve 2-2. Coloration: carapace and chelicerae reddish-brown. Endites and labium reddish-brown, sternum yellowish. Legs I and II: coxae and trochanters reddish-brown; femora reddish-brown, yellowish distally; tibiae, metatarsi and tarsi yellowish. Legs III and IV: coxae whitish, trochanters brownish; femora whitish prolaterally and retrolaterally, brownish ventrally and distally; patellae, tibiae, metatarsi and tarsi brownish. Abdomen dark gray dorsally, with diamond-shaped spot in middle, followed by four white transversal lines; white ventrally (Fig 3A – C). Palp: RTA with apical spur short, pointed; dorsal lobe long, excavated, with folded edges and oriented upwards; medial lobe smaller than dorsal lobe, pointed and oriented upwards; ventral lobe sub-squared, not excavated. Sperm duct with short loop (Fig 4A, B). Female. (MRAC MT 204.306–MRAC MT 177.640). Measurements: Total length 9.65-10.18, CL 4.32-4.28, CW 3.80-4.00.0, AL 5.33-5.90, AW 3.68-4.52, SL 2.08-2.14, SW 1.76-1.96. Eye diameters and interdistances: AME-AME 0.28, AME-ALE 0.32, ALE-ALE 1.56, PME-PME 0.44, PME-PLE 0.36, PLE-PLE 2.02. Length of leg segments: I 3.88+1.86+4.00+3.04+1.4=14.18; II 3.92+1.86+4.04+3.16+1.44=14.42; III 3.00+1.42+2.52+2.24+ 1.20=10.38; IV 3.56+1.48+3.28+3.20+1.30=12.82. Chelicerae with three spaced teeth on promargin, median tooth largest, distal tooth smallest; two teeth on retromargin, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-0-1, II do 0-1-0, III - IV do 0-1-0-1; tibiae: I-II ve 2-2-2-2-2-2, III ve 1 r -1 p -2, IV ve pl 0-1-0-1 rl 0-1-0-1; metatarsi: I pl 1-0-0 ve 2-2, II rl 1-0-0 ve 2-2, III pl 0-1-0-1 rl 0-1-0-1 ve 2-2, IV plv 4 rlv 3 pl 0-1-0 rl 0-1-0 ve 1 p -2-2. Coloration: carapace and chelicera dark red; endites, labium and sternum reddish. Legs I and II: coxae and trochanter reddish brown; femora dark red; tibia, metatarsus and tarsus reddish. Legs III and IV: coxae whitish, trochanter brownish; femur whitish prolaterally and retrolaterally, brownish ventrally and distally; tibia, metatarsus and tarsus brownish. Abdomen: pale gray with several spots dorsally, present two pair of white spots and four white transversal lines well defined dorsally (Fig. 3D – F). Epigynum: CDv long, slender, ST2 anteriorly positioned in relation to the ST1, similarly sized to ST1, with conspicuous gland ducts; CDd S-shaped (Figs 4D, 5C, D). Other material examined: CÔTE D’IVOIRE: Mankono, Ranch de la Marahoué, 08°27’N, 06°52’W, 12. III.1980, leg. J. Everts, riverine forest, 1♂ (MRAC 173.984); Bettié, forêt classeé de Mabi, 06°05’N, 03°30’W, dense forest, by hand, 26.XI.1993, R. Jocqué, leg. 1♀ (MRAC 177.640; SEM preparations MJR 574–576); same locality, Eco. Grappe 3, 24. III.1997, T. Steyn leg., 1♀ (MRAC 207.387); Appouesso, FC Bossematié, Forest, pitfall, station 1B, 12.II.1995, R. Jocqué and R. Tanoh leg., 1♀ (MRAC 204.306); same locality, station 5, found in leaf litter, 21. III.1997, T. Steyn leg., 1♂ (MRAC 207.386). DEMOCRATIC REPUBLIC OF THE CONGO: Eyolo Forest, ca. 2 km E of Lieki, 00°41.785’N, 24°14.512’E, 25–29. V.2010, A.H. Kirk-Spriggs leg. (Malaise traps, lowland evergreen swamp forest), 1♀ (IRSNB IG.34481). GUINEA: Mount Nimba, Gallery Forest of Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 3.X.2011, D. van den Spiegel & A. Henrard leg. (fogging 1, canopy of trees, understory of shrub layer), 1♂ (MRAC 238.050); same locality, Station de pompage Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 11.X.2011, D. van den Spiegel & A. Henrard leg. (sieving litter under “matete” [high grass], near route, open area), 2♂ (MRAC 237.965); same locality, Station de pompage Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 1.X.2011, D. van den Spiegel & A. Henrard leg. (sieving litter under “matete” [high grass], near route, open area), 1♂ (MRAC 237.984); same locality, Gba Valley, 07°40’N, 08°23’W, 880 m. a.s.l., 9.X.2011, D. van den Spiegel & A. Henrard leg. (beating, primary gallery forest, litter in trees and shrubs, at 1.5-3m above the floor, “chablis”), 2♂ (MRAC 238.090); same locality, Ziela, near Pierré-Richaud, 0742’N, 0821’W, 568 m. a.s.l., 20.II.2012, A. Henrard, C. Allard, P. Bimou & M. Sidibé leg. (sieving litter), 1♂ (MRAC 238.697). Distribution. Widespread across equatorial West and Central Africa (Map 1).

Carteronius teke Bonaldo & Bosselaers sp. nov. Figs 13C, D, 14C, D; Map 1 Type. ♀ holotype from Kivu, Rwankwi [01°19’S, 29°22’E], DEMOCRATIC REPUBLIC OF THE CONGO, VI.1946, J. Leroy leg. (MRAC 168.643). Etymology. The specific name is a noun in apposition referring to the Teke people, one of the three kingdoms that ruled Congo before the arrival of Europeans. Diagnosis. Females of Carteronius teke sp. nov. differ from all other species of Carteronius by the presence of completely straight epigynal transversal ridge (Fig. 14C). Description. Male. Unknown. Female. (MRAC 168.643) Measurements:Total length 13.45, CL 4.90, CW 4.39, AL 7.99, AW 5.24, SL 2.44, SW 2.15. Eye diameters and interdistances:AME 0.38, ALE 0.25, PLE 0.28, PME 0.25, AME-AME 0.36, AME-ALE 0.40, ALE-ALE 2.30, PME-PME 0.57, PME-PLE 0.67, PLE-PLE 2.81. Length of leg segments (sequence from femur to tarsus, and total): I 4.21+2.04+3.57+2.92+1.25=13.99; II 4.15+1.86+3.89+2.79 +1.19=13.88; III 2.83+1.43+2.52+2.20+1.04=10.02; IV 3.42+1.42+3.03+3.14+1.28=12.29. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: IV do 1-0-0; tibiae: I ve 1 p -2-2-2-2-2, II ve 0-0-2-2-2-2, III ve 0-0-1 p -0, IV ve 1 p -0-1 p -0; metatarsi: I ve 2-2-2-2, II ve 1 r -1 p -2-2, III ve 0-2-2-0, IV ve 0-2-2-0. Coloration: carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs I and II reddish-brown, III and IV dark yellow. Abdomen dorsally pale gray, ventrally white with darker band converging at spinnerets (Fig. 13C). Epigynum: CDv long, arched, ST2 tapering, medially located, gland ducts inconspicuous, much smaller than ST1; CDd folded ventrally (Fig. 14D). Other material examined. None. Distribution. Only known from the Democratic Republic of the Congo (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 359-361, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Carteronius gentilis (Simon, 1909) comb. nov. Figs 18, 19; Map 1 Procopius gentilis Simon, 1909: 382 (♂ holotype from “ Fernando Poo: Basilé”, leg. L. Fea, MNHN 22.254 —examined). Note. The male holotype, from Bioko Island, Equatorial Guinea, is here tentatively associated with a female from continental Africa (Cameroon). Bioko is a continental-shelf island that supports relatively low levels of endemism of angiosperms, bats, birds, reptiles, and amphibians when compared to oceanic islands of the Gulf of Guinea (Príncipe, São Tomé and Annobón) (Jones 2004). This island was separated from mainland Africa by the rise in sea level at the end of the last glacial period, approximately 10 000 years ago (Schabetsberger et al. 2004; PérezPérez & Yu 2021). Both the Bioko and Cameroon specimens share a strikingly similar dorsal abdominal pattern (Fig. 18A, F), as well as similarities in leg spination. Nevertheless, this association must be tested when additional samples come to light. From a strictly nomenclatural point of view, proposing this doubtful association is preferable to making available a possibly unnecessary specific name. Diagnosis. Males of C. gentilis comb. nov. differ from all other Carteronius species in having a bifid dorsal lobe on the RTA (Fig. 19A, B). Females resemble those of C. myene sp. nov. by the slightly curved transverse ridge (Figs 14A, 19C), but differ by the relatively small posterior sector and the extremely narrow copulatory duct in relation to the spermathecae (Fig. 19C, D). Description. Male (holotype). Measurements: Total length 8.03, CL 3.82, CW 3.57, AL 3.95, AW 2.80, SL 1.93, SW 1.81. Eye diameters and interdistances: AME 0.32, ALE 0.21, PLE 0.24, PME 0.26, AME-AME 0.21, AME-ALE 0.24, ALE-ALE 1.66, PME-PME 0.85, PME-PLE 0.52, PLE-PLE 2.07. Length of leg segments: I 4.13+ 1.74+4.05+3.10+1.22=14.24; II 4.02+1.70+3.95+3.20+1.26=14.13; III 2.61+1.26+2.18+2.15+1.00=9.20; IV 3.36+ 1.29+2.82+3.12+1.10=11.69. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-1 pl 0-1-0, II do 0-1-0, III do 0-1-2, IV do 0-1-2; tibiae: I ve 2-2-2-2-2-2, II ve 1 p -1 r -2-2-2-2, III ve 0-2-2-0, IV ve 0-0-0-1 r; metatarsi: I ve 0-2-2-0, II ve 0-2-2-0, III pl 0-1-1 rl 0-1-1 ve 2-2-1, IV pl 0-1-1 rl 0-1-1 ve 2-2-0. Coloration: carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs dark yellow, with femora I and II reddish-brown. Abdomen dorsally with yellowish area anteriorly and five transversal follicular bands in posterior half (Fig. 18A). Palp: RTA with apical spur approximately same size as ventral lobe, medial lobe short and pointed; ventral lobe sub-squared and excavated; sperm duct with long loop, retrolateral apical tegular process present, blunt; embolus long and thin, without projections (Figs 19A, B). Female (MRAC 162.128). Measurements: Total length 7.92, CL 3.26, CW 2.78, AL 4.62, AW 3.26, SL 1.57, SW 1.60. Eyes diameters and interdistances: AME 0.22, ALE 0.18, PLE 0.19, PME 0.19, AME-AME 0.63, AME-ALE 0.20, ALE-ALE 1.35, PME-PME 0.32, PME-PLE 0.38, PLE-PLE 1.60. Length of leg segments I 3.13+1.33+3.01+2.30+1.10=10.87; II 3.05+1.30+2.90+2.29+1.09=10.63; III 2.23+1.00+1.89+1.73+0.87=7.72; IV 2.74+1.06+2.30+2.40+0.93=9.43. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two teeth, inner tooth larger. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-2, IV do 0-1-2; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve 1 p -1 r -2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 p- 1 p -0-0; metatarsi: I ve 0-2- 2-0, II ve 0-2-2-0, III pl 1-0-0-1 rl 0-0-1 ve 2-2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 p- 1 p -1 r -1 r. Coloration: carapace and chelicerae dark reddish brown. Endites, labium and sternum orange red. Legs whitish with femora I and II reddish. Abdomen as in male (Fig. 18F). Epigynum: CDv long, straight, narrow; ST2 tappering, anteriorly located, gland ducts conspicuous, smaller than ST1; CDd large, S-shaped (Fig. 19D). Other material examined: CAMEROON: Mount Cameroun, near Buea, 04°12’N, 09°11’E, 1400 m.a.s.l., II-III.1981, Bosmans & Van Stalle leg. (montane forest with arable fields, V.H.F), 1♀ (MRAC 162.128). Distribution. Recorded from the island of Bioko (Equatorial Guinea) and Cameroon (Map 1).

Carteronius myene Bonaldo & Labarque sp. nov. Figs 13A, B; 14A, B; Map 1 Type. ♀ holotype from Parc National de Moukalaba-Doudou, Département de Ndougou, Province de Ogooué Maritime (02°35’S, 10°14’E), GABON, III–IV.2003, O.S.G. Olivier & M. Burger leg. (forêt marecaguese) (MRAC 220.827). Etymology. The specific name is a noun in apposition referring to the Myene people, which settled fishing communities along the coast of Gabon. Diagnosis. Females of Carteronius myene sp. nov. resemble those of C. gentilis comb. nov. by the slightly curved epigynal transversal ridge (Figs 14A, 19C), but can be recognized by the posterior sector relatively larger; spermathecae barely visible by transparency in anterior sector (Fig. 14A). Description. Male. unknown. Female. (MRAC 220.827). Measurements: Total length 11.29, CL 5.29, CW 4.85, AL 6.33, AW 4.59, SL 2.64, SW 2.30. Eye diameters and interdistances: AME 0.42, ALE 0.34, PLE 0.29, PME 0.32, AME-AME 0.32, AME-ALE 0.37, ALE-ALE 2.32, PME-PME 0.39, PME-PLE 0.62, PLE-PLE 2.71. Length of leg segments: I 4.68+2.28+4.48+3.46+1.34=16.24; II 4.96+2.29+4.27+3.58+1.41=16.58; III 3.59+1.71+2.83+2. 81+1.00=11.29; IV 4.03+1.69+3.43+3.44+1.15=13.74. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, proximal tooth largest; Leg spination: femora: II do 0-1-0, III do 0-1-0 ve 0-1-0, IV do 1-0-0; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve 2-2, IV ve 1-0-1; metatarsi: I ve 2-2-2-2, II ve 1 p -1 p -0-1 r -2, III pl 0-0-0-1 rl 0-0-0-1 ve 2-2-1, IV pl 2-0-1-0 ve 0-1-0 -0. Coloration: Carapace and chelicerae dark reddish-brown. Endites, labium and sternum dark reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen dark gray, with several small white spots, two pairs of small white spots in middle dorsally, and three faint chevrons posteriorly. Venter gray, with irregular white spots forming two transversal lines (Fig. 13A). Epigynum: CDv long, gently arched, ST2 tapering, anteriorly located, gland ducts present, approximately same size as ST1; CDd folded ventrally (Fig. 14B). Other material examined. None. Distribution. Only known from Gabon (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 358-359, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Carteronius sudanus (Karsch, 1880) comb. nov. Figs 2 – 6; Map 1 Mandane sudana Karsch, 1880: 377 (♂ holotype from Adafoah, Volta River Basin, Ghana, Ungar leg., ZMB 2143, examined). Carteronius helluo Simon, 1896: 400 (♂ holotype from Freetown, Sierra Leone, MNHN 10611, accidentally switched with the male holotype of Carteronius scriptus Simon, 1896, from Diego Soares, Madagascar, MNHN 14.625). syn. nov. Mandaneta sudana; Strand 1932: 140; Haddad & Bosselaers 2010: 7, figs 16–19; Ramírez 2014: 365, figs 20A, 69D. Medmassa laurenti Lessert, 1946: 211, fig. 10 (♂ lectotype and ♀ paralectotype from Eala, the Democratic Republic of the Congo, leg. H. J. Bredo, MRAC 12419-12420, one palp of the ♂ lectotype in Museum of Natural History, Geneva, not re-examined). Synonymized with Mandaneta sudana by Haddad & Bosselaers 2010: 7. Diagnosis. Males of C. sudanus comb. nov. resemble those of C. lumumba sp. nov. by the absence of a retrolateral apical tegular process and by the bifid embolus tip (Figs 4A, 16B), but differ by the chelicerae with a frontal anterior excavation (Fig. 2A), the wider proximal half of embolus, and by the short distal loop of the spermatic duct, which is restricted to the middle of the bulb (Fig. 4A). Females resemble those of C. arboreus sp. nov. by the strongly recurved epigynal transversal ridge (Figs 4C, 8C), but differ by the lateral plates of the posterior sector not being sclerotized (Fig. 4C). Description. Male. (MRAC MT 207.386). Measurements: Total length 9.21, CL 4.52, CW 4.00, AL 4.69, AW 2.72, SL 2.20, SW 1.94. Eye diameters and interdistances: AME 0.30, ALE 0.19, PLE 0.18, PME 0.23, AME-AME 0.32, AME-ALE 0.36, ALE-ALE 1.62, PME-PME 0.52, PME-PLE 0.62, PLE-PLE 2.18. Length of leg segments: I 4.36+1.86+4.44+3.52+1.62=15.80; II 4.52+2.04+4.56+3.60+1.58=16.30; III 3.16+1.36+2.84+2.36+1.18=10.90; IV 3.68+1.48+3.52+3.52+1.22=13.42. Chelicerae: promargin with three spaced teeth, median tooth largest, distal tooth smallest; retromargin with two teeth, subequal in size. Leg spination: femora: I do 0-0-1-0 pl 0-0-1, II do 0- 1-0, III - IV do 0-1-0-1; tibiae: I ve 2-2-2-2-2-2, II ve 2-2-2-2-2, III ve p 1- r 1-2, IV pl 1-0-1-1 rl 0-1-1; metatarsi: I-II ve 2-2, III pl 1-1-0 lv 0-1-0 ve 2-2, IV pl 0-1-0 lv 0-1-0 ve 2-2. Coloration: carapace and chelicerae reddish-brown. Endites and labium reddish-brown, sternum yellowish. Legs I and II: coxae and trochanters reddish-brown; femora reddish-brown, yellowish distally; tibiae, metatarsi and tarsi yellowish. Legs III and IV: coxae whitish, trochanters brownish; femora whitish prolaterally and retrolaterally, brownish ventrally and distally; patellae, tibiae, metatarsi and tarsi brownish. Abdomen dark gray dorsally, with diamond-shaped spot in middle, followed by four white transversal lines; white ventrally (Fig 3A – C). Palp: RTA with apical spur short, pointed; dorsal lobe long, excavated, with folded edges and oriented upwards; medial lobe smaller than dorsal lobe, pointed and oriented upwards; ventral lobe sub-squared, not excavated. Sperm duct with short loop (Fig 4A, B). Female. (MRAC MT 204.306–MRAC MT 177.640). Measurements: Total length 9.65-10.18, CL 4.32-4.28, CW 3.80-4.00.0, AL 5.33-5.90, AW 3.68-4.52, SL 2.08-2.14, SW 1.76-1.96. Eye diameters and interdistances: AME-AME 0.28, AME-ALE 0.32, ALE-ALE 1.56, PME-PME 0.44, PME-PLE 0.36, PLE-PLE 2.02. Length of leg segments: I 3.88+1.86+4.00+3.04+1.4=14.18; II 3.92+1.86+4.04+3.16+1.44=14.42; III 3.00+1.42+2.52+2.24+ 1.20=10.38; IV 3.56+1.48+3.28+3.20+1.30=12.82. Chelicerae with three spaced teeth on promargin, median tooth largest, distal tooth smallest; two teeth on retromargin, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-0-1, II do 0-1-0, III - IV do 0-1-0-1; tibiae: I-II ve 2-2-2-2-2-2, III ve 1 r -1 p -2, IV ve pl 0-1-0-1 rl 0-1-0-1; metatarsi: I pl 1-0-0 ve 2-2, II rl 1-0-0 ve 2-2, III pl 0-1-0-1 rl 0-1-0-1 ve 2-2, IV plv 4 rlv 3 pl 0-1-0 rl 0-1-0 ve 1 p -2-2. Coloration: carapace and chelicera dark red; endites, labium and sternum reddish. Legs I and II: coxae and trochanter reddish brown; femora dark red; tibia, metatarsus and tarsus reddish. Legs III and IV: coxae whitish, trochanter brownish; femur whitish prolaterally and retrolaterally, brownish ventrally and distally; tibia, metatarsus and tarsus brownish. Abdomen: pale gray with several spots dorsally, present two pair of white spots and four white transversal lines well defined dorsally (Fig. 3D – F). Epigynum: CDv long, slender, ST2 anteriorly positioned in relation to the ST1, similarly sized to ST1, with conspicuous gland ducts; CDd S-shaped (Figs 4D, 5C, D). Other material examined: CÔTE D’IVOIRE: Mankono, Ranch de la Marahoué, 08°27’N, 06°52’W, 12. III.1980, leg. J. Everts, riverine forest, 1♂ (MRAC 173.984); Bettié, forêt classeé de Mabi, 06°05’N, 03°30’W, dense forest, by hand, 26.XI.1993, R. Jocqué, leg. 1♀ (MRAC 177.640; SEM preparations MJR 574–576); same locality, Eco. Grappe 3, 24. III.1997, T. Steyn leg., 1♀ (MRAC 207.387); Appouesso, FC Bossematié, Forest, pitfall, station 1B, 12.II.1995, R. Jocqué and R. Tanoh leg., 1♀ (MRAC 204.306); same locality, station 5, found in leaf litter, 21. III.1997, T. Steyn leg., 1♂ (MRAC 207.386). DEMOCRATIC REPUBLIC OF THE CONGO: Eyolo Forest, ca. 2 km E of Lieki, 00°41.785’N, 24°14.512’E, 25–29. V.2010, A.H. Kirk-Spriggs leg. (Malaise traps, lowland evergreen swamp forest), 1♀ (IRSNB IG.34481). GUINEA: Mount Nimba, Gallery Forest of Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 3.X.2011, D. van den Spiegel & A. Henrard leg. (fogging 1, canopy of trees, understory of shrub layer), 1♂ (MRAC 238.050); same locality, Station de pompage Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 11.X.2011, D. van den Spiegel & A. Henrard leg. (sieving litter under “matete” [high grass], near route, open area), 2♂ (MRAC 237.965); same locality, Station de pompage Zié, 07°40’N, 08°22’W, 1250 m.a.s.l., 1.X.2011, D. van den Spiegel & A. Henrard leg. (sieving litter under “matete” [high grass], near route, open area), 1♂ (MRAC 237.984); same locality, Gba Valley, 07°40’N, 08°23’W, 880 m. a.s.l., 9.X.2011, D. van den Spiegel & A. Henrard leg. (beating, primary gallery forest, litter in trees and shrubs, at 1.5-3m above the floor, “chablis”), 2♂ (MRAC 238.090); same locality, Ziela, near Pierré-Richaud, 0742’N, 0821’W, 568 m. a.s.l., 20.II.2012, A. Henrard, C. Allard, P. Bimou & M. Sidibé leg. (sieving litter), 1♂ (MRAC 238.697). Distribution. Widespread across equatorial West and Central Africa (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 349-353, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035, {"references":["Karsch, F. (1880) Arachnologische Blatter (Decas I). Zeitschrift fur die gesammten Naturwissenschaft, 53, 373 - 409.","Simon, E. (1896) Descriptions d'arachnides nouveaux de la famille des Clubionidae. Annales de la Societe Entomologique de Belgique, 40 (9), 400 - 422. https: // doi. org / 10.5962 / bhl. part. 2026","Strand, E. (1932) Miscellanea nomenclatorica zoologica et palaeontologica, III, IV. Folia Zoologica et Hydrobiologica, 4, 133 - 147 + 193 - 196.","Haddad, C. R. & Bosselaers, J. (2010) A revision of the genus Medmassa Simon, 1887 (Araneae: Corinnidae) in the Afrotopical Region. Zootaxa, 2361 (1), 1 - 12. https: // doi. org / 10.11646 / zootaxa. 2361.1.1","Ramirez, M. J. (2014) The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History, 390, 1 - 374. https: // doi. org / 10.1206 / 821.1","Lessert, R. de (1946) Araignees du Congo Belge. Revue Suisse de zoologie, 58, 204 - 225."]}

Carteronius Simon Mandane Karsch, 1880: 337, pl. 12, fig. 4 (type species by monotypy, M. sudana). Preoccupied by Kinberg 1866, in Spionidae, Annelida: Polychaeta) Carteronius Simon, 1897: 85 (type species by original designation, C. helluo Simon, 1896). Mandaneta Strand, 1932: 140. Replacement name for Mandane Karsch. syn. nov. Notes on the identity of Carteronius. Simon (1896) described four species of Carteronius: C. helluo (the type species), C. vittiger, C. scriptus, and C. fuscus. The first three species were described from males only, and C. fuscus from a female. The type locality is Sierra Leone for C. helluo, Madagascar for C. vittiger and C. scriptus, and Mauritius for C. fuscus. Based on Simon’s original Latin descriptions, the male of C. helluo can be considered to differ significantly from the males of C. vittiger and C. scriptus: it has a strongly rugose cephalothorax, granulose chelicerae, anterior tibiae with six short ventral spine pairs, anterior metatarsi with four long ventral spine pairs, a male palpal tibia that is only slightly longer than the patella, a large, lamelliform, bifid retrolateral tibial apophysis and an oval, blunt cymbium. Carteronius vittiger and C. scriptus both have an almost smooth cephalothorax and smooth chelicerae, anterior tibiae with four long ventral spine pairs, anterior metatarsi with two long ventral spine pairs, a male palpal tibia that is longer than the patella, a small, sharp and simple retrolateral tibial apophysis, and a very elongate, curved cymbium that is much longer than the bulbus itself. Simon (1897) presented the genus description of Carteronius, transferred Cheiracanthium argenticomum Keyserling, 1877 to the new genus (pp. 79–80), and synonymized C. scriptus with C. argenticomus (p. 83). Upon consulting the MNHN collection (Paris) and studying the type specimens of Simon’s four species, it turned out that the vial of the type species C. helluo did not contain the specimen corresponding with Simon’s original description. Instead, this vial, labeled Clubionidae — Carteronius E.S. helluo E.S. 10.611 Sierra Leone: Free town, contained a smooth male specimen with four and two ventral spine pairs on the anterior tibiae and metatarsi, respectively, as well as a male palp with a long tibia and cymbium and a simple retrolateral tibial apophysis, as described in Simon (1896) for C. vittiger and C. scriptus (Figs 1A–E; both here transferred to Donuea, the latter under D. argenticoma). On the other hand, the C. scriptus vial, labeled Clubionidae — Carteronius E.S. scriptus E.S. 14.625 Diego Suares, contained a rugose male specimen with six and four ventral spine pairs on the anterior tibiae and metatarsi, as well a male palp with a short tibia, a blunt cymbium, and a bifid retrolateral tibial apophysis, as described in Simon (1896) for C. helluo (Figs 1F–J). It must be concluded that the type specimen of C. helluo has been accidentally switched with the type specimen of C. scriptus. After proper photographic documentation (Fig 1A–E), the specimens were re-switched to their original vials to avoid confusion in the future. The C. vittiger vial, labeled Clubionidae — Carteronius E.S. vittiger E.S. 10.188 Diego Suares, can still be assumed to contain the correct male type specimen, as it has a spider that corresponds in detail with Simon’s original (1896) description of this species. The C. vittiger type specimen is slightly smaller than the C. scriptus type, having a brown cephalothorax, with a darker eye region and two dark dentate bands, brown chelicerae, and a male palpal tibia that is longer than the patella. The abdomen of the C. vittiger type is completely bleached and can no longer be compared with Simon’s description of its markings. The C. scriptus type has lost its coloration over time, but it can be recognized by the more robust chelicerae and the more robust ventral spines on the anterior tibiae and a male palpal tibia that is only slightly longer than the patella. Diagnosis. Members of the genus Carteronius share with those of Bunyoronius gen. nov. the trilobulated RTA, with an apical spur inserted in the base of the ventral lobe, a sperm duct with a single ventral fold, and the basally widened, curved embolus surrounding the tegular margins. They can be readily recognized by the absence of modifications on the male palpal femur (with a large apical retrolateral apophysis in Bunyoronius gen. nov.); the ventral fold of sperm duct oriented prolaterally (retro-dorsally in Bunyoronius gen. nov.); a hook-shaped median apophysis inserted retro-apically in a deep, wide circular pit (vestigial in Bunyoronius gen. nov.); and embolus inserted retrolaterally with a narrow tip (basally inserted, with a wide tip in Bunyoronius gen. nov.) (Figs 4A, B, 8A, B, 10A, B). Females can be recognized by the epigynal plate divided by a single transversal ridge, delimiting posterior and anterior sectors (two lateral longitudinal ridges, delimiting a median atrium in Bunyoronius gen. nov.); epigynal posterior sector with two lateral plates; and copulatory openings oriented posteriorly (oriented anteriorly in Bunyoronius gen. nov.) (Figs 5A, B, 8C, D). Description. Medium to large-sized spiders, 6.03 – 13.45 mm in length. Carapace heavily sclerotized, surface granulate with scarce long setae; very broad, sub-oval, as long as wide, cephalic region well-demarcated posteriorly, swollen anterolaterally, higher than thoracic region; thoracic region abruptly depressed, posteriorly rounded, thoracic fovea present (Figs 2, 3A – E, 13B, D). Clypeus low, height less than AME diameter, generally nearly half AME diameter (Fig. 2). AER straight in frontal view, ALE oblique, eyes equidistant; PER procurved in dorsal view, only slightly wider than AER, eyes equidistant; AME largest, approximately two times ALE diameter, remaining eyes sub-equal in size. Eyes circular. Chilum present, entire, generally bilobed, with two geminated tubercles; with a large median tubercle only in females of C. lumumba sp. nov. and C. simoni sp. nov.. Chelicerae sclerotized, slightly longer than half the length of the carapace, frontal surface granulate, strongly geniculate in females, slightly geniculated in males, heavily modified in males of C. sudanus (frontally with anterior excavation and blunt median tubercle on prolateral surface, Fig. 2A) and C. lumumba sp. nov. (ventrally with retromarginal proximal tooth modified into a large keel and a proximal constriction on prolateral margin, Fig. 16A), basal boss evident, promargin usually with three teeth, retromargin with two teeth. Endites convergent, promargin anteriorly protruded, retromargin slightly excavated; labium sub-squared, nearly as long as wide, slightly longer than half an endite’s length, proximal lateral constrictions shallow. Sternum shield-shaped, slightly longer than wide; surface covered by small setae-bearing tubercles, precoxal and intercoxal sclerites present, margins well defined, especially anterolaterally. Leg formula: I.II.IV.III. Legs long, I–II more robust; femur I with one dorsal spine, tibiae I with five to seven pairs of ventral spines (Fig. 11C); metatarsus I with two or four pairs of ventral spines. Femora with short setae inserted on tubercles; scopulae sparse, present in all tarsi and distal third of metatarsi I–II, metatarsi III–IV with sparse cluster of long setae. Tarsal trichobothria with lowered distal plate below a transverse ridge (only C. sudanus and C. ashanti sp. nov. surveyed, Figs 6D, 11F). Abdomen oval, with distinctive chevron markings variable across species (Figs 3A, B; 7A, C, 15A, C, 17A, 18A, F); dorsal and ventral scuta absent. Spinnerets (only female of C. sudanus surveyed with SEM): ALS with two major ampullate gland spigots near the middle of the spinning field and several piriform glands spigots around; PMS with three cylindrical gland spigots, two minor ampullate glands spigots, and 2–3 aciniform gland spigots; PLS with two cylindrical gland spigots and several aciniform glands spigots (Fig. 6F–I). Male palp: retrolateral femoral apophysis absent, retrolateral tibial apophysis complex, with three lobes, ventral lobe with apical spur (Figs 4A, 8A); cymbium densely covered by setae, with a denser dorso-apical cluster of short chemosensory setae, forming a conspicuous patch (Fig. 12A); median apophysis hook-shaped; embolus basally widened, curved, surrounding tegular margins, with membranous tip (tip bifid in C. sudanus and C. lumumba sp. nov.), hyaline conductor present, lamelliform; retrolateral apical tegular process present in, C. arboreus sp. nov., C. ashanti sp. nov. and C. gentilis (Figs 8A, 10A, 19A). Epigynum characterized by the presence of a transverse ridge delimiting anterior and posterior sectors; posterior sector with a pair of rounded plates (inconspicuous in C. lumumba sp. nov.); two copulatory openings located posteriorly (Figs 4C, 14A, C, 16D, 19C); CD distinct between primary and secondary spermathecae; ventral sector of CD (distal to ST2 insertion) long, generally thick (narrow in C. gentilis). Secondary spermathecae generally placed anteriorly to the level of the anterior curve of the ventral sector of CD (posteriorly in C. myene sp. nov. and C. teke sp. nov.), commonly tapering (globose in C. arboreus sp. nov.); dorsal sector of CD (between ST1 and ST2, proximal to ST2 insertion) wide, generally S-shaped (nearly straight in C. arboreous sp. nov. and folded ventrally in C. myene sp. nov. and C. teke sp. nov.). ST1 appendiciform, posteriorly positioned, generally smaller than ST2. Fertilization ducts large in relation to ST1 size (Figs 4D, 5C, D, 8D, 14B, D, 16E). Distribution. West and Central Africa. Misplaced species. The three species presently included in Carteronius other than the type species belong to the genus Donuea (recently transferred to Corinnidae). They share with described representatives of the genus (the type species D. decorsei (Simon, 1903) and D. collustrata Bosselaers & Dierick, 2010 the large, highly modified median apophysis and the long, filiform embolus in the male palp (Fig. 1B–D), or the flat epigynal plate with a long, narrow atrium with anchoring lateral ridges that may be present in all representatives of that genus. - Carteronius argenticomus (Keyserling, 1877) = Donuea argenticoma (Keyserling, 1877) comb. nov. - Carteronius vittiger Simon, 1896 = Donuea vittiger (Simon, 1896) comb. nov. - Carteronius fuscus Simon, 1896 = Donuea fusca (Simon, 1896) comb. nov. Key to Carteronius species 1 Males (those of C. teke sp. nov., C. myene sp. nov. and C. simoni sp. nov. unknown)................................ 2 - Females (those of C. ashanti sp. nov. unknown)............................................................. 7 2(1) Chelicerae with anterior surface or teeth modified (Figs 2A, 16A); retrolateral apical tegular process absent (Figs 4A, 16B). 3 - Chelicerae unmodified, anterior surface and teeth unmodified; retrolateral apical tegular process present (Figs 8A, 10A)... 4 3(2) Chelicerae with frontal anterior excavation, teeth unmodified (Fig. 2A); proximal half of embolus relatively broad (Fig. 4A)................................................................................... C. sudanus comb. nov. - Chelicerae without frontal anterior excavation, retromarginal proximal tooth modified into a large keel (Fig. 16A); proximal half of embolus relatively narrow (Fig. 16B)................................................ C. lumumba sp. nov. 4(2) Apical third of embolus with a retrolaterally-directed process interlocking tegular edge (Figs 8A, 10A)................. 5 - Apical third of embolus without such a process (Fig. 19A).................................... C. gentilis comb. nov. 5(4) Median and dorsal lobes of RTA sharing the same base (Fig. 8B); embolar apices wide, sub-apical embolar process large (Fig. 8A)................................................................................. C. arboreus sp. nov. - Median and dorsal lobes of RTA completely separated at base (Fig. 10B); embolar apices not enlarged; sub-apical embolar process small (Fig. 10A).................................................................. C. ashanti sp. nov. 7(1) Transversal ridge of epigynum strongly recurved (Figs. 4C, 8C)................................................ 8 - Transversal ridge otherwise (Figs 14A, C, 16D)............................................................. 9 8(7) Lateral plates of posterior sector not sclerotized (Fig. 4C).................................... C. sudanus comb. nov. - Lateral plates of posterior sector sclerotized (Fig. 8C)......................................... C. arboreus sp. nov. 9(7) Lateral plates of posterior sector inconspicuous; central septum not rebordered laterally (Figs 16D, 17C)............... 10 - Lateral plates of posterior sector well defined, central septum of posterior sector rebordered laterally (Figs 14A, C)...... 11 10(9) Copulatory duct convergent in ventral view (Fig. 16D)........................................ C. lumumba sp. nov. - Copulatory duct divergent in ventral view (Fig. 17C)............................................ C. simoni sp. nov. 11(9) Epigynal transversal ridge gently recurved (Figs 14A, 19).................................................... 12 - Epigynal transversal ridge straight (Fig. 14C).................................................... C. teke sp. nov. 12(11) Posterior sector relatively small, nearly one quarter as long as the anterior sector; spermathecae visible by transparency in anterior sector (Fig. 19C).............................................................. C. gentilis comb. nov. - Posterior sector relatively large, nearly half as long as the anterior sector; spermathecae barely visible by transparency in anterior sector (Fig. 14A).................................................................. C. myene sp. nov., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 345-348, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035, {"references":["Karsch, F. (1880) Arachnologische Blatter (Decas I). Zeitschrift fur die gesammten Naturwissenschaft, 53, 373 - 409.","Kinberg, J. G. H. (1866) Annulata Nova. Continuatio. [various errantia & sedentaria]. Ofversigt af Koniglich Vetenskapsakademiens forhandlingar, Stockholm, 22 (4), 239 - 258.","Simon, E. (1897) Histoire naturelle des araignees. Tome Second. Deuxieme Edition. Roret, Paris, 192 pp.","Simon, E. (1896) Descriptions d'arachnides nouveaux de la famille des Clubionidae. Annales de la Societe Entomologique de Belgique, 40 (9), 400 - 422. https: // doi. org / 10.5962 / bhl. part. 2026","Strand, E. (1932) Miscellanea nomenclatorica zoologica et palaeontologica, III, IV. Folia Zoologica et Hydrobiologica, 4, 133 - 147 + 193 - 196.","Keyserling, E. (1877) Einige Spinnen von Madagascar. Verhandlungen der Kaiserlich-Koniglichen Zoologisch-Botanischen Gesellschaft in Wien, 27, 85 - 96.","Simon, E. (1903) Descriptions d'arachnides nouveaux de Madagascar, faisant partie des collections du Museum. Bulletin du Museum d'Histoire Naturelle, 9, 133 - 140.","Bosselaers, J., Dierick, M., Cnudde, V., Masschaele, B., Hoorebeke, L. van & Jacobs, P. (2010) High-resolution X-ray computer tomography of an extant new Donuea (Araneae: Liocranidae) species in Madagascan copal. Zootaxa, 2427 (1), 25 - 35. https: // doi. org / 10.11646 / zootaxa. 2427.1.3"]}

Carteronius arboreus Bonaldo & Haddad sp. nov. Figs 7, 8; Map 1 Types. ♂ holotype from Bas Congo, Mayombe, Luki Forest Reserve (05°37’S, 13°05’E), 28.IX.2007, DEMOCRATIC REPUBLIC OF THE CONGO, D. de Bakker & J. P. Michiels leg. (sieving along trail in primary rainforest) (MRAC 223.475). ♀ paratype, same locality and collectors, 10.XI.2006 (Fog 3, primary rainforest) (MRAC 220.925). Etymology. The specific name is a Latin adjective meaning arboreal, in reference to the fact that most of the known specimens were collected by canopy fogging. Diagnosis. Males of Carteronius arboreus sp. nov. are similar to those of C. ashanti sp. nov. by the presence of a sub-apical embolar process (Figs 8A, 10A), but differ by the dorsal lobe of the RTA, which is broad and retrolaterally oriented, and the rounded medial lobe, which is small in relation to the dorsal lobe, and share the same base (Fig. 8B). In C. ashanti sp. nov., the dorsal lobe is spoon-shaped and the medial lobe longer and fang-shaped, with its own base. Females resemble those of C. sudanus comb. nov. in the strongly recurved epigynal transversal ridge (Figs 4C, 8C), but differ by the lateral plates of the epigynal posterior sector being strongly sclerotized (Fig. 8C). Description. Male. (MRAC 223475). Measurements: Total length 6.03, CL 3.16, CW 2.71, AL 2.95, AW 2.24, SL 1.39, SW 1.49. Eye diameters and interdistances: AME 0.28, ALE 0.17, PLE 0.18, PME 0.16, AME-AME 0.65, AME-ALE 0.16, ALE-ALE 1.34, PME-PME 0.33, PME-PLE 0.37, PLE-PLE 1.57. Length of leg segments: I 3.08+1.17+2.67+2.29+1.22=10.43; II 3.10+1.24+3.10+2.24+1.27=10.95; III 2.07+0.93+1.61+1.65+0.96=7.22; IV 2.55+0.94+2.11+2.25+1.05=8.90. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-0-1-0, II do 0-1-0, III do 0-1-0, IV do 0-1-0; tibiae: I ve 2-2-2-2-2, II ve 1 p -1 r -1 p -2-2-2, III ve 1 r -1 p -2, IV rl 0-1-0-1 ve 1 p -0-1 p; metatarsi: I ve 2-2, II ve 2-2, III pl 0-1-0-1 rl 0-1-0-1 ve 2-2, IV pl 0-1-0-1 rl 0-1-0-1 ve 1 r -1 p -1 r -0. Coloration: carapace and chelicerae reddish-brown. Endites, labium and sternum reddish-brown. Legs I and II: coxae and trochanters reddish-brown; femora reddish-brown, yellowish distally; tibiae, metatarsi and tarsi yellowish. Legs III and IV yellowish. Abdomen dark gray dorsally, with two well defined white bands; white ventrally (Fig. 7A). Palp: RTA with apical spur short, curved and pointed, dorsal lobe with apical edges bent ventrally, ventral lobe rounded and excavated. Tegulum with short retrolateral apical tegular process, spermatic duct with long loop (Fig. 8A, B). Female. (MRAC 220925). Measurements: Total length 10.58, CL 4.10, CW 3.86, AL 6.37, AW 4.91, SL 1.97, SW 1.93. Eye diameters and interdistances: AME 0.31, ALE 0.21, PLE 0.21, PME 0.18, AME-AME 0.25, AME-ALE 0.29, ALE-ALE 1.75, PME-PME 0.45, PME-PLE 0.49, PLE-PLE 2.13. Length of leg segments: I 3.87+1.81+3.70+2.88+1.28=13.54; II 3.70+1.73+3.66+2.99+1.31=13.39; III 2.75+1.32+2.17+2.08+0.91=9.23; IV 3.25+1.31+2.76+2.84+0.98=11.14. Chelicerae: promargin with three spaced teeth, median tooth largest; retromargin with two spaced teeth, subequal in size. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-0-1 p, IV do 0-1-0-1 p; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-1 r -1 p -1 r -1 p -1 r, III ve 1 p -1 r -2, IV rl 0-1-0-1 ve 1p-0-1 p; metatarsi: I ve 2-0-2-0, II ve 2-2-2, III pl 0-1-0-1 rl 0-0-0-1 ve 2-2-1, IV pl 0-1-0-1 rl 1-0-1 ve 1 p -1 r -1 p - 0-0. Coloration: Carapace and chelicerae dark reddish-brown. Endites, labium and sternum reddish-brown. Legs reddish-brown, with femora I and II darker. Abdomen gray, dorsum with scattered small white spots, denser in middle, forming irregular white longitudinal band; posteriorly with large triangular white spot. Ventrally gray with two irregular lateroventral white bands (Fig. 7C). Epigynum: CDv slight folded posteriorly, ST2 globose, anteriorly located, larger than ST1, CDd almost straight (Fig. 8D). Other material examined. DEMOCRATIC REPUBLIC OF THE CONGO: Bas Congo, Mayombe, Luki Forest Reserve (05°37’S, 13°05’E), 18.IX.2007, D. de Bakker & J.P. Michiels leg. (Fog 5, old secondary forest), 1♀ (MRAC); same data but 22.IX.2007, 1♀ (MRAC); same data but 30.IX.2007, 1♂ (MRAC). Distribution. Only known from the Democratic Republic of the Congo (Map 1)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 353-356, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Bunyoronius femoralis Bonaldo, Ramírez & Haddad sp. nov. Figs 20–23; Map 1 Types. ♂ holotype from Budongo Forest, 01°45’N, 31°25’E, 1200 m.a.s.l., UGANDA, 15-25.I.1997, leg. T. Wagner (collected by fogging in Rinorea beniensis, swamp forest), 1♂ (ZMFK Ar-23935). Paratypes: same locality and collector as the holotype, all collected by fogging in Rinorea beniensis; 11-20.VII.1995 (secondary forest), 1♂ (ZMFK Ar-23936); 11-20.VII.1995 (secondary forest) 1♂, 1♀ (ZMFK Ar-23937); 19-30.vi.1995 (primary forest), 1♂ (ZMFK Ar-23938); 21-31.VII.1995 (swamp forest), 2♂ (ZMFK Ar-23939); 5-15.I.1997 (swamp forest), 1♂ (ZMFK Ar-23940); 15-25.I.1997 (secondary forest), 1♂, (ZMFK Ar-23941). Etymology. The specific name is a Latin adjective highlighting the large male palpal apical retrolateral femoral apophysis. Diagnosis. As for the genus. Description. Male (Holotype). Measurements: Total length 7.37, CL 3.63, CW 3.10, AL 3.73, AW 2.56, SL 1.64, SW 1.59. Eye diameters and interdistances: AME 0.27, ALE 0.14, PLE 0.14, PME 0.18, AME-AME 0.22, AME-ALE 0.21, ALE-ALE 1.14, PME-PME 0.38, PME-PLE 0.42, PLE-PLE 1.56. Length of leg segments: I 2.79 +1.04+2.98+2.30+1.26=13.35; II 2.91+1.01+2.96+2.32+1.23=10.43; III 2.08+0.86+1.64+1.74+0.79=7.11; IV 2.36 +0.88+2.17+2.39+0.84=8.86. Chelicerae: promargin with three teeth; retromargin with two teeth, proximal twice as large as the distal. Leg spination: I do 0-0-1-0 pl 0-0-1-0, II do 0-0-1-0, III do 0-1-0-1, IV do 0-1-0-2; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-2-2-2, III pl 0-1-0-0 rl 0-0-1-0 ve 0-1-1-0, IV pl 0-1-0-1 rl 1-1-0-1 ve 1-0-0-1; metatarsi: I ve 2-2, II ve 2-2, III do 0-2-0-0 ve 2-2, IV pl 0-1-0-0 rl 1-0-1-0 ve 0-1-0-0. Coloration: Carapace, chelicerae, endites, labium and sternum reddish-brown (Fig. 20B). Legs yellowish, with femora I and II reddish-brown. Abdomen pale, with gray spots forming folicular shape in posterior region (Fig. 20A). Palp: Retrolateral apical femoral apophysis present, bifid; retrolateral tibial apophysis complex, with three lobes, ventral lobe with apical spurn; apical spur flat, smaller than ventral lobe, medial lobe very small, pointed; ventral lobe sub-rectangular, distally excavated; cymbium densely covered in setae, with dense dorso-apical cluster of short setae, forming conspicuous cymbial scopula (Fig. 23E); median apophysis absent, insertion area vestigial, represented by small unsclerotized window; sperm duct with two long loops, one retrolateral, directed proximally, and one ventral, directed distally; retrolateral apical tegular process absent. Conductor lamelliform, inserted retro-apically. Embolus wide, robust, not tapering distally, with apical sclerotized projection (Figs 22 A-C, 23). Female (ZMB Ar-23937). Measurements: Total length 6.92, CL 3.34, CW 3.02, AL 3.63, AW 2.62, SL 1.65, SW 1.57. Eye diameters and interdistances: AME 0.24, ALE 0.14, PLE 0.13, PME 0.17, AME-AME 0.21, AME-ALE 0.21, ALE-ALE 1.10, PME-PME 0.39, PME-PLE 0.39, PLE-PLE 1.49. Length of leg segments: I 2.98+1.10+2.93+ 2.09+1.17=10.80; II 3.04+1.04+2.81+2.21+1.21=10.31; III 2.20+?+?+?+?=?; IV 2.52+0.82+2.20+2.35+0.95=8.84. Chelicerae: promargin with three teeth; retromargin with two teeth, proximal twice as large as distal. Leg spination: femora: I do 0-1-0 pl 0-1-0, II do 0-1-0, III do 0-1-0-1, IV do 0-1-0-1; tibiae: I ve 2-2-2-2-2-2-2, II ve 2-2-2-2-2-2, III broken, IV ve 1-0-0 rl 0-1-0-0; metatarsi: I ve 2-2, II ve 2-2, III broken, IV ve 1-1-1-0-0 do 0-2-0-2. Coloration: carapace, chelicerae, endites, labium and sternum reddish-brown (Fig. 20D). Legs yellowish, with femora I and II reddish-brown. Abdomen pale, with irregular gray spots (Fig. 20C). Epigynum with two lateral curved ridges, forming median atrium. Two copulatory openings located anteriorly; CDv short, obliquely inserted; ST2 globous, smaller than ST1, anteriorly positioned; CDd large and convoluted; ST1 reniform, FD long and robust (Fig. 22D, E). Other material examined. RWANDA: Ibanda Makera, Rusumo, 02°09’S, 30°55’E, 1350 m.a.s.l., X.1995, T. Wagner leg. (Fogging Teclea nobilis, gallery forest), 1♂ (ZMFK Ar-23942). CENTRAL AFRICAN REPUBLIC: Prefecture Sangha-Mbaér, Parc National Dzanga-Ndoki, 37.9 km 169 S of Lidjombo, 02°22’14N, 16°10’21E, 360 m. a.s.l., 20–28. V.2001, B.L. Fisher leg. (rainforest, beating low vegetation), 2 imm. 1♀ (CAS, CASENT 9033197)., Published as part of Eb. Bonaldo, Ramírez, Martín J., Om. Labarque, Shimano, Yulie, Silva-Junior, Cláudio J. & Haddad, Charles R., 2022, Switching identities: a revision of the Afrotropical spider genus Carteronius Simon 1897 (Araneae, Corinnidae), senior synonym of Mandaneta Strand, 1932, with a new genus of the Pronophaea group, pp. 343-373 in Zootaxa 5205 (4) on pages 368-371, DOI: 10.11646/zootaxa.5205.4.3, http://zenodo.org/record/7307035

Genus Afrodrassex gen. nov. Type species. Afrodrassex balrog sp. nov. Etymology. The genus name is a contraction referring to the currently known distribution in the Afrotropical Region, and Leptodrassex, to which it is related. Gender masculine. Diagnosis. Afrodrassex gen. nov. can be distinguished from all other Leptodrassinae by the distinctive genitalic structure: females have very long copulatory ducts running around the periphery of the epigyne before entering the spermathecae posteriorly (Figs 34, 39), while the male palps have a large curved anterior tegular process, lack a median apophysis, and have a long embolus associated with a large membranous conductor (Figs 36–38, 41–43). Description. Small pale spiders (Figs 1, 2, 7–10), females 2.23–3.20 mm and males 1.85–2.75 mm in length; carapace creamy-white to yellow; carapace oval, eye region narrow, broadest between coxae II and III, without fovea (Fig. 11); posterior margin straight or slightly concave; carapace gradually elevated from eye region, highest at 3/4 its length, with steep posterior slope; carapace smooth and matte, densely covered in feathery setae, with long straight setae in eye region (Figs 11–14). All eyes surrounded by black rings, pigment continuous between anterior eyes (Figs 7–10); AER procurved in anterior view, slightly recurved in dorsal view (Figs 12–14); clypeus height slightly larger than AME diameter; AME largest, separated by approximately 1/2 their diameter, separated from ALE by 1/8 ALE diameter; PER strongly procurved in dorsal view (Figs 13, 14); PME oval and flattened, PLE round, PME slightly larger than PLE; PME separated from each other and from PLE by distance between 1/2 to equal to their diameter; ALE and PLE almost touching (Figs 13, 14); MOQ narrower posteriorly than anteriorly, anterior width slightly larger than MOQ length. Cheliceral dentition: promargin with three teeth, usually middle tooth largest, distal tooth smallest, a tiny denticle; retromargin with two subequal teeth, larger than promarginal teeth; endites with slightly depressed lateral margins, distal margins rounded, with distinct serrula and maxillar hair tuft; labium trapezoid, rounded anteriorly, slightly longer than wide. Pleural bars weakly sclerotised, isolated; sternum oval, approximately 1¼ times longer than broad, broadest at coxa II, surface smooth, sparsely covered in straight setae; precoxal triangles present, intercoxal sclerites present between all coxal pairs. Abdomen oval, as broad as or slightly broader than carapace, dorsal scutum absent in both sexes (Figs 7–10); dorsum with single pair of sigilla, usually indistinct; dorsum and sides densely covered in feathery setae, with scattered fine plumose setae (Figs 15, 16), venter only with fine plumose setae (Fig. 16). Spinnerets (only observed in A. balrog sp. nov.; spinnerets of male partly retracted and obscured by silk threads): ALS of female with two major ampullate gland spigots anteriorly, two large piriform gland spigots mesally, two slender modified piriform gland spigots lateral to anterior piriform gland spigot, and two tartipores posteriorly (Fig. 17); PMS of female with two large minor ampullate gland spigots mesally, one small cylindrical gland spigot posteriorly, one tartipore, and five small aciniform gland spigots peripherally (Fig. 18); PLS of female (partly retracted) with only one small minor ampullate gland spigot anteriorly, one large cylindrical gland spigot mesally, and three aciniform gland spigots posteriorly (Fig. 19); ALS of male with only two large piriform gland spigots distinguishable (Fig. 20); PMS of male with only one posterior minor ampullate gland spigot and three peripheral aciniform gland spigots distinguishable (Fig. 21); PLS of male with single large anterior minor ampullate gland spigot and two aciniform gland spigots distinguishable (Fig. 22). Leg formula 4123 or 4213; legs densely covered in feathery setae, with scattered straight plumose setae between them, feathery setae sparse on tarsi; patellae with narrow indentation and small lyriform organ on retrolateral side, with single proximal and distal erect long seta dorsally on patellae III and IV that are usually missing, presumably easily damaged and lost during preservation; metatarsi with well-developed dorsal stopper distally; tarsi with sparse chemosensory setae, two pairs of dorsal trichobothria, oval tarsal organ and dense claw tufts (Figs 23–26); tarsal claws with at least three small ventral teeth (Fig. 26). Female epigyne with shallow paired ovoid atria, separated by median septum (Figs 27, 34, 39), with atria frequently filled with secretory plugs; internally with extremely lengthened copulatory ducts, with spermathecae posteromedially positioned, with posteriorly-directed fertilization ducts. Male palpal femur and patella without apophyses, except A. catharinae sp. nov., with a small ventral patellar denticle (Figs 41, 43); palpal patella with retrolateral lyriform organ (Fig. 31); palpal tibia with retrolateral apophysis singular, variable in shape (Figs 29, 38, 41, 43); cymbium narrower than tegulum, with dense setae distally on dorsal surface (Fig. 28); tegulum generally ovoid, with slender embolus originating proximally or prolaterally; embolus free of subtegulum, associated basally with large membranous conductor; embolus extending towards or around distal end of tegulum, closely associated with prolateral groove in cymbium (Fig. 30) and conductor (Fig. 31), and at its distal end with deep groove in apical tegular process (Figs 30–33, 36–38, 41–43); median apophysis absent., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 3-4, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Genus Leptodrassex Murphy, 2007 Type species. Leptodrassus simoni Dalmas, 1919, by original designation. Diagnosis. Leptodrassex was defined by Murphy (2007) as a group of small, pale spiders usually 2–4 mm in length, with small teeth on the chelicerae (2–4 promarginal and 2 or 3 retromarginal teeth), with males lacking a dorsal scutum and the AME larger than the other eyes. The two new species described in this paper are very consistent in the presence of all of these characters, but the genitalic morphology differs slightly from the three currently known species from the Mediterranean, being more similar to the undescribed species from Kenya that Murphy (2007) included in his book. Leptodrassex can be separated from Leptodrassus by the cheliceral dentition, with the latter having two large angular translucent teeth on the promargin and four or five small conical teeth on the retromargin (Murphy 2007). It can be distinguished from Leptopilos by the genitalic structure, with females of the latter having an epigyne with an anterior hood and males having a palp with several laminae (Levy 2009). It can be separated from Neodrassex by the female genitalic structure, with the latter possessing a large, divided atrium and paired posterior epigynal processes (Ott 2012, 2013), which are lacking in Leptodrassex. Lastly, it differs from Afrodrassex gen. nov. by the shorter embolus and presence of a median apophysis, and the short copulatory ducts of females. Description. Small pale spiders (Figs 3, 4, 45–47), females 1.80–3.60 mm and males 1.65–2.80 mm in length; carapace creamy-white to yellow; carapace oval, eye region narrow, broadest between coxae II and III, without fovea; posterior margin straight or slightly concave; carapace gradually elevated from eye region, highest at 3/4 its length, with steep posterior slope; carapace smooth and matte, densely covered in feathery setae, with long straight setae in eye region. All eyes surrounded by black rings, pigment continuous between anterior eyes (Figs 45–47); AER procurved in anterior view, slightly recurved in dorsal view; clypeus height slightly larger than AME diameter; AME largest, separated by approximately 1/2 to 3/4 their diameter, separated from ALE by 1/8 ALE diameter; PER strongly procurved in dorsal view; PME oval and flattened, PLE round, PME approximately 3/4 times PLE diameter; PME separated from each other and from PLE by distance between 1/2 to equal to their diameter; ALE and PLE almost touching; MOQ narrower anteriorly than posteriorly, anterior width slightly larger than MOQ length. Cheliceral dentition (southern African species): promargin with three teeth, usually middle tooth largest, distal tooth smallest, a tiny denticle; promargin with escort seta and rake setae (Fig. 48); retromargin with two or three subequal teeth and retromarginal escort seta (Fig. 49); endites with slightly depressed lateral margins, distal margins rounded, with distinct serrula and maxillar hair tuft (Fig. 50); serrula teeth with undulating sides (Fig. 51); labium trapezoid, slightly longer than wide, with rounded anterior margin. Pleural bars weakly sclerotised, isolated; sternum oval, approximately 1¼ times longer than broad, broadest at coxa II, surface smooth, sparsely covered in straight setae; precoxal triangles present, intercoxal sclerites present between all coxal pairs.Abdomen oval, as broad as or slightly broader than carapace, dorsal scutum absent in both sexes (Figs 45–47); dorsum with single pair of sigilla, usually indistinct; dorsum and sides densely covered in feathery setae (Fig. 52), with scattered fine plumose setae, venter only with fine plumose setae (Fig. 53). Spinnerets (only observed in L. murphyi sp. nov.): ALS of female with two major ampullate gland spigots anteromesally, two large piriform gland spigots mesally, and three adjacent slender modified piriform gland spigots (Fig. 54); PMS of female with two large minor ampullate gland spigots mesally, one small cylindrical gland spigot posteriorly, one tartipore, and several small aciniform gland spigots peripherally (Fig. 55); PLS of female with only one large cylindrical gland spigot medially, two small minor ampullate gland spigots anteriorly, and six aciniform gland spigots peripherally (Fig. 56); ALS of male with two small major ampullate gland spigots anteromesally, two large piriform gland spigots medially, two slender modified piriform gland spigots adjacent to the anterior piriform gland spigot, and posterior nubbin (Fig. 57); PMS of male with two medial minor ampullate gland spigots, two tartipores, and several peripheral aciniform gland spigots (Fig. 58); PLS of male with single large minor ampullate gland spigot, one anterior tartipore, one posterior nubbin, and five peripheral aciniform gland spigots (Fig. 59). Leg formula 4213; legs densely covered in feathery setae, with scattered straight plumose setae between them, feathery setae sparse on tarsi (Figs 63–67); femora with distal retrolateral lyriform organ (Figs 60, 61); patellae with narrow indentation and lyriform organ on retrolateral side (Figs 62–64), with single proximal and distal erect long seta dorsally on patellae III and IV, sometimes lost during preservation; metatarsi with well-developed dorsal stopper distally (Fig. 66); tarsi with sparse chemosensory setae, two pairs of dorsal trichobothria, followed by single median trichobothrium, oval tarsal organ, and dense claw tufts (Figs 67–70); tarsal claws with three small ventral teeth (Fig. 70). Female epigyne with shallow paired ovoid atria, separated by narrow median septum, with atria frequently filled with secretory plugs (Figs 71–73); internal structure with short copulatory ducts, with spermathecae laterally positioned, with mesally-directed fertilization ducts. Male palpal femur and patella without apophyses, palpal patella with retrolateral lyriform organ (Figs 74, 75); palpal tibia with small prolateral and dorsal apophyses (Fig. 76) and dorsal and ventral retrolateral apophyses (Figs 76–78); cymbium pear-shaped, with dense setae distally on dorsal surface; tegulum generally ovoid, with very slender embolus originating proximally, entering groove in large membranous prolateral subtegulum, leading embolus to large prolateral distal apical tegular process with fine groove (Figs 76–79); retrolateral tegular process shorter, slightly curved (Fig. 77); median apophysis present in Palaearctic species and L. murphyi sp. nov., hook- (e.g. Murphy 2007: fig. 513) or spike-like (Figs 76–78)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 11-12, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932, {"references":["Murphy, J. (2007) s. n. In: Gnaphosid genera of the world. Vols. 1 & 2. British Arachnological Society, St Neots, Cambridgeshire, pp. i - xii + 1 - 92 & pp. i - ii + 93 - 605.","Dalmas, R. de (1919) Catalogue des araignees du genre Leptodrassus (Gnaphosidae) d'apres les materiaux de la collection E. Simon au Museum d'Histoire naturelle. Bulletin du Museum National d'Histoire Naturelle de Paris, 25, 243 - 250.","Levy, G. (2009) New ground-spider genera and species with annexed checklist of the Gnaphosidae (Araneae) of Israel. Zootaxa, 2066, 1 - 49. https: // doi. org / 10.11646 / zootaxa. 2066.1.1","Ott, R. (2012) Neodrassex, a new genus of the Leptodrassex group (Araneae, Gnaphosidae) from South America. Iheringia, Serie Zoologia, 102, 343 - 350. https: // doi. org / 10.1590 / S 0073 - 47212012000300015","Ott, R. (2013) Three new species of Neodrassex (Araneae, Gnaphosidae) from Brazil. Iheringia, Serie Zoologia, 103, 381 - 387. https: // doi. org / 10.1590 / S 0073 - 47212013000400008"]}

Afrodrassex balrog sp. nov. Figures 1, 2, 7, 8, 11–38 Type material. Holotype ♀: SOUTH AFRICA: Free State: Bloemfontein, Langenhoven Park, 29°05.105’S, 26°09.563’E, 1420 m a.s.l., 9.III.2015, leg. C. Haddad (on walls of house at night) (NCA 2014/1936). Paratypes: Same data as holotype but 26.I.2015, 1♂ (NCA 2014/1939); Same data as holotype but 18.X.2020, 3♀ (NCA 2020/703). SOUTH AFRICA: Northern Cape: Richtersveld National Park, Sendelingsdrift, 28°07.805’S, 16°53.503’E, 9.X.2015, leg. P.J. Goede (home on wood), 1♀ (NCA 2016/3487); Rooipoort Nature Reserve, 28°38.220’S, 24°16.800’E, 23.III.2013, leg. M. Stiller (canopy fogging, Acacia tortilis), 1♀ (NCA 2015/4269). Limpopo: Little Leigh, 22°56.518’S, 29°52.735’E, 21.III.2006, leg. F. Maanda (Kirkia wilmsi, above knee searching), 1♀ (NCA 2009/719). Etymology. The species name is a noun in apposition of the fictional character referred to as the “Balrog”, a demon from the Lord of the Rings trilogy by author J.R.R. Tolkien. In Peter Jackson’s movies based on the books, the Balrog is depicted as wielding a long whip of fire, reminiscent of the very long embolus of this species. Diagnosis. Afrodrassex balrog sp. nov. females can be distinguished from A. catharinae sp. nov. by the small copulatory openings and transversely oval spermathecae (Figs 27, 34, 35) compared to the large copulatory openings and globular spermathecae in A. catharinae sp. nov. (Figs 39, 40), and males by the single elongate spike-like retrolateral tibial apophysis in retrolateral view (Figs 29, 38) compared to the small sharp tooth-like apophysis of A. catharinae sp. nov. (Figs 41, 43). Description. Female (holotype, NCA 2014/1936). Colouration (Fig. 7): carapace and chelicerae creamyyellow; endites and labium cream, labium slightly darker; sternum cream, margins brown; femora cream, remaining segments creamy-yellow. Abdomen cream dorsally and ventrally, with grey mottling on dorsum posteriorly above spinnerets. Measurements: CL 1.11, CW 0.90, AL 1.49, AW 1.16, TL 2.73. Eye diameters and interdistances: AME 0.11, ALE 0.08 PME 0.08, PLE 0.07, AME–AME 0.04, AME–ALE 0.01, PME–PME 0.08, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.71, 0.35, 0.56, 0.51, 0.38 = 2.51; II 0.76, 0.38, 0.44, 0.52, 0.24 = 2.34; III 0.62, 0.33, 0.44, 0.41, 0.24 = 2.04; IV 1.02, 0.41, 0.73, 0.87, 0.30 = 3.33. Leg spination: femora: I do 2, II do 1, III do 1 rl 1, IV do 2 rl 1; patellae: spineless; tibiae: I plv 2 rlv 2, II plv 1 rlv 2, III pl 2 rl 1 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1-2 rlv 1, III pl 3 rl 1 plv 1 rlv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur do 1, patella spineless, tibia pl 2 plv 1, tarsus pl 2 rl 1 plv 2 rlv 1. Epigyne with large figure-6-shaped ridges, coursing around periphery of epigynal plate, with tiny copulatory openings originating centrally at start of ridges (Figs 27, 34); copulatory ducts narrow, following path of external ridges, on lateral sides continuing posteriorly, looping mesally before entering transverse oval spermathecae on their lateral margins; fertilization ducts on posterior margin of spermathecae, directed posterolaterally (Fig. 35). Male (paratype, NCA 2014/1939). Colouration (Fig. 8): similar to female, but grey mottling dorsally on abdominal posterior extending to middle. Measurements: CL 0.97, CW 0.81, AL 0.95, AW 0.63, TL 1.85. Eye diameters and interdistances: AME 0.10, ALE 0.07, PME 0.08, PLE 0.06, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.07, PME–PLE 0.05, ALE–PLE 0.01. Leg measurements: I 0.76, 0.33, 0.62, 0.52, 0.38 = 2.61; II 0.78, 0.35, 0.65, 0.60, 0.40 = 2.78; III 0.67, 0.32, 0.43, 0.51, 0.27 = 2.20; IV 0.98, 0.38, 0.71, 0.86, 0.30 = 3.23. Leg spination: femora: I do 1, II do 1, III do 1, IV do 2 rl 1; patellae spineless; tibiae: I rlv 1, II spineless, III pl 1 rl 1 plv 1, IV pl 1 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 2 plv 1 rlv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: spineless. Palp: tibia broader than long, partly obscured by tegulum and membranous conductor, with elongate, slightly curved spike-like retrolateral apophysis (Figs 29, 38); tegulum largely obscured by massive bean-shaped membranous conductor; embolus long and very slender, originating prolaterally, looping proximally, then dorsally and distally above cymbium, with tip looping around ventral aspect of cymbium (Figs 30–33, 36–38); apical tegular process large, bending towards retrolateral side of palp, with split tip and deep groove along distal margin (Fig. 33); median apophysis absent. Additional material examined. SOUTH AFRICA: Free State: Bloemfontein, Langenhoven Park, 29°05.105’S, 26°09.563’E, 1420 m a.s.l., 3.X.2020, leg. C. Haddad (on ceiling at night), 1♂ (S.E.M. preparations); Same locality, 29°05.384’S, 26°09.392’E, 23.VII.2020, leg. R. Booysen (in garden on plants at night), 1 imm. 1 subadult ♂ 1♀ (S.E.M. preparations). ANGOLA: Malanje: Malange [09°32’S, 16°20’E], 11.IX.1949, leg. B. Malkin, 1 imm. 1♀ (CAS, CASENT 9058549). Variation. Total length: females 2.23–3.10 (average 2.71, n = 6); males: only the single paratype described. Habitat and biology. All seven examined females (including the S.E.M. specimen, epigyne cleared) had plugged epigynes (Fig. 27). This species has been recorded from three biomes (desert, grassland and savanna), although the majority of the specimens were collected inside houses and gardens in central South Africa, where they were active at night. Distribution. Widely distributed in South Africa, but only known from four localities (Fig. 44). The precise locality of the specimen from Angola is unclear, as coordinates are missing from the specimen label. According to the global gazetteer, there are three towns in Angola called Malange: one in the Kwanza Sul Region (09°57’S, 14°55’E), one in the Lunda Norte Region (08°05’S, 19°00’E), and one in the Malanje Region (09°32’S, 16°20’E). Considering the similarity in the town name and that of the latter region, we have plotted the latter co-ordinates on the distribution map (Fig. 44)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 7-9, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Genus Leptopilos Levy, 2009 Type species. Drassus tenerrimus O. Pickard-Cambridge, 1872, by original designation. Diagnosis. Leptopilos can be recognised from other Leptodrassinae by the presence of an anterior median hood in the female epigyne, and a male palp armed with closely grouped, often pointed distal laminae (tegular processes), and a single retrolateral tibial apophysis (Levy 2009). Description. Small pale spiders (Figs 5, 6, 87–92), females 2.15–3.80 mm and males 1.77–2.98 mm in length; carapace creamy-white to yellow; carapace oval, eye region narrow, broadest between coxae II and III, without fovea (Fig. 93); posterior margin slightly concave; carapace gradually elevated from eye region, highest at approximately 3/4 its length, with steep posterior slope; carapace smooth and matte, densely covered in feathery setae (Figs 93, 94), with long straight setae in eye region. All eyes surrounded by black rings, pigment continuous between anterior eyes (Figs 87–92); AER procurved in anterior view, slightly recurved in dorsal view (Fig. 94); clypeus height slightly larger than AME diameter; AME largest, separated by approximately 1/2 their diameter, separated from ALE by 1/8 ALE diameter; PER strongly procurved in dorsal view (Fig. 94); PME oval and flattened, PLE round, PME slightly larger than PLE; PME separated from each other by approximately 3/4 their diameter, from PLE by approximately 1/2 PME diameter; ALE and PLE almost touching (Fig. 94); MOQ slightly narrower posteriorly than anteriorly, anterior width slightly larger than MOQ length. Chelicerae: with promarginal escort seta and rake setae, and single retromarginal escort seta (Fig. 96); cheliceral dentition (southern African species): promargin with one tooth; retromargin with two teeth, proximal tooth larger than distal (Figs 97, 98); endites with slightly depressed lateral margins, distal margins rounded, with distinct serrula and maxillar hair tuft; serrula teeth with weakly undulating lateral margins (Fig. 99); labium trapezoid, slightly longer than wide, with rounded anterior margin. Pleural bars weakly sclerotised, isolated; sternum shield-shaped, approximately 1¼ times longer than broad, broadest at coxa II, surface smooth, sparsely covered in straight setae (Fig. 100); precoxal triangles present, intercoxal sclerites present between all coxal pairs. Abdomen oval, slightly broader than carapace, dorsal scutum absent in both sexes (Figs 87–92); dorsum with single pair of indistinct sigilla; dorsum and sides densely covered in feathery setae, with scattered fine plumose setae (Fig. 101), venter only with fine plumose setae. Spinnerets (only observed in L. digitus sp. nov.): ALS of female with two major ampullate gland spigots anteriorly, two large piriform gland spigots mesally, and two slender modified piriform gland spigots, one anterior and one posterior to anterior piriform gland spigot (Fig. 102); PMS of female with two large minor ampullate gland spigots mesally, two small cylindrical gland spigots posteriorly, single mesal tartipore, and eight small aciniform gland spigots peripherally (Fig. 103); PLS of female with two small minor ampullate gland spigots anteriorly, one large cylindrical gland spigot mesally, single mesal nubbin, and nine aciniform gland spigots peripherally (Fig. 104); ALS of male with one large major ampullate gland spigot with adjacent nubbin anteromesally, one large piriform gland spigot mesally, and three slender modified piriform gland spigots, two anterior and one posterior to anterior piriform gland spigot (Fig. 105); PMS of male with one posterior minor ampullate gland spigot, one tartipore and nubbin anterior to it, one posterior tartipore, and seven peripheral aciniform gland spigots (Fig. 106); PLS of male with single large anterior minor ampullate gland spigot, one posterior nubbin, and five aciniform gland spigots peripherally (Fig. 107). Leg formula 4123; legs densely covered in feathery setae, with scattered straight plumose setae between them, feathery setae sparse on tarsi (Figs 108–110); patellae with narrow indentation and lyriform organ on retrolateral side (Figs 108, 109), with single distal erect long seta dorsally on all patellae; metatarsi with well-developed dorsal stopper distally (Fig. 110); tarsi with sparse chemosensory setae, three pro- and retrolateral dorsal trichobothria in alternating arrangement, oval tarsal organ and dense claw tufts (Figs 110–113); tarsal claws with three ventral teeth (Fig. 113). Female epigyne with median anterior hood (Figs 121, 131) or pair of anterior ridges (Fig. 126), with atria frequently filled with secretory plugs (Fig. 114); internally with short copulatory ducts, initially directed laterally, then looping anteriorly and mesally, entering teardrop-shaped spermathecae anteriorly, with posteriorly-directed fertilization ducts. Male palpal femur and patella without apophyses; palpal patella with retrolateral lyriform organ (Fig. 117); palpal tibia with single retrolateral apophysis (Figs 115, 116, 118); cymbium ovoid, with dense setae distally on dorsal surface (Fig. 115); tegulum generally ovoid, with very slender embolus originating proximally, entering prolateral groove in subtegulum, leading embolus to distal apical tegular process, with embolus tip in narrow retrolateral groove (Figs 119, 120); retrolateral tegular process closely associated with apical tegular process (Figs 119, 124, 129, 134); median apophysis hook-like, originating retrolaterally (Fig. 119)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on page 19, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932, {"references":["Levy, G. (2009) New ground-spider genera and species with annexed checklist of the Gnaphosidae (Araneae) of Israel. Zootaxa, 2066, 1 - 49. https: // doi. org / 10.11646 / zootaxa. 2066.1.1"]}

Leptodrassex murphyi sp. nov. Figures 46–79, 82–86 Leptodrassex sp. Rodrigues & Rheims, 2020: figs 8H, 19A, 22C, 24B, 28B, 29B. Type material. Holotype ♀: SOUTH AFRICA: KwaZulu-Natal: Ndumo Game Reserve, Main Camp, 26°54.516’S, 32°18.861’E, 13.VI.2005, leg. C. Haddad (grass litter) (NCA 2005/969). Paratypes: Together with holotype, 1♂ (NCA 2005/969); MOZAMBIQUE: Gaza: Bilene, Praia do Bilene, 25°16’S, 33°18’E, 27 m a.s.l., 20.XII.2007, leg. C. Haddad, R. Lyle & R. Fourie (leaf litter, coastal forest), 1♀ (NMBA 11318). Inhambane: Bartholomew Dias Point, 21°16’S, 35°07’E, 5 m a.s.l., 10.XII.2007, leg. C. Haddad, R. Lyle & R. Fourie (leaf litter, mangroves), 3♂ 5♀ (NMBA 11244); Vilankulos, Casa Chibububo, 22°01’S, 35°19’E, 3 m a.s.l., 12.XII.2007, leg. C. Haddad, R. Lyle & R. Fourie (leaf litter, coastal bush), 1♂ (NMBA 11358). Maputo: Near Marracuene, Blue Anchor Inn, 25°35’S, 32°40’E, 50 m a.s.l., 28.XI.2007, leg. C. Haddad & R. Lyle (leaf litter, savanna), 1♂ 2♀ (NMBA 11414). SOUTH AFRICA: Free State: Bloemfontein, Shelleyvale-Rayton road, 29°04’S, 26°12’E, 21.V.2015, leg. C. Haddad & N. Jolintini (Sorghum bicolor tussocks), 2♂ (NCA 2015/2512); Luckhoff district, Farm Bankfontein, 30°04.980’S, 24°54.170’E, 22.I.2015, leg. C. Haddad (base of grass tussocks, wetland), 1♂ (NCA 2015/1651). Etymology. This species is named for the late John Murphy, who described the genus and included in his book an undescribed species from Kenya that closely resembles this species (Murphy 2007). Diagnosis. The female of this species most closely resembles that of L. capensis sp. nov., but can be recognised by the more strongly bent copulatory ducts and the larger copulatory openings (compare Figs 80 and 82). Males most closely resemble those of the undescribed Leptodrassex sp. 1 of Murphy (2007), but have a much shorter retrolateral tegular process and shorter dorsal retrolateral tibial apophysis (compare Figs 84–86 with Murphy 2007: fig. 515). Description. Female (holotype, Ndumo, NCA 2005/969). Colouration (Fig. 46): carapace, endites and chelicerae creamy-yellow, sternum creamy-white, margins yellow-brown at coxae; femora creamy-white, remaining segments creamy-yellow. Abdomen creamy-white dorsally and ventrally. Measurements: CL 0.99, CW 0.83, AL 1.57, AW 0.67, TL 2.70. Eye diameters and interdistances: AME 0.10, ALE 0.09, PME 0.07, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.06, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.71, 0.35, 0.57, 0.48, 0.37 = 2.48; II 0.76, 0.37, 0.63, 0.53, 0.40 = 2.69; III 0.62, 0.33, 0.44, 0.41, 0.24 = 2.04; IV 1.01, 0.40, 0.79, 0.84, 0.25 = 3.29. Leg spination: femora: I and II spineless, III do 1 rl 1, IV do 1 rl 1; tibiae: I plv 2 rlv 2, II rlv 2, III pl 3 rl 2 plv 2, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 2 rl 2 plv 1 vt 3; palp: femur do 2, patella spineless, tibia pl 2 plv 1, tarsus pl 1 rl 1 plv 2 rlv 4 vt 2. Epigyne with recurved hemispherical ridges anteriorly, continuing along midline into posterior half, forming bean-shaped atria (Figs 71–73), narrowly separated by median septum; copulatory openings oblique, procurved, situated in anterior part of atria; copulatory ducts S-shaped, broad initially but narrowing quickly, initially directed posteriorly, curving laterally at midpoint, with sharp lateral bend before entering ovoid lateral spermathecae; fertilization ducts on posteromesal margin of spermathecae, directed mesally (Figs 82, 83). Male (paratype, Ndumo, NCA 2005/969). Colouration (Fig. 47): similar to female, slightly paler. Measurements: CL 0.86, CW 0.71, AL 1.08, AW 0.67, TL 1.98. Eye diameters and interdistances: AME 0.09, ALE 0.05, PME 0.08, PLE 0.05, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.03, PME–PLE 0.03, ALE–PLE 0.01. Leg measurements: I 0.68, 0.30, 0.56, 0.46, 0.34 = 2.34; II 0.71, 0.32, 0.58, 0.48, 0.36 = 2.45; III 0.59, 0.30, 0.40, 0.38, 0.24 = 1.91; IV 0.95, 0.35, 0.72, 0.76, 0.25 = 3.03. Leg spination: femora: I and II spineless, III do 1, IV do 1 rl 1; tibiae: I plv 2 rlv 2, II rlv 2, III pl 2 rl 1 plv 1, IV pl 2 rl 2 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 2 rl 2 plv 1 vt 3; palp: femur do 2 plv 1-2, patella spineless, tibia plv 1, tarsus plv 2 rlv 2. Palp: tibia longer than broad, with small lobate prolateral apophysis, small dorsal apophysis, slightly larger ventral retrolateral apophysis, and slender ventrally curved dorsal tibial apophysis (Figs 76–78, 84–86); tegulum ovoid, with large curved apical tegular process, smaller spike-like retrolateral tegular process, and slender straight median apophysis (Figs 77, 85); fine laminate conductor hidden behind apical tegular process (Fig. 77); embolus very slender, originating proximally and entering groove in subtegulum, continuing along prolateral margin distally, before entering groove in apical tegular process (Figs 77–79). Additional material examined. SOUTH AFRICA: Free State: Amanzi Private Game Reserve, 28°35.785’S, 26°26.335’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Vachellia karroo woodland), 1♀ (NCA 2013/3379); Same locality, Obstacle course, 28°35.994’S, 26°25.650’E, 30.XII.2020, leg. C. Haddad (base of grass tussocks), 2♂ 1♀ (S.E.M. preparations); Bloemfontein, Free State National Botanical Gardens, 29°02’S, 26°12’E, 8.VI.2015, leg. C. Haddad & N. Jolintini (Hyparrhenia hirta tussocks), 1♀ (NCA 2015/2521); Same locality, 21.V.2015, leg. C. Haddad & N. Jolintini (H. hirta tussocks), 2♂ (NCA 2015/2503); Brandfort, Florisbad Research Station, 28°46’S, 26°05’E, 21.XII.1987 – 5.I.1988, leg. L. Lotz (pitfalls), 1♀ (NMBA 8435); Gariep Dam Nature Reserve, 30°35’S, 25°32’E, 1340 m a.s.l., 10.IV.2017, leg. M. Morake & N. Tshabalala (sifting leaf litter, Nama Karoo veld), 1♀ (NCA 2019/896); Harrismith, Platberg Nature Reserve, 28°16.842’S, 29°12.024’E, 2040 m a.s.l., 13.XI.2015 – 26.I.2016, leg. C. Haddad, D. Fourie & Z. Mbo (pitfall traps, alpine grassland), 1♂ 3♀ (NCA 2015/2300); Sterkfontein Dam Nature Reserve, 28°24.925’S, 29°02.529’E, 1700 m a.s.l., 11.XI.2015, leg. C. Haddad (under rocks, shore of dam), 1♀ (NCA 2015/2125). Gauteng: Pretoria, Pretoria National Botanical Gardens, 25°44’S, 28°16’E, 6.X–24.XI.2007, leg. E. Kassimatis (pitfalls), 1♀ (NCA 2010/2262). KwaZulu-Natal: Enseleni Nature Reserve, 28°41.350’S, 31°59.900’E, 12.X.2020, leg. R. Booysen & R. Steenkamp (hand collecting, grass tussocks), 2♂ (NCA 2020/727); iSimangaliso Wetlands Park, Mission Rocks picnic site, 28°15.879’S, 32°28.922’E, 90 m a.s.l., 29.XI.2015, leg. C. Haddad (base of grass tussocks, coastal forest), 1♀ (NCA 2015/2257). Limpopo: Soutpansberg Mountains, Lajuma Mountain Retreat, 23°02.306’S, 29°26.633’E, 6.II.2008, leg. R. Lyle & R. Fourie (beats, Afromontane forest), 1♀ (NCA 2008/494). Northern Cape: Benfontein Nature Reserve, 28°49’S, 24°49’E, 4.I.2006, leg. R. Lyle (pitfalls, dry savanna), 1♀ (NCA 2006/1128); Kathu, Farm Pniel, 28°35.420’S, 24°31.967’E, 31.X.2005, leg. R. Lyle (pitfalls, dry savanna), 1♀ (NCA (2006/1093). Variation. Total length: females 1.80–3.10 (average 2.44, n = 22); males 1.65–2.28 (average 1.96, n = 11). Habitat and biology. Approximately 80% of the examined females had plugged epigynes. This species occupied a broad range of biomes (Nama Karoo, grassland, savanna, forest, Indian Ocean Coastal Belt), where it was predominantly sampled from the ground by pitfalls, litter sifting, beneath rocks or from grass tussocks, and only rarely from woody vegetation by beating. Distribution. Widely distributed in southern Mozambique and the eastern half of South Africa (Fig. 44)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 17-19, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932, {"references":["Rodrigues, B. V. B. & Rheims, C. A. (2020) Phylogenetic analysis of the subfamily Prodidominae (Arachnida: Araneae: Gnaphosidae). Zoological Journal of the Linnean Society, 190, 654 - 708. https: // doi. org / 10.1093 / zoolinnean / zlaa 013","Murphy, J. (2007) s. n. In: Gnaphosid genera of the world. Vols. 1 & 2. British Arachnological Society, St Neots, Cambridgeshire, pp. i - xii + 1 - 92 & pp. i - ii + 93 - 605."]}

Leptopilos vasivulva sp. nov. Figures 91, 92, 131–135 Type material. Holotype ♀: BOTSWANA: North-East: Near Francistown, Selkirk Mine, 21°19.494’S, 27°45.030’E, 30.XI–6.XII.2007, leg. D.H. Jacobs (pitfalls) (NCA 2010/1835). Paratypes: Together with holotype, 2♂ 3♀ (NCA 2010/1835); Same data as holotype, 1♂ 1♀ (NCA 2009/3027); Same data as holotype but 30.X–6.XI.2007, 3♀ (NCA 2010/642). SOUTH AFRICA: Limpopo: Blouberg Nature Reserve, 22°59.332’S, 29°06.397’E, leg. S. Foord & E. Stam (pitfalls, Sclerocarrea birrea), no date, 1♀ (NCA 2009/1779); Vhembe Biosphere Reserve, Baries Farm, 22°28.672’S, 29°24.316’E, 6.XII.2012, leg. C. Schoeman (pitfalls, Musina mopane bushveld), 1♀ (NCA 2015/1240). Etymology. The species name is a contraction of the Greek words vasis and vulva, referring to the vase-shaped margins of the female’s epigynal atrium. Diagnosis. The females of L. vasivulva sp. nov. can be distinguished from congeners by the unique vaseshaped atrium and a single median curved marking on the anterior hood (Fig. 131). Males of L. vasivulva sp. nov. are similar to L. butleri sp. nov., but have longer apical and retrolateral tegular processes and a distally constricted retrolateral tibial apophysis (compare Figs 124 and 125 with Figs 134 and 135). Description. Female (holotype, NCA 2010/1835). Colouration (Fig. 91): carapace and chelicerae creamyyellow; endites and labium creamy-yellow, labium slightly darker; sternum creamy-yellow, margins yellow-brown; femora creamy-yellow, remaining segments pale yellow; abdomen creamy-grey dorsally and ventrally. Measurements: CL 1.13, CW 0.89, AL 1.90, AW 1.21, TL 3.40. Eye diameters and interdistances: AME 0.11, ALE 0.09, PME 0.09, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.06, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.92, 0.38, 0.65, 0.62, 0.48 = 3.05; II 0.78, 0.37, 0.54, 0.53, 0.40 = 2.62; III 0.67, 0.35, 0.44, 0.49, 0.32 = 2.27; IV 1.16, 0.44, 0.84, 1.00, 0.33 = 3.77. Leg spination: femora: I do 1, II do 1, III pl 1 do 2 rl 1, IV pl 1 do 3 rl 1; patellae spineless; tibiae: I and II spineless, III pl 2 rl 2 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I and II spineless, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur do 2, patella spineless, tibia plv 1, tarsus plv 3 rlv 3. Epigyne with strongly curved hemispherical anterior hood, with small recurved median marking; atrium vase-shaped, broad anteriorly and narrowed posteriorly (Fig. 131); copulatory openings anterolaterally in atrium, longitudinal; copulatory ducts short, initially directed laterally, looping anteriorly, then posteriorly, entering ovoid lateral spermathecae on their anterior margin; fertilization ducts on posterior margin of spermathecae, directed posteriorly (Fig. 132). Male (paratype, NCA 2010/1835). Colouration (Fig. 92): similar to female, slightly darker. Measurements: CL 1.05, CW 0.86, AL 1.43, AW 0.84, TL 2.63. Eye diameters and interdistances: AME 0.11, ALE 0.08, PME 0.10, PLE 0.08, AME–AME 0.04, AME–ALE 0.01, PME–PME 0.04, PME–PLE 0.03, ALE–PLE 0.01. Leg measurements: I 0.90, 0.40, 0.76, 0.70, 0.51 = 3.27; II 0.81, 0.37, 0.60, 0.59, 0.43 = 2.80; III 0.66, 0.32, 0.48, 0.50, 0.33 = 2.29; IV 1.16, 0.41, 0.86, 1.02, 0.35 = 3.80. Leg spination: femora: I do 2, II do 2, III do 2 rl 1, IV pl 1 do 2 rl 1; patellae spineless; tibiae: I and II spineless, III pl 2 rl 2 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I and II spineless, III pl 2 rl 1 plv 1 vt 3; IV pl 2 rl 2 plv 1 vt 3; palp: femur pl 1 do 3, patella spineless, tibia with several long ventral setae, tarsus with several distal setae ventrally. Palpal tibia slightly longer than broad, broadened distally, without prolateral apophysis, with single triangular retrolateral apophysis with constricted tip (Fig. 135); tegulum ovoid, with slender, gradually curved median apophysis with hooked tip (Fig. 134); embolus very slender, originating proximally and entering groove in membranous conductor, continuing along prolateral margin in S-shaped groove (Fig. 133), before entering groove in apical tegular process, partly hidden behind retrolateral tegular apophysis. Additional material examined. ZIMBABWE: Bulawayo, Hillside, 20°10’S, 28°35’E, VI.1999, leg. M. FitzPatrick, 11♂ 14♀ (NMZ/ A16735); Same locality, VIII.1999, leg. M. FitzPatrick, 6 imm. 6♂ 14♀ (NMZ/ A16718); Same locality, III–IV.1999, leg. M. FitzPatrick (pitfall traps), 2 imm. 2♂ 11♀ (NMZ/ A14033). Variation. Total length: females 2.80–3.60 (average 3.23, n = 9); males 2.15–2.80 (average 2.53, n = 3). Habitat and biology. All nine examined females (100%) from South Africa had plugged epigynes. This species was only recorded from pitfall traps in savanna habitats. Distribution. Only known from a small area in northern Limpopo Province, South Africa, and south-western Zimbabwe, and south-eastern Botswana (Fig. 136)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 28-30, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Leptopilos digitus sp. nov. Figures 89, 90, 93–120, 126–130 Type material. Holotype ♀: SOUTH AFRICA: Northern Cape: Namaqua National Park, Koeroebees, 30°08.683’S, 17°42.177’E, 240 m a.s.l., 27.III.2022, leg. C. Haddad & R. Booysen (leaf litter, dry river bed) (NMBA 18534). Paratypes: SOUTH AFRICA: Free State: Erfenis Dam Nature Reserve, 28°30.134’S, 26°48.427’E, 22.XI– 23.XII.2005, leg. C. Haddad (pitfalls, burnt site 2), 1♂ (NMBA 13573). Northern Cape: Namaqua National Park, Koeroebees, 30°08.683’S, 17°42.177’E, 240 m a.s.l., 27.III.2022, leg. C. Haddad, R. Booysen, L. Malope & S. Sibisi (leaf litter, dry river bed), 2♂ 3♀ (NMBA 18533); Prieska district, Green Valley Nuts, 29°34’S, 22°55’E, 950 m a.s.l., 15.I.2001, leg. C. Haddad (canopy fogging, pistachio orchard), 1♂ (NCA 2004/196). Western Cape: Tankwa Karoo National Park, Tanqua Guesthouse, 32°23.911’S, 19°50.713’E, 355 m a.s.l., 19.I.2021, leg. C. Haddad & R. Booysen (hand collecting, at night around houses), 1♀ (NCA 2021/1016). Etymology. The species name is Latin for finger or toe (digit), referring to the shape of the apical tegular sclerite. Diagnosis. This species is very distinct in the genus by the short finger-like apical tegular process (Fig. 129) and the short, slender dorsal tibial apophysis (Figs 120, 130) of the male palp. Females can be distinguished from congeners by the paired, weakly sclerotized recurved ridges anteriorly in the epigyne, as opposed to a single median hood, and the paired longitudinal ridges forming the lateral margins of the atrium (Fig. 126). Description. Female (holotype, Namaqua, NMBA 18534). Colouration (Fig. 89): carapace and chelicerae creamy-white; endites and labium pale yellow-brown; sternum creamy-white, margins brown at coxae; femora creamy-white, remaining segments creamy-yellow, metatarsi and tarsi slightly darker; abdomen white dorsally and ventrally. Measurements: CL 1.06, CW 0.87, AL 1.71, AW 1.25, TL 2.80. Eye diameters and interdistances: AME 0.10, ALE 0.07, PME 0.08, PLE 0.07, AME–AME 0.05, AME–ALE 0.01, PME–PME 0.06, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.86, 0.40, 0.67, 0.57, 0.44 = 2.94; II 0.81, 0.37, 0.62, 0.56, 0.43 = 2.79; III 0.67, 0.35, 0.49, 0.48, 0.30 = 2.29; IV 1.05, 0.40, 0.84, 0.97, 0.30 = 3.56. Leg spination: femora: I do 1, II do 1, III do 1, IV do 2 rl 1; patellae spineless; tibiae: I plv 1, II spineless, III pl 2 rl 2 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 rlv 1 vt 3, IV pl 3 rl 2 plv 2 vt 3; palp: femur do 1, patella pl 1 do 1, tibia plv 1, tarsus pl 1 rl 1 plv 3 rlv 2. Epigyne with paired, recurved, weakly sclerotized ridges anteriorly, longitudinal ridges laterally forming margins of atrium, with further funnel-shaped ridges medially (Fig. 126); copulatory openings small, beneath anterolateral corners of funnel-shaped ridges; copulatory ducts short, looping laterally, then anteriorly and posteriorly, entering teardrop-shaped spermathecae along their anterior margin (Fig. 127); fertilization ducts on posterior margin of spermathecae, directed posteriorly. Male (paratype, Prieska, NCA 2004/196). Colouration (Fig. 90): carapace and chelicerae light yellow; endites and labium creamy-yellow; sternum creamy-yellow, margins brown at coxae; femora to tibiae creamy-yellow, metatarsi and tarsi slightly darker; abdomen creamy-white dorsally and ventrally. Measurements: CL 1.05, CW 0.79, AL 1.81, AW 1.02, TL 2.98. Eye diameters and interdistances: AME 0.11, ALE 0.08, PME 0.09, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.05, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.87, 0.38, 0.67, 0.57, 0.51 = 3.00; II 0.83, 0.37, 0.62, 0.56, 0.48 = 2.86; III 0.65, 0.32, 0.48, 0.46, 0.33 = 2.24; IV 1.06, 0.41, 0.83, 0.95, 0.33 = 3.58. Leg spination: femora: I do 1, II do 1, III do 1 rl 1, IV pl 1 do 2 rl 1; tibiae: I plv 1 rlv 1, II spineless, III pl 2 rl 1 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I rlv 1, II rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 2 rl 2 plv 1 vt 3; palp: femur do 1, patella and tibia spineless, tarsus plv 1 rlv 1. Palpal tibia very slightly longer than broad, with small spike-like dorso-retrolateral apophysis (Figs 115, 116, 130); tegulum oval, with narrow finger-like prolateral apical tegular process, lobate mesal retrolateral tegular process with additional smaller subtriangular lobe, and slender strongly curved median apophysis (Fig. 129); embolus very slender, originating proximally and entering S-shaped groove in prolateral subtegulum, continuing along prolateral margin distally (Fig. 128), before entering groove in apical tegular process. Additional material examined. SOUTH AFRICA: Northern Cape: Namaqua National Park, Koeroebees, 30°08.683’S, 17°42.177’E, 240 m a.s.l., 14.I.2021, leg. C. Haddad, R. Booysen, R. Christiaan & A. Stander (leaf litter, dry river bed), 1♀ (NCA 2021/726; epigyne lost in preparation); Same locality, 27.III.2022, leg. C. Haddad, R. Booysen, L. Malope & S. Sibisi (leaf litter, dry river bed), 2♂ 2♀ (S.E.M. preparations). Variation. Total length: females 2.78–3.32 (average 2.98, n = 7); males 2.05–2.98 (average 2.36, n = 4). Habitat and biology. All of the females collected had the epigyne plugged (Fig. 114). Recorded from the Succulent Karoo, Nama Karoo and Grassland biomes. At the type locality, it was a common species collected in the litter of short shrubs in a dry river bed, with adults far more common in autumn (March) than midsummer (January) or winter (July). Distribution. Widespread in the western half of South Africa (Fig. 136)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 26-27, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Leptodrassex capensis sp. nov. Figures 45, 80, 81 Type material. Holotype ♀: SOUTH AFRICA: Western Cape: Fisherhaven, near Hermanus, 34°21.430’S, 19°07.557’E, 30.IX.2007, leg. C. Haddad & R. Lyle (sifting leaf litter) (NCA 2008/360). Paratype: SOUTH AFRICA: Western Cape: Outeniquastrand, near George, 34°02.754’S, 22°17.037’E, 7.I.2015, leg. C. Haddad (base of grass tussocks), 1♀ (NCA 2015/1755). Etymology. This species is named for terra typica, the Western Cape Province of South Africa, from which it is known. Diagnosis. Females of L. capensis sp. nov. are most similar to those of L. murphyi sp. nov., but differ by the smaller copulatory openings and the slight curvature of the copulatory ducts (Fig. 80), which are respectively larger and strongly curved in the latter species (Fig. 82). Male unknown. Description. Female (holotype, NCA 2008/360). Colouration (Fig. 45): carapace and chelicerae creamy-yellow; endites and labium cream; sternum cream, margins yellow-brown at coxae; femora cream, remaining segments creamy-yellow. Abdomen cream dorsally and ventrally. Measurements: CL 1.04, CW 0.85, AL 1.52, AW 1.02, TL 2.85. Eye diameters and interdistances: AME 0.09, ALE 0.08, PME 0.09, PLE 0.08, AME–AME 0.07, AME–ALE 0.02, PME–PME 0.06, PME–PLE 0.05, ALE–PLE 0.01. Leg measurements: I 0.76, 0.34, 0.57, 0.48, 0.35 = 2.50; II 0.79, 0.37, 0.61, 0.54, 0.39 = 2.70; III 0.63, 0.33, 0.44, 0.40, 0.25 = 2.05; IV 1.00, 0.57, 0.76, 0.82, 0.22 = 3.37. Leg spination: femora: I and II spineless, III do 1, IV do 1 rl 1; patellae spineless; tibiae: I plv 2 rlv 2, II rlv 2, III pl 2 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1-4 rlv 1-3, II plv 0-1 rlv 1-2, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur and patella spineless, tibia pl 2 plv 1 rlv 1, tarsus pl 2 plv 2 rlv 1. Epigyne with weakly sclerotized recurved hemispherical ridges anteriorly, continuing along midline into posterior half, forming oval atria; copulatory openings transverse, situated in anteromesal part of atria; copulatory ducts narrow and initially directed posteriorly, bending laterally and diverging, with slight lateral bend before entering ovoid lateral spermathecae along their anterior margin; fertilization ducts on posteromesal margin of spermathecae, directed mesally (Figs 80, 81). Additional material examined. None. Variation. The single female paratype is 2.57 mm in total length. Habitat and biology. Both of the known females had plugged epigynes. This species was sampled from the ground (leaf litter and grass tussocks) in the Fynbos and Thicket biomes of South Africa. Distribution. Only known from two localities in the coastal parts of south-western South Africa (Fig. 44)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on page 16, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Leptopilos butleri sp. nov. Figs 87, 88, 121–126 Type material. Holotype ♀: BOTSWANA: Ngamiland: Okavango Delta, Shakawe, 18°21.960’S, 21°50.829’E, 28.XI.2006, leg. C. Haddad (leaf litter, dry savanna) (NCA 2007/1030). Paratypes: SOUTH AFRICA: Free State: Amanzi Private Game Reserve, 28°35.980’S, 26°24.935’E, 1– 30.IX.2012, leg. V. Butler (pitfall traps, Digitaria eriantha grazing), 7♂ 1♀ (NCA 2013/3295); Same locality, 28°35.506’S, 26°25.340’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Searsia burchellii woodland), 8♂ 4♀ (NCA 2013/3163). Etymology. The species is named for Vivian Butler, who collected most of the known material during a yearlong pitfall study in grassland biotopes (Haddad & Butler 2018). Diagnosis. The females of L. butleri sp. nov. can be easily recognized by the broad hemispherical anterior hood that is as wide as the lateral margins of the spermathecae (only as broad as the inner margins of the spermathecae in the type species, L. tenerrimus (O. Pickard-Cambridge, 1872); see Levy 2009: fig. 18), with a pair of dark n-shaped markings at their anterior margin (Fig. 121). It differs from L. vasivulva sp. nov., which has a vase-shaped atrium and a single median curved marking on the anterior hood (compare Figs 121 and 131). Males of L. butleri sp. nov. have shorter apical and retrolateral tegular processes than L. vasivulva sp. nov., with the retrolateral tibial apophysis of the latter species also constricted at its distal end (compare Figs 124 and 125 with Figs 134 and 135). Description. Female (holotype, Shakawe, NCA 2007/1030). Colouration (Fig. 87): carapace and chelicerae yellow; endites creamy-yellow, labium slightly darker; sternum creamy-yellow, margins brown at coxae; femora creamy-yellow, remaining segments yellow; abdomen cream dorsally and ventrally. Measurements: CL 1.16, CW 0.90, AL 1.79, AW 1.11, TL 2.98. Eye diameters and interdistances: AME 0.11, ALE 0.09, PME 0.10, PLE 0.08, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.05, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.91, 0.38, 0.67, 0.62, 0.46 = 3.04; II 0.81, 0.38, 0.61, 0.59, 0.43 = 2.82; III 0.71, 0.35, 0.52, 0.51, 0.31 = 2.40; IV 1.27, 0.46, 0.92, 1.05, 0.33 = 4.03. Leg spination: femora: I do 2, II do 2, III pl 1 do 3 rl 1, IV pl 1 do 3 rl 1; patellae spineless; tibiae: I plv 1 rlv 1, II rlv 1, III pl 2 rl 2 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur pl 1 do 2, patella spineless, tibia pl 1 plv 1, tarsus pl 1 rl 1 plv 3 rlv 3. Epigyne with broad hemispherical hood anteriorly, with paired n-shaped markings on either side of midline (Fig. 121); copulatory openings small, beneath posterior margin of hood, separated by distance equal to spermatheca width; copulatory ducts narrow, U-shaped initially, curving laterally, then looping anteriorly before entering almost globose spermathecae along their anterior margin (Fig. 122); fertilization ducts on posterior margin of spermathecae, directed posteriomedially. Male (paratype, Amanzi, NCA 2013/3163). Colouration (Fig. 88): similar to female, carapace slightly paler, abdomen slightly darker. Measurements: CL 0.92, CW 0.72, AL 1.06, AW 0.76, TL 2.08. Eye diameters and interdistances: AME 0.10, ALE 0.07, PME 0.08, PLE 0.07, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.03, PME–PLE 0.03, ALE–PLE 0.01. Leg measurements: I 0.81, 0.32, 0.60, 0.53, 0.41 = 2.67; II 0.71, 0.30, 0.54, 0.46, 0.38 = 2.39; III 0.62, 0.27, 0.42, 0.41, 0.28 = 2.00; IV 1.04, 0.37, 0.76, 0.87, 0.30 = 3.34. Leg spination: femora: I do 1, II do 1, III do 3 rl 1, IV pl 1 do 3 rl 1; patellae spineless; tibiae: I and II spineless, III pl 2 rl 1 plv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I spineless, II rlv 0-1, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur pl 1 do 2, patella spineless, tibia rlv 2, tarsus plv 1 with several modified distal setae ventrally. Palpal tibia slightly longer than broad, with single triangular retrolateral apophysis (Fig. 125); tegulum ovoid, with slender straight median apophysis with curved tip (Fig. 124); embolus very slender, originating proximally and entering groove in subtegulum, continuing along prolateral margin in S-shaped groove (Fig. 123), before entering groove in apical tegular process, partly hidden behind retrolateral tegular apophysis. Additional material examined. SOUTH AFRICA: Free State: Amanzi Private Game Reserve, 28°35.805’S, 26°25.190’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Digitaria eriantha grazing), 8♂ 2♀ (NCA 2013/3310); Same locality, 28°35.765’S, 26°25.455’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, mesa, plateau), 1♂ (NCA 2013/3236); Same data but 1–31.X.2012, 1♀ (NCA 2014/1541); Same locality, 28°35.565’S, 26°25.550’E, 1– 31.X.2012, leg. V. Butler (pitfall traps, mesa, plateau), 18♂ 4♀ (NCA 2014/1562); Same locality, 28°35.506’S, 26°25.340’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Searsia burchellii woodland), 1♀ (NCA 2013/3162); Same data but 1–30.IV.2013, 1♀ (NCA 2013/3581); Same locality, 28°35.975’S, 26°25.435’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Tarchonanthus camphoratus woodland), 1♂ (NCA 2013/3321); Same locality, 28°35.785’S, 26°26.335’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, Vachellia karroo woodland), 1♂ 1♀ (NCA 2013/3380). Gauteng: Bronkhortspruit, Ezemvelo Nature Reserve, Farm Elandsfontein, 25°42’20.7’’S, 28°56’27.0’’E, 1403 m a.s.l., 8.III.2005, leg. R. Koko (pitfalls, koppie crest), 1♀ (NCA 2016/3183); Wonderboom, Tswaing Nature Reserve, Farm Tswaing, 25°24’15.1’’S, 28°03’34.9’’E, 1116 m a.s.l., 9.XI.2004, leg. R. Koko (baited pitfall, open woodland), 1♀ (NCA 2014/3235), 1♀ (NCA 2014/3252). KwaZulu-Natal: Tembe Elephant Park, Viewing Tower, Open woodland/sand, 27°02.225’S, 32°24.905’E, 115 m a.s.l., 3.XII.2015, leg. C. Haddad, R. Booysen, V. & N. Butler (hand collecting), 1♀ (NCA 2016/2045). Limpopo: Vyeboom, 23°18.534’S, 30°24.486’E, 24.I.2015, leg. E. Mauda (pitfalls), 1♂ (NCA 2016/245); Vhembi Biosphere Reserve, Mara Research Station, 23°02’S, 29°39’E, 7.XI.2012, leg. C. Schoeman (pitfalls, Makhado sweet bushveld), 1♂ (NCA 2015/1172). Northern Cape: Prieska district, Green Valley Nuts, 29°34.924’S, 22°54.376’E, 28.I.2009, leg. C. Haddad (base of grass tussocks), 1♂ (NCA 2009/1474). ZIMBABWE: Bulawayo, Hillside, 20°10’S, 28°35’E, III–IV.1999, leg. M. FitzPatrick (pitfall traps), 2♀ (NMZ/ A14034); Mtshelele Dam, 20°35’S, 28°36’E, 1–3.XII.2004, leg. M. FitzPatrick, 1♀ (NMZ/ A15180); Pomongwe, 20°32’S, 28°30’E, XII.2004, leg. M. FitzPatrick, 1♂ (NMZ/ A15250); Rowallan Park, 20°25’S, 28°31’E, VII.2005, leg. M. FitzPatrick, 1♀ (NMZ/ A15864). Variation. Total length: females 2.15–3.08 (average 2.51, n = 13); males 1.77–2.23 (average 2.03, n = 21). Habitat and biology. The extent of epigyne plugging was not quantified for this species. It occupies Nama Karoo, grassland and savanna habitats, and was predominantly sampled from the ground by pitfalls, litter sifting, beneath rocks or from grass tussocks, and only rarely from woody vegetation by beating. It represented 1.11% of the ground-dwelling spider fauna in a pitfall survey in the grassland biome, but was almost exclusively sampled there from Searsia burchellii closed evergreen shrubland and cultivated Digitaria eriantha pastures (Haddad & Butler 2018). Distribution. Widely distributed in South Africa, Zimbabwe and Botswana (Fig. 136)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 23-26, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932, {"references":["Haddad, C. R. & Butler, V. P. (2018) Ground-dwelling spider assemblages in contrasting habitats in the central South African Grassland Biome. Koedoe, 60, a 1482. https: // doi. org / 10.4102 / koedoe. v 60 i 1.1482","Levy, G. (2009) New ground-spider genera and species with annexed checklist of the Gnaphosidae (Araneae) of Israel. Zootaxa, 2066, 1 - 49. https: // doi. org / 10.11646 / zootaxa. 2066.1.1"]}

Afrodrassex catharinae sp. nov. Figures 9, 10, 39–43 Type material. Holotype ♀: SOUTH AFRICA: KwaZulu-Natal: Ndumo Game Reserve, Crocodile farm, 26°54.426’S, 32°19.185’E, 12.I.2007, leg. C. Haddad (under rocks) (NCA 2007/3068). Paratypes: SOUTH AFRICA: KwaZulu-Natal: Ndumo Game Reserve, Nyamiti Pan, 26°53.409’S, 32°17.576’E, 35 m a.s.l., 3.XII.2019, leg. C. Haddad & V. Swart (canopy fogging, Pappea capensis), 2♂ 1♀ (NCA 2020/270). Etymology. The species is named for Catharine Hanekom, Regional Ecologist of Ezemvelo KZN Wildlife for the Maputaland Region, in recognition of more than twenty years of logistical support to the senior author during arachnid surveys in northern KwaZulu-Natal Province. Diagnosis. This species is most similar to A. balrog sp. nov., but females can be distinguished by their large copulatory openings and globular spermathecae (Figs 39, 40), which are small and transversely oval, respectively, in A. balrog sp. nov. (Figs 34, 35). Males of A. catharinae sp. nov. have a small sharp tooth-like retrolateral tibial apophysis in lateral view (Figs 41, 43), compared to an elongate spike-like retrolateral tibial apophysis in A. balrog sp. nov. (Figs 29, 38). Description. Female (holotype, NCA 2007/3068). Colouration (Fig. 9): carapace and chelicerae yellow; labium and endites creamy-yellow; sternum cream, margins yellow-brown at coxae; femora creamy-yellow, remaining segments yellow; abdomen cream dorsally and ventrally. Measurements: CL 1.16, CW 0.98, AL 1.73, AW 1.17, TL 3.20. Eye diameters and interdistances: AME 0.11, ALE 0.08, PME 0.09, PLE 0.08, AME–AME 0.05, AME–ALE 0.01, PME–PME 0.08, PME–PLE 0.04, ALE–PLE 0.01. Leg measurements: I 0.78, 0.38, 0.59, 0.56, 0.41 = 2.72; II 0.87, 0.40, 0.65, 0.64, 0.44 = 3.00; III 0.72, 0.37, 0.49, 0.59, 0.30 = 2.47; IV 1.11, 0.44, 0.81, 0.95, 0.33 = 3.64. Leg spination: femora: I do 2, II do 2, III do 2 rl 1, IV do 3 rl 1; patellae: spineless; tibiae: I plv 2 rlv 2, II plv 2 rlv 2, III pl 2 rl 1 plv 2, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 2 plv 1 vt 3; palp: femur do 1, patella spineless, tibia pl 2 plv 1, tarsus pl 1 rl 1 plv 4 rlv 2. Epigyne with elongate curved 6-shaped copulatory openings at anterior of epigynal plate (Fig. 39); copulatory ducts narrow, initially directed posteriorly, looping mesally and anteriorly before following path of faint external ridges around periphery of epigyne, laterally with distinct kink before continuing posteriorly, bending mesally before entering small globular spermathecae on their ventral surface; fertilization ducts on anterolateral margin of spermathecae, directed posteriorly (Figs 39, 40). Male (paratype, NCA 2020/270). Colouration (Fig. 10): as for female. Measurements: CL 1.16, CW 0.94, AL 1.19, AW 0.84, TL 2.35. Eye diameters and interdistances: AME 0.11, ALE 0.08, PME 0.09, PLE 0.09, AME–AME 0.06, AME–ALE 0.01, PME–PME 0.07, PME–PLE 0.05, ALE–PLE 0.01. Leg measurements: I missing; II 0.90, 0.41, 0.76, 0.67, 0.44 = 3.18; III 0.79, 0.37, 0.55, 0.62, 0.33 = 2.66; IV 1.24, 0.44, 0.88, 1.02, 0.37 = 3.95. Leg spination (leg I derived from second male paratype in NCA 2020/270): femora: I do 2, II do 2, III do 3 rl 1, IV pl 1 do 3 rl 1; patellae spineless; tibiae: I plv 2 rlv 2, II plv 2 rlv 2, III pl 2 rl 1 plv 2 rlv 1, IV pl 2 rl 2 plv 1 vt 2; metatarsi: I plv 1 rlv 1, II plv 1 rlv 1, III pl 3 rl 1 plv 1 vt 3; IV pl 3 rl 2 plv 1 vt 3; palp: femur do 1, other segments spineless. Palp: patella with small proximal ventral bump-like apophysis; tibia slightly longer than broad, with small sharp tooth-like apophysis, with additional tiny basal tooth (Figs 41, 43); tegulum ovoid, with large curved apical tegular process distally; median apophysis absent; conductor massive, translucent and comma-shaped, covering most of ventral aspect of tegulum; embolus originating proximally on tegulum, curling proximally and prolaterally, curving gradually along prolateral margin of tegulum, bending sharply near distal end of cymbium (Figs 41–43). Additional material examined. None. Variation. Total length: females 3.03–3.20 (average 3.12, n = 2); males 2.35–2.75 (average 2.55, n = 2). Habitat and biology. Both of the known females had plugged epigynes. This species was sampled from the ground and canopies of a short tree (Pappea capensis) in savanna habitats. Distribution. Only known from the type locality in northern KwaZulu-Natal, South Africa (Fig. 44)., Published as part of Haddad, Charles R. & Booysen, Ruan, 2022, The ground spider genera Leptodrassex Murphy, 2007 and Leptopilos Levy, 2009 (Araneae: Gnaphosidae) in southern Africa, including the description of a new genus and seven new species, pp. 1-32 in Zootaxa 5194 (1) on pages 9-10, DOI: 10.11646/zootaxa.5194.1.1, http://zenodo.org/record/7141932

Orthobula arca sp. nov. Figs 3, 4, 59–63 Diagnosis. Females of O. arca sp. nov. can be easily recognized from congeners by the large arch-shaped anterior ridge, with lateral extensions flanking the CO (Fig. 59). Males can be easily distinguished from congeners by the distinct bend of the embolus at approximately 2/3 its length (Figs 61–63). Etymology. From the Latin for “arch”, referring to the shape of the anterior margin of the epigynal atrium. Female (holotype, Deelhoek, NCA 2021/1248). Measurements: CL 0.93, CW 0.70, AL 1.03, AW 0.79, TL 2.00 (1.60–2.30), PERW 0.29, MOQAW 0.12, MOQPW 0.17, MOQL 0.14. Length of leg segments: I 0.59 + 0.22 + 0.51 + 0.44 + 0.27 = 2.03; II 0.51 + 0.21 + 0.38 + 0.38 + 0.25 = 1.73; III 0.42 + 0.20 + 0.30 + 0.35 + 0.22 = 1.49; IV 0.56 + 0.22 + 0.47 + 0.52 + 0.29 = 2.06. Colour: carapace orange-brown, with mottled black margins and faint black mottled patch centrally; chelicerae yellow-brown; endites and labium yellow-brown, cream distally; sternum yellow-orange, with yellow-brown margins; palps yellow-brown; legs with femora, patellae and tibiae yellow-brown, other leg segments yellow; patellae and tibiae I with black mottling dorsally and laterally, on II–IV on retrolateral side only; abdomen dark grey dorsally, with pair of cream longitudinal patches anterolaterally and oblique cream markings at midpoint, merging medially to form X; venter cream, with faint mottling on epigastric plate, faint mottled grey patch in front of spinnerets; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 3. Epigyne with lateral CO in broad arch-shaped epigynal ridge, with posterolateral extensions (Fig. 59); CO short, directed posteriorly, entering round SBB at centre of epigyne; CD approximately same length as width of SBB; SBB separated by slightly less than their width; BU subtriangular, with boomerang-shaped accessory gland on their anteromesal surface; SBB connected to ST I by short looping ducts, entering broad, transversely oval ST I on their mesal surface (Fig. 60). Male (paratype, Erfenis Dam Nature Reserve, NCA 2009/3636). Measurements: CL 0.81, CW 0.64, AL 0.68, AW 0.58, TL 1.53 (1.46–1.70), PERW 0.26, MOQAW 0.10, MOQPW 0.14, MOQL 0.13. Length of leg segments: I 0.47 + 0.18 + 0.40 + 0.35 + 0.23 = 1.63; II 0.44 + 0.17 + 0.33 + 0.32 + 0.23 = 1.49; III 0.37 + 0.17 + 0.25 + 0.29 + 0.19 = 1.17; IV 0.51 + 0.18 + 0.41 + 0.44 + 0.25 = 1.79. Colour: carapace orange-brown, with mottled black margins and indistinct black mottled patch centrally; chelicerae yellow-brown; endites and labium yellow-brown, cream distally, labium darker in proximal half; sternum yellow-orange, with yellow-brown margins and faint black mottling; palps yellow-brown; legs with femora, patellae and tibiae yellow-brown, other leg segments yellow; patellae and tibiae I with black mottling dorsally and laterally, on II–IV on retrolateral side only; abdomen black dorsally, with pair of cream subtriangular patches anterolaterally and transverse cream band at midpoint, latter slightly projecting anteriorly along midline; venter cream, with small mottled grey patch in front of spinnerets; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 5 rlv 5, II plv 4 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 3. Palpal femur with small, finger-like, curved RFA (Fig. 63); tibia with small, simple triangular apophysis; retrodistal tegular ridge weakly developed, restricted to retrolateral surface, not curving around ventrally to base of embolus (Figs 62, 63); basal section of embolus leaning slightly prolaterally, with distinct bend at 2/3 its length, tip directed slightly retrolaterally (Figs 61–63). Type material. Holotype ♀: SOUTH AFRICA: Free State: Bloemfontein district, Deelhoek farm, 28°51’S, 26°07’E, 1265 m a.s.l., 17.XI.2001, leg. C. Haddad (Eucalyptus leaf litter) (NCA 2021 /1248). Paratypes: SOUTH AFRICA: Free State: Benfontein Nature Reserve, 28°50.452’S, 24°49.473’E, 13.III.2010, leg. C. Haddad (base of grass tussocks), 2♂ 2♀ (NCA 2010 /317); Bethulie district, Tussen-die-Riviere Nature Reserve, 30°29’S, 26°11’E, 15.X.2008, leg. L. Lotz & C. Haddad (under rocks, riparian woodland edge), 1♀ (NMBA 12755); Bloemfontein district, Hopefield farm, 28°52.949’S, 26°09.336’E, 1290 m a.s.l., 11.XI.2001, leg. C. Haddad (sweepnetting, Eragrostis grass), 1♀ (NCA 2021 /1246); Bloemfontein district, Deelhoek farm, 28°51’S, 26°07’E, 1265 m a.s.l., 21.X.2001, leg. C. Haddad (sweeps, Themeda grassland), 1♀ (MHBU); Bloemfontein, Free State National Botanical Gardens, 29°02.892’S, 26°12.662’E, 1390 m a.s.l., 26.II–13.III.2011, leg. J.A. Neethling (pitfalls, open grassland), 4♂ 6♀ (MHBU); Brandfort district, Florisbad Research Station, 28°46’S, 26°05’E, 1250 m a.s.l., 8.XII.1987, leg. L.N. Lotz (pitfall traps), 1♀ (NMBA 8627); Erfenis Dam Nature Reserve, Site 1, 28°28.892’S, 26°48.508’E, 31.VIII–30.IX.2009, leg. R. Fourie & A. Grobler (pitfall traps, grassland near trench), 1♂ (NCA 2009 /3636); Kroonstad district, Koffielaagte farm, 27°29’S, 27°28’E, 1425 m a.s.l., 4.I.2002, leg. C. Haddad (sweeps in grassland), 1♀ (NCA 2021 /1247). Other material examined. SOUTH AFRICA: Free State: Amanzi Private Game Reserve, 28°35.845’S, 26°26.280’E, 1–30.IX.2012, leg. V. Butler (pitfall traps, open grassland), 1♀ (NCA 2013 /3355); Same locality, 28°36.365’S, 26°25.030’E, 1–30.IV.2013, leg. V. Butler (pitfall traps, Acacia karroo woodland), 1♂ 3♀ (NCA 2013 /3551); Same locality, 28°35.430’S, 26°26.290’E, 1–31.I.2013,leg. V. Butler (pitfall traps, vegetation around dam), 1♂ (NCA 2013 /4307); Same locality, 28°36.722’S, 26°26.148’E, 24.XI.2013 – 20.I.2014, leg. C. Haddad & V. Butler (pitfalls, open woodland), 1♂ (NCA 2014 /1077); Same locality, 28°36.712’S, 26°26.186’E, 1450 m a.s.l., 10.X.2017, leg. C. Haddad (base of grass tussocks), 1♀ (NCA 2017 /933); Same locality, 28°35’S, 26°25’E, 19–23.I.2014, leg. AFRAS colloquium delegates, 1♀ (NCA 2014 /1028); Bloemfontein, Free State National Botanical Gardens, 29°02.892’S, 26°12.662’E, 1390 m a.s.l., 24.IX–8.X.2009, leg. C. Haddad (pitfalls, Rhus lancea woodland), 1♀ (NCA 2009 /3490); Same locality, 27.X–16.XI.2009, leg. C. Haddad (pitfall traps), 1♀ (NMBA 13989); Same locality, 9.XII.2009 – 4.I.2010, leg. C. Haddad (pitfall traps, grassland), 1♂ 1♀ (NMBA 15103), 3♂ 7♀ (NMBA 15024); Same locality, 20.X–9.XI.2013, leg. C. Haddad (open pitfalls, grassland), 3♂ 3♀ (NCA 2017 /1555); Same locality, XII.2014, leg. C. Haddad (open pitfalls, grassland), 2♀ (NCA 2015 /1845); Same locality, VI.2016, leg. C. Haddad & L. Whitehead (Hyparrhenia hirta grass tussocks), 1♀ (NCA 2016 /3060); Erfenis Dam Nature Reserve, Site 1, 28°28.892’S, 26°48.508’E, 30.IX–28.X.2009, leg. R. Fourie & A. Grobler (pitfall traps, grassland near trench), 3♀ (NCA 2009 /3579), 1♀ (NMBA 14359); Erfenis Dam Nature Reserve, Site 8, 28°29.804’S, 26°48.503’E, 31.VIII–30.IX.2009, leg. R. Fourie & A. Grobler (pitfall traps, Themeda grassland), 3♀ (NCA 2009 /3655); Same data but 30.IX–28.X.2009, 1♀ (NCA 2009 /3604), 2♀ (NMBA 14485); Erfenis Dam Nature Reserve, Burned site 1, 28°30.373’S, 26°48.437’E, 21.IX–22.X.2005, leg. C. Haddad, S. Otto & R. Poller (pitfall traps, grassland), 1♀ (NMBA 13923); Same data but 22.X–22.XI.2005, leg. C. Haddad, S. Otto & R. Poller, 3♀ (NMBA 13924); Same data but 23.XII.2005 – 23.I.2006, leg. C. Haddad, 1♂ 1♀ (NMBA 13925); Same data but 24.II–27.III.2006, leg. S. Otto & R. Poller, 3♂ (NMBA 13926); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 1♂ 1♀ (NMBA 13927); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 3♀ (NMBA 13928); Same data but 26.V–28.VI.2006, leg. S. Otto & R. Poller, 1♂ 1♀ (NMBA 13929); Same data but 28.VI–28.VII.2006, leg. C. Haddad & R. Lyle, 1♂ (NMBA 13930); Erfenis Dam Nature Reserve, Burned site 2, 28°30.134’S, 26°48.427’E, 22.X–22.XI.2005, leg. C. Haddad, S. Otto & R. Poller (pitfall traps, grassland), 1♂ 1♀ (NMBA 13931); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 1♀ (NMBA 13933); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 1♀ (NMBA 13932); Erfenis Dam Nature Reserve, Burned site 3, 28°29.990’S, 26°48.486’E, 21.IX–22.X.2005, leg. C. Haddad, S. Otto & R. Poller, 1♀ (NMBA 13935); Same data but 22.X–22.XI.2005, leg. C. Haddad, S. Otto & R. Poller (pitfall traps, grassland), 2 imm. 4♂ (NMBA 13934); Same data but 23.XII.2005 – 23.I.2006, leg. C. Haddad, S. Otto & R. Poller, 2♂ (NMBA 13936); Same data but 24.I–24.II.2006, leg. C. Haddad & R. Lyle, 1♂ (NMBA 13937); Same data but 24.II–27.III.2006, leg. S. Otto & R. Poller, 5♀ (NMBA 13938); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 5♂ 5♀ (NMBA 13939); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 1 imm. 2♂ 6♀ (NMBA 13940); Same data but 28.VII–1.IX.2006, leg. C. Haddad, S. Otto & R. Poller, 1♂ 1♀ (NMBA 13941); Erfenis Dam Nature Reserve, Unburned site 1, 28°29.888’S, 26°48.488’E, 21.IX–22.X.2005, leg. C. Haddad, S. Otto & R. Poller (pitfall traps, grassland), 1♂ (NMBA 13900); Same data but 22.X–22.XI.2005, leg. C. Haddad, S. Otto & R. Poller, 2 imm. 2♂ 2♀ (NMBA 13901); Same data but 22.XI– 23.XII.2005, leg. C. Haddad, 11♂ 7♀ (NMBA 13902); Same data but 23.XII.2005 – 23.I.2006, leg. C. Haddad, S. Otto & R. Poller, 6♂ 3♀ (NMBA 13903); Same data but 23.I–24.II.2006, leg. C. Haddad & R. Lyle, 4♂ 2♀ (NMBA 13904); Same data but 24.II–27.III.2006, leg. S. Otto & R. Poller, 2 imm. 5♂ 3♀ (NMBA 13905); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 1 imm. 1♀ (NMBA 13906); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 1♂ 2♀ (NMBA 13907); Erfenis Dam Nature Reserve, Unburned site 2, 28°29.706’S, 26°48.281’E, 21.IX–22.X.2005, leg. C. Haddad, S. Otto & R. Poller (pitfall traps, grassland), 2♂ 1♀ (NMBA 13908); Same data but 22.X–22.XI.2005, leg. C. Haddad, S. Otto & R. Poller, 1 imm. 4♂ 2♀ (NMBA 13909); Same data but 22.XI–23.XII.2005, leg. C. Haddad, 2♂ 2♀ (NMBA 13910); Same data but 23.XII.2005 – 23.I.2006, leg. C. Haddad, S. Otto & R. Poller, 1 imm. 3♂ 4♀ (NMBA 13911); Same data but 24.II–27.III.2006, leg. S. Otto & R. Poller, 2♂ (NMBA 13912); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 7♂ 5♀ (NMBA 13913); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 6♂ 2♀ (NMBA 13914); Same data but 1–30.IX.2006, leg. C. Haddad, S. Otto & R. Poller, 1♂ (NMBA 13915); Erfenis Dam Nature Reserve, Unburned site 3, 28°29.741’S, 26°48.065’E, 22.XI–23.XII.2005, leg. C. Haddad (pitfall traps, grassland), 6♂ 6♀ (NMBA 13916); Same data but 23.XII.2005 – 23.I.2006, leg. C. Haddad, S. Otto & R. Poller, 1♂ 1♀ (NMBA 13917); Same data but 23.I–24.II.2006, leg. C. Haddad & R. Lyle, 2♀ (NMBA 13918); Same data but 24.II–27.III.2006, leg. S. Otto & R. Poller, 5♂ 2♀ (NMBA 13919); Same data but 27.III–28.IV.2006, leg. C. Haddad & R. Lyle, 2♂ 3♀ (NMBA 13920); Same data but 28.IV–26.V.2006, leg. C. Haddad & R. Lyle, 2♂ 2♀ (NMBA 13921); Same data but 1–30.IX.2006, leg. C. Haddad, S. Otto & R. Poller, 1♀ (NMBA 13922); Sandveld Nature Reserve, 27°44.043’S, 25°45.805’E, 2–30.X.2009, leg. R. Fourie & A. Grobler (pitfall traps, grassland), 4♀ (NCA 2009 /3566); Same locality, 2.IX–2.X.2009, leg. R. Fourie & A. Grobler (pitfall traps, grassland), 1♀ (NMBA 15107). Gauteng: Johannesburg, Roodepoort, Ruimsig, 26°08’S, 27°51’E, 20.V.2000, leg. A. Leroy (in house), 1♀ (NCA 2007 /3834); Pretoria, Rietondale Research Campus, 25°43.780’S, 28°13.862’E, 16.VII.1998, leg. J. Nkwana (pitfall traps), 1♂ (NCA 2007 /1151). KwaZulu-Natal: Sani Pass Elevational Transect, 30°11’00.6’’S, 30°09’07.8’’E, 900 m a.s.l. (8a), leg. University of Pretoria students (pitfall traps), 1♀ (NCA 2011 /774); Sani Pass, 29°39’S, 29°27’E, IX.2006, leg. D. Prentice (pitfall traps), 1♀ (NCA 2008 /1982). Habitat and biology. This species was regularly collected from pitfall traps in the Grassland Biome of central South Africa. In contrast with its congeners, this species was far more abundant in open grasslands (3.8 %; Haddad et al. 2015) than in leaf litter (absent in Butler & Haddad 2011; Haddad et al. 2019) or shrubland habitats (0.13 %; Haddad & Butler 2018). On rare occasions, it was collected by sweeping grass (e.g. Haddad 2005). In a study investigating the impacts of controlled burning in central South African grasslands (Haddad et al. 2015), 180 adult O. arca sp. nov. were collected from the Erfenis Dam Nature Reserve. The proportion of males (n = 92) to females (n = 88) was almost identical. Considering its dominance as a component of Corinnidae (n = 234) in the study (its placement at the time), it likely contributed to the observed significant impact of fire on this family, as 126 individuals were sampled from the three unburnt control sites compared to only 54 individuals sampled at the three burnt sites. Regarding adult phenology (Fig. 64), the species occurred throughout the year, but was most common from late spring to late autumn, and was rare during winter. Distribution. Widespread in the Grassland Biome of central South Africa (Fig. 82)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on pages 364-367, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677, {"references":["Haddad, C. R., Foord, S. H., Fourie, R. & Dippenaar-Schoeman, A. S. (2015) Effects of a fast-burning spring fire on the grounddwelling spider assemblages (Arachnida: Araneae) in a central South African grassland habitat. African Zoology, 50, 281 - 292. https: // doi. org / 10.1080 / 15627020.2015.1088400","Butler, V. P. & Haddad, C. R. (2011) Spider assemblages associated with leaf litter of three tree species in central South Africa (Arachnida: Araneae). African Journal of Ecology, 49, 301 - 310. https: // doi. org / 10.1111 / j. 1365 - 2028.2011.01265. x","Haddad, C. R., De Jager, L. & Foord, S. H. (2019) Habitats and cardinal directions are key variables structuring spider leaf litter assemblages under Searsia lancea. Pedobiologia, 73, 10 - 19. https: // doi. org / 10.1016 / j. pedobi. 2019.01.002","Haddad, C. R. & Butler, V. P. (2018) Ground-dwelling spider assemblages in contrasting habitats in the central South African Grassland Biome. Koedoe, 60, a 1482. https: // doi. org / 10.4102 / koedoe. v 60 i 1.1482","Haddad, C. R. (2005) Ecology of spiders (Arachnida: Araneae) inhabiting Themeda triandra Forskal grassland in semi-arid South Africa. Navorsinge van die Nasionale Museum, Bloemfontein, 21, 25 - 36."]}

Orthobula milloti Caporiacco, 1949 Figs 9, 10, 90–95 Orthobula milloti Caporiacco, 1949: 441, fig. 75a–c (♂ and ♀ syntypes: KENYA: Nairobi [01°15’S, 36°50’E], 1500 m a.s.l., 1944, leg. Toschi & Meneghetti, deposited in MSNVR—not examined). Remark. Although we were unable to examine the syntypes of O. milloti, we were able to redescribe a topotypical female and match that to other specimens from East Africa, both males and females. Diagnosis. Females of O. milloti share with those of O. aethiopica sp. nov. a broad, semicircular anterior ridge in the epigyne, but can be distinguished by the short copulatory ducts that are approximately equal in length to the width of the SBB (Fig. 90), while twice as long as the width of SBB in O. aethiopica sp. nov. (Fig. 54). Males of O. milloti most closely resemble those of O. radiata in the short, straight spike-like embolus, but can be distinguished by the shape of the RFA, which is simple and finger-like in O. milloti (Fig. 95), while bifid in O. radiata (Fig. 100). Female (Nairobi, BMNH). Measurements: CL 1.15, CW 0.86, AL 1.04, AW 0.98, TL 2.52, PERW 0.24, MOQAW 0.10, MOQPW 0.13, MOQL 0.11. Length of leg segments: I 0.47 + 0.21 + 0.44 + 0.41 + 0.21 = 1.74; II 0.43 + 0.17 + 0.33 + 0.35 + 0.29 = 1.57; III 0.35 + 0.16 + 0.27 + 0.33 + 0.18 = 1.29; IV 0.48 + 0.18 + 0.42 + 0.47 + 0.22 = 1.77. Colour: carapace dark brown, with faint black mottling; chelicerae dark brown; endites and labium dark brown, creamy-grey distally; sternum dark brown, margins slightly darker; palps brown, distal ends of segments with yellow rings; legs with femora, patellae and tibiae I and II brown, of III and IV yellow-brown, all with black mottling laterally; distal ends of femora and tibiae, and basal end of patellae, with yellow rings; metatarsi yellow-brown, with faint mottling; tarsi yellow; abdomen black dorsally, with pair of cream longitudinal patches anterolaterally and transverse cream band at midpoint; venter creamy-grey; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 6, II plv 5 rlv 5; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 3. Epigyne with lateral CO in broad semi-circular anterior epigynal ridge (Fig. 90); CD very short, initially directed mesally, curving sharply before entering round SBB at centre of epigyne; CO separated by approximately SBB diameter, SBB separated by approximately ¼ their width; CD approximately as long as SBB width; BU ovoid, almost round, with teardrop-shaped accessory gland on their anteromesal surface; SBB connected to ST I by short looping longitudinal ducts, entering transversely oval ST I on their mesal surface; ST I almost touching along their mesal surfaces (Fig. 91). Male (Pugu Forest Reserve, MRAC 159243). Measurements: CL 0.81, CW 0.66, AL 0.83, AW 0.65, TL 1.64, PERW 0.32, MOQAW 0.12, MOQPW 0.16, MOQL 0.14. Length of leg segments: I 0.62 + 0.24 + 0.53 + 0.49 + 0.27 = 2.15; II 0.54 + 0.21 + 0.42 + 0.42 + 0.27 = 1.86; III 0.43 + 0.19 + 0.33 + 0.39 + 0.26 = 1.60; IV 0.57 + 0.21 + 0.49 + 0.57 + 0.31 = 2.15. Colour: carapace deep brown, pits with darker edges, lateral margins black; chelicerae brown, with faint black mottling; endites and labium brown, lighter distally; sternum orange-brown, with sparse mottling, margins darker; palps yellow-brown; legs yellow-brown, anterior femora darker; all femora, patellae and tibiae with distal mottled patches laterally; abdomen black, with pair of dark yellow-brown oval anterolateral markings; venter brown, with dense black mottling laterally, epigastric plate deep brown; spinnerets yellow-brown. Leg spination: femora and patellae: spineless; tibiae: I plv 5 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Palpal femur with curved, finger-like RFA (Fig. 95); tibia with small, simple triangular RTA; retrodistal tegular ridge short, not curving around ventrally to base of embolus (Fig. 92) embolus short, straight, spike-like (Figs 92–94). Material examined. KENYA: Nairobi, grounds of National Museum [01°16’S, 36°48’E], 12.IV.1988, leg. A. Russell-Smith (in tall grass), 1♀ (BMNH). TANZANIA: Pwani Region: Kisarawe district, Pugu Forest Reserve, 09°59’S, 39°07’E, 11.XI.1979, leg. K.M. Howell, 1♂ (MRAC 159243). Lindi Region: Lindi District, Litipo Forest Reserve, 10°02’S, 39°29’E, VII–IX.1993, leg. Frontier Tanzania, 1♀ (ZMUC). Tanga Region: Mlola, Mafia, 07°53’S, 39°50’E, X–XI.1990, leg. Frontier Tanzania (evergreen coastal thicket on clay, coral rag), 2♂ (ZMUC). Habitat and biology. A rare ground-dwelling spider sampled in urban natural habitat fragments, and forest and savanna habitats. Distribution. Only known from western Kenya and eastern Tanzania (Fig. 101)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on pages 374-375, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677, {"references":["Caporiacco, L. di (1949) Aracnidi della colonia del Kenya raccolti da Toschi e Meneghetti negli anni 1944 - 1946. Commentationes Pontificia Academia Scientiarum, 13, 309 - 492."]}

Genus Orthobula Simon, 1897 Orthobula Simon, 1897a: 152; Deeleman-Reinhold 2001: 431. Type species. Orthobula impressa Simon, 1897, by original designation. Diagnosis. Orthobula shares with Capobula the large number of paired ventral spines on the anterior tibiae and metatarsi, and the absence of ventral cusps in both sexes. In somatic morphology, Afrotropical species of Orthobula have pits across the entire carapace surface, including along the midline (Figs 13–16), while absent along the midline in Capobula (Haddad et al. 2021: figs 11, 14) and some Asian species of Orthobula. Female Orthobula can be distinguished by the posteriorly positioned spermathecae (as opposed to laterally positioned in Capobula), while male Orthobula can be recognized by the considerably inflated palpal tegulum (only as wide as cymbium in Capobula), the narrow, finely coiled embolus (longer, uncoiled and retrolaterally directed in Capobula), and the absence of a patellar apophysis (present in Capobula). Description. Small spiders, 1.30–2.70mm in length; carapace orange-brown to wine-red, frequently with mottled black markings (Figs 1–12); carapace oval, broadest at coxae II, eye region narrow; fovea absent, only a short shallow depression (Fig. 13); posterior margin slightly concave; carapace surface finely wrinkled, with series of deep pits forming striate patterns over entire surface (Figs 13–16), each pit with central pore, often plugged with unknown exudate (Fig. 14); surface sparsely covered with scattered fine curved setae, particularly in eye region (Figs 13–16). All eyes surrounded by black rings, weakly developed around PME (Figs 1–12); AER procurved in anterior view, recurved in dorsal view (Figs 15, 16); AME very slightly larger than ALE; AME separated by approximately ½ their diameter, nearly touching ALE (Figs 15, 16); PER recurved in dorsal view, PME oval, PLE round; PME and PLE similar in diameter; median ocular quadrangle narrower anteriorly than posteriorly, length and posterior width approximately equal. Chilum indistinct, a tiny transverse sclerite; cheliceral promargin with three teeth, separated by approximately their basal width, proximal tooth smallest, median tooth slightly larger than distal; retromargin with two subequal teeth, set close together; cheliceral escort seta present (Figs 17, 18); fang with distinct serrula; endites convergent, notched laterally (Fig. 17), with distinct serrula comprising sharp, ventrally curved denticles (Fig. 19); dense maxillar hair tuft on mesal margins (Figs 17–19); labium trapezoidal, slightly wider than long. Pleural bars sclerotised, isolated; sternum shield-shaped, longer than broad, surface smooth with deep pits without pores, sparsely covered in long straight setae (Figs 20, 21); precoxal triangles present, intercoxal sclerites present between all coxal pairs. Leg formula 4123, legs I and IV often very similar in length; sparsely covered in long fine setae; femora I with mesal convex curvature, all femora strongly constricted proximally, with ventral setal bases tuberculate (Fig. 22); patellar indentation on retrolateral side narrow, with lyriform organ at proximal end; anterior legs with strong paired ventral spines, chemosensory setae and trichobothria on tibiae, metatarsi and tarsi (Figs 23–33); metatarsi with single long distal trichobothrium (Fig. 29) and dorsal stopper (Fig. 30), metatarsi III and IV with ventral preening comb at distal end; tarsi usually with single or pair of short trichobothria at 1/3 their length and long pair at 2/3 their length (Figs 31, 32, 38); trichobothria with sunken distal plate, distal margin of hood overlapping plate, hood with two curved ridges, roughly concentric (Fig. 28); tarsal organ oval, very slightly elevated from integument, surface finely wrinkled, opening oval and distally placed (Fig. 39); paired tarsal claws short, with four intermediate teeth and moderately dense tenant setae forming claw tufts in between (Figs 34–37); palpal claw simple, sharply curved distally. Abdomen oval, clearly larger in females than males, with dorsal scutum in males only; dorsum with very sparse fine setae and two pairs of sigilla, prominent in females, barely distinguishable on scutum in males (Figs 1–12); abdominal markings variable, predominantly composed of cream markings forming a X on dark grey or black background (Figs 1–12), sometimes reduced in O. calceata and O. marusiki sp. nov. (Figs 70–81); venter without post-epigastric sclerites, with large ventral sclerite in males that is wider than long, absent in females; small, weakly sclerotised inframamillary sclerite present in females, indistinct in males. Spinnerets (details observed in O. radiata only): female (Figs 40, 42–44): ALS of female with one major ampullate gland spigot and nine piriform gland spigots; PMS of female with one small minor ampullate gland spigot anteriorly and five large cylindrical gland spigots; PLS of female with two large cylindrical gland spigots anteriorly and four aciniform gland spigots posteriorly; male spinnerets (Fig. 41) similarly short and stout, but spigots retracted and not examined in detail. Female epigyne with small paired CO positioned anterolaterally in epigyne (Fig. 45), usually in sclerotised anterior ridges, with CD directed posteriorly towards midline (e.g. Figs 54, 59, 65); CD passing round, sclerotised bases of BU centrally; BU large, membranous, oval or teardrop-shaped, directed anterolaterally, each with small an teromesal accessory gland; short ducts connecting base of BU to transversely oval posterior ST I; fertilization ducts originating on posteromedian surface of ST I (e.g. Figs 55, 60). Male palpal femur with small, finger-like RFA, with single (e.g. Fig. 63) or bifid (e.g. Figs 46, 47, 58) tip; palpal patella without apophysis, but with retrolateral lyriform organ (Figs 49, 50); palpal tibia with small triangular RTA (Figs 48, 51); tegulum subtriangular, greatly expanded proximally on prolateral side, tapering distally, base nearly twice as broad as cymbium, with sperm duct running down retrolateral side of tegulum in ventral view, with single proximal loop (e.g. Fig. 57); embolus with broad transverse ridge at base, spike-like, with fine coils along its length (Fig. 52); tegulum without any other structures., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on page 358, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677, {"references":["Simon, E. (1897 a) Histoire naturelle des araignees. Tome Second. Premier Fascicule. Librairie Encyclopedique de Roret, Paris, 192 pp.","Deeleman-Reinhold, C. L. (2001) Forest spiders of South East Asia: with a revision of the sac and ground spiders (Araneae: Clubionidae, Corinnidae, Liocranidae, Gnaphosidae, Prodidomidae and Trochanterriidae [sic]). Brill, Leiden, 591 pp. https: // doi. org / 10.1163 / 9789004475588","Haddad, C. R., Jin, C., Platnick, N. I. & Booysen, R. (2021) Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae). Zootaxa, 4942 (1), 41 - 71. https: // doi. org / 10.11646 / zootaxa. 4942.1.2"]}

Orthobula sp. indet. Remarks. Prior to the untimely passing of the third author, he sent the first author a comprehensive list of material that he had examined and identified, mainly originating from MRAC and CAS. We were able to confirm most of the records, as initial morphospecies photo plates sent to him to aid identification corresponded to the geographical limitations seen amongst other material examined by the first author, e.g. O. calceata in West Africa versus O. radiata in southern Africa and O. milloti in western Kenya. However, some specimens from Central Africa fall into a potential convergence zone between these three species (Fig. 102), and we thought it risky to try and attribute them to either of these three species at this point. We include them here as undetermined specimens to record the occurrence of the genus in these four countries, but these specimens should be reexamined when they are available again to clarify their identification and provide new distribution data for the particular species. Material examined. D.R. CONGO: Cataractes: Lukango region, Lubero–Butembo, Kasumbira Forest, 00°01’S, 29°15’E, 2150 m a.s.l., XII.1974 – I.1975, leg. M. Lejeune (moss, litter under ferns), 4♂ 2♀ (MRAC 168018, 168144, 168178, 168193). North Kivu: Butembo, 00°09’N, 29°17’E, 1740 m a.s.l., III.1965, leg. R. Celis, 1♀ (MRAC 128346); Makayoba camp, Semliki River, 00°44’S, 29°43’E, 12.VII.1968, leg. R. Lejeune, 3♀ (MRAC 135741); “Vallée de la Moyenne”, Semliki River, 00°44’S, 29°43’E, 9.VIII.1968, leg. R. Lejeune, 1♀ (MRAC 135578); Visiki, NW Butembo, 00°12’N, 29°15’E, 1100 m a.s.l., VI.1974, leg. M. Lejeune, 2♀ (MRAC 155599); Same locality, Visiki Forest, 22.XII.1971, leg. M. Lejeune, 1♂ (MRAC 140796); Same locality, Visiki Forest, 00°12’S, 29°15’E, 27.XII.1971, leg. R. Lejeune, 1♀ (MRAC 140822). MALAWI: Northern: Nkhata Bay, Mount Nkwadzi, N Chintheche, 11°41’S, 34°15’E, 10.I.1976, leg. R. Jocqué, 1♀ (MRAC 148075); Nyika Plateau, Chitipa, Chisanga Falls, 10°40’S, 33°50’E, 2200 m a.s.l., 9.XII.1981, leg. R. Jocqué (Brachystegia woods), 1♀ (MRAC 156111); Nyika Plateau, Kaperekezi entrance gate, Western Escarpment road, 10°31’S, 33°39’E, 1450 m a.s.l., 8.XII.1981, leg. R. Jocqué (Brachystegia litter), 1♀ (MRAC 156057); Nyika Plateau, Western Escarpment road, 10°18’S, 33°30’E, 1650 m a.s.l., 8.XII.1981, leg. R. Jocqué (under stones in Brachystegia woods), 1♀ (MRAC 155695). RWANDA: Eastern: Akagera National Park, highway 34, 7 km toward highway 32, 01°40’S, 30°35’E, 16.XI–2.XII.1985, leg. R. Jocqué, Nsengimana & Michiels (pitfall), 1♂ (MRAC 165469); Same locality, 1 km N Lake Ihema fishery, 01°55’S, 30°45’E, 16.XI.1985, leg. R. Jocqué, Nsengimana & Michiels (under dead trunk), 1♂ (MRAC 164909); Same locality, Lake Ihema fishery, 01°55’S, 30°45’E, 7.XII.1985, leg. R. Jocqué, Nsengimana & Michiels, 1♀ (MRAC 164803); Same locality, Lake Kivumba, 01°43’S, 30°45’E, 15.XI–4.XII.1985, leg. R. Jocqué, Nsengimana & Michiels (savanna), 1♀ (MRAC 165648); Kibungo, 02°09’S, 30°32’E, 28.XI.1985, leg. R. Jocqué, Nsengimana & Michiels (sifted litter, eucalypt plantation), 1♂ (MRAC 165393). Southern: Butare, 02°36’S, 29°44’E, 6.XI.1985, leg. R. Jocqué, Nsengimana & Michiels (dense grass tufts), 1♀ (MRAC 164935). UGANDA: Eastern: Busaga District, Kamuli, Tabu River marsh, 00°45’N, 33°30’E, II.1967, leg. J. Ruabunesa, 1♀ (MRAC 131564); Busaga District, Luuka, Mt. Kisunyi, 00°40’N, 33°18’E, II.1967, leg. J. Ruabunesa, 1♀ (MRAC 131589). Western: Rubirizi, Queen Elizabeth National Park [00°10’N, 30°05’E], 16.IV.1971, leg. W. Maher (grassland), 1♀ (CNC)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on page 379, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677

Orthobula marusiki sp. nov. Figs 7, 8, 76–81, 83–89 Diagnosis. Females of O. marusiki sp. nov. can be recognized by the transverse anterior ridge, comprising two slightly recurved parts, which are slightly separated from the broad copulatory openings (Figs 83, 85). Males can be easily recognized from continental congeners by the much longer, slightly curved embolus and by the shape of the bifid RFA, with the proximal and distal branches converging and almost touching (Figs 87–89). Etymology. Named for arachnologist Yura Marusik, who collected the holotype, in recognition of his massive contribution to spider taxonomy, including papers on the genus under study here. Female (holotype, Abuko, NCA 2018/33). Measurements: CL 1.16, CW 0.86, AL 1.35, AW 1.15, TL 2.39 (1.65–2.70), PERW 0.38, MOQAW 0.15, MOQPW 0.21, MOQL 0.18. Length of leg segments: I 0.81 + 0.32 + 0.73 + 0.62 + 0.31 = 2.79; II 0.71 + 0.30 + 0.57 + 0.55 + 0.29 = 2.42; III 0.59 + 0.27 + 0.46 + 0.52 + 0.28 = 2.12; IV 0.79 + 0.29 + 0.68 + 0.76 + 0.35 = 2.87. Colour: carapace deep orange-brown, with black mottling around margins and sides; chelicerae yellow-brown; endites and labium yellow-brown, cream distally; sternum orange-brown, darker around margins; palps dark yellow-brown; legs with femora, patellae and tibiae I and II orange-brown, with dense black mottling laterally; distal ends of femora and tibiae with yellow rings; metatarsi and tarsi I and II yellow; legs III and IV deep yellow, femora with broad orange-brown band medially, patellae and tibiae with black stripe on retrolateral side only; abdomen dark grey dorsally, with pair of cream longitudinal patches anterolaterally and pair of transverse cream patches at midpoint; posterior half with seven fine grey chevrons, narrowing posteriorly; sides and venter pale grey; spinnerets creamy-grey. Leg spination: femora and patellae: spineless; tibiae: I plv 7 rlv 6, II plv 6 rlv 5; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with broad CO positioned slightly posterior to broad, transverse anterior epigynal ridge, comprising two slightly recurved ridges merging mesally (Figs 83, 85); CD directed posteriorly, with slightly concave curvature, entering round SBB in posterior half of epigyne; SBB separated by slightly less than half their width; CD approximately twice as long as width of SBB; BU large, ovoid, with lollipop-shaped accessory gland on their anteromesal surface; SBB connected to ST I by short, strongly curving ducts, entering transversely oval ST I near their centre (Figs 84, 86). Male (paratype, Mbigni, ZFMK 8896). Measurements: CL 0.93, CW 0.73, AL 0.88, AW 0.70, TL 1.84 (1.40– 2.00), PERW 0.30, MOQAW 0.13, MOQPW 0.17, MOQL 0.16. Length of leg segments: I 0.67 + 0.24 + 0.60 + 0.48 + 0.29 = 2.28; II 0.56 + 0.22 + 0.44 + 0.43 + 0.27 = 1.92; III 0.49 + 0.21 + 0.37 + 0.40 + 0.27 = 1.74; IV 0.62 + 0.22 + 0.56 + 0.59 + 0.32 = 2.31. Colour: carapace orange, with black mottling on sides; chelicerae pale yellow-brown; endites and labium yellow-brown, cream distally; sternum pale orange, with darker margins; palps yellow-brown; legs creamy-yellow, anterior femora darker, with mottled patches on sides distally; anterior patellae and tibiae with mottling dorsally and laterally; anterior metatarsi with mottled ring distally; femora III with prolateral mottled stripe and distal patch retrolaterally; patellae and tibiae III with retrolateral stripe; femora, patellae and tibiae IV with mottled retrolateral stripes; abdomen dark grey dorsally, with indistinct, slightly lighter patches anterolaterally and at midpoint; venter cream, with faint grey mottling; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 7 rlv 6, II plv 5 rlv 4; metatarsi: I plv 3 rlv 3, II plv 3 rlv 3; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Palpal femur with small bifid RFA, proximal branch sharply hook-shaped, distal branch thumb-like, converging and nearly touching (Figs 87, 89); tibia with elongate, simple triangular apophysis; retrodistal tegular ridge strongly developed, curving around ventrally, ending close to base of embolus (Fig. 88); embolus long, slender, curving slightly in ventral view (Figs 87–89). Type material. ♀ Holotype: GAMBIA: Banjul: Abuko Nature Reserve, 13°23.472’N, 16°39.003’W, 17.X.2008, leg. Y. Marusik (litter, primary gallery forest) (NCA 2018 /33). Paratypes: BURKINA FASO: Cascades: 4 km NE of Dérégoué, 10°46’N, 04°04’W, 10.I–25.II.1983, leg. Schreiber & Paulus (gallery forest at Koba River), 19♂ 2♀ (ZFMK 8935). CAMEROON: Adamaoua: ca. 30 km NE Tignere, Mbigni, 07°33’N, 12°53’E, 27.II–15.III.1984, leg. Kunze & Nagel, 1♂ (ZFMK 8896). COTE D’IVOIRE: Woroba: Touba [08°17’N, 07°41’W], III–IV.1994, leg. A. Russell-Smith (quadrat sample, upland rice), 1♀ (BMNH). NIGERIA: Oyo: Ibadan [07°22’N, 03°54’E], I.I.T.A., VII.1973, A. Russell-Smith (fallow bush, COP R site), 1♀ (BMNH). Other material examined. BURKINO FASO: Cascades: 4 km NE Dérégué, gallery forest at Koba River, 10°47.4’N, 04°04.8’W, 14.I–18.II.1984, leg. H. Schreiber & M. Biegel (BF), 7♂ (ZFMK 8929), 4 imm. 2♂ 1♀ (ZFMK 8930), 6♂ 1♀ (ZMFK 8933), 1♂ (ZFMK 8934), 1♀ (ZFMK 8936); Same data but savanna, 2♂ 1♀ (ZFMK 8932); 10 km SE Tiéfora, gallery forest at Sinlo River, 10°34.4’N, 04°29.7’W, 11.I–19.II.1982, leg. Nagel, Schreiber, Kalmund (BF), 2♂ (ZFMK 8931). CAMEROON: Adamaoua: ca. 20 km S Minim, 06°49’N, 12°52’E, 1200 m a.s.l., XI–XII.1979, leg. Flacke & Nagel, 1 imm. 1♂ (ZFMK 8881); ca. 30 km NE Tignere, 07°34’N, 12°50’E, 1000 m a.s.l., III.1983, leg. Flacke, Horras & Nagel, 1♂ (ZFMK 8907), 2 imm. 2♂ 1♀ (ZFMK 8909); Tchabal Mbabo, SW slope, 07°25’N, 12°49’E, 1.IV.1983, 1500 m a.s.l., leg. R. Bosmans & Van Stalle (gallery forest litter), 3♀ (MRAC 162235); Same locality, 1400 m a.s.l., 11.IV.1983, leg. R. Bosmans & Van Stalle (gallery forest litter), 1♀ (MRAC 162214); Same locality, 1200 m a.s.l., 8.IV.1983, leg. R. Bosmans & Van Stalle (gallery forest litter), 2♂ (MRAC 162249); Same locality, 7.IV.1983, 1300 m a.s.l., leg. R. Bosmans & Van Stalle (dry grassland), 1♂ 4♀ (MRAC 162211); Same locality, 1600 m a.s.l., 10.IV.1983, leg. R. Bosmans & Van Stalle (transition between gallery forest and grassland), 1♂ 2♀ (MRAC 162211). CENTRAL AFRICAN REPUBLIC: Sangha- Mbaéré: Dzanga-Ndoki National Park, 37.9 km 169°S Lidjombo, 02°22’14’’N, 16°10’21’’E, 360 m a.s.l., 20–28.V.2001, leg. B. Fisher (beating low vegetation, rainforest), 3♀ (CAS, CASENT 9068197). COTE D’IVOIRE: Lacs: Kossou, 06°57’N, 04°58’W, 23.V.1975, leg. R. Jocqué (savanna), 1♂ (MRAC 152938). Sassandra-Marahoué: Bouaflé, Klébo, 06°52’N, 06°08’W, I–III.1981, leg. J. Everts, 1♂ 1♀ (MRAC 166428); Same locality, II.1981, leg. J. Everts (pitfalls), 2♂ (MRAC 166385); Bouaflé, Tuyankro, 06°59’N, 05°39’W, 5.II.1981, leg. J. Everts (pitfalls), 1♂ 1♀ (MRAC 159532). Savanes: Badenou / M’Bengué, 09°50’N, 05°50’W, 7.XI–18.XII.1982, leg. Dister, Huschens, Sumkeller (H 2 O), 2 imm. 4♂ 3♀ (ZFMK 8958), 1♂ 1♀ (ZFMK 8965); Same locality, 19.XI–9.XII.1984, leg. Fliedner & Krüger (BF), 4♂ 2♀ (ZFMK 8957); Near Korhogo, 09°27’N, 05°38’W, I.1980, leg. J. Everts (pitfalls, gallery forest), 2♂ 2♀ (MRAC 169841); Same data but I.1980, 1♂ (MRAC 169839); Same data but III.1980, 1♂ (MRAC 169840); Same data but V.1980, 1♂ 2♀ (MRAC 169843); Riv. Badeni / Tawara, 25km N, 09°52’N, 05°39’W, 24.I–11.II.1983, leg. Krüger & Summkeller, 1♂ 3♀ (ZFMK 8950); Riv. Bagoé / Guingéréni, 09°32’N, 06°36’W, 1–16.III.1984, leg. Krüger & Petermann (BF 2 Sav D), 1♂ (ZFMK 8973); Same locality, 1–16.III.1984, leg. Krüger & Petermann (BF 2 Sav D), 1♂ (ZFMK 8973); Same locality, 5–19.III.1984, leg. Krüger & Petermann (BF 5 Gw-D), 8♂ (ZFMK 8976); Séguébé, SE Sirasso, 09°11’N, 06°03’W, X–XII.1983, leg. G. Rosner & R. Summkeller (BF FS), 1 imm. 2♂ 3♀ (ZFMK 8953); Sirasso, 09°16’N, 06°05’W, 7.XI–18.XII.1982, leg. Dister, Huschens & Sumkeller (FS), 3♂ (ZFMK 8952). NIGERIA: Oyo: Ibadan, 07°22’N, 03°54’E, I.I.T.A., 7.III.1974, leg. A. Russell-Smith (riverine woodland), 1♂ (MRAC 160339). TOGO: Centrale: Between Bassari and Sokodé, 01°01’N, 09°01’E, V–VII.1984, leg. P. Dauben, 1♀ (MRAC 169838). Habitat and biology. A common ground-dwelling spider sampled from pitfalls and litter in various forest and savanna habitats. Distribution. Widespread in West Africa and the western Congo Basin (Fig. 101). Variation. Most of the O. marusiki sp. nov. specimens examined lacked comprehensive abdominal markings, with the majority having a uniform grey abdomen (Figs 76, 79). A smaller proportion, including the holotype female (Fig. 7), have variable degrees of cream patches forming a reduced form of the typical X-marking of Orthobula (e.g. Figs 77, 78, 80, 81)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on pages 371-374, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677

Orthobula radiata Simon, 1897 Figs 11–53, 96–100 Orthobula radiata Simon, 1897b: 499 (Holotype ♂: SOUTH AFRICA: Gauteng: Hammanskraal [25°24’S, 28°17’E], deposited in MNHN—examined). Remark. The locality indicated on the label of the holotype is Hammanskraal, as indicated in the bibliography above, but in the publication (Simon 1897b) it is indicated as Pretoria (25°42’S, 28°13’E), with about 50 km separating the localities. Diagnosis. Females of O. radiata have a similar general epigyne structure to O. calceata, but lack the anterior ridges seen in the latter species (Fig. 65), with these being replaced by shallow depressions (Figs 45, 96), and also by the short CD. Males of O. radiata most closely resemble those of O. milloti, particularly in the short, straight spike-like embolus, but can be distinguished by presence of a bifid (Fig. 100) as opposed to a simple finger-like RFA (Fig. 95). Female (Pretoria, NCA 2008/1981). Measurements: CL 1.09, CW 0.83, AL 1.35, AW 1.08, TL 2.55 (1.60– 2.55), PERW 0.36, MOQAW 0.15, MOQPW 0.18, MOQL 0.16. Length of leg segments: I 0.73 + 0.27 + 0.65 + 0.57 + 0.29 = 2.51; II 0.63 + 0.25 + 0.51 + 0.49 + 0.26 = 2.14; III 0.52 + 0.24 + 0.40 + 0.44 + 0.25 = 1.85; IV 0.66 + 0.25 + 0.60 + 0.67 + 0.33 = 2.51. Colour: carapace deep orange-brown, with black mottling around margins, faint on sides; chelicerae orangebrown; endites orange-brown, labium darker brown, both cream distally; sternum orange, margins orange-brown; palps yellow-brown; legs with femora, patellae and tibiae orange-brown, with dense black mottling laterally; distal ends of femora and tibiae, and basal end of patellae, with yellow rings; metatarsi and tarsi yellow, metatarsi with faint black mottling; abdomen black dorsally, with pair of oval cream anterolateral patches and transverse cream band at midpoint, merging centrally to form X; posterior half with six slender transverse chevrons, last four merging centrally; venter creamy-grey; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with lateral CO in shallow depressions (Figs 45, 96); CD curving posteromesally, entering slightly ovoid SBB at centre of epigyne; SBB almost touching along their mesal surfaces; CD similar in length to width of SBB; BU broad, ovoid, with boomerang-shaped accessory gland on their anteromesal surface; SBB connected to ST I by short looping ducts, entering obliquely-orientated oval ST I on their posteromesal surface (Fig. 97). Male (Pretoria, NCA 2008/1980). Measurements: CL 0.86, CW 0.70, AL 0.95, AW 0.73, TL 1.85 (1.48–2.05), PERW 0.30, MOQAW 0.12, MOQPW 0.16, MOQL 0.15. Length of leg segments: I 0.59 + 0.22 + 0.52 + 0.46 + 0.26 = 2.09; II 0.53 + 0.21 + 0.41 + 0.39 + 0.24 = 1.78; III 0.43 + 0.19 + 0.30 + 0.38 + 0.22 = 1.52; IV 0.57 + 0.20 + 0.49 + 0.56 + 0.29 = 2.11. Colour: carapace deep orange-brown, mottled around margins, with black mottled marking centrally; chelicerae orange-brown; endites and labium orange-brown, cream distally; sternum orange, margins orange-brown; palps orange-brown, with faint black mottling; legs with femora, patellae and tibiae I and II orange-brown, metatarsi and tarsi yellow-brown, patellae and tibiae with black mottling dorsally and laterally; femora, patellae and tibiae III and IV yellow-brown, metatarsi and tarsi yellow, patellae and tibiae with black retrolateral stripe; abdomen black dorsally, with pair of oblique cream anterolateral markings, converging and merging centrally with transverse cream band; venter and spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 4 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Palpal femur with small bifid RFA, proximal branch sharply hook-shaped, distal branch small, tooth-like (Fig. 100); tibia with small, simple triangular apophysis; retrodistal tegular ridge strongly developed, curving around ventrally, ending close to base of embolus (Fig. 99) embolus narrow, straight, spike-like, with slightly constricted tip (Figs 98–100). Other material examined. BOTSWANA: Ngamiland: Maxwee, 19°30’S, 23°40’E, 28.VIII.1975, leg. A. Russell-Smith (grassland), 16♂ 7♀ (MRAC 154603); Near Maun, Manxunyane Lagoon [19°54’S, 23°28’E], 1.IV.1976, leg. F. Wanless & A. Russell-Smith (grassland, ground layer), 1♀ (BMNH); Okavango, Maun, Island Safari Lodge [19°55’S, 23°30’E], 21.IX.1975, leg. A. Russell-Smith (riverine forest), 1♀ (BMNH); Okavango, Maun, Thamalakane River [19°59’S, 23°30’E], 11.IX.1976, leg. A. Russell-Smith (Setaria grassland), 1♂ 4♀ (BMNH); Okavango, Maxwee [19°30’S, 23°40’E], V–VI.1976, leg. A. Russell-Smith (Mopane woodland grass litter), 1♀ (BMNH); Okavango, Moremi Game Reserve, Mboma Lagoon [19°11’S, 23°16’E], 19.XI.1977, leg. A. Russell- Smith (moist Hyparrhenia grassland), 1♀ (BMNH); Okavango, Moremi Game Reserve, Mboma Island, P. Smith’s Camp [19°11’S, 23°16’E], 20.VI.1977, leg. A. Russell-Smith (wet Setaria grassland), 2♀ (BMNH); Okavango, Pom Pom, 19°35’S, 22°50’E, 15.VIII.2001, leg. M. Dangerfield (riparian pitfall), 1♂ 1♀ (NCA 2001 /418). MOZAMBIQUE: Gaza: Bilene, Praia do Bilene, 27 m a.s.l., 25°15.649’S, 33°17.659’E, 20.XII.2007, leg. C. Haddad, R. Lyle & R. Fourie (leaf litter, coastal forest), 1♂ 2♀ (NCA 2008 /213). Inhambane: Morrungulo, Morrungulo Resort, 12 m a.s.l., 23°13.983’S, 35°29.587’E, 6.XII.2007, leg. C. Haddad, R. Lyle & R. Fourie (leaf litter, dune forest), 4♂ (NCA 2008 /187). Maputo: Near Marrucuene, Blue Anchor Inn, 50 m a.s.l., 25°35.124’S, 32°39.568’E, 28.XI.2007, leg. C. Haddad & R. Fourie (sifting leaf litter, savanna), 1♂ 2♀ (NCA 2008 /166). NAMIBIA: Kavango East: Popa Falls, 18°07’S, 21°35’E, 24.X.1987, leg. R. Jocqué (densely vegetated banks of rapids), 1♀ (MRAC 168515). Kunene: Nimmerrus, 19°28’S, 15°48’E, 17.IV.1984, leg. C. Schultz, 1♀ (NCA 85 /53). SOUTH AFRICA: Gauteng: Johannesburg, Roodepoort, Poortview, 26°05.240’S, 27°51.449’E, 9.VII.2005, leg. A. Leroy (on kitchen shelf), 1♂ (NCA 2011 /871); Pretoria National Botanical Garden, 29°44’S, 28°16’E, 6.X–24.XI.2007, leg. E. Kassimatis (pitfall traps), 2♂ 3♀ (NCA 2008 /1979); Same data but 8–22.IX.2007, 1♂ (NCA 2008 /1980); Same data but 23.IX–5.X.2007, 1♀ (NCA 2008 /1981); Pretoria, Rietondale Research Station, 25°43’S, 28°14’E, 16.V.1998, leg. J. Nkwana (pitfall traps), 1♂ (NCA 2007 /1151); Roodeplaat Dam Nature Reserve, 25°38’S, 28°21’E, 28.IV.1983, leg. A. van den Berg (sweepnet, long grass), 1♂ (NCA 84 /190); Same locality, 10.V.2000, leg. A. Leroy (in house), 1♀ (NCA 2007 /3834). KwaZulu-Natal: 15km N of Richard’s Bay, 28°46.842’S, 32°02.298’E, 27.XI.1995, leg. T. Wassenaar (sweeps, rehabilitated coastal forest), 1♂ (NCA 96 /504); Drakensberg Mountains, Monk’s Cowl, 29°02.790’S, 29°24.235’E, 1465 m a.s.l., 3.XI.2012, leg. J. Neethling (sifting leaf litter, Afromontane forest), 1♂ (NCA 2019 /1000); Durban, Kloof, Kranzkloof Nature Reserve, 29°46.182’S, 30°49.744’E, 440 m a.s.l., 24.VII.2002, leg. C. Haddad (leaf litter on hillside), 1♂ 1♀ (NCA 2021 /1244); eThekwini [Durban], Buffelsdraai township, 29°38’S, 30°58’E, 5.II.2019, leg. S.P. Mntambo (pitfalls, Chromolaena odorata project), 1♂ (NCA 2019 /956); iSimangaliso Wetlands Park, Eastern Shores Nature Reserve, 28°21.341’S, 32°25.062’E, 20 m a.s.l., 30.XI.2015, leg. C. Haddad (hand collecting), 1♂ (NCA 2015 /2557); Same locality, Gwala-Gwala Forest, 28°23.042’S, 32°24.436’E, 21.IV.2006, leg. C. Haddad (leaf litter, coastal forest), 1♂ (NCA 2006 /724); Same locality, Mission Rocks picnic site, 28°15.811’S, 32°28.953’E, 90 m a.s.l., 29.XI.2015, leg. R. Booysen, V. & N. Butler (sifting leaf litter, coastal forest), 1♂ 1♀ (NCA 2015 /2281); Same locality, near Mission Rocks, 28°14.118’S, 32°29.332’E, 14.V.2012, leg. J. Neethling & C. Luwes (leaf litter, wetland bush patch), 1♀ (NCA 2012 /3954); Ithala Game Reserve, Ntshodwe Camp, 27°32.699’S, 31°16.911’E, 27.I.2014, leg. C. Haddad & Z. Mbo (sifting leaf litter, short forest), 2♀ (NCA 2013 /4937); Same data but 29.I.2014, leg. C. Haddad, 4♀ (NCA 2013 /5004); Same locality, 29.I.2014, leg. C. Haddad (base of grass tussocks), 1♂ 1♀ (NCA 2013 /5015); Kosi Bay Nature Reserve, 26°57.767’S, 32°48.981’E, 15.IV.2006, leg. C. Haddad (leaf litter, coastal forest), 2♀ (NCA 2006 /758); Lake Sibaya, east side, 27°22’S, 32°43’E, 25.VI.1967, leg. B.H. Lamoral (forest floor litter), 2♀ (NMSA 22040); Luneburg, road to Paulpietersburg, 27°20.322’S, 30°29.563’E, 23.V.2012, leg. J. Neethling & C. Luwes (leaf litter, bush patch), 2♀ (NCA 2012 /3953); Mtunzini, Twin Streams farm (I.F. Garland), 28°57’S, 31°46’E, 19–20.I.1984, leg. T. & C. Griswold, P. Croeser & P. Reavell (sifting leaf litter), 1♂ 1♀ (NMSA 18876); Ndumo Game Reserve, Broadleaf woodland, near Croc Farm, 26°54.431’S, 32°19.045’E, 25.VI.2006, leg. C. Haddad & F. Jordaan (under rocks), 1♀ (NCA 2006 /1263); Same locality, Pongola River floodplain, 26°53.384’S, 32°19.097’E, 16.I.2006, leg. C. Haddad (riverine forest leaf litter), 1♀ (NCA 2006 /790); Same locality, Pongola River floodplain, 26°53.362’S, 32°18.892’E, 6.II.2005, leg. C. Haddad (leaf litter, floodplain), 2♂ 1♀ (NCA 2005 /41); Same locality, Shokwe Pan, 26°52.340’S, 32°12.745’E, 45 m a.s.l., 2.XII.2018, leg. J. Neethling (leaf litter, Ficus sycomorus forest), 1♀ (NCA 2019 /196); Same locality, Start of game count transect 8, 26°50.182’S, 32°13.138’E, 11.VII.2004, leg. C. Haddad (leaf litter, floodplain reeds), 1♂ 2♀ (MHBU); Newcastle district, Moorfield farm, 27°52.552’S, 29°42.495’E, 1780 m a.s.l., 29.VI.2015, leg. R. Booysen (leaf litter, lowland grassland), 1♀ (NCA 2017 /647); Newcastle district, Normandien Farms, 27°55.363’S, 29°49.767’E, 1300 m a.s.l., 12–16.VII.2015, leg. R. Booysen & R. Vlok (pitfalls, Eucalyptus plantation), 1♀ (NCA 2017 /692); Same locality, 13.VII.2015, leg. R. Booysen & A. Booysen (leaf litter, Eucalyptus plantation), 1♂ (NCA 2017 /708); Same locality, 23.I.2016, leg. R. Booysen et al. (day active search, Eucalyptus plantation), 3♀ (NCA 2017 /851); Ngome State Forest, 27°49’S, 31°26’E, I.1993, leg. M. van der Merwe (pitfall traps, grassland), 2♂ (NCA 94 /439); Ophathe Game Reserve, Rocky mountainside, 28°23.202’S, 31°24.077’E, 505 m a.s.l., 1.X.2008, leg. C. Haddad (sifting leaf litter), 1 imm. 2♂ 3♀ (NCA 2008 /4040); Same locality, Ombesanoni River Bed, 28°23.727’S, 31°23.643’E, 455 m a.s.l., 2.X.2008, leg. C. Haddad (sifting leaf litter), 4♀ (NCA 2008/4203); Pietermaritzburg [29°35’S, 30°25’E], IX.1951, leg. R.F. Lawrence, 9♀ (NMSA 5675); Same locality, Prestbury [29°36’S, 30°20’E], 11.V.1988, leg. W. Starega (in house at night), 1♂ (NMSA 18880); Same locality, Scottsville [29°37’S, 30°23’E], VII.1951, leg. R.F. Lawrence, 1♂ 4♀ (NMSA 5642); Pietermaritzburg, Clarendon Nature Reserve [29°36’S, 30°20’E], 15.IV.1976, leg. A. Russell-Smith (in garden), 1♂ 2♀ (BMNH); St Lucia Game Reserve, Makakatana Bay, 28°14.217’S, 32°24.633’E, 88 m a.s.l., 1–2.IV.2001, leg. M.J. Ramírez, 1♀ (MACN-Ar 42434); Tembe Elephant Park, Open woodland/sand, near offices, 27°03’S, 32°25’E, 12.I.2002, leg. C. Haddad (sweeps, grass and herbs), 1♀ (NCA 2007 /3578); Same locality, Open woodland/sand, near offices, 27°03’S, 32°25’E, 17.I.2002, leg. C. Haddad (active searching, under logs), 1♂ (NCA 2007 /3179); Same locality, 15.III.2003, leg. A. Honiball (sweeps, grass and herbs), 1 imm. 4♂ 6♀ (NCA 2004 /264); Same locality, 27°02.547’S, 32°25.407’E, 115 m a.s.l., 9–12.IV.2001, leg. M.J. Ramírez, 1♀ (MACN-Ar 39884), 1♀ (MACN-Ar 39894); Umgeni Valley Nature Reserve, near Howick, 29°28’S, 30°14’E, 11.V.2003, leg. C. Haddad (leaf litter, tree base), 2♂ 3♀ (NCA 2008 /577); Vernon Crookes Nature Reserve, near Umzinto, 30°16.085’S, 30°36.718’E, 458 m a.s.l., 26.XI.2012, leg. J. Neethling (leaf litter, Afromontane forest), 1♂ (NCA 2021 /1243); Same locality, 30°16.250’S, 30°36.402’E, 485 m a.s.l., 9–12.X.2020, leg. R. Booysen & R. Steenkamp (hand collecting), 1♂ (NCA 2021 /1202). Limpopo: Blouberg Nature Reserve, 22°59.255’S, 29°07.950’E, 864 m a.s.l., 30.XI.2005, leg. E. Stam (leaf litter, Spirostachys africana), 1♀ (NCA 2008 /2767); Same locality, 23°00.110’S, 29°03.855’E, 903 m a.s.l., 25.III.2006, leg. F. Maanda (leaf litter, Philenoptera violaceae), 1♀ (NCA 2008 /2770); Lajuma Mountain Retreat, 23°02.414’S, 29°26.687’E, 6.II.2008, leg. C. Haddad (base of grass tussocks), 1♀ (NCA 2008 /507); Same locality, 3.V.1997, leg. M. van der Merwe (pitfall traps), 1♂ (NCA 98 /25); Same data but 29.VI.1997, 1♂ (NCA 98 /26); Same locality, woodland island 1a, 23°01.906’S, 29°26.141’E, 17.XI.2004, leg. M. Mafadza (pitfalls), 1♀ (NCA 2005 /2008); Same locality, woodland island 2, 23°01.921’S, 29°26.193’E, 23.XI.2004, leg. M. Mafadza (active searching), 1♂ 1♀ (NCA 2005 /2005); Same locality, woodland island 4, 23°01.894’S, 29°26.123’E, 28.XI.2004, leg. M. Mafadza (active searching), 1♀ (NCA 2005 /2006); Same locality, woodland island 4, 23°01.894’S, 29°26.123’E, 28.XI.2004, leg. M. Mafadza (sifting leaf litter, Ignota, sample 2), 2♂ 4♀ (NCA 2005 /2007); Little Leigh, 22°56.057’S, 29°53.610’E, 1086 m a.s.l., 23.XI.2005, leg. B. van der Waal (leaf litter, Burkea africana), 1♂ (NCA 2008 /2768); Same locality, 22°56.175’S, 29°53.155’E, 1086 m a.s.l., 22.III.2006, leg. S. Foord (leaf litter, Pterocarpus rotundifolius), 1♂ 1♀ (NCA 2008 /2769); Same locality, 22°56.175’S, 29°53.155’E, 1086 m a.s.l., 25.XI.2005, leg. E. Stam (leaf litter, P. rotundifolius), 2♀ (NCA 2008 /2771); Same locality, 22°56.175’S, 29°53.155’E, 1086 m a.s.l., 25.XI.2005, leg. S. Foord (leaf litter, P. rotundifolius), 2♀ (NCA 2008 /2772); Venetia Mines, V4, 22°26’S, 29°18’E, 25.III.2008, leg. S. Foord (leaf litter sifting), 1♀ (NCA 2008 /2871). Mpumulanga: 11 km SE Pilgrim’s Rest [ca. 24°59’S, 30°48’E], 1400 m a.s.l., 11–31.XII.1985, leg. S. & J. Peck (flight intercept-malaise, relict native forest edge), 4♂ 2♀ (AMNH); Badplaas, 25°57’03’’S, 30°34’00’’E, 26–29.III.2001, 1100 m a.s.l., leg. D. & S. Ubick (at night, in litter under riparian trees), 1♀ (CAS, CAS 9068193); Carolina District, mile 52, road from Barberton to Carolina, 25°55’S, 30°30’E, X.1961, leg. N. Leleup (under rocks), 1♀ (MRAC 131880); Embuleni Nature Reserve, Badplaas, 25°57’12’’S, 30°33’15’’E, 28.III.2001, 1100 m a.s.l., leg. D. & S. Ubick (grassveld savannah, in wooded areas), 1♂ 1♀ (CAS, CASENT 9068194); Same locality, 28.III.2001, leg. M. Ramirez, 1♀ (NCA 2002 /124); Same locality, 28.III.2001, leg. A. Russell-Smith (in litter in woodland), 2 imm. 4♂ 6♀ (BMNH); Same locality, 25°56’S, 30°34’E, 28.III.2001, leg. M.J. Ramírez, 1♂ (MACN-Ar 39881), 1♀ (MACN-Ar 42067); Kruger National Park, Mopani, Tsendze, 23°41.100’S, 31°31.080’E, XI.2008, leg. B. Reynolds (litter sifting), 1♀ (NCA 2010 /2741); Marble Hall, 25°15’S, 29°15’E, 15.V.2002, leg. M. Mellett (pitfall traps, Bt cotton), 1♀ (NCA 2004 /1295). North West: Brits district, Jacksonstuin, 25°44’S, 27°45’E, IX.1960, leg. N. Leleup (litter), 3♀ (MRAC 140372); Venterskroon district, Thabela Thabeng Mountain Retreat, 26°51.825’S, 28°17.819’E, 1.IX–1.X.2009, leg. R. Fourie & A. Grobler (pitfall traps, woodland grassland), 1♀ (NCA 2009 /3619); Same data but 1–29.X.2009, 5♂ 1♀ (NCA 2009 /3550); Same locality, 26°51.828’S, 28°17.805’E, 1–29.X.2009, leg. R. Fourie & A. Grobler (pitfalls, Vaal River bank), 2♀ (NCA 2009 /3560). Northern Cape: Upington, Duine-in-die-Weg Guest Farm, 28°34’S, 21°45’E, 835 m a.s.l., 22.I.2018, leg. H. Badenhorst (hand collecting, sample 14 Ks), 2♂ 1♀ (NCA 2020 /68). ZAMBIA: Southern: Wildlives Game Farm, near Choma, Hunter’s Camp, 16°58.957’S, 26°36.973’E, 3.XII.2006, leg. C. Haddad, J. Parau & F. Jordaan (leaf litter), 4♀ (NCA 2007 /482); Same locality, BFA study plot, 9–14.XII.1994, leg. F. Nyathi (small pitfall traps), 1♀ (NMZ A11815). Muchinga: Mpika, thermal source of Kanyama, 11°50’S, 31°26’E, VII.1960, 1700 m a.s.l., leg. N. Leleup, 1♀ (MRAC 115299). ZIMBABWE: Harare, Highlands, 19 Walmer drive, 17°48’S, 31°05’E, XII.2008, leg. M. Cumming (in garden), 1♀ (NCA 2010 /392); Same data but 15.III.2005, 1♂ (NCA 2010 /4649). Habitat and biology. A common ground-dwelling spider sampled from pitfalls and litter in various biomes in southern Africa, including arid Nama Karoo, grasslands, forests, thickets and savannas. Specimens were regularly sampled near ants’ nests (e.g. Monomorium and Tetramorium), particularly under logs and in leaf litter. Distribution. Widespread in southern Africa (Fig. 101)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on pages 376-379, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677, {"references":["Simon, E. (1897 b) Etudes arachnologiques. 27 e Memoire. XLII. Descriptions d'especes nouvelles de l'ordre des Araneae. Annales de la Societe Entomologique de France, 65, 465 - 510."]}

Orthobula marusiki sp. nov. Figs 7, 8, 76–81, 83–89 Diagnosis. Females of O. marusiki sp. nov. can be recognized by the transverse anterior ridge, comprising two slightly recurved parts, which are slightly separated from the broad copulatory openings (Figs 83, 85). Males can be easily recognized from continental congeners by the much longer, slightly curved embolus and by the shape of the bifid RFA, with the proximal and distal branches converging and almost touching (Figs 87–89). Etymology. Named for arachnologist Yura Marusik, who collected the holotype, in recognition of his massive contribution to spider taxonomy, including papers on the genus under study here. Female (holotype, Abuko, NCA 2018/33). Measurements: CL 1.16, CW 0.86, AL 1.35, AW 1.15, TL 2.39 (1.65–2.70), PERW 0.38, MOQAW 0.15, MOQPW 0.21, MOQL 0.18. Length of leg segments: I 0.81 + 0.32 + 0.73 + 0.62 + 0.31 = 2.79; II 0.71 + 0.30 + 0.57 + 0.55 + 0.29 = 2.42; III 0.59 + 0.27 + 0.46 + 0.52 + 0.28 = 2.12; IV 0.79 + 0.29 + 0.68 + 0.76 + 0.35 = 2.87. Colour: carapace deep orange-brown, with black mottling around margins and sides; chelicerae yellow-brown; endites and labium yellow-brown, cream distally; sternum orange-brown, darker around margins; palps dark yellow-brown; legs with femora, patellae and tibiae I and II orange-brown, with dense black mottling laterally; distal ends of femora and tibiae with yellow rings; metatarsi and tarsi I and II yellow; legs III and IV deep yellow, femora with broad orange-brown band medially, patellae and tibiae with black stripe on retrolateral side only; abdomen dark grey dorsally, with pair of cream longitudinal patches anterolaterally and pair of transverse cream patches at midpoint; posterior half with seven fine grey chevrons, narrowing posteriorly; sides and venter pale grey; spinnerets creamy-grey. Leg spination: femora and patellae: spineless; tibiae: I plv 7 rlv 6, II plv 6 rlv 5; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with broad CO positioned slightly posterior to broad, transverse anterior epigynal ridge, comprising two slightly recurved ridges merging mesally (Figs 83, 85); CD directed posteriorly, with slightly concave curvature, entering round SBB in posterior half of epigyne; SBB separated by slightly less than half their width; CD approximately twice as long as width of SBB; BU large, ovoid, with lollipop-shaped accessory gland on their anteromesal surface; SBB connected to ST I by short, strongly curving ducts, entering transversely oval ST I near their centre (Figs 84, 86). Male (paratype, Mbigni, ZFMK 8896). Measurements: CL 0.93, CW 0.73, AL 0.88, AW 0.70, TL 1.84 (1.40– 2.00), PERW 0.30, MOQAW 0.13, MOQPW 0.17, MOQL 0.16. Length of leg segments: I 0.67 + 0.24 + 0.60 + 0.48 + 0.29 = 2.28; II 0.56 + 0.22 + 0.44 + 0.43 + 0.27 = 1.92; III 0.49 + 0.21 + 0.37 + 0.40 + 0.27 = 1.74; IV 0.62 + 0.22 + 0.56 + 0.59 + 0.32 = 2.31. Colour: carapace orange, with black mottling on sides; chelicerae pale yellow-brown; endites and labium yellow-brown, cream distally; sternum pale orange, with darker margins; palps yellow-brown; legs creamy-yellow, anterior femora darker, with mottled patches on sides distally; anterior patellae and tibiae with mottling dorsally and laterally; anterior metatarsi with mottled ring distally; femora III with prolateral mottled stripe and distal patch retrolaterally; patellae and tibiae III with retrolateral stripe; femora, patellae and tibiae IV with mottled retrolateral stripes; abdomen dark grey dorsally, with indistinct, slightly lighter patches anterolaterally and at midpoint; venter cream, with faint grey mottling; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 7 rlv 6, II plv 5 rlv 4; metatarsi: I plv 3 rlv 3, II plv 3 rlv 3; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Palpal femur with small bifid RFA, proximal branch sharply hook-shaped, distal branch thumb-like, converging and nearly touching (Figs 87, 89); tibia with elongate, simple triangular apophysis; retrodistal tegular ridge strongly developed, curving around ventrally, ending close to base of embolus (Fig. 88); embolus long, slender, curving slightly in ventral view (Figs 87–89). Type material. ♀ Holotype: GAMBIA: Banjul: Abuko Nature Reserve, 13°23.472’N, 16°39.003’W, 17.X.2008, leg. Y. Marusik (litter, primary gallery forest) (NCA 2018 /33). Paratypes: BURKINA FASO: Cascades: 4 km NE of Dérégoué, 10°46’N, 04°04’W, 10.I–25.II.1983, leg. Schreiber & Paulus (gallery forest at Koba River), 19♂ 2♀ (ZFMK 8935). CAMEROON: Adamaoua: ca. 30 km NE Tignere, Mbigni, 07°33’N, 12°53’E, 27.II–15.III.1984, leg. Kunze & Nagel, 1♂ (ZFMK 8896). COTE D’IVOIRE: Woroba: Touba [08°17’N, 07°41’W], III–IV.1994, leg. A. Russell-Smith (quadrat sample, upland rice), 1♀ (BMNH). NIGERIA: Oyo: Ibadan [07°22’N, 03°54’E], I.I.T.A., VII.1973, A. Russell-Smith (fallow bush, COP R site), 1♀ (BMNH). Other material examined. BURKINO FASO: Cascades: 4 km NE Dérégué, gallery forest at Koba River, 10°47.4’N, 04°04.8’W, 14.I–18.II.1984, leg. H. Schreiber & M. Biegel (BF), 7♂ (ZFMK 8929), 4 imm. 2♂ 1♀ (ZFMK 8930), 6♂ 1♀ (ZMFK 8933), 1♂ (ZFMK 8934), 1♀ (ZFMK 8936); Same data but savanna, 2♂ 1♀ (ZFMK 8932); 10 km SE Tiéfora, gallery forest at Sinlo River, 10°34.4’N, 04°29.7’W, 11.I–19.II.1982, leg. Nagel, Schreiber, Kalmund (BF), 2♂ (ZFMK 8931). CAMEROON: Adamaoua: ca. 20 km S Minim, 06°49’N, 12°52’E, 1200 m a.s.l., XI–XII.1979, leg. Flacke & Nagel, 1 imm. 1♂ (ZFMK 8881); ca. 30 km NE Tignere, 07°34’N, 12°50’E, 1000 m a.s.l., III.1983, leg. Flacke, Horras & Nagel, 1♂ (ZFMK 8907), 2 imm. 2♂ 1♀ (ZFMK 8909); Tchabal Mbabo, SW slope, 07°25’N, 12°49’E, 1.IV.1983, 1500 m a.s.l., leg. R. Bosmans & Van Stalle (gallery forest litter), 3♀ (MRAC 162235); Same locality, 1400 m a.s.l., 11.IV.1983, leg. R. Bosmans & Van Stalle (gallery forest litter), 1♀ (MRAC 162214); Same locality, 1200 m a.s.l., 8.IV.1983, leg. R. Bosmans & Van Stalle (gallery forest litter), 2♂ (MRAC 162249); Same locality, 7.IV.1983, 1300 m a.s.l., leg. R. Bosmans & Van Stalle (dry grassland), 1♂ 4♀ (MRAC 162211); Same locality, 1600 m a.s.l., 10.IV.1983, leg. R. Bosmans & Van Stalle (transition between gallery forest and grassland), 1♂ 2♀ (MRAC 162211). CENTRAL AFRICAN REPUBLIC: Sangha- Mbaéré: Dzanga-Ndoki National Park, 37.9 km 169°S Lidjombo, 02°22’14’’N, 16°10’21’’E, 360 m a.s.l., 20–28.V.2001, leg. B. Fisher (beating low vegetation, rainforest), 3♀ (CAS, CASENT 9068197). COTE D’IVOIRE: Lacs: Kossou, 06°57’N, 04°58’W, 23.V.1975, leg. R. Jocqué (savanna), 1♂ (MRAC 152938). Sassandra-Marahoué: Bouaflé, Klébo, 06°52’N, 06°08’W, I–III.1981, leg. J. Everts, 1♂ 1♀ (MRAC 166428); Same locality, II.1981, leg. J. Everts (pitfalls), 2♂ (MRAC 166385); Bouaflé, Tuyankro, 06°59’N, 05°39’W, 5.II.1981, leg. J. Everts (pitfalls), 1♂ 1♀ (MRAC 159532). Savanes: Badenou / M’Bengué, 09°50’N, 05°50’W, 7.XI–18.XII.1982, leg. Dister, Huschens, Sumkeller (H 2 O), 2 imm. 4♂ 3♀ (ZFMK 8958), 1♂ 1♀ (ZFMK 8965); Same locality, 19.XI–9.XII.1984, leg. Fliedner & Krüger (BF), 4♂ 2♀ (ZFMK 8957); Near Korhogo, 09°27’N, 05°38’W, I.1980, leg. J. Everts (pitfalls, gallery forest), 2♂ 2♀ (MRAC 169841); Same data but I.1980, 1♂ (MRAC 169839); Same data but III.1980, 1♂ (MRAC 169840); Same data but V.1980, 1♂ 2♀ (MRAC 169843); Riv. Badeni / Tawara, 25km N, 09°52’N, 05°39’W, 24.I–11.II.1983, leg. Krüger & Summkeller, 1♂ 3♀ (ZFMK 8950); Riv. Bagoé / Guingéréni, 09°32’N, 06°36’W, 1–16.III.1984, leg. Krüger & Petermann (BF 2 Sav D), 1♂ (ZFMK 8973); Same locality, 1–16.III.1984, leg. Krüger & Petermann (BF 2 Sav D), 1♂ (ZFMK 8973); Same locality, 5–19.III.1984, leg. Krüger & Petermann (BF 5 Gw-D), 8♂ (ZFMK 8976); Séguébé, SE Sirasso, 09°11’N, 06°03’W, X–XII.1983, leg. G. Rosner & R. Summkeller (BF FS), 1 imm. 2♂ 3♀ (ZFMK 8953); Sirasso, 09°16’N, 06°05’W, 7.XI–18.XII.1982, leg. Dister, Huschens & Sumkeller (FS), 3♂ (ZFMK 8952). NIGERIA: Oyo: Ibadan, 07°22’N, 03°54’E, I.I.T.A., 7.III.1974, leg. A. Russell-Smith (riverine woodland), 1♂ (MRAC 160339). TOGO: Centrale: Between Bassari and Sokodé, 01°01’N, 09°01’E, V–VII.1984, leg. P. Dauben, 1♀ (MRAC 169838). Habitat and biology. A common ground-dwelling spider sampled from pitfalls and litter in various forest and savanna habitats. Distribution. Widespread in West Africa and the western Congo Basin (Fig. 101). Variation. Most of the O. marusiki sp. nov. specimens examined lacked comprehensive abdominal markings, with the majority having a uniform grey abdomen (Figs 76, 79). A smaller proportion, including the holotype female (Fig. 7), have variable degrees of cream patches forming a reduced form of the typical X-marking of Orthobula (e.g. Figs 77, 78, 80, 81).

Orthobula calceata Simon, 1897 Figs 5, 6, 65–75 Orthobula calceata Simon, 1897b: 499 (Holotype ♀: SIERRA LEONE: leg. Mocquerys, deposited in MNHN—examined). Orthobula calceata Ramírez, 2014: 212, figs 140E, 165E–F, 180A. Diagnosis. Females of O. calceata have a similar general epigyne structure to those of O. radiata, but can be distinguished by the curved anterolateral ridges flanking the CO and the very narrow separation of the ST I and SBB (Fig. 65), with the ridges replaced by shallow depressions, and the ST I and SBB separated by approximately ½ SBB diameter, in O. radiata (Figs 45, 96). Males of O. calceata most closely resemble those of O. aethiopica sp. nov. in their palpal structure, particularly the slight undulation of the embolus in ventral view (Figs 57, 68), but can be distinguished by the shape of the RFA, which is short and hook-shaped in O. calceata (Fig. 69) but longer and bifid in O. aethiopica sp. nov. (Fig. 58). Female (Badenou, ZFMK 8947). Measurements: CL 0.93, CW 0.71,AL 1.09,AW 0.99, TL 2.08 (1.60–2.25), PERW 0.29, MOQAW 0.13, MOQPW 0.17, MOQL 0.16. Length of leg segments: I 0.63 + 0.24 + 0.56 + 0.50 + 0.27 = 2.20; II 0.54 + 0.22 + 0.44 + 0.40 + 0.27 = 1.87; III 0.43 + 0.21 + 0.34 + 0.37 + 0.25 = 1.60; IV 0.60 + 0.22 + 0.53 + 0.59 + 0.29 = 2.23. Colour: carapace bright yellow-orange, with black mottling around margins and sides, black mottled patch centrally; chelicerae yellow-brown; endites and labium yellow-brown, cream distally; sternum yellow-orange, with black mottled margins; palps yellow, tibiae with black mottling prolaterally and dorsally; legs with femora, patellae and tibiae orange-brown, with dense black mottling laterally; distal ends of anterior femora and tibiae, and proximal ends of patellae, with yellow rings; posterior femora, patellae and tibiae yellow, with pro- and retrolateral mottled markings in distal half of femora, only retrolaterally on patellae and tibiae; all metatarsi yellow, tarsi creamy-yellow; abdomen grey dorsally, with pair of longitudinal cream anterolateral patches and paired cream transverse patches at midpoint, merging in square medially; faint cream patch above spinnerets; venter cream, with small mottled grey patch in front of spinnerets; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 4 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 2, II plv 2 rlv 2. Epigyne with lateral CO in curved anterolateral ridges (Fig. 65); CO initially curving posteromesally, then bending sharply and diverging slightly before entering round SBB at centre of epigyne; SBB almost touching along their mesal surfaces, also almost touching anterior surface of ST I; CD approximately 1½ times width of SBB; BU ovoid, wrinkled, with finger-shaped accessory gland on their anteromesal surface; SBB connected to ST I by very short looping ducts, entering transversely oval ST I on their anteromesal surface (Fig. 66). Male (Bagoé River, ZFMK 8962). Measurements: CL 0.80, CW 0.63, AL 0.82, AW 0.61, TL 1.70 (1.30–1.70), PERW 0.28, MOQAW 0.11, MOQPW 0.15, MOQL 0.14. Length of leg segments: I 0.52 + 0.21 + 0.48 + 0.40 + 0.26 = 1.87; II 0.46 + 0.19 + 0.38 + 0.33 + 0.25 = 1.61; III 0.40 + 0.17 + 0.27 + 0.30 + 0.21 = 1.35; IV 0.51 + 0.25 + 0.44 + 0.46 + 0.27 = 1.93. Colour: carapace yellow-orange with dense black mottling around margins and on sides, with distinct patch centrally; chelicerae yellow-orange; endites and labium yellow-orange, cream distally, proximal half of labium with black mottling; sternum yellow-orange, margins darker, with dense black mottling around margins; palps yellowbrown; legs yellow, anterior femora slightly darker; anterior femora with distal mottled patches laterally, patellae and tibiae with black mottling dorsally and laterally; posterior femora with distal mottled patch retrolaterally, extending as stripe along patellae and tibiae; abdomen black dorsally, with pair of cream oval anterolateral markings and transverse cream band at midpoint, latter expanded medially along anterior side; venter cream, epigastric plate with dense mottling laterally; lateral grey colouration extending as ring around spinnerets. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 4 rlv 3; metatarsi: I plv 4 rlv 4, II plv 4 rlv 3; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Palpal femur with small hook-shaped RFA (Figs 67, 69); tibia with small, simple triangular apophysis; retrodistal tegular ridge weakly developed, only curving slightly to ventral surface, not close to base of embolus (Fig. 68); embolus narrow, undulating slightly in ventral view, with weak curve at half its length (Figs 67–69). Other material examined. CAMEROON: Adamaoua: ca. 20 km S Minim, 06°49’N, 12°52’E, 1200 m a.s.l., III.1981, leg. W. Flacke & P. Nagel, 1♀ (ZFMK 10429); Same locality, XI–XII.1980, leg. W. Flacke & P. Nagel, 1♂ (ZFMK 10445); Tchabal Mbabo, SW slope, 07°25’N, 12°49’E, 1600 m a.s.l., 10.IV.1983, leg. R. Bosmans & Van Stalle (sweeping, transition between gallery forest and grassland), 1♂ (MRAC 162251). Northwest: Mount Oku, 06°15’N, 10°26’E, 8.III.1983, 2350 m a.s.l., leg. R. Bosmans & Van Stalle (pitfalls), 1♂ (MRAC 162239); Ndop plateau, near Koutoupi, W slope, Massif de Mbam, 05°54’N, 10°44’E, 31.III.1983, 1100 m a.s.l., leg. R. Bosmans & Van Stalle (forest litter), 1♂ (MRAC 162210). COTE D’IVOIRE: Gôh-Djiboua: Gagnoa [06°08’N, 05°57’W], 26.VI.1996, leg. A. Russell-Smith (pitfall, upland rice), 1♂ 1♀ (BMNH). Lacs: Kossou, 06°57’N, 04°58’W, 24.XII.1974, leg. R. Jocqué (savanna), 1♂ (MRAC 152488). Savanes: Bagoé / Guingéréni, 09°32’N, 06°36’W, 14.II–13.III.1983, leg. Fliedner & Krüger, 3 imm. 4♂ 4♀ (ZFMK 8945); Same locality, 14.II–13.III.1983, leg. Krüger & Summkeller (BF), 1 imm. 11♂ 4♀ (ZFMK 8971); Same locality, 18.II–13.III.1983, leg. Krüger & Summkeller (BF/GW), 5♂ (ZFMK 8972); Same locality, 21.II.1983, Krüger & Sumkeller (wasser bis 15:30), 1♂ (ZFMK 8962); Same locality, III–IV.1984, leg. Krüger & Petermann (BF 1 GwD+P), 12♂ 2♀ (ZFMK 8974); Same locality, 6–19.III.1984, leg. Krüger & Petermann, 1♂ (ZFMK 8939); Same locality, 6–19.III.1984, leg. Krüger & Petermann (BF 6 Gw), 3♂ 2♀ (ZFMK 8948); Near Korhogo, 09°27’N, 05°38’W, I.1980, leg. J. Everts (pitfalls, gallery forest), 1♂ 1♀ (MRAC 169841); Same data but II.1980, 2♂ 1♀ (MRAC 169.839); Same data but III.1980, 1♂ 1♀ (MRAC 169840); Same data but V.1980, 3♀ (MRAC 169843); Riv. Badeni / Tawara, 25 km N, 09°52’N, 05°39’W, 24.I–11.II.1983, leg. Krüger & Summkeller, 2 imm. 1♂ 1♀ (ZFMK 8951); Riv. Badenou /M’bengué, 09°50’N, 05°50’W, 19.XI–9.XII.1984, leg. Krüger & Sumkeller (BF), 1♀ (ZFMK 8987); Same locality, 7.XI–18.XII.1982, leg. Dister, Huschens & Sumkeller (FH), 4 imm. 3♂ 3♀ (ZFMK 8959); Same locality, XI.1983, leg. R. Summkeller (oben), 1♂ 1♀ (ZFMK 8947); Same locality, XII.1983, leg. R. Summkeller (hang quer), 2♂ (ZFMK 8949); Same locality, XI–XII.1983, leg. G. Rosner & R. Summkeller, 1 imm. 1♀ (ZFMK 8997), 2♀ (ZFMK 8961). Sassandra-Marahoué: Bouaflé, Klébo, 06°52’N, 06°08’W, I.1981, leg. J. Everts (pitfall), 1♂ (MRAC 166405); Same data but II.1981, 1♂ 1♀ (MRAC 166385). Vallée du Bandama: Bouaké [07°42’N, 05°02’W], 9.XI.1995, leg. A. Russell-Smith (rice, pitfalls, WARDA), 3♂ 5♀ (BMNH). Woroba: Touba [08°17’N, 07°41’W], 24.VIII.1995, leg. A. Russell-Smith (pitfall, rice), 1♂ (BMNH). D.R. CONGO: Équateur: Mabali, Bikoro District, 00°53’S, 18°07’E, X.1959, leg. N. Leleup, 1♀ (MRAC 114565). EQUATORIAL GUINEA: Kié-Ntem: Micomeseng, 02°08’N, 10°37’E, 13–17.VII.1989, leg. M. Alderweireldt (pitfall, field), 1♀ (MRAC 170228). GHANA: Greater Accra: Achimota [05°37’N, 00°13’W], V.1961, leg. C. Hinckley (on outside and in house), 1♀ (MCZ 16418). NIGERIA: Kwara: Borgu Game Reserve, 10°15’N, 04°10’E, 5–6.VIII.1973, leg. A. Russell-Smith (flood debris on river bank), 1♀ (MRAC 160343). Habitat and biology. A common ground-dwelling spider sampled from pitfalls and litter in various forest and savanna habitats. Distribution. Widespread in West Africa and the western parts of the Congo Basin (Fig. 82); largely sympatric with O. marusiki sp. nov. (Fig. 101) and often collected together in the same samples. Variation. All of the examined O. calceata examined have some abdominal markings in both sexes, with the majority of specimens showing “complete” markings (Figs 72, 75), and only a few individuals of each sex a reduction in their extent (e.g. Figs 70, 71, 73, 74)., Published as part of Haddad, Charles R., Jin, Chi & Platnick, Norman I., 2022, A revision of the spider genus Orthobula Simon, 1897 (Araneae: Trachelidae) in the Afrotropical Region. I. Continental species, pp. 355-382 in Zootaxa 5133 (3) on pages 367-370, DOI: 10.11646/zootaxa.5133.3.3, http://zenodo.org/record/6522677, {"references":["Simon, E. (1897 b) Etudes arachnologiques. 27 e Memoire. XLII. Descriptions d'especes nouvelles de l'ordre des Araneae. Annales de la Societe Entomologique de France, 65, 465 - 510.","Ramirez, M. J. (2014) The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History, 390, 1 - 374. https: // doi. org / 10.1206 / 821.1","Haddad, C. R., Foord, S. H., Fourie, R. & Dippenaar-Schoeman, A. S. (2015) Effects of a fast-burning spring fire on the grounddwelling spider assemblages (Arachnida: Araneae) in a central South African grassland habitat. African Zoology, 50, 281 - 292. https: // doi. org / 10.1080 / 15627020.2015.1088400"]}

Spiders are an ancient and extremely diverse animal order. They show a considerable diversity of genome sizes, karyotypes and sex chromosomes, which makes them promising models to analyse the evolution of these traits. Our study is focused on the evolution of the genome and chromosomes in haplogyne spiders with holokinetic chromosomes. Although holokinetic chromosomes in spiders were discovered a long time ago, information on their distribution and evolution in these arthropods is very limited. Here we show that holokinetic chromosomes are an autapomorphy of the superfamily Dysderoidea. According to our hypothesis, the karyotype of ancestral Dysderoidea comprised three autosome pairs and a single X chromosome. The subsequent evolution has frequently included inverted meiosis of the sex chromosome and an increase of 2n. We demonstrate that caponiids, a sister clade to Dysderoidea, have enormous genomes and high diploid and sex chromosome numbers. This pattern suggests a polyploid event in the ancestors of caponiids. Holokinetic chromosomes could have arisen by subsequent multiple chromosome fusions and a considerable reduction of the genome size. We propose that spider sex chromosomes probably do not pose a major barrier to polyploidy due to specific mechanisms that promote the integration of sex chromosome copies into the genome.

Jäger, Peter, Azevedo, Guilherme H. F., Baehr, Barbara, Bonaldo, Alexandre B., Haddad, Charles R., Harms, Danilo, Hormiga, Gustavo, Labarque, Facundo M., Muster, Christoph, Ramírez, Martín J., Santos, Adalberto J. (2021): Twenty years, eight legs, one concept: describing spider biodiversity in Zootaxa (Arachnida: Araneae). Zootaxa 4979 (1): 131-146, DOI: https://doi.org/10.11646/zootaxa.4979.1.14

Despite progress in genomic analysis of spiders, their chromosome evolution is not satisfactorily understood. Most information on spider chromosomes concerns the most diversified clade, entelegyne araneomorphs. Other clades are far less studied. Our study focused on haplogyne araneomorphs, which are remarkable for their unusual sex chromosome systems and for the co-evolution of sex chromosomes and nucleolus organizer regions (NORs); some haplogynes exhibit holokinetic chromosomes. To trace the karyotype evolution of haplogynes on the family level, we analysed the number and morphology of chromosomes, sex chromosomes, NORs, and meiosis in pholcids, which are among the most diverse haplogyne families. The evolution of spider NORs is largely unknown.Our study is based on an extensive set of species representing all major pholcid clades. Pholcids exhibit a low 2n and predominance of biarmed chromosomes, which are typical haplogyne features. Sex chromosomes and NOR patterns of pholcids are diversified. We revealed six sex chromosome systems in pholcids (X0, XY, XThe evolution of cytogenetic characters was largely derived from character mapping on a recently published molecular phylogeny of the family. Based on an extensive set of species and mapping of their characters, numerous conclusions regarding the karyotype evolution of pholcids and spiders can be drawn. Our results suggest frequent autosome-autosome and autosome-sex chromosome rearrangements during pholcid evolution. Such events have previously been attributed to the reproductive isolation of species. The peculiar X

Synemosyna myrmeciaeformis (Taczanowski, 1871) Figs 3B, E, 4B, E, 5A���B, 8A���D Janus myrmeciaeformis Taczanowski, 1871: 125, pl. 4 fig. 9. Simonella peckhami Mello-Leit��o, 1933: 56. Simonella myrmeciaeformis ��� Peckham et al. 1889: 253, pl. 12 fig. 8. ��� Peckham & Peckham 1892: 81, pl. 7 fig. 4. ��� Simon 1901: 509, figs 605���609. Synemosyna myrmeciformis ��� Galiano 1966: 367, figs 18���20, 38���39. Type deposit Holotype ♂ in PAS; Janus myrmeciaeformis Taczanowski, 1871 (examined). Diagnosis Tibial apophysis of male palp finger-like, with distinct median bend, somewhat laterally flattened; bulb large, obliquely oval, 70% of cymbium length; embolus originating prolaterally, with complete revolution around bulb, tip directed retrodistally; epigyne with transverse atrium with subtriangular excavation anteriorly (Fig. 5A); copulatory openings located posterolaterally in atrium, copulatory ducts directed anteriorly, with S-shaped loop at level of spermathecae, entering globular spermathecae posteriorly (Fig. 5B), Material examined Holotype FRENCH GUIANA ��� ♂; Saint Laurent du Maroni; R. Jelski leg.; PAS. Other material BOLIVIA ��� La Paz Dept ��� 1 ♂, 2 ♀♀; Villa Teresa; 16.201�� S, 67.829�� W; 6 Apr. 2016; R. Perger leg.; IBSI-Ara 0756 ��� 3 ♂♂; same collection data as for preceding; 17 Jan. 2018; R. Perger leg.; IBSI- Ara 1022 ��� 2 ♂♂, 6 ♀♀, 1 imm.; same collection data as for preceding; CBF. ��� Cochabamba Dept ��� 1 ♂; Villa Tunari; 16.9844�� S, 65.4094�� W; 10���11 Jul. 2018; R. Perger leg.; IBSI-Ara 00763 ��� 2 ♂♂; same collection data as for preceding; 6 Dec. 2017; R. Perger leg.; IBSI-Ara 1032 ��� 1 ♂, 6 ♀♀; same collection data as for preceding; CBF. ��� Beni Dept ��� 3 ♂♂, 9 ♀♀; Riberalta; 11.0163�� S, 65.9958�� W; 21���23 Jan. 2018; R. Perger leg.; CBF. Comparisons The shape of the tibial apophysis and the embolus with complete circular revolution are shared with S. americana (Peckham & Peckham, 1885) (Mexico to Venezuela) and S. petrunkevichi (Chapin, 1922) (USA, Cuba). However, both species can be separated by a smaller bulb (65% of cymbium length in S. americana and 50% in S. petrunkevichi). In addition to differences in genitalic characters, S. americana and S. petrunkevichi have the carapace not or only slightly narrowed between the cephalic and thoracic areas. The carapace narrowed between cephalic and thoracic areas, and the globular and small spermathecae of S. myrmeciaeformis, are shared with S. paraenesis Galiano, 1967 (Brazil, French Guiana). Females of the latter can be distinguished from those of S. myrmeciaeformis by the spermathecae located in a protuberance in the epigastric area and the copulatory ducts entering the spermathecae laterally, while entering the spermathecae posteriorly in S. myrmeciaeformis (Fig. 5B). Additionally, the tibial apophysis of the male palp of S. paraensis is bifurcate. Variation Three different color forms were observed, displaying a different geographic pattern: orange (in all three locations) (Fig. 8A), orange forms with black cephalic part (Villa Tunari, Cochabamba Dept, and Riberalta, Beni Dept) (Fig. 8B), or dark brown (Villa Teresa, La Paz Dept) forms (Fig. 8C���D). There was no apparent sex-related difference in body color. However, there may be an ontogenetic change in body color. Orange variants were observed within a generally smaller size range of 3.1���6.7 mm (n = 11), dark brown individuals in a range of 6.3���8.05 mm (n = 10), and orange forms with a black cephalic part in a range of 5.84���8.00 mm (n = 20). The dark brown and orange forms with black cephalic part are described here for the first time. Geographical and ecoregion distribution (Fig. 7) Synemosyna myrmeciaeformis is known from Venezuela (type locality), French Guiana, Brazil (Peckham & Peckham 1892; Galiano 1966) and Bolivia (Cutler 1981a; present study). This species was found in ecoregions of the Amazon biome north of 16�� S (Fig. 7). The distributional data of S. myrmeciaeformis refers to Cordillera La Costa montane forest, Guianan moist forest, and Uatuma- Trombetas moist forest (ecoregion regionalization according to Olson et al. 2011). In the present study, the species was collected in Sub-Andean Southwest Amazon forest (Villa Tunari, Riberalta) and Bolivian Yungas forest (Villa Teresa). Remarks Taczanowski (1871) described S. myrmeciaeformis based on a male. Peckham & Peckham (1892) described the female, although Galiano (1966) mentioned that no female could be found among the material examined by Peckham & Peckham (1892). Cutler & M��ller (1991) stated that the female of S. myrmeciaeformis was still considered to be unknown. The females collected in the current study are consistent with the description of Peckham & Peckham (1892), which we here consider to be valid. The female is illustrated here for the first time (Figs 4B, E, 5A���B, 8 A-C). With a maximum BL of 8.05 mm, S. myrmeciaeformis is the largest Bolivian species of Synemosyna collected in this study (maximum BL of S. aurantiaca 7.0 mm and of S. nicaraguaensis 5.85 mm)., Published as part of Perger, Robert, Rubio, Gonzalo D. & Haddad, Charles R., 2021, On ant-like Synemosyna Hentz, 1846 spiders from Bolivia, with indirect evidence for polymorphic mimicry complexes (Araneae: Salticidae: Simonellini), pp. 67-88 in European Journal of Taxonomy 748 (1) on pages 74-76, DOI: 10.5852/ejt.2021.748.1343, http://zenodo.org/record/4745113, {"references":["Taczanowski L. 1871. Les araneides de la Guyane francaise. Horae Societatis entomologicae Rossicae 8: 32 - 132.","Mello-Leitao C. F. de. 1933. Ensaio sobre as myrmarachninas do Brasil. Boletim do Museu Nacional de Rio de Janeiro 9: 39 - 102.","Peckham G. W., Peckham E. G. & Wheeler W. H. 1889. Spiders of the subfamily Lyssomanae. Transactions of the Wisconsin Academy of Sciences, Arts and Letters 7: 222 - 256.","Peckham G. W. & Peckham E. G. 1892. Ant-like spiders of the family Attidae. Occasional Papers of the Natural History Society of Wisconsin 2 (1): 1 - 84. Available from https: // www. biodiversitylibrary. org / part / 82774 [accessed 5 Mar. 2021].","Simon E. 1901. Histoire naturelle des Araignees. Deuxieme edition, tome second. Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 51973","Galiano M. E. 1966. Salticidae (Araneae) formiciformes V. Revision del genero Synemosyna Hentz, 1846. Museo Argentino de Ciencias Naturales Bernardino Rivadavia 1: 339 - 380.","Mello-Leitao C. F. de 1917. Aranhas novas ou pouco conhecidas de Thomisidas e Salticidas brasileiras. Archivos da Escola Superior de Agricultura e Medicina Veterinaria, Rio de Janeiro 1: 117 - 153.","Cutler B. 1993. A new Middle American Synemosyna (Araneae: Salticidae). Revista nicaraguense de entomologia 24: 1 - 4.","Peckham G. W. & Peckham E. G. 1885. On some new genera and species of the Attidae, from Madagascar and Central America. Proceedings of the Natural History Society of Wisconsin 1885: 23 - 42.","Galiano M. E. 1967. Salticidae (Araneae) formiciformes. VIII. Nuevas descripciones. Physis. Revista de la Sociedad Argentina de Ciencias Naturales 27: 27 - 39.","Cutler B. 1981 a. On a collection of antlike jumping spiders from Bolivia (Araneae: Salticidae). Studies on Neotropical Fauna and Environment 16: 51 - 55. https: // doi. org / 10.1080 / 01650528109360581","Olson D. M., Dinerstein E., Wikramanayake E. D., Burgess N. D., Powell G. V. N., Underwood E. C., D'Amico J. A., Itoua I., Strand H. E., Morrison J. C., Loucks C. J., Allnutt T. F., Ricketts T. H., Kura Y., Lamoreux J. F., Wettengel W. W., Hedao P. & Kassem K. R. 2011. Terrestrial ecoregions of the world: a new map of life on Earth. BioScience 51 (11): 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2","Cutler B. & Muller H. G. 1991. The spider genus Synemosyna in northern Colombia (Araneae: Salticidae). Studies on Neotropical Fauna and Environment 26: 171 - 177. https: // doi. org / 10.1080 / 01650529109360850"]}

Synemosyna aurantiaca (Mello-Leit��o, 1917) Figs 3A, D, 4A, D, 6A���D Simonella aurantiaca Mello-Leit��o, 1917: 138. Simonella mastigostyla Mello-Leit��o, 1917: 140, fig. 2. Simonella claustrorum Mello-Leit��o, 1933: 62, pl. 1 fig. 6. Simonella ypsilon Piza, 1937: 311, pl. 1 figs 3���4, pl. 2 fig. 5. Simonella aurantiaca ��� Mello-Leit��o 1933: 58, pl. 1 fig. 2. Simonella mastigostyla ��� Mello-Leit��o 1933: 59, pl. 1 fig. 4, pl. 2 fig. 8. Synemosyna aurantiaca ��� Galiano 1966: 348, figs 8���11, 44���46, 49, 61. ��� Cutler & M��ller 1991: 174, figs 13���15. Type deposit The types of S. aurantiaca and its synonyms were destroyed in a recent fire (World Spider Catalog 2020; A. Kury, unpublished). Diagnosis Tibial apophysis of male palp triangular, shark fin-like and laterally flattened; bulb narrowing distally, 1.2 times longer than wide; embolus originating prolaterally, curling around retrodistal end of cymbium, bending back sharply along dorsal cymbial surface towards base, before bending back near retrolateral edge to tip; anterior margin of epigynal atrium slightly procurved; copulatory openings located posteromedially, copulatory ducts loop anteriorly back to level of spermathecae, entering kidney-shaped spermathecae anterolaterally. Material examined BOLIVIA ��� Santa Cruz Dept ��� 2 ♂♂; Bermejo; 18.136�� S, 63.631�� W; 13 Jan. 2016; R. Perger leg.; IBSI-Ara 0735 ��� 4 ♂♂, 8 ♀♀, 1 imm.; Cotoca; 17.7736�� S, 63.065�� W; 11 Jul. 2018; R. Perger leg.; CBF ��� 2 ♂♂, 3 ♀♀; Santa Rosa de la Mina; 16.5391�� S, 62.4622�� W; 9���13 Sep. 2016; R. Perger leg.; CBF. ��� Chuquisaca Dept ��� 1 ♂, 2 ♀♀; Teja Huasi; 18.9475�� S, 65.1369�� W; 18 Dec. 2017; R. Perger leg.; CBF. ��� Tarija Dept ��� 1 ♂, 4 ♀♀; Arambulo; 21.809�� S, 64.232�� W; 9 Jan. 2016; R. Perger leg.; IBSI-Ara 0773 ��� 4 ♂♂, 5 ♀♀; same collection data as for preceding; CBF. ARGENTINA ��� Misiones Province ��� 1 ♀; General Manuel Belgrano Dept, San Antonio; 26.019�� S, 53.791�� W; 11 Nov. 2011; G.D. Rubio leg.; IBSI-Ara 0182 ��� 1 ♀; Karadya Bio-Reserve; 25.859�� S, 53.961�� W; 7 Apr. 2016; J.E. Baigorria leg.; IBSI-Ara 0612 ��� 1 ♂; same collection data as for preceding; 15 Sep. 2016; IBSI-Ara 0783 ��� 1 ♂; Bernardo de Irigoyen; 26.247�� S, 53.639�� W; 31 Oct. 2016; G.D. Rubio leg.; IBSI-Ara 0867. Comparisons The triangular, shark fin-like tibial apophysis and the embolus without complete revolution are also found in S. decepiens (O. Pickard-Cambridge, 1896) (Mexico, Guatemala) (Cutler 1985). However, in the latter, the bulb broadens distally and is 1.5 times longer than wide. In S. invemar Cutler & M��ller, 1991 (Colombia), the spermathecae are kidney-shaped but the tubes do not loop back to the level of the spermathecae and enter the spermathecae posteriorly (Cutler & M��ller 1991). Variation The following forms were collected in sympatry in all locations: orange-reddish, light brown, and dark brown to blackish forms (Fig. 6A���D). All orange-reddish forms were smaller than 5.55 mm (n = 9) and all dark-brown to blackish variants longer than 5.8 mm (n = 8), indicating an ontogenetic change in body color with light brown transitional forms. There was no apparent sex-related difference in body color. Geographical and ecoregion distribution (Fig. 7) This species has been recorded from Brazil (Galiano 1966; Raizer 2004; Podgaiski et al. 2007; Rodrigues et al. 2009, 2016), Uruguay (Laborda Turri��n 2016), Argentina (Galiano 1966; Zapata & Grismado 2015) and Bolivia (present study). Cutler & Edwards (2002) recorded S. aurantiaca from Trinidad Island (Lesser Antilles). The taxonomic status of these specimens remains to be determined. Synemosyna aurantiaca is distributed in mostly semi-deciduous forests south of 18�� (Fig. 7). According to the biogeographic regionalization of Olson et al. (2011), previous records refer to the following ecoregions: Pantanal, Humid Pampas, Parana flooded Savanna, Uruguayan Savanna, Alto Paran�� Atlantic forest, Serra do Mar coastal forests and Araucaria moist forest. In the present study, S. aurantiaca was sampled in Inter-Andean Bolivian Tucuman dry forest (Teja Huasi), Bolivian Tucuman forests (Arambulo), and subhumid, semi-deciduous Chiquitano forest (Bermejo, Cotoca, Santa Rosa de la Mina, Santiago de Chiquitos). Remarks Galiano (1966) collected co-occurring yellow/orange and dark forms in Paran�� de las Palmas and Canal 6, Buenos Aires province, Argentina. Oliveira (1988) reported similar forms from Brazil. Both authors mentioned that the species lacks sexual dichromatism, which is supported by the observations in the present study. Previous records of this species referred to a comparably narrow range in southeast Brazil and northeast Argentina. Synemosyna aurantiaca is reported here for the first time from Bolivia. Teja Huasi (Chuquisaca Dept), located at an elevation of 2000 m a.s.l., is the highest reported elevation for any species of the tribe Simonellini. The record from Bermejo (Santa Cruz Dept) is the northern- and westernmost record of this species, extending the distributional range more than 900 km to the northwest of the previously reported northernmost location in Mato Grosso do Sul (Fazenda S��o Bento, Brazil). The record from Colombia (Cutler & M��ller 1991) likely refers to another species, as the illustrated female does not only have a different epigyne (as already stated by Cutler & M��ller 1991), but also pear-shaped spermathecae (kidney-shaped in S. aurantiaca); a bipartite dorsal scutum on the anterior half of the abdomen (one entire scutum in S. aurantiaca); a narrower carapace, resulting in a ratio length/width of 2.7 (~ 2.1 in S. aurantiaca), with concave lateral borders (convex in S. aurantiaca), and the middle part as narrow as the distance between the last posterior eyes (broader in S. aurantiaca)., Published as part of Perger, Robert, Rubio, Gonzalo D. & Haddad, Charles R., 2021, On ant-like Synemosyna Hentz, 1846 spiders from Bolivia, with indirect evidence for polymorphic mimicry complexes (Araneae: Salticidae: Simonellini), pp. 67-88 in European Journal of Taxonomy 748 (1) on pages 72-74, DOI: 10.5852/ejt.2021.748.1343, http://zenodo.org/record/4745113, {"references":["Mello-Leitao C. F. de 1917. Aranhas novas ou pouco conhecidas de Thomisidas e Salticidas brasileiras. Archivos da Escola Superior de Agricultura e Medicina Veterinaria, Rio de Janeiro 1: 117 - 153.","Mello-Leitao C. F. de. 1933. Ensaio sobre as myrmarachninas do Brasil. Boletim do Museu Nacional de Rio de Janeiro 9: 39 - 102.","Piza Jr. S. de T. 1937. Novas especies de aranhas myrmecomorphas do Brasil e consideracoes sobre o seu mimetismo. Revista do Museu Paulista 23: 307 - 319.","Galiano M. E. 1966. Salticidae (Araneae) formiciformes V. Revision del genero Synemosyna Hentz, 1846. Museo Argentino de Ciencias Naturales Bernardino Rivadavia 1: 339 - 380.","Cutler B. & Muller H. G. 1991. The spider genus Synemosyna in northern Colombia (Araneae: Salticidae). Studies on Neotropical Fauna and Environment 26: 171 - 177. https: // doi. org / 10.1080 / 01650529109360850","World Spider Catalog. 2020. World Spider Catalog. Version 19.5. Natural History Museum Bern. Available from http: // wsc. nmbe. ch [accessed 10 Sep. 2020].","Cutler B. 1985. Taxonomic notes on Neotropical species in the genus Synemosyna (Araneae: Salticidae). Studies on Neotropical Fauna and Environment 20: 83 - 91. https: // doi. org / 10.1080 / 01650528509360674","Raizer J. 2004. Comunidade de aranhas em capoes de mata das sub-regioes Miranda e Abobral no pantanal sul-mato-grossense. PhD Thesis, Universidade Estadual de Campinas.","Podgaiski L. R., Ott R., Lopes Rodrigues E. N., Buckup E. H. & Leao Marques M. A. 2007. Araneofauna (Arachnida; Araneae) do Parque Estadual do Turvo, Rio Grande do Sul, Brasil. Biota Neotropica 7 (2): 1 - 15. https: // doi. org / 10.1590 / S 1676 - 06032007000200023","Rodrigues E. N. L., Mendonca J. M. S. & Ott J. 2009. Spider diversity in a rice agroecosystem and adjacent areas in southern Brazil. Revista Colombiana de Entomologia 35 (1): 89 - 97.","Rodrigues E. N. L., Rodrigues P. E. S., Mendonca J. M. S. & Milton S. 2016. Spider species composition in the tree-shrub strata of riparian forests and its microhabitats in southern Brazil. Zoologia (Curitiba) 33 (3): e 20150102. https: // doi. org / 10.1590 / S 1984 - 4689 zool- 20150102","Laborda Turrion A. 2016. Comunidad de aranas islenas en el rio Uruguay: la diversidad en un corredor biologico. Tesis de maestria, Universidad de la Republica (Uruguay), Facultad de Ciencias.","Zapata L. V. & Grismado C. J. 2015. Lista sistematica de aranas (Arachnida: Araneae) de la Reserva Ecologica Costanera Sur (Ciudad Autonoma de Buenos Aires, Argentina), con notas sobre su taxonomia y distribucion. Revista del Museo Argentino de Ciencias Naturales 17 (2): 183 - 211. Available from https: // ri. conicet. gov. ar / handle / 11336 / 46712 [accessed 5 Mar. 2021].","Cutler B. & Edwards G. B. 2002. The jumping spiders (Araneae: Salticidae) of Trinidad and Tobago. Living World. Journal of the Trinidad and Tobago Field Naturalists' Club 2002: 39 - 44.","Olson D. M., Dinerstein E., Wikramanayake E. D., Burgess N. D., Powell G. V. N., Underwood E. C., D'Amico J. A., Itoua I., Strand H. E., Morrison J. C., Loucks C. J., Allnutt T. F., Ricketts T. H., Kura Y., Lamoreux J. F., Wettengel W. W., Hedao P. & Kassem K. R. 2011. Terrestrial ecoregions of the world: a new map of life on Earth. BioScience 51 (11): 933 - 938. https: // doi. org / 10.1641 / 0006 - 3568 (2001) 051 [0933: TEOTWA] 2.0. CO; 2","Oliveira P. S. 1988. Ant-mimicry in some Brazilian salticid and clubionid spiders (Araneae: Salticidae, Clubionidae). Biological Journal of the Linnean Society 33: 1 - 15. https: // doi. org / 10.1111 / j. 1095 - 8312.1988. tb 00443. x","Taczanowski L. 1871. Les araneides de la Guyane francaise. Horae Societatis entomologicae Rossicae 8: 32 - 132.","Cutler B. 1993. A new Middle American Synemosyna (Araneae: Salticidae). Revista nicaraguense de entomologia 24: 1 - 4."]}

Genus Synemosyna Hentz, 1846 Synemosyna Hentz, 1846: 367. Simonella Peckham & Peckham, 1885: 23 (considered a junior synonym by Galiano 1966: 341). Type species Synemosyna formica Hentz, 1846 (by original designation). Diagnosis (modified from Perger & Rubio 2020a) Species of Synemosyna can be distinguished from those of Fluda Peckham & Peckham, 1892 and Erica Peckham & Peckham, 1892 by a carapace laterally constricted (continuous in the latter genera) and an epigyne with a single opening (2���3 in the latter genera), and from those of Sympolymnia Perger & Rubio, 2020 by a fusiform abdomen that is longer than the carapace, the carapace with at the most one lateral white patch, and spermathecae small, globular, pear- or kidney-shaped (abdomen ovate, carapace with 2 lateral white patches and spermathecae lung-shaped in Sympolymnia). Cylistella Simon, 1901 has a rounded, beetle-like habitus without constrictions and is the morphologically most distinct group within this tribe. Key to adults of Bolivian species of Synemosyna 1. Constriction between cephalic and thoracic parts weakly defined laterally, only slightly narrower than both parts (Figs 3A, C, 4A, C).................................................................................................. 2 ��� Constriction between cephalic and thoracic parts distinct laterally, considerably narrower than both parts (Figs 3B, 4B)........................................................... S. myrmeciaeformis (Taczanowski, 1871) 2. Males................................................................................................................................................. 3 ��� Females............................................................................................................................................. 4 3. Tibial apophysis of male palp with obtuse dorsal tooth............ S. aurantiaca (Mello-Leit��o, 1917) ��� Tibial apophysis of male palp bifurcate (Fig. 5E)............................ S. nicaraguaensis Cutler, 1993 4. Copulatory ducts entering spermathecae anteriorly, approximately four times longer than spermathecae.............................................................................. S. aurantiaca (Mello-Leit��o, 1917) ��� Copulatory ducts enter spermathecae posteriorly, approximately twice the length of spermathecae (Fig. 5D)........................................................................................... S. nicaraguaensis Cutler, 1993, Published as part of Perger, Robert, Rubio, Gonzalo D. & Haddad, Charles R., 2021, On ant-like Synemosyna Hentz, 1846 spiders from Bolivia, with indirect evidence for polymorphic mimicry complexes (Araneae: Salticidae: Simonellini), pp. 67-88 in European Journal of Taxonomy 748 (1) on page 71, DOI: 10.5852/ejt.2021.748.1343, http://zenodo.org/record/4745113, {"references":["Peckham G. W. & Peckham E. G. 1885. On some new genera and species of the Attidae, from Madagascar and Central America. Proceedings of the Natural History Society of Wisconsin 1885: 23 - 42.","Galiano M. E. 1966. Salticidae (Araneae) formiciformes V. Revision del genero Synemosyna Hentz, 1846. Museo Argentino de Ciencias Naturales Bernardino Rivadavia 1: 339 - 380.","Perger R. & Rubio G. D. 2020 a. Sympolymnia, a new genus of Neotropical ant-like spider, with description of two new species and indirect evidence for transformational mimicry (Araneae, Salticidae, Simonellini). Zoosystematics and Evolution 96 (2): 781 - 795. https: // doi. org / 10.3897 / zse. 96.55210","Peckham G. W. & Peckham E. G. 1892. Ant-like spiders of the family Attidae. Occasional Papers of the Natural History Society of Wisconsin 2 (1): 1 - 84. Available from https: // www. biodiversitylibrary. org / part / 82774 [accessed 5 Mar. 2021].","Simon E. 1901. Histoire naturelle des Araignees. Deuxieme edition, tome second. Roret, Paris. https: // doi. org / 10.5962 / bhl. title. 51973","Taczanowski L. 1871. Les araneides de la Guyane francaise. Horae Societatis entomologicae Rossicae 8: 32 - 132.","Mello-Leitao C. F. de 1917. Aranhas novas ou pouco conhecidas de Thomisidas e Salticidas brasileiras. Archivos da Escola Superior de Agricultura e Medicina Veterinaria, Rio de Janeiro 1: 117 - 153.","Cutler B. 1993. A new Middle American Synemosyna (Araneae: Salticidae). Revista nicaraguense de entomologia 24: 1 - 4."]}

Synemosyna nicaraguaensis Cutler, 1993 Figs 3C, F, 4C, F, 5C���F, 9A���D Synemosyna nicaraguaensis Cutler, 1993: 2, figs 1���2. Type deposit Holotype ♀ in MEL (examined). Diagnosis Bulb small (60% of cymbium length) (Fig. 5F); tibial apophysis bifurcate, ventral lobe larger; epigyne with complete, sclerotized transverse hood with strongly procurved anterior margin; female spermathecae large (width of each 35���45% of maximum width of epigyne between sclerotized rims), pear-shaped, and accessory glands as long as spermatheca diameter (Fig. 5D). Material examined Holotype NICARAGUA ��� ♀; R��o San Juan, El Castillo [El Castillo de Concepci��n, 11��01��� N, 84��25��� W]; 30 Jul. 1989; F. Reinboldt leg.; MEL. Other material BOLIVIA ��� Santa Cruz Dept ��� 25 ♂♂, 37 ♀♀; La Guardia; 17.883�� S, 63.317�� W; Sep. 2015 ��� Aug. 2017; R. Perger leg.; CBF ��� ��� 2 ♂♂, 2 ♀♀; same collection data as for preceding; IBSI-Ara 0724 ��� 1 ♀; Buena Vista; 17.465�� S, 63.696�� W; 21 Jan. 2016; R. Perger leg.; IBSI-Ara 0725 ��� 2 ♀♀; same collection data as for preceding; CBF ��� 5 ♂♂, 8 ♀♀; Santa Mar��a La Antigua; 17.3719�� S, 63.6563�� W; 10���11 Apr. 2018; R. Perger leg.; CBF. ��� La Paz Dept ��� 2 ♂♂, 6 ♀♀; Villa Teresa; 16.201�� S, 67.829�� W; 17 Jan. 2018; R. Perger leg.; IBSI-Ara 1023 ��� 3 ♂♂, 8 ♀♀; same collection data as for preceding; 16���18 Jan. 2018; R. Perger leg.; CBF. Comparisons The bifurcated tibial apophysis is only shared with S. maddisoni Cutler, 1985 and S. paraensis Galiano, 1967. However, both species can be separated from S. nicaraguaensis by a larger bulb (65% of cymbium length in S. maddisoni and 80% in S. paraensis) and the different size and shape of the lobes of the retrolateral tibial apophysis (cf. Fig. 5F with Cham��-V��zquez et al. 2020: figs 14���15). The female habitus (Fig. 4C) and the almost round spermathecae (Fig. 5D) are shared with the female of S. maddisoni. Females of S. maddisoni can be distinguished from those of S. nicaraguaensis by an epigyne with a U-shaped flap, which is wide and short in S. nicaraguaensis (compare Fig. 5C with Cham��-V��zquez et al. 2020: fig. 11). The habitus of females of S. nicaraguaensis resembles that of S. hentzi Peckham & Peckham, 1892 (Peckham & Peckham 1892: pl. 7 fig. 2). Synemosyna hentzi was described very briefly based on a single female that was collected in Chapada dos Guimar��es, close to Cuiab��, Mato Grosso state, Brazil, and maintained in the Herbert H. Smith collection (Peckham & Peckham 1892), but the type specimen appears to be lost (Galiano 1971). No description or illustration of the genitalia was provided, and no further specimens of this species were recorded in subsequent works. Considering the original description (Peckham & Peckham 1892), S. hentzi differs from S. nicaraguaensis by a broader, less elongated carapace, 2 times longer than wide (2.5 in S. nicaraguaensis), with the lateral borders parallel (with a slightly narrower constriction in S. nicaraguaensis) and without a dorsal constriction (present in S. nicaraguaensis), the AER only slightly curved (strongly in S. nicaraguaensis), and the ALE and AME touching each other (separated in S. nicaraguaensis). Description Male (IBSI-Ara 0724) Total BL: 4.60. Carapace length 2.05; width: 0.81. Integument smooth, shiny, with simple, separate white setae, denser and longer on cephalic area (particularly around AME), posterior constriction of thoracic area, and on transverse area of anterior part of abdomen. Carapace slender and elongated, cephalic portion slightly longer than wide, followed by slightly narrower constriction, slight impression behind cephalic area (resembling anterior pronotal margin); dense white scales posterolateral of cephalic area, thoracic part about as long as cephalic part, of same width, globose, posteriorly constricted, constriction concave in lateral view. Row of AME and ALE recurved, AME touching each other, separated from ALE, ALE at anterior edges of cephalic area, rectangle of ALE and PE 1.5 times as wide as long. Chelicerae with 2 promarginal and 5 retromarginal teeth. Sternum dark brown. Coxae and trochanters translucent whitish; legs relatively long, 4312; femora stout, remaining segments slender, femora I���III translucent yellowish, I and III somewhat darker distally, IV proximally and distally as in anterior part of abdomen, central half translucent yellowish; tibia I simple, with moderately long setae. Abdomen length: 2.50; width: 0.70, proximal half broadening in distal direction in dorsal view, proximal and distal half of abdomen separated by concave constriction, distal part of abdomen acuminate oval. Dorsal scutum almost extending along the whole length of abdomen, rectangular proximally, merging into disc-shaped, heavily sclerotized plate on distal half of anterior abdominal part, proximal part of plate surrounded by band of short white setae; scutum narrowing at level of abdominal constriction and widening again on distal half of abdomen, terminating shortly before abdominal apex; epigastric scutum folded over dorsolateral border of anterior part of abdomen, covering somewhat lateral sides of scutum. Bulb about 60% of the cymbium length, ovoid, narrowing proximally, tibial apophysis bifurcate, ventral tooth larger; embolus thin, arising from the basal side of the bulb, without complete circular revolution, angular, surrounding bulb prolaterally towards the apical cymbial groove. Variation Four different color morphs were observed in males (Fig. 9B���D, Table 1), with no apparent ontogenetic change in body color. The forms included orange-reddish, completely light brown or dark brown forms (all locations), and variants with dark orange carapace and black abdomen (La Guardia and Villa Teresa). In a sample of 51 females, 92% were orange with a pair of black spots on the posterior half of the abdomen (Fig. 9A), and 8% dark brown. Orange males with dark spots on their abdomen were not observed. The comparably even distribution of color forms among males, the high number of orange females, and the lack of orange forms with black abdominal spots in males indicate sex-specific polychromatism. Geographical and ecoregion distribution (Fig. 7) Synemosyna nicaraguaensis is known from Nicaragua (type location) and Bolivia (present study). In Bolivia, it was collected in the following forest types: Bolivian Yungas forest (Villa Teresa), Sub-Andean Southwest Amazon moist forest (Buena Vista), and Sub-Andean subhumid semi-deciduous Chiquitano forest (La Guardia, Santa Mar��a la Antigua). Remarks The localities in Nicaragua and Bolivia are more than 3500 km apart, which is remarkable for such a small arthropod, and would suggest a certain degree of reproductive isolation between both populations. However, the epigyne (examined from drawings by Cutler 1993) and external morphology (examined from photographs of the type) are similar in the type specimen of S. nicaraguaensis and the Bolivian material. Another species with a wide distributional range, S. americana, occurs from Mexico to Venezuela (Cutler 1981b). The other two Bolivian species of Synemosyna have also comparably wide distribution ranges (Fig. 7). The widest collection locations of S. aurantiaca are about 2200 km apart, and of S. myrmeciaeformis about 3100 km. Molecular studies are needed to test whether the low grade of morphological differentiation in these species represents a high gene flow., Published as part of Perger, Robert, Rubio, Gonzalo D. & Haddad, Charles R., 2021, On ant-like Synemosyna Hentz, 1846 spiders from Bolivia, with indirect evidence for polymorphic mimicry complexes (Araneae: Salticidae: Simonellini), pp. 67-88 in European Journal of Taxonomy 748 (1) on pages 76-80, DOI: 10.5852/ejt.2021.748.1343, http://zenodo.org/record/4745113, {"references":["Cutler B. 1993. A new Middle American Synemosyna (Araneae: Salticidae). Revista nicaraguense de entomologia 24: 1 - 4.","Taczanowski L. 1871. Les araneides de la Guyane francaise. Horae Societatis entomologicae Rossicae 8: 32 - 132.","Cutler B. 1985. Taxonomic notes on Neotropical species in the genus Synemosyna (Araneae: Salticidae). Studies on Neotropical Fauna and Environment 20: 83 - 91. https: // doi. org / 10.1080 / 01650528509360674","Galiano M. E. 1967. Salticidae (Araneae) formiciformes. VIII. Nuevas descripciones. Physis. Revista de la Sociedad Argentina de Ciencias Naturales 27: 27 - 39.","Chame-Vazquez D., Cutler B. & Ibarra-Nunez G. 2020. Further taxonomic notes on the jumping spider Synemosyna maddisoni Cutler, 1985 (Araneae: Salticidae). Arthropoda Selecta 29 (4): 475 - 480. https: // doi. org / 10.15298 / arthsel. 29.4.10","Peckham G. W. & Peckham E. G. 1892. Ant-like spiders of the family Attidae. Occasional Papers of the Natural History Society of Wisconsin 2 (1): 1 - 84. Available from https: // www. biodiversitylibrary. org / part / 82774 [accessed 5 Mar. 2021].","Galiano M. E. 1971. Salticidae (Araneae) formiciformes. XII. Descripcion del allotypus de Synemosyna paraensis Galiano, 1967. Revista de la Sociedad Entomologica Argentina 33: 133 - 135.","Mello-Leitao C. F. de 1917. Aranhas novas ou pouco conhecidas de Thomisidas e Salticidas brasileiras. Archivos da Escola Superior de Agricultura e Medicina Veterinaria, Rio de Janeiro 1: 117 - 153.","Cutler B. 1981 b. Synemosyna bicolor is the female of Synemosyna americana (Araneae, Salticidae). Journal of Arachnology 9: 342 - 343."]}

Capobula ukhahlamba spec. nov. Figs 10, 73, 74 Type material: Holotype ♀: SOUTH AFRICA: KwaZulu-Natal: Champagne Castle, Hlathikulu Forest, 29°02.366’S, 29°23.421’E, 1560 m a.s.l., 20.I.2011, leg. C. Haddad (base of grass tussocks, grassland-forest ecotone) (NCA 2018 /35). Paratypes: SOUTH AFRICA: KwaZulu-Natal: Cathedral Peak Forest Station, 75 km WSW Estcourt, 1380 m a.s.l., 14.XII.1979, leg. S. & J. Peck (Leucosidea boulder bush scrub, night sweeping), 1♀ (AMNH); Didima National Park, Cathedral Peak area, Rainbow Gorge, 33.5 km WSW of Winterton, 28°57.600’S, 29°13.635’E, 1518 m a.s.l., 19.I.2011, leg. H. Wood & C. Griswold (sifting leaf litter, Afromontane forest), 1♀ (CAS, CASENT 9043022, SA11-041); Same locality, 28°56.982’S, 29°13.874’E, 1400 m a.s.l., 19.I.2011, leg. C. Haddad (base of grasses and ferns), 1♀ (MHBU); Royal Natal National Park, 38.6 km W of Bergville, 28°41.137’S, 28°57.425’E, 1403 m a.s.l., 21.I.2011, leg. H. Wood, C. Haddad & C. Griswold (general collecting in ferns and grass tussocks), 1♀ (CAS, CASENT 9043423, SA11-046); Sani Pass, IX.2006, leg. D. Prentice (pitfall traps), 1♀ (NCA 2008 /1982); Sani Pass elevational project, Ixopo, 30°11.010’S, 30°09.130’E, 900 m a.s.l., 1.I.2009, leg. Univ. of Pretoria students (pitfall traps, site 8a), 1♀ (NCA 2011 /774). Diagnosis. Females can be easily distinguished by the large circular ridges surrounding the copulatory openings (Fig. 73), the oval rather than teardrop-shaped primary spermathecae (Fig. 74), and the presence of a cream Y-shaped marking on the abdominal dorsum (Fig. 10), which is absent in the other species (Figs 3–9). Male unknown. Remarks. Although the male of this species is unknown and the female has an inverted Y-shaped cream marking that it absent in congeners, we place this species in Capobula and not Orthobula because of the absence of pits along the dorsal midline of the carapace and the anterolaterally-positioned spermathecae; name in apposition. Etymology. Taken from the name of the Ukhahlamba-Drakensberg Transfrontier Conservation Park, a conservation initiative between South Africa and Lesotho, which includes the distribution range of this species. Ukhahlamba is the isiZulu word meaning “barrier of spears”; noun in apposition. Female (holotype, NCA 2018/35). Measurements: CL 0.89, CW 0.67, AL 1.06, AW 0.89, TL 2.15 (2.03–2.15), PERW 0.39, MOQAW 0.12, MOQPW 0.16, MOQL 0.15. Length of leg segments: I 0.54 + 0.22 + 0.48 + 0.41 + 0.27 = 1.92; II 0.48 + 0.22 + 0.37 + 0.37 + 0.25 = 1.69; III 0.41 + 0.19 + 0.33 + 0.33 + 0.24 = 1.50; IV 0.57 + 0.24 + 0.48 + 0.54 + 0.30 = 2.13. Colour: carapace orange-brown, pits and lateral margins brown; chelicerae yellow-brown; endites and labium yellow-brown proximally, distal third cream; sternum yellow, pit margins and lateral margins yellow-brown; palps and legs creamy-yellow; abdomen dark grey dorsally and laterally, dorsally with diverging cream Y-shaped marking in anterior half, narrow cream chevrons in posterior half; venter cream, with faint grey mottling; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with tiny copulatory openings in round epigynal ridges (Fig. 73); copulatory ducts initially curving medially, looping posteriorly, then laterally, before entering anterolateral primary spermathecae along mesal margin; bursae small and spherical (Fig. 74). Habitat and biology. Distributed in montane grasslands at elevations between 900–1600 m a.s.l. Distribution. Only known from KwaZulu-Natal, South Africa, with most records from the Drakensberg Mountains (Fig. 75)., Published as part of Haddad, Charles R., Jin, Chi, Platnick, Norman I. & Booysen, Ruan, 2021, Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae), pp. 41-71 in Zootaxa 4942 (1) on pages 59-60, DOI: 10.11646/zootaxa.4942.1.2, http://zenodo.org/record/4596097

Genus Capobula gen. nov. Type species: Orthobula infima Simon, 1896a Diagnosis. Capobula gen. nov. can be recognized from all other trachelid genera except Orthobula by the combination of a large number of paired ventral spines on the anterior tibiae, metatarsi and tarsi, the absence of ventral cusps in both sexes, the deep pits on the carapace, and the presence of a large ventral sclerite on the abdomen of males. Although Spinotrachelas Haddad, 2006 and Poachelas also have heavily spined anterior legs, both have ventral cusps on the anterior legs, lack carapace pits, and lack a ventral abdominal scutum (Haddad 2006; Haddad & Lyle 2008). Female Capobula gen. nov. can be recognized by the laterally positioned, teardrop-shaped primary spermathecae that are in the same transverse plane as the copulatory openings, while in Orthobula the spermathecae are oval and positioned posteriorly, close to the midline of the epigyne, and far from the anteriorly positioned copulatory openings (e.g. Marusik et al. 2013: figs 19–22). Further, Capobula gen. nov. lack pits along the midline of the carapace (Figs 3–11), which are present in Orthobula (e.g. Marusik et al. 2013: figs 1, 4; Ramírez 2014: fig. 6A). Capobula gen. nov. can be separated from Orthobula by the male palpal tegulum being pear-shaped and approximately as broad as the cymbium, and the embolus with a broad base, clear bend near its midpoint, and tip directed retrodistally (e.g. Fig. 52). In Orthobula the tegulum is subtriangular and strongly swollen basally towards the prolateral side, and the embolus usually straight, narrow, finely coiled and directed distally (e.g. Marusik et al. 2013: figs 11–16). Etymology. The genus name is a combination of Cape, referring to the Western and Eastern Cape Provinces of South Africa from where the majority of the species have been recorded, and Orthobula, to which it is closely related. Gender is feminine. Description. Small spiders, 1.70–2.35 mm in total length; carapace bright orange to deep red-brown (Figs 3–10); carapace oval, broadest at middle of coxae II, eye region narrow, fovea indistinct, a short shallow depression (Fig. 11); posterior margin very slightly concave; carapace surface finely granulose, with series of deep pits forming striae, pits absent from midline of carapace (Figs 11, 12), each pit with central pore; surface sparsely covered with scattered fine curved setae (Figs 12, 13). All eyes surrounded by black rings (Figs 3–10); anterior eye row procurved in anterior view, recurved in dorsal view, anterior median eyes approximately ¾ anterior lateral eye diameter; anterior median eyes separated by approximately ½ their diameter, nearly touching anterior lateral eyes (Figs 13, 14); posterior eye row recurved in dorsal view, posterior median eyes oval, posterior lateral eyes round; posterior median eyes approximately 1¼ times posterior lateral eye diameter; MOQ narrower anteriorly than posteriorly, length and posterior width approximately equal. Chilum indistinct, a tiny transverse sclerite; cheliceral promargin and retromargin with two teeth each, cheliceral escort seta present (Figs 15, 16); fang with distinct serrula (Fig. 16); endites convergent, not sexually dimorphic, slightly notched laterally (Fig. 17), with distinct serrula comprising sharp, ventrally curved denticles (Fig. 18); dense maxillar hair tuft on mesal margins (Fig. 17); labium trapezoidal, slightly wider than long. Pleural bars sclerotised, isolated; sternum shield-shaped, longer than broad, surface smooth with deep pits without pores, sparsely covered in long straight setae (Figs 19, 20); precoxal triangles present, intercoxal sclerites present between all coxal pairs. Leg formula 4132, sparsely covered in long fine setae; femora I with mesal convex curvature, all femora strongly constricted proximally, with sparse ventral tubercles in basal half (Figs 21, 22); patellae with small lyriform organ on prolateral side (Fig. 23), patellar indentation on retrolateral side narrow, with lyriform organ at proximal end (Figs 24, 25); anterior legs with strong paired ventral spines on tibiae, metatarsi and tarsi (Figs 26–29); metatarsi IV with sparse chemosensory setae and trichobothria dorsally, ventral preening brush at distal end (Fig. 30); tarsi with sparse tactile hairs, few dorsal trichobothria and chemosensory setae (Figs 31, 32); trichobothria with sunken distal plate, distal margin of hood overlapping plate, hood with four curved ridges, roughly concentric (Fig. 33); tarsal organ oval, very slightly elevated from integument, surface finely wrinkled, opening oval and distally placed (Figs 31, 34); paired tarsal claws short, with two teeth and moderately dense tenant setae forming claw tufts in between (Figs 35, 36); palpal claw simple, sharply curved distally (Fig. 37). Abdomen oval, clearly larger in females than males, with dorsal scutum in males only; dorsum with very sparse fine setae and two pairs of sigilla, prominent in females, barely distinguishable on scutum in males (Figs 3–10, 38); venter without post-epigastric sclerites, with large ventral sclerite in males that is wider than long, absent in females; small, weakly sclerotized inframamillary sclerite present in females, indistinct in males. Spinnerets (observed in C. infima comb. nov. only): female (Figs 39, 41–43): anterior lateral spinnerets of female each with one major ampullate gland spigot, one nubbin and approximately ten piriform gland spigots; posterior median spinnerets of female each with one small minor ampullate gland spigot and five large cylindrical gland spigots; posterior lateral spinnerets of female each with two large cylindrical gland spigots and approximately ten aciniform gland spigots; male (Figs 40, 44–46): anterior lateral spinnerets of male each with one major ampullate gland spigot and five piriform gland spigots; posterior median spinnerets of male each with only one minor ampullate gland spigot and one tartipore evident; posterior lateral spinnerets of male each with only a single tartipore discernible. Female epigyne with small paired copulatory openings positioned anteriorly in epigyne (Fig. 47), in same transverse plane as teardrop-shaped laterally positioned primary spermathecae (e.g. Figs 54, 59); copulatory ducts directed posteriorly along midline, with membranous anterior bursae originating near initial part of ducts; posteriorly, copulatory ducts bend laterally to enter spermathecae along their mesal margin (e.g. Figs 55, 60); secondary spermathecae not distinguished. Male palpal femur, patella and tibia all with ventral apophyses (Figs 48–52), patella with lyriform organ associated with apophysis (Figs 49, 50); tegulum pear-shaped, broadest basally, tapering distally, slightly broader than cymbium, with narrow sperm duct running down centre of tegulum in ventral view, with single basal loop (e.g. Fig. 57); embolus distal, curving retrodistally (Fig. 53), tegulum without any other structures. Composition. Capobula infima (Simon, 1896a) comb. nov. (type species) and four new species, C. capensis spec. nov., C. montana spec. nov., C. neethlingi spec. nov. and C. ukhahlamba spec. nov. Distribution. Only known from South Africa and the enclave of Lesotho.

Capobula capensis spec. nov. Figs 3–4, 54–58 Type material. Holotype ♀: SOUTH AFRICA: Western Cape: Cederberg Wilderness Area, Pakhuis Pass, 12.6 km 70° E of Clanwilliam, 32°08.419’S, 19°01.034’E, 773 m a.s.l., 9.X.2011, leg. L. Almeida, C. Griswold & T. Meikle (general collecting, Restionaceae fynbos) (CAS, CASENT 9043323, SA11-109). Paratypes: SOUTH AFRICA: Western Cape: Cederberg, leg. G. Smith, 1♂ (MRAC 133613); Cederberg, 32°21’S, 19°10’E, VII.1962, leg. N. Leleup (litter), 1♂ 5♀ (MRAC 131897); Cederberg Wilderness Area, Pakhuis Pass, 12.6 km 70° E of Clanwilliam, 32°08.419’S, 19°01.034’E, 773 m a.s.l., 9.X.2011, leg. L. Almeida, C. Griswold & T. Meikle (general collecting, Restionaceae fynbos), 1♂ (CAS, CASENT 9043335, SA11-112); Same locality, 13.5 km 76° E of Clanwilliam, 32°09.014’S, 19°01.793’E, 900 m a.s.l., 9.X.2011, leg. L. Almeida, C. Griswold & T. Meikle (general collecting, Restionaceae fynbos), 1♀ (CAS, CASENT 9043334, SA11-112); Same locality, VII.1962, leg. N. Leleup (litter), 4♂ 35♀ (MRAC 131979); Klein-Dassenberg Road, Farm Trinity, 33°34.94’S, 18°36.08’E, 25–27.XII.2012, leg. D. Ubick, 1♂ (CAS, CASENT 9055498); 17 km N Citrusdal, road on E bank of Olifants River, 32°35’S, 19°01’E, 7.I.1989, leg. R. Jocqué, 1♀ (MRAC 169745); Touws River [33°20’S, 20°01’E], 1953, leg. J. Gordon, 5♂ 9♀ (MRAC 127791). Diagnosis. Females are distinct from congeners by the M-shaped atrial margins, with the copulatory openings positioned in the anterolateral corners (Fig. 54). Males of this species share with C. infima comb. nov. the broad, ridge-like retrolateral tibial apophysis, but can be distinguished by the spike-shaped femoral apophysis and short, stout distal section of the embolus that is directed retro-distally (Figs 57, 58), while in C. infima comb. nov. the femoral apophysis is lobate and the distal section of the embolus long, narrow and curved (Figs 62, 63). Males are also similar to C. montana spec. nov., but have a clearly thicker embolus (compare Figs 57 and 68). Etymology. Named for the Western Cape Province, from which it originates; Latin adjective. Female (holotype, CAS, CASENT 9043323). Measurements: CL 0.90, CW 0.70, AL 1.24, AW 1.06, TL 2.35 (1.90–2.35), PERW 0.30, MOQAW 0.11, MOQPW 0.16, MOQL 0.17. Length of leg segments: I 0.67 + 0.27 + 0.57 + 0.51 + 0.28 = 2.30; II 0.59 + 0.23 + 0.48 + 0.44 + 0.27 = 2.01; III 0.51 + 0.21 + 0.33 + 0.43 + 0.25 = 1.73; IV 0.68 + 0.25 + 0.54 + 0.60 + 0.29 = 2.36. Colour: carapace deep orange-brown, pits and lateral margins brown; chelicerae yellow-brown; endites and labium yellow-brown proximally, distal ends cream; sternum bright yellow-orange, pit margins yellow-brown, lateral margins deep yellow-brown; palps creamy-yellow; legs with femora I and II pale yellow-brown, becoming progressively paler on III and IV, remaining segments creamy-yellow; abdomen dark grey dorsally and laterally, dorsally with seven fine cream chevrons in posterior half to spinnerets; venter paler mottled grey; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 3. Epigyne with small copulatory openings positioned anterolaterally in shallow M-shaped ridges (Fig. 54); copulatory ducts short and C-shaped, initially curving mesally, then laterally before entering teardrop-shaped lateral primary spermathecae along their interior margin; bursae large, elongate-oval, almost twice the size of primary spermathecae (Fig. 55). Male (paratype, CAS, CASENT 9043335). Measurements: CL 0.83, CW 0.60, AL 0.77, AW 0.60, TL 1.61 (1.58–1.82), PERW 0.27, MOQAW 0.10, MOQPW 0.14, MOQL 0.15. Length of leg segments: I 0.57 + 0.22 + 0.46 + 0.41 + 0.27 = 1.93; II 0.50 + 0.22 + 0.38 + 0.39 + 0.25 = 1.74; III 0.41 + 0.21 + 0.29 + 0.37 + 0.24 = 1.52; IV 0.56 + 0.22 + 0.46 + 0.51 + 0.26 = 2.01. Colour: carapace deep yellow, pits pale yellow-brown, lateral margins dark yellow-brown; chelicerae pale yellow-brown; endites yellow-brown medially, cream proximally and distally; labium yellow-brown, cream at distal end; sternum bright creamy-yellow, pit margins yellow, lateral margins yellow-brown; palps and legs creamy-yellow; abdomen with yellow dorsal scutum covered in dark grey mottling, cream with grey mottling laterally; venter with creamy-yellow epigastric and ventral sclerites, latter with faint grey mottling; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 5, II plv 5 rlv 4; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 3. Palpal femur with spike-like retrolateral apophysis; patella with very faint retrolateral ridge; tibia with retrolateral tibial apophysis triangular in ventral view, broad and ridge-like in retrolateral view; embolus short and stout, tip directed retrodistally (Figs 56–58). Habitat and biology. Mainly collected from litter or by hand in fynbos habitats. Distribution. Only known from a few localities in the Western Cape, South Africa (Fig. 75)., Published as part of Haddad, Charles R., Jin, Chi, Platnick, Norman I. & Booysen, Ruan, 2021, Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae), pp. 41-71 in Zootaxa 4942 (1) on pages 52-54, DOI: 10.11646/zootaxa.4942.1.2, http://zenodo.org/record/4596097

Capobula neethlingi spec. nov. Figs 9, 71, 72 Type material. Holotype ♀: SOUTH AFRICA: Western Cape: George, Saasveld Pass, 33°58.198’S, 22°31.778’E, 149 m a.s.l., 7.XII.2012, leg. J. Neethling (leaf litter, indigenous forest) (NCA 2019 /1002a). Paratypes: together with holotype, 1♀ (NCA 2019 /1002b). SOUTH AFRICA: Western Cape: Laingsburg, Anysberg Nature Reserve, 33°27.300’S, 20°34.862’E, 735 m a.s.l., 8.IX–8.X.2015, leg. Z. Mbo (pitfall traps, karoo veld) (NCA 2016 /2464); Swartberg Nature Reserve, Gamkaskloof, 33°21’S, 21°41’E, 15.I.2001, leg. Z. van der Walt (on ground), 2♀ (NCA 2002 /198). Diagnosis. Females of C. neethlingi spec. nov. have a similar epigyne to that of C. montana spec. nov., but can be distinguished by the V- rather than J-shaped ridges containing the copulatory openings, the slightly separated copulatory ducts (touching medially in C. neethlingi spec. nov.), and the bursae that are separated by approximately half their diameter, while almost touching in C. neethlingi spec. nov. (compare Figs 71 and 72 with Figs 65 and 66). Male unknown. Etymology. Named for arachnologist Jan Andries Neethling, who collected the holotype; name in genitive case. Female (holotype, NCA 2019/1002). Measurements: CL 0.84, CW 0.64, AL 1.02, AW 0.87, TL 1.95 (1.80– 1.98), PERW 0.30, MOQAW 0.12, MOQPW 0.15, MOQL 0.16. Length of leg segments: I 0.60 + 0.24 + 0.52 + 0.45 + 0.25 = 2.06; II 0.54 + 0.23 + 0.41 + 0.38 + 0.24 = 1.80; III 0.46 + 0.21 + 0.33 + 0.38 + 0.23 = 1.61; IV 0.59 + 0.24 + 0.49 + 0.56 + 0.27 = 2.15. Colour: carapace deep orange-brown, with black mottling at centre, pits and lateral margins with black edges; chelicerae deep yellow-brown, with faint black mottling; endites and labium yellow-brown, cream at distal ends; sternum bright orange, pits slightly darker, lateral margins orange-brown; palps pale yellow-brown; legs with femora and tibiae I and II pale orange-brown, yellow-brown distally; patellae, metatarsi and tarsi I and II, and legs III and IV yellow-brown; abdomen dark grey dorsally and laterally, dorsally with eight fine cream chevrons in posterior half to spinnerets; venter slightly paler mottled grey; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 6, II plv 6 rlv 5; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with small copulatory openings in J-shaped epigynal ridges (Fig. 71); copulatory ducts initially curving medially, then posteriorly, looping dorsally and anterolaterally before entering teardrop-shaped primary spermathecae along their interior margin; bursae subtriangular, apices converging mesally, similar in size to primary spermathecae (Fig. 72). Habitat and biology. This species was recorded in two very contrasting biotopes, viz. moist Afromontane Forest and xeric Nama Karoo. Distribution. Only known from three localities in the Western Cape Province, South Africa (Fig. 75).

A new genus of the spider family Trachelidae L. Koch, 1872 from the Afrotropical Region is described. Capobula gen. nov. is represented by five species, known from South Africa and Lesotho only. Adults of both sexes of Orthobula infima Simon, 1896a, which is widely distributed in the Western Cape, South Africa, are described for the first time, and this species is transferred to Capobula gen. nov. as its type species. Four new species are described: C. capensis spec. nov. and C. neethlingi spec. nov. (South Africa: Western Cape), C. montana spec. nov. (Lesotho and South Africa: Eastern Cape, Free State and KwaZulu-Natal) and C. ukhahlamba spec. nov. (South Africa: KwaZulu-Natal). A phylogenetic analysis based on the cytochrome oxidase subunit I (COI) gene, including 14 genera of Trachelidae, one genus of Clubionidae Wagner, 1887 and three genera of Phrurolithidae Banks, 1892, supports the placement of Capobula gen. nov. in Trachelidae, with Orthobula Simon, 1897 as its likely closest relative.

Genus Capobula gen. nov. Type species: Orthobula infima Simon, 1896a Diagnosis. Capobula gen. nov. can be recognized from all other trachelid genera except Orthobula by the combination of a large number of paired ventral spines on the anterior tibiae, metatarsi and tarsi, the absence of ventral cusps in both sexes, the deep pits on the carapace, and the presence of a large ventral sclerite on the abdomen of males. Although Spinotrachelas Haddad, 2006 and Poachelas also have heavily spined anterior legs, both have ventral cusps on the anterior legs, lack carapace pits, and lack a ventral abdominal scutum (Haddad 2006; Haddad & Lyle 2008). Female Capobula gen. nov. can be recognized by the laterally positioned, teardrop-shaped primary spermathecae that are in the same transverse plane as the copulatory openings, while in Orthobula the spermathecae are oval and positioned posteriorly, close to the midline of the epigyne, and far from the anteriorly positioned copulatory openings (e.g. Marusik et al. 2013: figs 19–22). Further, Capobula gen. nov. lack pits along the midline of the carapace (Figs 3–11), which are present in Orthobula (e.g. Marusik et al. 2013: figs 1, 4; Ramírez 2014: fig. 6A). Capobula gen. nov. can be separated from Orthobula by the male palpal tegulum being pear-shaped and approximately as broad as the cymbium, and the embolus with a broad base, clear bend near its midpoint, and tip directed retrodistally (e.g. Fig. 52). In Orthobula the tegulum is subtriangular and strongly swollen basally towards the prolateral side, and the embolus usually straight, narrow, finely coiled and directed distally (e.g. Marusik et al. 2013: figs 11–16). Etymology. The genus name is a combination of Cape, referring to the Western and Eastern Cape Provinces of South Africa from where the majority of the species have been recorded, and Orthobula, to which it is closely related. Gender is feminine. Description. Small spiders, 1.70–2.35 mm in total length; carapace bright orange to deep red-brown (Figs 3–10); carapace oval, broadest at middle of coxae II, eye region narrow, fovea indistinct, a short shallow depression (Fig. 11); posterior margin very slightly concave; carapace surface finely granulose, with series of deep pits forming striae, pits absent from midline of carapace (Figs 11, 12), each pit with central pore; surface sparsely covered with scattered fine curved setae (Figs 12, 13). All eyes surrounded by black rings (Figs 3–10); anterior eye row procurved in anterior view, recurved in dorsal view, anterior median eyes approximately ¾ anterior lateral eye diameter; anterior median eyes separated by approximately ½ their diameter, nearly touching anterior lateral eyes (Figs 13, 14); posterior eye row recurved in dorsal view, posterior median eyes oval, posterior lateral eyes round; posterior median eyes approximately 1¼ times posterior lateral eye diameter; MOQ narrower anteriorly than posteriorly, length and posterior width approximately equal. Chilum indistinct, a tiny transverse sclerite; cheliceral promargin and retromargin with two teeth each, cheliceral escort seta present (Figs 15, 16); fang with distinct serrula (Fig. 16); endites convergent, not sexually dimorphic, slightly notched laterally (Fig. 17), with distinct serrula comprising sharp, ventrally curved denticles (Fig. 18); dense maxillar hair tuft on mesal margins (Fig. 17); labium trapezoidal, slightly wider than long. Pleural bars sclerotised, isolated; sternum shield-shaped, longer than broad, surface smooth with deep pits without pores, sparsely covered in long straight setae (Figs 19, 20); precoxal triangles present, intercoxal sclerites present between all coxal pairs. Leg formula 4132, sparsely covered in long fine setae; femora I with mesal convex curvature, all femora strongly constricted proximally, with sparse ventral tubercles in basal half (Figs 21, 22); patellae with small lyriform organ on prolateral side (Fig. 23), patellar indentation on retrolateral side narrow, with lyriform organ at proximal end (Figs 24, 25); anterior legs with strong paired ventral spines on tibiae, metatarsi and tarsi (Figs 26–29); metatarsi IV with sparse chemosensory setae and trichobothria dorsally, ventral preening brush at distal end (Fig. 30); tarsi with sparse tactile hairs, few dorsal trichobothria and chemosensory setae (Figs 31, 32); trichobothria with sunken distal plate, distal margin of hood overlapping plate, hood with four curved ridges, roughly concentric (Fig. 33); tarsal organ oval, very slightly elevated from integument, surface finely wrinkled, opening oval and distally placed (Figs 31, 34); paired tarsal claws short, with two teeth and moderately dense tenant setae forming claw tufts in between (Figs 35, 36); palpal claw simple, sharply curved distally (Fig. 37). Abdomen oval, clearly larger in females than males, with dorsal scutum in males only; dorsum with very sparse fine setae and two pairs of sigilla, prominent in females, barely distinguishable on scutum in males (Figs 3–10, 38); venter without post-epigastric sclerites, with large ventral sclerite in males that is wider than long, absent in females; small, weakly sclerotized inframamillary sclerite present in females, indistinct in males. Spinnerets (observed in C. infima comb. nov. only): female (Figs 39, 41–43): anterior lateral spinnerets of female each with one major ampullate gland spigot, one nubbin and approximately ten piriform gland spigots; posterior median spinnerets of female each with one small minor ampullate gland spigot and five large cylindrical gland spigots; posterior lateral spinnerets of female each with two large cylindrical gland spigots and approximately ten aciniform gland spigots; male (Figs 40, 44–46): anterior lateral spinnerets of male each with one major ampullate gland spigot and five piriform gland spigots; posterior median spinnerets of male each with only one minor ampullate gland spigot and one tartipore evident; posterior lateral spinnerets of male each with only a single tartipore discernible. Female epigyne with small paired copulatory openings positioned anteriorly in epigyne (Fig. 47), in same transverse plane as teardrop-shaped laterally positioned primary spermathecae (e.g. Figs 54, 59); copulatory ducts directed posteriorly along midline, with membranous anterior bursae originating near initial part of ducts; posteriorly, copulatory ducts bend laterally to enter spermathecae along their mesal margin (e.g. Figs 55, 60); secondary spermathecae not distinguished. Male palpal femur, patella and tibia all with ventral apophyses (Figs 48–52), patella with lyriform organ associated with apophysis (Figs 49, 50); tegulum pear-shaped, broadest basally, tapering distally, slightly broader than cymbium, with narrow sperm duct running down centre of tegulum in ventral view, with single basal loop (e.g. Fig. 57); embolus distal, curving retrodistally (Fig. 53), tegulum without any other structures. Composition. Capobula infima (Simon, 1896a) comb. nov. (type species) and four new species, C. capensis spec. nov., C. montana spec. nov., C. neethlingi spec. nov. and C. ukhahlamba spec. nov. Distribution. Only known from South Africa and the enclave of Lesotho., Published as part of Haddad, Charles R., Jin, Chi, Platnick, Norman I. & Booysen, Ruan, 2021, Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae), pp. 41-71 in Zootaxa 4942 (1) on pages 46-51, DOI: 10.11646/zootaxa.4942.1.2, http://zenodo.org/record/4596097, {"references":["Simon, E. (1896 a) Descriptions d'arachnides nouveaux de la famille des Clubionidae. Annales de la Societe Entomologique de Belgique, 40, 400 - 422.","Haddad, C. R. (2006) Spinotrachelas, a new genus of tracheline sac spiders from South Africa (Araneae: Corinnidae). African Invertebrates, 47, 85 - 93.","Haddad, C. R. & Lyle, R. (2008) Three new genera of tracheline sac spiders from southern Africa (Araneae: Corinnidae). African Invertebrates, 49, 37 - 76. https: // doi. org / 10.5733 / afin. 049.0204","Marusik, Y. M., Ozkutuk, R. S. & Kunt, K. B. (2013) On the identity and distribution of the poorly known spider Orthobula charitonovi (Mikhailov, 1986) (Aranei: Corinnidae). Arthropoda Selecta, 22, 157 - 162.","Ramirez, M. J. (2014) The morphology and phylogeny of dionychan spiders (Araneae: Araneomorphae). Bulletin of the American Museum of Natural History, 390, 1 ‾ 374. https: // doi. org / 10.1206 / 821.1"]}

Capobula neethlingi spec. nov. Figs 9, 71, 72 Type material. Holotype ♀: SOUTH AFRICA: Western Cape: George, Saasveld Pass, 33°58.198’S, 22°31.778’E, 149 m a.s.l., 7.XII.2012, leg. J. Neethling (leaf litter, indigenous forest) (NCA 2019 /1002a). Paratypes: together with holotype, 1♀ (NCA 2019 /1002b). SOUTH AFRICA: Western Cape: Laingsburg, Anysberg Nature Reserve, 33°27.300’S, 20°34.862’E, 735 m a.s.l., 8.IX–8.X.2015, leg. Z. Mbo (pitfall traps, karoo veld) (NCA 2016 /2464); Swartberg Nature Reserve, Gamkaskloof, 33°21’S, 21°41’E, 15.I.2001, leg. Z. van der Walt (on ground), 2♀ (NCA 2002 /198). Diagnosis. Females of C. neethlingi spec. nov. have a similar epigyne to that of C. montana spec. nov., but can be distinguished by the V- rather than J-shaped ridges containing the copulatory openings, the slightly separated copulatory ducts (touching medially in C. neethlingi spec. nov.), and the bursae that are separated by approximately half their diameter, while almost touching in C. neethlingi spec. nov. (compare Figs 71 and 72 with Figs 65 and 66). Male unknown. Etymology. Named for arachnologist Jan Andries Neethling, who collected the holotype; name in genitive case. Female (holotype, NCA 2019/1002). Measurements: CL 0.84, CW 0.64, AL 1.02, AW 0.87, TL 1.95 (1.80– 1.98), PERW 0.30, MOQAW 0.12, MOQPW 0.15, MOQL 0.16. Length of leg segments: I 0.60 + 0.24 + 0.52 + 0.45 + 0.25 = 2.06; II 0.54 + 0.23 + 0.41 + 0.38 + 0.24 = 1.80; III 0.46 + 0.21 + 0.33 + 0.38 + 0.23 = 1.61; IV 0.59 + 0.24 + 0.49 + 0.56 + 0.27 = 2.15. Colour: carapace deep orange-brown, with black mottling at centre, pits and lateral margins with black edges; chelicerae deep yellow-brown, with faint black mottling; endites and labium yellow-brown, cream at distal ends; sternum bright orange, pits slightly darker, lateral margins orange-brown; palps pale yellow-brown; legs with femora and tibiae I and II pale orange-brown, yellow-brown distally; patellae, metatarsi and tarsi I and II, and legs III and IV yellow-brown; abdomen dark grey dorsally and laterally, dorsally with eight fine cream chevrons in posterior half to spinnerets; venter slightly paler mottled grey; spinnerets cream. Leg spination: femora and patellae: spineless; tibiae: I plv 6 rlv 6, II plv 6 rlv 5; metatarsi: I plv 4 rlv 4, II plv 4 rlv 4; tarsi: I plv 3 rlv 3, II plv 3 rlv 2. Epigyne with small copulatory openings in J-shaped epigynal ridges (Fig. 71); copulatory ducts initially curving medially, then posteriorly, looping dorsally and anterolaterally before entering teardrop-shaped primary spermathecae along their interior margin; bursae subtriangular, apices converging mesally, similar in size to primary spermathecae (Fig. 72). Habitat and biology. This species was recorded in two very contrasting biotopes, viz. moist Afromontane Forest and xeric Nama Karoo. Distribution. Only known from three localities in the Western Cape Province, South Africa (Fig. 75)., Published as part of Haddad, Charles R., Jin, Chi, Platnick, Norman I. & Booysen, Ruan, 2021, Capobula gen. nov., a new Afrotropical dark sac spider genus related to Orthobula Simon, 1897 (Araneae: Trachelidae), pp. 41-71 in Zootaxa 4942 (1) on pages 58-59, DOI: 10.11646/zootaxa.4942.1.2, http://zenodo.org/record/4596097