Revision of the Afrotropical Crab-Spider Genus Parabomis Kulczyński, 1901 (Araneae: Thomisidae)

Zootaxa 4899 (1): 161–174 ISSN 1175-5326 (print edition) https://www.mapress.com/j/zt/ Article ZOOTAXA Copyright © 2020 Magnolia Press ISSN 1175-5334 (online edition) https://doi.org/10.11646/zootaxa.4899.1.8 http://zoobank.org/urn:lsid:zoobank.org:pub:A0C2249D-1099-4F9E-80F2-04A3770E8744

Revision of the Afrotropical crab-spider genus Parabomis Kulczyński, 1901 (Araneae: Thomisidae)

ANNA S. DIPPENAAR-SCHOEMAN1,2,3* & STEFAN H. FOORD4 1ARC-Plant Health and Protection, Private Bag X134, Queenswood 0121, South Africa 2Department of Zoology, Centre for Invasion Biology, University of Venda, Thohoyandou 0950, South Africa 3 �[email protected]; https://orcid.org/0000-0003-1532-1379 4 �[email protected]; https://orcid.org/0000-0002-9195-2562 *Corresponding author.

Abstract

The Afrotropical spider genus Parabomis Kulczyński, 1901 is revised. Members of Parabomis are some of the smallest thomisids known, and occur from Eritrea in the north of Africa to South Africa in the south, but are absent from Madagascar. Prior to this study, three species were known, namely P. levanderi Kulczyński, 1901 (Eritrea, ♂), P. martini Lessert, 1919 (Tanzania, ♂♀) and P. anabensis Lawrence, 1928 (Namibia, ♀). Parabomis anabensis sp. nov. is here recognized as a junior synonym of P. martini and four new species are described: P. elsae sp. nov. from South Africa (♂♀), P. megae sp. nov. from Zimbabwe (♂♀), P. pilosus sp. nov. from Botswana (♂♀) and P. wandae sp. nov. from Ghana (♂♀). A key to the six species is provided. The monotypic Afrotropical genus Felsina Simon, 1895, only known from its type species, F. granulum Simon, 1895, resembles Parabomis closely is known only from juveniles.

Key words: Africa, Bominae, Felsina, new species, taxonomy

Introduction

Members of the subfamily Bominae (Araneae: Thomisidae) are a unique group of nine thomisid genera character- ized by their small, globular bodies and short, stout legs (Ono 1988). Five of the nine Bominae genera are known from the Afrotropical Region and three have been revised: Avelis Simon, 1895 (Dippenaar-Schoeman 1986a), Ho- lopelus Simon, 1886 (Dippenaar-Schoeman 1986b) and Thomisops Karsch, 1879 (Dippenaar-Schoeman 1989). The fourth genus, Felsina Simon, 1895, is monotypic and was described from Senegal. According to Kulczyński (1901), Parabomis Kulczynski, 1901 and Felsina differ only in the shape of the clypeus and the presence of setae on the promargin of the chelicerae in Parabomis. The fifth genus, Parabomis, is revised here. The type species, P. levanderi Kulczyński, 1901, was based on a male from Eritrea; the second species, P. martini Lessert, 1919, was described from both sexes from Tanzania; and the third species, P. anabensis Lawrence, 1928, was described from Namibia based on females only. Members of Parabomis are rare in collections, and because of their small size they may easily be overlooked in the field. They are mainly collected by sweeping and beating grass, low shrubs and trees. However, during recent surveys in the Afrotropical Region they have also been sampled from canopy fogging trees. None of the species have been rede- scribed and several new species have been collected. This paper therefore aims to redescribe the known species and describe new species.

Material and methods

This revision is based on more than 120 specimens obtained from the following collections (curators given in brack- ets): FMNH—Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland (P. Cardoso); MCZ—

Accepted by Y. Marusik: 17 Nov. 2020; published: 30 Dec. 2020 161 Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, U.S.A. (G. Giribet); MHNG— Museum d’ Histoire Naturelle, Geneva (P. Schwendinger); MNHN—Muséum National d’Histoire Naturelle Paris, France (C. Rollard); MRAC—Musée Royal de l’Afrique Centrale, Tervuren, Belgium (R. Jocqué); NCA—National Collection of Arachnida, ARC-Plant Health and Protection, Pretoria, South Africa (P. Marais); NMBA—National Museum, Bloemfontein, South Africa (L. Lotz); NMSA—KwaZulu-Natal Museum, Pietermaritzburg, South Af- rica (A. Ndaba, K. Williams); NMZ—National Museum of Zimbabwe, Bulawayo, Zimbabwe (M. Fitzpatrick); SAM—Iziko South African Museum, Cape Town, South Africa (M. Cochrane, D. Larsen); SMF—Naturmuseum und Forschungsinstitut Senckenberg, Frankfurt am Main, Germany (P. Jäger); ZFMK—Zoologisches Forschungs- museum Alexander Koenig, Bonn, Germany (B. Huber). All specimens were preserved and examined in 70% ethanol. Measurements are given in millimetres and were taken using a Leica EZ4D stereomicroscope. Body measurements were taken of the holotypes and paratypes, as well as the smallest and largest specimens of both sexes to provide a size range. Leg segments were measured for leg I, II, III, IV on the lateral side of each segment and the sum given as the total length of each leg. Abbreviations used in text: AER—anterior eye row; AME—anterior median eyes; ALE—anterior lateral eyes; CL—carapace length; CW—carapace width; MOQ—median ocular quadrangle; PER—posterior eye row; PME—posterior median eyes; PLE—posterior lateral eyes; RTA—retrolateral tibial apophysis; TL—total length; VTA—ventral tibial apophysis. The techniques used for specimen preparation and study follow that of Dippenaar- Schoeman (1983), and terminology follows Ono (1988).

Taxonomy

Thomisidae Sundevall, 1833

Parabomis Kulczyński, 1901

Parabomis Kulczyński, 1901: 3, 30, figs 25–27; Simon 1903: 1011; Lessert 1919: 181; Ono 1988: 207; Dippenaar-Schoeman 2014: 244.

Type species. Parabomis levanderi Kulczyński, 1901.

Diagnosis. Parabomis shares with the other four African genera of the Bominae the globular body shape, small size and very short legs without spines. They share with Thomisops and Holopelus the peg-like setae on the promar- gins of the chelicerae, which are absent in Avelis and Felsina. It can be distinguished from Avelis, Thomisops and Holopelus by the high carapace (carapace height > carapace length), the very broad sloping clypeus, as well as the eyes that are grouped far apart, > 0.6 × clypeus width (Fig. 1). Parabomis male palps differ from those of the other African genera in having both a ventral apophysis and strongly developed retrolateral apophysis on the tibia. The marginally sclerotized atrium of Parabomis differs from Holopelus and Thomisops in the absence of a hood, and from Avelis in the absence of a median septum. Description. Female. Total length varies between 3.1–3.6 mm. Carapace cream to brown, sometimes tinted with green (Fig. 3), frequently mottled with white in eye region and on clypeus, with darker patches laterally, as seen in live specimens (Figs 1–3). Carapace as wide as long, anteriorly only slightly narrower than posteriorly; thoracic region high and convex, with distinct posterior declivity; declivity edge with semicircular carina with row of polyp-like tubercles, each bearing a short club-shaped seta; integument granulose, with scattered small tubercles, bearing very short, thick, flat-lying setae, more numerous laterally (very distinct in P. pilosus sp. nov. (Fig. 1)). Eyes small; both eye rows recurved, with AER shorter than PER and strongly recurved; lateral eyes on small tubercles, with posterior tubercle slightly larger than anterior tubercle; AME closer to ALE than to each other; ALE same size as AME; PME closer to PLE than to each; PME slightly smaller than PLE; MOQ much wider than long, slightly wider anteriorly than posteriorly. Clypeus distinct, strongly sloping, protruding slightly over cheliceral base; edge curved. Chelicerae flattened anteriorly; promargin bearing densely packed bristle-like setae. Labium longer than wide. Sternum cream to dark; narrow-oval, prolonged posteriorly to form an obtuse point between legs IV; integument granulose. Legs same color as carapace, varying from cream to pale brown, in some species mottled with white, with articulation areas of each leg segment frequently with distinct white rims; legs short and thick; patellae

162 · Zootaxa 4899 (1) © 2020 Magnolia Press Dippenaar-Schoeman & Foord nearly as long as metatarsi; tibiae longer than metatarsi; macro-setae absent, legs bearing short hair-like setae; tarsi with numerous teeth on paired claw; metatarsi and tarsi in some species with strong erect setae ventrally. Abdomen creamish white to brown, with darker hue in dorsal depressions (Figs 1–3); rounded, wider than long; shape varying from ball-shaped (when swollen with food or eggs) (Fig. 17) to slightly flattened, with lateral and transverse striae sometimes forming deep depressions; striae bearing small tubercles, each with translucent club-shaped seta; abdomen closely pressed against cephalothorax, fitting into posterior declivity; ventral region pale. Epigyne with marginally sclerotized atrium, of which shape varies from sub-oval, triangular to bell-shaped; spermathecae simple, longer than wide, bilobed sac-like structures, short, simple copulatory ducts, with intromittent orifices uncovered and situated in anterior part.

FIGURES 1–3. Habitus of live Parabomis spp. 1–2 P. pilosus sp. nov., female; 3 P. martini Lessert, 1919, female. 1–2 courtesy of J. Wilkinson; 3 courtesy of L. Wiese.

Male: Total length varies between 1.7–2.03 mm. Males resemble females but differ as follows: smaller in size; carapace and legs darker than female, usually uniformly brown; abdomen dorsum darker, shield-like in appearance, white laterally; dorsal depressions dark; legs slightly more slender than females; tibiae, metatarsi and tarsi bearing long dense setae ventrally. Palp: femur and patella not modified; tibia with two apophyses, VTA smaller than RTA, both fused; RTA about same length as tibia, apex pointed; shape varying between species; cymbium as long as wide, tutaculum absent; tegulum round, lacking apophyses; embolus filiform at tip, originating at 9h00 position. Distribution. Parabomis is endemic to the Afrotropical Region and was previously known from Bioko Island (Fernando Poo), Cameroon, Eritrea, Equatorial Guinea (Spanish Guinea), Namibia and Tanzania. New records: Bot- swana, Ghana, Guinea, Kenya, Malawi, Nigeria, Rwanda, South Africa, Zanzibar and Zimbabwe.

Key to the species of Parabomis

1 Males ...... 2 - Females...... 7 2 Male palp with RTA divided (Fig. 9)...... P. levanderi - RTA single, not divided ...... 3 3 RTA apex blunt (Fig. 23); carapace bearing numerous club-shaped setae...... P. pilosus sp. nov. - RTA apex pointed (Fig.18); carapace with few club-shaped setae...... 4 4 RTA apex directed laterad (Fig. 12)…...... 5 - RTA apex directed dorsad (Fig. 18)...... 6 5 RTA apex sharply pointed (Fig. 12); abdomen profile round...... P. martini - RTA base broad, beak-like (Fig. 28); abdomen profile hump-shaped...... P. wandae sp. nov. 6 RTA apex sharply pointed (Fig. 18)...... P. megae sp. nov. - RTA apex claw-like (Fig. 6)...... P. elsae sp. nov. 7 Atrium triangular-shaped (Fig. 13)...... P. martini - Atrium round to oval or bell-shaped ...... 8 8 Carapace with numerous club-shaped setae (Fig. 22); atrium bell-shaped, anterior edges of atrium thickened (Fig. 24) ...... P. pilosus sp. nov. - Carapace with few scattered club-shaped setae, atrium not bell-shaped...... 9 9 Atrium egg-shaped, ventral border thickened (Fig. 29)...... P. wandae sp. nov. - Atrium elongate-oval, lateral border thickened...... 10 10 Carapace and legs strongly mottled (Fig. 5)...... P. elsae sp. nov. - Carapace and legs uniform (Fig. 17) ...... P. megae sp. nov.

Etymology. Named after Elsa van Niekerk (Pretoria South Africa), the graphic artist at Agricultural Research Coun- cil (ARC)—Plant Health and Protection, in recognition of her support for spider research at the ARC. Diagnosis. Males and females of this species are recognized by their mottled bodies and copulatory organs (Figs 4, 5). Parabomis elsae sp. nov. females resemble those of P. megae sp. nov. in the shape of the epigyne, but the colour differs by being strongly mottled in P. elsae sp. nov. In males of both species the RTA apex is directed dorsad, but in P. elsae sp. nov. the RTA apex is claw-like (Fig. 6). Description. Holotype male. Measurements: TL 1.8; CL 0.85; CW 0.95. Legs: I 1.42 (0.51, 0.12, 0.30, 0.27, 0.22) II; 1.42 (0.63, 0.21, 0.28, 0.20, 0.10); III 1.08 (0.34, 0.12, 0.32, 0.20, 0.10); IV 1.32 (0.47, 0.15, 0.32, 0.19, 0.19). Carapace fawn, with 3 faint yellowish bands, one medially other 2 radiating from PLE to posterior edge (Fig. 4); darker patches laterally; integument granulose, with scattered small tubercles, with scattered short, thick, flat-lying setae, more numerous laterally; carapace slightly wider than long. Chelicerae brown; mouthparts paler brown. Sternum pale brown; integument granulose. Legs brown, same as in carapace; femora and tibiae slightly darker; articulation area of joints with strong white rim. Abdomen dorsum shield-like; with dark brown patches, with paler transverse patches in between, sides grey, speckled with white. Male palp with RTA tip claw-like curving upwards; VTA small fused with RTA (Fig. 6). Female. Measurements: TL 3.36; CL 1.36; CW 1.36. Legs: I 1.93 (0.68, 0.29, 0.43, 0.25, 0.28); II 1.84 (0.60, 0.27, 0.44, 0.25, 0.28); III 1.23 (0.42, 0.24, 0.26, 0.12, 0.19); IV 1.74 (0.56, 0.46, 0.29, 0.31, 0.12). Resembles male. Carapace fawn, with strong mottled appearance; 3 faint yellowish bands, one medially other two radiating; darker patches laterally; clypeus and eye area with three infused medial bands; clypeus paler (Fig. 5); carapace as wide as long; integument granulose, with scattered small tubercles. Chelicerae pale yellow, mouthparts yellowish brown. Sternum yellowish brown; integument granulose. Legs brown, strongly mottled with white, base of setae brown giving legs spotted appearance; articulation area of joints with strong white rim. Abdomen dorsum creamish white, base of abdominal setae dark, giving abdomen strong spotted appearance, especially anteriorly. Epigyne atrium wider than long, egg-shaped, with lateral edges slightly thickened (Fig. 7); intromittent orifices uncovered and situated in anterior part; spermatheca with two lobes (Fig. 8). Type material. Holotype ♂: SOUTH AFRICA: Gauteng Province: Roodeplaat Dam Nature Reserve (West), 17 km NE of Pretoria, 28º22’S, 25º37’E, leg. A.S. Dippenaar-Schoeman, 5.XI.1981 (beating Vachelia trees) (NCA 82/492). Paratypes: SOUTH AFRICA: Gauteng Province, Roodeplaat Dam Nature Reserve (West), 17 km NE of Preto- ria, 28º22’S, 25º37’E, leg. A.S. Dippenaar-Dippenaar, 27.XI.1979 (beating flowering Acacia trees, spider collected from yellow flowers), 1♀ (NCA 80/277); Same locality, leg. A.S. Dippenaar-Schoeman, 17.XII.1981 (sweeping grass), 1♂ (NCA 81/1091); Same locality, leg. National Collection of Arachnida staff, 22.IX.1988 (beating Acacia trees), 1♂, 2 imm. (NCA 92/607). Other material examined. SOUTH AFRICA: Gauteng Province: Enkeldoorn, 25°24’S, 28°40’E, leg. A.S. Dippenaar-Schoeman, 6.VIII.1974 (sweeping grass), 4imm. (NCA 76/298); Roodeplaatdam Nature Reserve, 28º22’S, 25º37’E, leg. M. Stiller, 29.IX.1980 (sweeping grass), 1imm. (NCA 1992/608); Same locality, leg. M. Stiller, 13.VIII.1980 (sweeping grass), 1imm. (NCA 1981/862); Same locality, leg. A.S. Dippenaar-Schoeman, 27.XI.1980 (sweeping grass), 2imm. (NCA 1999/34). KwaZulu-Natal Province: iSimangaliso Wetland Park, Hell’s Gate, 28º01.2’S, 32º21.6’E, leg. J. Esterhuizen, 1.XI.2004 (blue trap), 1♂ (NCA 2020/547); Same locality data, 3.VIII.2008 (blue trap), 1♂ (NCA 2020/548); Same locality data, 20.IX.2004 (blue trap), 2♂, 1imm. (NCA 2020/549); iSimangaliso Wetland Park, False Bay Park, 27º58.8’S 32º21.72’E, leg. M. Hamer, 6.XI.2004 (tree beating), 1♂, 1imm. (NCA 2006/58). Limpopo Province: Rust de Winter, 25°13’S, 28°29’E, leg. A.S. Dippenaar- Schoeman, 30.IV.1981 (beating grass), 1♀, 1imm. (NCA 99/35); Polokwane Nature Reserve, 23º58’S, 29º28’E, leg. T. Khoza & M. Modiba, 3.II.2006 (riverine and sweet thorn ticket, tree beating), 1♂, 1♀ (NCA 2008/2204). Mpumalanga Province: Loskopdam Nature Reserve, 25°46’S, 29°23’E, leg. A.S. Dippenaar-Schoeman, 2.IV.1973 (sweeping grass), 1imm. (NCA 2009/455). Distribution. South Africa: Gauteng, Limpopo, KwaZulu-Natal, and Mpumalanga Provinces (Fig. 32). Biology/Habitat. Specimens have been mainly collected beating flowering Vachellia trees and sweeping grass in the Savanna Biome. Adults were sampled from September to December.

Parabomis levanderii Kulczyński, 1901: 3, 30, figs 25–27 (Holotype ♂: ERITREA: Ghinda, 15°27’N, 39°05’E, leg. K.M. Levander, FMNH—examined).

Diagnosis. Males resemble those of P. martini in colour and the shield-like abdomen, but in P. levanderi the dorsum has two longitudinal series of black spots dorsally in depressions, which are absent in P. levanderi. Parabomis levanderi is the only male where the palp has the RTA two-pronged, with the one lobe longer and slightly hooked (Fig. 9). Description. Holotype male. Measurements: TL 1.9; CL 1.1; CW 1.1. Legs: I 2.17 (0.65, 0.35, 0.47, 0.36, 0.34); II 2.14 (0.65, 0.34, 0.46, 0.36, 0.33); III (missing); IV 1.64 (0.52, 0.29, 0.34, 0.23, 0.26). Carapace chestnut brown, with dark patches laterally; posterior declivity yellow; clypeus mottled, with paler longitudinal infusions; carapace as wide as long, with deep posterior declivity; edge of declivity bearing numerous small tubercles; integument with scattered tubercles, each bearing club-shaped seta. Mouthparts and chelicerae paler than carapace. Ster- num dark reddish brown, with narrow white margin, integument granulose. Legs: coxae dark; trochanters yellow; anterior femora black; patella yellow; anterior tibiae dark reddish brown; tarsi dark red; each segment distinctly rimmed with white. Abdomen: dorsum white, with shield-like dark area. Male palp with small fused VTA, RTA two-pronged (Fig. 9).

FIGURES 4–9. Parabomis elsae sp. nov. (4–8) and Parabomis levanderi Kulczyński, 1901 (9). 4 Male habitus, dorsal view; 5 Female habitus, dorsal view; 6, 9 Left male palp, ventral view; 7 Epigyne, ventral view; 8 Epigyne, dorsal view. Scales: 1 mm.

Parabomis martini Lessert, 1919 Figures 3, 10–14

Parabomis martini Lessert, 1919: 116, pl. 2, figs 6–8, 15 (Lectotype ♂ and 1♀ paralectotype, here designated: TANZANIA: Kilimandjaro Region: Kibonoto (Kibognoto), 03°12’S, 37°07’E, MNHN—examined; 2♀ paralectotypes: KENYA: Mt Meru, Ngare na Nyuki [now Nanyuki], 03°05’S, 39°51’E, MGNH—examined); Millot 1942: 33, fig. 15A–B. Parabomis anabensis Lawrence, 1928: 246, pl. 22, fig. 29 (Holotype ♀: NAMIBIA: Kaoko Otavi, 18°18’S, 13°41’E, found on tree Grewia monticola, SAM 7035—examined); 1♀ paratype: NAMIBIA: Anabib, 18º10’S, 12º31’E, SAM 6690—examined). syn nov.

FIGURES 10–14. Parabomis martini Lessert, 1919. 10 Male habitus, dorsal view; 11 Female habitus, dorsal view; 12 Left male palp, ventral view; 13 Epigyne, ventral view; 14 Epigyne, dorsal view. Scales: 1 mm.

166 · Zootaxa 4899 (1) © 2020 Magnolia Press Dippenaar-Schoeman & Foord Diagnosis. Males resemble those of P. wandae sp. nov., with the RTA apex directed laterad in both species. They differ by the RTA in P. wandae sp. nov. having a broad base and beak-like form (Fig. 28), compared to sharply pointed tip of P. martini. The males also differ in the profile of the abdomen, which is round in P. martini and hump- shaped in P. wandae sp. nov. In the females, the atrium is subtriangular (Fig. 13) in P. martini and egg-shaped in P. wandae sp. nov. Remark. Parabomis anabensis Lawrence, 1928 from Namibia is known only from a female, which resembles P. martini closely in all aspects and is here regarded as its junior synonym. Description. Female. Measurements: TL 3.6; CL 1.3; CW 1.5. Legs: I 1.91 (0.68, 0.29, 0.42, 0.25, 0.27); II 1.84 (0.61, 0.28, 0.43, 0.24, 0.28); III 1. 21 (0.42, 0.22, 0.26, 0.12, 0.19); IV 1.72 (0.55, 0.46, 0.28, 0.31, 0.12). Cara- pace fawn, with clypeus and eye region strongly infused with white; sometimes with green hue (Fig. 3); carapace lateral sides slightly darker; carapace wider than long; chelicerae, mouthparts yellowish white. Eyes as in genus description. Sternum yellowish. Legs fawn to brown, infused with white; femora darker, especially legs II and III; articulation area of joints with strong white rim. Abdomen fawn to white with faint spots; depressions on dorsum bearing two brownish triangular indentations (Fig. 11). Epigyne with atrium subtriangular (Fig. 13); spermathecae with two lobes, ventral lobe with folds (Fig. 14); intromittent orifices uncovered and situated in anterior part. Male. Measurements: TL 1.7; CL 0.9; CW 1.0. Legs: I 1.43 (0.51, 0.13, 0.30, 0.27, 0.22); II 1.43(0.63, 0.22, 0.28, 0.20, 0.10); III 1.09 (0.34, 0.12, 0.32, 0.21, 0.10); IV 1.32 (0.47, 0.15, 0.32, 0.19, 0.19). Male smaller than the female and darker in colour (Fig. 10). Carapace dark brown, with darker patches laterally; chelicerae, mouthparts, sternum dark to yellowish brown. Sternum dark to yellowish brown, granulose. Legs brown, with coxae and femora darker; articulation area of joints with strong white rim. Abdomen dorsum shield-like, white with brown hue, with two longitudinal series of black spots dorsally in depressions; venter creamish white. Palp with VTA small and triangular, RTA with broad base, apex sharply pointed, directed laterad (Fig.12). Other material examined. SOUTH AFRICA: Eastern Cape Province: Farm Gretna, 6 km SW of Graham- stown, 31°30’S, 26°38’E, leg. P. Croeser, 7.XI.1978 (on flower Pterocelastrus tricuspidatus), 1♂ (NCA 92/604); Alexandria, 46 km SSW of Grahamstown, 33°43’S, 26°22’E, leg. P. Croeser, 21.X.1978 (coastal dune forest, in undergrowth), 1♀ (NCA 92/606); Grahamstown, 33°18’S, 26°31’E, leg. P. Croeser, 20.X.1979, 1♂ (NCA 2009/261); Hankey, Baviaanskloof, 33°39’S, 24°27’E, leg. L. Lotz, 26.II.2008 (beating grass), 1♀ (NMBA 11531); East Lon- don, Pineapple Research Station, 33°01’S, 27°55’E, leg. A.S. Dippenaar-Schoeman, 7.XII.1977 (sweeping undergrowth), 1♂ (NCA 97/480); Prentjiesberg, 31°12’S, 28°17’E, leg. M. Hamer, 10.XII.2005 (sweeping grass), 1♂ (NCA 2007/750); Thyspunt, 12 km WNW of Cape St Francis, 34º17’S, 24º43’E, leg. Linda Wiese, 11.II.2012 (sweeping), 1♀ (NCA 2013/1147); Same locality data, 1♀ (NCA 2012/5636). Gauteng Province: Roodeplaatdam Nature Reserve, 28º22’S, 25º37’E, leg. A.S. Dippenaar-Schoeman, 27.XI.1979 (beating trees), 5♀ (NCA 1980/276, 1980/292); Same locality, leg. E.A. Ueckermann 29.XI.1979 (beating Euclea crispa), 3♀, 1 imm. (NCA 1980/275, 1981/472); Same locality, leg. E. A. Ueckermann 27.XI.1979 (beating trees), 1♀ (NCA 1980/274, 1981/431); Wal- ter Sisulu National Botanical Gardens, 26º05’S, 27º50’E, leg. A. Leroy, 22.IX.2008 (in flower head), 1♀ (NCA 2009/183). KwaZulu-Natal Province: Tembe Elephant Park, 27º02’S, 32º25’E, leg. R. Harris, 22.XI.1989 (sweeping grass), 1♀ (NCA 1999/32); Umgeni Valley, 29º28’S, 30º16’E, leg. A. Leroy, 7.I.1997 (deep forest), 1♀ (NCA 1999/33); Dukuduku Forest, 28º22’S, 32º14’E, leg. M. Filmer 7.XI.1991 (forest), 1♀ (NCA 1991/662); Sodwana Bay, 27º31’S, 32º41’E, leg. C. Car, 19.v.1981 (dune forest), 1imm. (SAM/Aran 880 C329); Spioenkop Dam, 30 km SW Ladysmith, 28º41’S, 29º25’E, leg. C. & T. Griswold, 9. XII.1985 (sweeping mixed grassland), 1♀ (NMSA 23718). Limpopo Province: Nylsvley Nature Reserve, 24°39’S, 28°42’E, leg. G. Ferriera, 1.II.1976 (beating Vitex rehmanni tree), 1imm. (NCA 1978/470); Same locality, leg. G. Ferriera, 1.I.1977 (beating Dombeya sp. tree), 1♀ (NCA 1978/515); Mabula Private Game Reserve, 24°44’S, 27°55’E, leg. M. Braack, 17.XII.1989 (in flower head), 1♀ (NCA 1991/484); Waterberg, near Vaalwater, farm Klipfontein, 28°18’S, 24°08’E, leg. C. Car, 4.XII.1979, 1♂ (NMZ/A432); Same locality, leg. C.A. Car, 6.XII.1979 (on grass), 1♂ (NMZ/A 364); Weltevreden Nature Reserve, 24°39’S, 27°58’E, leg. H. Kilian, 16.VIII.2006, 1♀ (NCA 2009/184); Tshulu, 22°57’S, 30°58’E, leg. S. Foord, 19.II.2008 (branch beating), 1♀ (NCA 2009/530); Makalali Nature Reserve, 29°09’S, 30°42’E, leg. C. Whitmore, 1.XII.1999, 1♂ (NCA 2009/262). Mpumalanga Province: 2D Ranch near Loskopdam, 25°25’S, 29°21’E, leg. A. Leroy, 27.II.1994, 1♀ (NCA 2009/263). North West Province: Kgaswane Mountain Reserve, 25°46’S, 27°20’E, leg. S. Stiemie, 5.II.1980 (sweeping grass), 1♀ (NCA 1980/250); Same locality, leg. A.S. Dippenaar-Schoeman, 20.XI.1981 (sweeping grass), 1♂ (NCA 1996/302); Same locality, leg. E. Ueckermann, 11.XII.1979 (beating Buddleia saligna), 1♂ (NCA 1981/478); Same locality, leg. M.K.P. Meyer, 7.XI.1979 (beating Euclea crispa),

Revision of Parabomis Zootaxa 4899 (1) © 2020 Magnolia Press · 167 1♂ (NCA 1981/469); Farm Steelpoort, on Krugersdorp road, 25°45’S, 27°51’E, leg. A.S. Dippenaar-Schoeman, 12.XI.1976 (beating trees), 1♀ (NCA 1976/1965); Same locality, leg. A.S. Dippenaar-Schoeman, 25.XI.1982 (beating trees), 1♀, 1imm. (NCA 1992/605); Meyer farm, Magaliesberg, near Rustenburg, 25°47’S, 27°16’E, leg. J. Leroy, 7.XII.1997 (beating Celtis africana tree), 1♀ (NCA 2009/259); Zeerust, 25°33’S, 26°06’E, leg. N. Hahn, 2.IV.2010 (branch beating), 1♀ (NCA). GUINEA: Kindia, 10°09’N, 12°31’W, VII.1937, 1♀ (MNHN); Dalaba, 10°41’N, 12°15’W, VIII.1937, 1♀ (MNHN). MALAWI: Chintheche, 11°50’S, 34°13’E, leg. R. Jocqué, 3.II.1976, 1♀ (MRAC 148.050). NAMIBIA: Grootfontein, 19º35’S, 18º07’E, leg. R.W. Tucker, no date, 1♀ (SAM 5065). RWANDA: Ihema, 01°52’S, 30°46’E, leg. R. Jocqué, Nsengimara and Michiels, 8.XII.1985, 1♀ (MRAC 165.827); Same locality, 14.XI-XII.1985, 1♀ (MRAC 165.292). TANZANIA: Zanzibar (no exact locality), leg. C. Cooke, 1863, 1♀ (MCZ); Mkomazi Game Reserve, Camp Ibayi, 04°37’S, 38°04’E, leg. A. Russell-Smith, 1.XI.1996, 1♂ (NCA 1997/860). ZAMBIA: Choma, Wildlife Game Farm, Hunters Camp, 16°58’S, 26°36’E, leg. C. Haddad, 4.XII.2006 (beating short shrubs), 1♂ (NCA 2007/518). ZIMBABWE: Sibizini Dam, Doddieburn Ranch, 2129A4, leg. J. Minshull, 11.XII.1985, 1♀ (NMZ A4227). Distribution. Tanzania, Kenya, Namibia. New records: Guinea, Malawi, Rwanda, South Africa, Zambia and Zimbabwe (Fig. 31). Biology/Habitat. Parabomis martini has been collected from a variety of habitats in the Grassland and Savanna Biomes, ranging from grass, trees in thorny bushveld, and the undergrowth of coastal dune forest. Tree species that they have been collected from include Buddleia saligna, Euclea crispa, Dombeya sp., Vitex rehmanni and Ptero- celastrus tricuspidatus (Dippenaar-Schoeman et al., 2009). In Namibia, P. anabensis was collected from the tree Grewia monticola, and according to Lawrence (1928) the spider closely resembles its flowers.

Parabomis megae sp. nov. Figures 15–20

Etymology. Named in honour of Ms Meg Cumming (Harare, Zimbabwe), in recognition of her contribution to our knowledge of Zimbabwean spiders. Diagnosis. The species is distinguished from other congeners by the male palp having the RTA straight, with the apex directed dorsally, while directed laterad in others, except for P. elsae sp. nov. It differs from P. elsae sp. nov. by the sharply pointed tip of the apex (Fig. 18). Parabomis megae sp. nov. resembles the female of P. elsae sp. nov. in the shape of the epigyne, but the colour differs, being uniform in P. megae sp. nov. and strongly mottled in P. elsae sp. nov. (Fig. 19). Description. Male (holotype). Measurements: TL 1.90; CL 0.95; CW 1.03. Legs: I 1.44 (0.52, 0.13, 0.30, 0.27, 0.22); II 1.44 (0.63, 0.22, 0.28, 0.20, 0.11); III 1.09 (0.34, 0.12, 0.32, 0.21, 0.10); IV 1.32 (0.47, 0.15, 0.32, 0.19, 0.19). Carapace dark brown (Fig. 15); eyes circled with black; chelicerae dark brown; mouthparts yellowish brown; palp dark brown; carapace slightly wider than long; strongly granulated, bearing numerous small tubercles, each with translucent seta. Sternum brown; granulose. Eyes as for genus. Legs: coxae, trochanters and tibiae brown, with blackish hue; femora I-IV dark brown, with femora I and II usually darker than rest; patellae yellowish brown; tibiae, metatarsi and partly tarsi of legs I and II clothed with numerous fine erect setae. Abdomen dorsum white with brown hue, shield-like with scattered black spots. Male palp with RTA straight, tip directed dorsally and apex pointed (Fig. 18). Female. Measurements: TL 3.2; CL 1.1; CW 1.4. Legs: I 1.83 (0.67, 0.29, 0.42, 0.25, 0.27); II 1.84 (0.61, 0.28, 0.43, 0.24, 0.28); III 1.61 (0.41, 0.22, 0.26, 0.12, 0.70); IV 1.71 (0.55, 0.46, 0.27, 0.31, 0.12). Carapace fawn, in some specimens darker laterally (Figs 16-17); strongly granulated, bearing numerous small tubercles, each with translucent seta; eye region infused with white; eyes sometimes circled with black; chelicerae creamish white. Sternum granulated, bearing tubercles with seta. Legs creamish white, with distinct white bands at joints; femora frequently darker. Abdomen fawn to white, with scattered dark setae giving it spotted appearance; depressions on dorsum darker, bearing two brownish markings (Fig. 16). Epigyne with atrium oval (Fig. 19); spermathecae with short ducts (Fig. 20). Type material. Holotype ♂: ZIMBABWE: Doddieburn Dam, Doddieburn Ranch, 21°28’S, 29°24’E, leg. J. Minshull, 12.XII.1985 (NMZ/A4208a). Paratypes: ZIMBABWE: Doddieburn Dam, Doddieburn Ranch, 21°28’S, 29°24’E, leg. J. Minshull,

168 · Zootaxa 4899 (1) © 2020 Magnolia Press Dippenaar-Schoeman & Foord 12.XII.1985, 1♀, 3♂, 4imm. (NMZ/A4208b); Tshowe River rapids, Matetsi Safari Area, 18°31’S, 28°40’E, leg. J. Minshull, 6.XII.1988, 1♂ (NMZ A6916); Walmer Drive, Harare, 17°49’S, 31°05’E, leg. M. Cumming, 14.VI.2004 (spider ballooning), 1♀ (NCA 2012/327); Same locality data, leg. M. Cumming, 15.I.2004 (sweeping grass and herbs in garden), 1♀, 3imm. (NCA 2012/326); Same locality data, leg. M. Cumming, 1.III.2004, 1♀ (NCA 2009/ 258).

FIGURES 15–20. Parabomis megae sp. nov. 15 Male habitus, dorsal view; 16 Female habitus, dorsal view; 17 Female habitus, dorsal view; 18 Left palp, ventral view; 19 Epigyne, ventral view; 20 Epigyne, dorsal view. Scales: 1 mm.

Other material examined. ZIMBABWE: Humani Ranch, Store Camp, 20°29’S, 32°15’E, leg. Falcon Col- lege, 6.IV.1987, 1imm. (NMZ A 5882); Doddieburn Dam, Doddieburn Ranch, 21°28’S, 29°24’E, leg. E. Nyathi, 20.IV.1986, 2 imm. (NMZ/A4661); Same locality, leg. J. Minshull, 11.XII.1985, 1 imm. ♀ (NMZ 4227); Chelo Farm Homestead, 2029A3, leg. F. Nyathi, 5.X.1988, 5 imm. (NMZ 715), 4 imm. (NMZ/A7190); Namakukwe, Zambezi confluence, 17°51’S, 25°23’E, leg. Falcon College, 26.VIII.1986, 1 imm. (NMZ 4794); Sabi-Lundi confluence, Gomarezhou National Park, 21°23’S, 33°50’E, leg. J. Minshull, 21.IV.1985, 1 imm. (NMZ 3289); Tshowe River rapids, Matetsi Safari Area 18°31’S, 28°40’E, leg. J. Minshull, 4.XII.1988 (NMZ 6895); Katombora Camp- site, 17°50’S, 25°24’E, leg. Falcon College, 1.X.1986, 1 imm. (NMZ 4720), 1 imm. NMZ 5040, 1 imm NMZ 5079, 1 imm. NMZ 5086); 1.5 km NW of Kazuma Forestry camp, 1825B3, leg. Falcon College, 15.IV.1988, 2 imm. (NMZ/A 6704); Farm 2 km E. Glenmore, 2028C, leg. F.Nyathi, 28.IX.1988, 1 imm. (NMZ/A 7094); Sengwa Plot, leg. M. Cumming, 2.XII.2002 (beating Trichelias tree), 1♂ (NCA 2009/456). Distribution. Known only from Zimbabwe (Fig. 32). Biology/Habitat. Specimens were sampled sweeping grass and herbs. Most of the adult specimens were sampled from December to March.

Etymology. The name refers to the numerous setae on the carapace of the species. Diagnosis. The male of this species differs from all the other congeners by the blunt RTA (Fig. 23) vs. pointed in other species. In the female, the epigyne atrium is bell-shaped (Fig. 24) and contrasts with that of P. martini, which is subtriangular. Carapace and abdomen bearing rows of flat-lying translucent club-shaped setae that are longer and denser than in other species, especially on the carapace declivity (Fig. 22). Description. Male (holotype). TL 1.7; CL 0.9; CW 1.0. Legs: I 1.43 (0.51, 0.13, 0.30, 0.27, 0.22); II 1.43 (0.63, 0.22, 0.28, 0.20, 0.10); III 1.09 (0.34, 0.12, 0.32, 0.21, 0.10); IV 1.32 (0.47, 0.15, 0.32, 0.19, 0.19). Carapace dark brown, granular, with numerous white club-shaped setae (Fig. 21); carapace edge white; eyes circled with black; chelicerae dark brown; mouthparts yellowish brown; carapace wider than long. Sternum orange brown, granulose, bearing numerous strong flat-lying translucent setae. Legs with coxae, trochanters and tibiae brown, with white articulations. Abdomen broad, dorsum dark brown, shield-like, with pale patches; granular. Palp: RTA not pointed but with blunt broad apex (Fig. 23); VTA absent.

FIGURES 21–25. Parabomis pilosus sp. nov. 21 Male habitus, dorsal view; 22 Female habitus, dorsal view; 23 Left palp, ventral view; 24 Epigyne, ventral view; 25 Epigyne, dorsal view. Scales: 1 mm.

Female. TL 2.6; CL 1.05; CW 1.3; CH 1.4; AW 2.1; AL 2.1. Legs: Legs: I 1.88 (0.65, 0.29, 0.42, 0.25, 0.27); II 1.81 (0.60, 0.26, 0.43, 0.24, 0.28); III 1.20 (0.42, 0.21, 0.26, 0.12, 0.19); IV 1.7 (0.55, 0.46, 0.26, 0.31, 0.12). Cara- pace fawn with clypeus and eye region infused with brown, lateral edges darker brown; chelicerae and mouthparts yellowish white; posterior declivity clothed with numerous white flat-lying club-shaped setae (Figs 1-2, 22). Ster-

170 · Zootaxa 4899 (1) © 2020 Magnolia Press Dippenaar-Schoeman & Foord num yellowish white. Legs fawn to brown infused with white; femora darker, especially legs II and III; articulate area of each leg segment with white rim. Abdomen fawn to white, with faint spots; depressions on abdomen darker, bearing two brownish triangular to round patches. Epigyne: atrium bell-shaped (Fig. 24); spermathecae oval, spermathecal ducts short, glandular lobes anteriorly of spermathecae (Fig. 25). Type material. Holotype ♂: BOTSWANA: Okavango Delta, Shakawe Fishing Camp, 18°26’S, 21°54’E, leg. J. van Niekerk, 1.XII.2001 (active searching) (NCA 2006/868). Paratypes: BOTSWANA: Okavango Delta, Xugana Island, 130 km NNW of Maun, 19°04’S, 23°03’E, leg. B.H. Lamoral, 17–25.XI.1980, 1♀ (NMSA 23719); Okavango Delta, 30 km N of Lechwee Camp, approx. between 18°40’S–19°00’S and 23°00’E–23°45’E, leg. B. Lamoral, 17-20.XI.1979, 1♂ (NMSA); Okavango Delta, N of Lechwee Camp and Khwai River Lodge, between 18°40’S–19°00’S and 23°00’E–23°45’E, leg. B. Lamoral, 18- 20.XI.1979, 2♂ (NMSA); Okavango Delta, near Shakawe, Lesideng Research Camp, 18°25’S, 21°53’E, leg. R. Lyle, 10-20.XII.2006 (beating shrubs in riverine forest), 1♀ (NCA 2020/550). Other material examined. BOTSWANA: Okavango Delta, Xugana Island, 130 km NNW of Maun, 19°04’S, 23°03’E, leg. F. Nyathi, 17-25.XI.1980, 1♀ (NMZ/A 70941). SOUTH AFRICA: Limpopo Province: Tshipise, Farm Alicedale (citrus), 22°42’S, 30°14’E, leg. J. Wilkinson, 12.IV.2016 (on tree in silk retreat, see Figs 1-2), 1♀ (SANSA Virtual Museum). Distribution. Known from Botswana and northern South Africa (Fig. 32). Biology/Habitat. Specimens were sampled by sweeping grass and herbs. In Tshipise, a female was sampled from a silk retreat made between leaves.

Parabomis wandae sp. nov. Figures 26–30

Etymology. This species is named in honour of Prof Wanda Wesołowska (Wrocław, Poland), in recognition of her significant contributions to the taxonomy of African and Palaearctic Salticidae. Diagnosis. Male of this species differ from congeners in the shape of the RTA that is almost as long as the width of the bulb, with the tip directed laterad (Fig. 28). The RTA base is broad and the apex is beak-like, in contrast to the sharply pointed apex of P. martini. The abdomen has a hump that is more distinct in males, while in P. martini the abdomen profile is round. The atrium of the female epigyne is egg-shaped, as wide as long, with the ventral edge thickened (Fig. 29), and the spermathecal ducts are long and curved (Fig. 30). Description. Holotype male. Measurements: TL 1.90; CL 0.91; CW 0.95. Legs: I 1.45 (0.52, 0.12, 0.31, 0.28, 0.22); II 1.45 (0.63, 0.23, 0.29, 0.20, 0.10); III 1.08 (0.34, 0.12, 0.32, 0.20, 0.10); IV 1.31 (0.46, 0.15, 0.32, 0.19, 0.19). Carapace: dark brown, darker laterally, strongly granulated, bearing numerous small tubercles, each with translucent seta (Fig. 26); eyes circled with black; chelicerae brown; mouthparts yellowish brown. Sternum brown, granulate. Legs dark; coxae, trochanters and tibiae brown, with blackish hue; femora I–IV dark brown, with femora I–II usually darker than rest; patellae yellowish brown; all legs with distinct white bands at joints distinct when viewed from below. Abdomen dorsum with brown scutum, with scattered black patches; in lateral view with distinct hump; white laterally and pale below. Palp with large beak-like RTA (Fig. 28); tip directed laterad with small VTA. Female. Measurements: TL 2.26; CL 0.97; CW 1.10. Legs: I 1.90 (0.67, 0.29, 0.42, 0.25, 0.27), II 1.82 (0.60, 0.27, 0.43, 0.24, 0.28); III 1.21 (0.42, 0.22, 0.26, 0.12, 0.19); IV 1.42 (0.55, 0.16, 0.28, 0.31, 0.12). Resembles the male but larger in size and paler in colour (Fig. 27). Carapace fawn to brown, with dark patch laterally; eyes circled with black; eye region infused with white; chelicerae fawn; integument strongly granulated, bearing numerous small tubercles, each with translucent seta. Sternum granulated, bearing tubercles with setae. Legs same colour as prosoma; white with distinct white bands at joints; femora darker. Abdomen white with dark patches, scattered dark setae giving it spotted appearance; abdomen with hump less distinct than in male. Epigyne with sub-oval atrium, slightly wider than long, edges slightly thickened (Fig. 29); spermathecae slightly longer than wide, with long curving spermathecal ducts (Fig. 30). Type material. Holotype ♂: GHANA: Kakum Forest, 05°21’N, 01°23’W, leg. R. Jocqué et al., 17.XI.2005 (fogging secondary forest) (MRAC 218.012). Paratypes: GHANA: Kakum Forest, °21’N, 01°23’W, 21.XI.2005 (fogging primary forest), leg. R. Jocqué et

Revision of Parabomis Zootaxa 4899 (1) © 2020 Magnolia Press · 171 al., 1♀ 2♂ (MRAC 218052); Same locality data, leg. R. Jocqué, et al., 15.XI.2005 (fogging secondary forest), 1♂ (MRAC 217.977). KENYA: Kakamega Forest, 00°22’N, 34°50’E, leg. W. Freund, 10.IX.2002 (canopy fogging), 1♀ (ZFMK AR133). Other material examined. GHANA: Kakum Forest, 05°21’N, 01°23’W, leg. R. Jocqué et al., 22.XI.2005 (fogging secondary forest), 1♂ (MRAC 218.058). KENYA: Kakamega Forest, 00°22’N, 34°50’E, leg. W. Freund, 10.IX.2002 (canopy fogging), 1♂ (ZFMK AR129), 1♂ (ZFMK AR051), 1♂ (ZFMK AR052), 1♂ (ZFMK AR053), 1♂ (ZFMK AR054), 1♂ (ZFMK AR132). IVORY COAST: Appouesso, FC Bossematié station 5, 06°36’N, 03°27’W, leg. R. Jocqué & L. Baert, 15.X.1993 (sweeping), 1♂ (MRAC 205.437); Bettie, forest close to Mabi, leg. R. Jocqué, 3.XI.1993 (sweeping forest edge), 1♀ (MRAC 177.635). RWANDA: Ibanda Makera, 02°20’S, 29°39’E, 1.X.1993 (fogging montane forest), 1♀, 4♂ (ZFMK, thom 12).

FIGURES 26–30. Parabomis wandae sp. nov. 26 Male habitus, dorsal view; 27 Female habitus, dorsal view; 28 Left palp, ventral view; 29 Epigyne, ventral view; 30 Epigyne, dorsal view. Scales: 1 mm.

Distribution. Known from Ghana, Kenya, Rwanda and Ivory Coast (Fig. 32). Biology/Habitat. Collected from primary and secondary forest while fogging the canopy of trees. One male was collected in the Ivory Coast while sweeping vegetation. Adults were sampled from September to November.

172 · Zootaxa 4899 (1) © 2020 Magnolia Press Dippenaar-Schoeman & Foord FIGURES 31–32. 31 Records of the previously known Parabomis species, P. levanderi Kulczyński, 1901 (red circles) and P. martini Lessert, 1919 (blue triangles); 32 Records of new Parabomis species in the Afrotropical Region.

Discussion

The Bominae genus from the Afrotropical region, Felsina is represented by type species, Felsina granulum Simon, 1895, described from Senegal and Sierra-Leone. Specimens from two bottles labelled as Felsina granulum in the MNHN were examined. The one bottle contains two females from Kharane, Senegal (MNHN 14111) but both specimens were immature. The second bottle (MNHN 14377), without locality data, contains 16 specimens but all of them are juveniles. These might have been the specimens reported from Sierra Leone. Simon (1907) mentioned specimens from Cameroon but no other material of the species was found in the museum. Based on the juveniles, Felsina and Parabomis might be synonymous, but the lack of adult material of Felsina means that no final decision can be taken at this stage.

Acknowledgements

We thank the curators of the various museums for the loan of material that made this study possible, as well as An- nette van den Berg and Robin Lyle (both Pretoria, South Africa) of the National Collection of Arachnida of the Ag- ricultural Research Council for their support in preparing the manuscript; the photographers Linda Wiese (Jeffrey’s Bay, South Africa) and John Wilkinson (Tshipise, South Africa) for images of the living spiders. We acknowledge support from the NRF Incentive Funding and the Agricultural Research Council.

References

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