Research Article |
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Corresponding author: Charles R. Haddad ( haddadcr@ufs.ac.za ) Academic editor: Galina N. Azarkina
© 2025 Charles R. Haddad.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Haddad CR (2025) And they just keep coming: four new genera of dark sac spiders from southern Africa (Araneae, Trachelidae). African Invertebrates 66(1): 19-64. https://doi.org/10.3897/AfrInvertebr.66.139299
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As part of ongoing revisions of the Afrotropical Trachelidae, four new genera are described from southern Africa: Foordana gen. nov., with F. distincta sp. nov. from South Africa (Eastern Cape, Free State, KwaZulu-Natal and Western Cape) as the type species, F. flavipoda sp. nov. from the Free State, F. kasouga sp. nov. from the Eastern Cape and KwaZulu-Natal, and a fourth undescribed species from Zimbabwe; the monotypic Mushimane gen. nov., with M. tswibilinki sp. nov. from KwaZulu-Natal as the type species; Namaquella gen. nov., with N. arida sp. nov. from the Northern Cape as the type species and N. samanthae sp. nov. from the Western Cape; and Rukuluk gen. nov. from South Africa, with R. gramineus sp. nov. from the Northern Cape as the type species and a second undescribed species from KwaZulu-Natal known only from juveniles.
Grass, new species, tree canopies
The last two decades have seen a massive resurgence in taxonomic work on dark sac spiders, Trachelidae, with more than 68% of the genera (17) and 43% of the global species richness (124 spp.) described during this period (
As continuous sampling has been undertaken in South Africa during the last three decades as part of the South African National Survey of Arachnida (
The material examined in this study is deposited in the following collections (curators in parentheses):
BMNH British Museum of Natural History, London, U.K. (J. Beccaloni).
NCA National Collection of Arachnida, ARC–Plant Health and Protection, Pretoria, South Africa (R. Lyle).
NMBA National Museum, Bloemfontein, South Africa (J. Neethling).
All specimens studied for descriptions and measurements were observed in 70% ethanol using a Nikon SMZ800 stereomicroscope. The epigynes of one or more female paratypes were dissected with 0-size insect pins and then cleared overnight by maceration in a pancreatin solution (
All measurements are presented in millimetres (mm) and are provided for the type specimens indicated in the descriptions. Eye arrangements are described for the anterior and dorsal views of the anterior and posterior eye rows, respectively. Leg measurements are given as the sequence from the femur to the tarsus, and the total. The following abbreviations are used in the text:
Ac aciniform gland spigot(s)
AER anterior eye row
AL abdomen length
ALE anterior lateral eye(s)
AME anterior median eye(s)
AW abdomen width
CL carapace length
CW carapace width
Cy cylindrical gland spigot(s)
dRTA dorsal retrolateral tibial apophysis
FL fovea length
MOQ median ocular quadrangle [l - length; AW - anterior width; PW - posterior width]
mAmp minor ampullate gland spigot(s)
MAmp major ampullate gland spigot(s)
Nu nubbin
PER posterior eye row
Pi piriform gland spigot(s)
PLE posterior lateral eye(s)
plv prolateral ventral
PME posterior median eye(s)
rlv retrolateral ventral
RTA retrolateral tibial apophysis
SL sternum length
ST I primary spermatheca
ST II secondary spermatheca
SW sternum width
Ta tartipore
TL total length
vRTA ventral retrolateral tibial apophysis
vt ventral terminal
For the type species of Foordana gen. nov., Mushimane gen. nov., Namaquella gen. nov. and Rukuluk gen. nov., material for scanning electron microscopy was dehydrated overnight in 100% ethanol, critical point dried in liquid carbon dioxide and then mounted onto aluminium stubs using double-sided tape. Thereafter, the samples were sputter coated three times with iridium for two minutes each in an argon chamber and then studied using a JEOL JSM-IT200 scanning electron microscope at 5 kV.
Various somatic characters of males and females of the type species of each new genus, as well as the dorsal habitus of most species, were taken using a Nikon D5-L3 camera system attached to a Nikon SMZ800 stereomicroscope. Depending on the structure, between five and 30 digital images were taken and stacked using the CombineZM imaging software (http://www.hadleyweb.pwp.blueyonder.co.uk) to increase the depth of field. For Mushimane tswibilinki sp. nov. and Rukuluk gramineus sp. nov., the cleared female epigynes were temporarily mounted on slides, stained with crystal blue, examined under a Nikon Eclipse 80i compound microscope and imaged with an attached Nikon DS-Fi2 camera. A series of images was taken and stacked as described above. The distribution map was created using the online mapping software SimpleMappr (
The relationships of the four new genera described here were evaluated using mitochondrial cytochrome oxidase c subunit I (COI) sequence data. Representative species of each genus of Afrotropical Trachelidae currently available on the Barcode of Life Data system (BOLD;
The material included in this COI analysis (Table
Summary of the species and South African sampling localities of specimens sequenced for the cytochrome c oxidase subunit 1 (COI) gene included in the phylogenetic analysis. Moggridgea loistata (Migidae) was used as the outgroup to root the tree.
| Species | Locality | Process ID | Catalog Num | COI-5P Seq. Length |
|---|---|---|---|---|
| Moggridgea loistata ♀ | Betty’s Bay | SPIZA1821-24 | NMBA18901 | 658 bp |
| Afroceto africana ♀ | Namaqua National Park | SPIZA613-21 | NCA 2021/214 | 658 bp |
| Afroceto martini ♂ | Ndumo Game Reserve | SPIZA391-19 | NCA 2019/584 | 658 bp |
| Afroceto plana ♀ | Ndumo Game Reserve | SPIZA388-19 | NCA 2019/582 | 658 bp |
| Afroceto sp. 1 ♂ | Nigramoep Slow Living Gues Farm | SPIZA905-21 | NCA 2021/526 | 658 bp |
| Afroceto sp. 2 ♂ | Akkerendam Nature Reserve | SPIZA1173-21 | NCA 2021/947 | 658 bp |
| Capobula infima ♂ | Marloth Nature Reserve | SPIZA2119-24 | NMBA 19184 | 658 bp |
| Capobula montana ♂ | Bankfontein Farm | SPIZA402-19 | NCA 2019/887 | 658 bp |
| Capobula neethlingi ♂ | Jonkershoek Nature Reserve | SPIZA1805-24 | NMBA 18895 | 658 bp |
| Coronarachne denticulata ♂ | Ndumo Game Reserve | SPIZA427-19 | NCA 2019/592 | 618 bp |
| Falcaranea gladius ♀ | Ndumo Game Reserve | SPIZA425-19 | NCA 2019/593 | 658 bp |
| Falcaranea maputensis ♂ | Ndumo Game Reserve | SPIZA422-19 | NCA 2019/758 | 658 bp |
| Foordana distincta ♀ | Queenstown | SPIZA1276-21 | NCA 2021/1044 | 658 bp |
| Fuchiba aquilonia ♀ | Tembe Elephant Park | SPIZA1603-23 | NMBA 18813 | 657 bp |
| Fuchiba capensis ♂ | Garcia Nature Reserve | SPIZA2257-24 | NMBA 19264 | 658 bp |
| Fuchibotulus haddadi ♀ | Golden Gate Highlands National Park | SPIZA652-21 | NCA 2021/244 | 658 bp |
| Fuchibotulus kigelia ♂ | Bankfontein Farm | SPIZA652-21 | NCA 2021/244 | 658 bp |
| Jocquestus schenkeli ♂ | Ndumo Game Reserve | SPIZA399-19 | NCA 2019/581 | 658 bp |
| Mushimane tswibilinki ♀ | Ndumo Game Reserve | SPIZA423-19 | NCA 2019/757 | 621 bp |
| Mushimane tswibilinki ♂ | Ndumo Game Reserve | SPIZA424-19 | NCA 2019/757 | 658 bp |
| Namaquella arida ♂ | Akkerendam Nature Reserve | SPIZA931-21 | NCA 2021/229 | 658 bp |
| Namaquella arida ♀ | Akkerendam Nature Reserve | SPIZA932-21 | NCA 2021/229 | 658 bp |
| Orthobula radiata ♂ | Vernon Crookes Nature Reserve | SPIZA1399-21 | NCA 2021/1202 | 614 bp |
| Patelloceto secutor ♂ | Ndumo Game Reserve | SPIZA405-19 | NCA 2019/590 | 658 bp |
| Planochelas haddadi ♀ | Ndumo Game Reserve | SPIZA430-19 | NCA 2019/999 | 596 bp |
| Poachelas montanus ♀ | Kogelberg Biosphere Reserve | SPIZA1975-24 | NMBA 18989 | 658 bp |
| Poachelas striatus ♀ | Platberg Nature Reserve | SPIZA365-19 | NCA 2019/422 | 658 bp |
| Spinotrachelas sp. 1 ♀ | Marloth Nature Reserve | SPIZA2120-24 | NMBA 19185 | 658 bp |
| Thysanina absolvo ♂ | Bloemfontein | SPIZA637-21 | NCA 2021/233 | 658 bp |
| Trachelas canariensis ♂ | uKhahlamba Drakensberg Mountains | SPIZA1349-21 | NCA 2021/1106 | 658 bp |
| Trachelas pusillus ♂ | Tembe Elephant Park | SPIZA1600-23 | NMBA 18810 | 658 bp |
| Rukuluk gramineus ♂ | Witsand Nature Reserve | SPIZA1505-23 | NMBA 18661 | 658 bp |
| Rukuluk sp. 1 Tembe imm. | Tembe Elephant Park | SPIZA410-19 | NCA 2019/760 | 658 bp |
| Rukuluk sp. 1 Tembe imm. | Tembe Elephant Park | SPIZA411-19 | NCA 2019/760 | 658 bp |
| Thysanina gracilis ♂ | Richtersveld National Park | SPIZA875-21 | NCA 2021/491 | 658 bp |
| Thysanina transversa ♂ | Karkkloof Canopy Tours | SPIZA1381-21 | NCA 2021/1135 | 658 bp |
The phylogenetic analysis was performed on the BOLD platform using the “Sequence analysis” tool, using the Kimura 2 Parameter distance model, Neighbour-joining algorithm and a minimum overall overlap of 200 bp between sequences. The sequences were aligned using Muscle (
Trachelidae Simon, 1897
Foordana distincta sp. nov.
Foordana gen. nov. superficially represent Afroceto by their size and Thysanina Simon, 1910 sensu stricto by the lack of leg spines, but can be recognized from the former by the heavily scopulate tibiae, metatarsi and tarsi of the anterior legs (Figs
Small spiders, 4.72–5.60 mm in length; carapace bright orange to deep red-brown; carapace oval, broadest at coxae II, gradually narrowed towards eye region (Figs
General habitus of live Foordana distincta sp. nov., female from Komani, Eastern Cape (A–C), Namaquella arida sp. nov., male from Akkerendam Nature Reserve, Northern Cape (D–F), and Rukuluk gramineus sp. nov., female from Witsand Nature Reserve, Northern Cape (G–I). A, D, G Dorsal view B, E, H Anterior view C, F Lateral view I Dorsolateral view. Photos by Ruan Booysen.
Digital microscope photographs of somatic morphology of Foordana distincta sp. nov., male (A–E) and female (F–J). A, F habitus, dorsal view B, G same, lateral view C, H same, ventral view D, I leg I, prolateral view E, J metatarsus and tarsus IV. Scale bars: 1.0 mm (A–C, F–H); 0.5 mm (D, E, I, J).
Scanning electron micrographs of Foordana distincta sp. nov., female A carapace, dorsal view B same, surface texture and setae C eye region, dorsal view D chelicerae, arrows indicating promarginal teeth E mouthparts F detail of endite serrula G sternum H patellar indentation, leg IV I same, detail of lyriform organ.
Scanning electron micrographs of Foordana distincta sp. nov., male A tibia I, ventral view B same, detail of cusps and scopulae C metatarsus I, ventral view D same, detail of cusps and scopulae E tarsus I, ventral view F same, detail of cusps and scopulae G–I detail of cusp structure on the tibia (G), metatarsus (H) and tarsus (I), arrows indicating triad of pores at base of cusp on proximal side.
Scanning electron micrographs of Foordana distincta sp. nov., female A tibia I, ventral view B metatarsus I, ventral view C same, detail of scopulate setae, arrows indicating ovoid pores between scopulate setae D same, detail of ovoid pores E metatarsus IV, lateral view F tarsus I, ventral view G same, lateral view, arrow indicating position of tarsal organ H tarsus IV, detail of claw tuft and claws I tarsus I, base of dorsal trichobothrium J same, detail of tarsal organ.
Scanning electron micrographs of Foordana distincta sp. nov., female A spinnerets, general view B anterior lateral spinnerets C posterior median spinnerets D posterior lateral spinnerets. Abbreviations: Ac – aciniform gland spigot(s); Cy – cylindrical gland spigot(s); mAmp – minor ampullate gland spigot; MAmp – major ampullate gland spigot; Pi – piriform gland spigot(s); Ta – tartipore.
Foordana distincta sp. nov., male (A–C) and female (D, E). A schematic representation of cusp arrangement on legs I and II B palp, ventral view C same, retrolateral view D epigyne, ventral view E same, dorsal view. Abbreviations: CD – connecting duct; Cd – copulatory duct; CO – copulatory opening; dRTA – dorsal retrolateral tibial apophysis; EM – embolus; FD – fertilization duct; SD – sperm duct; ST I – primary spermatheca; ST II – secondary spermatheca; vRTA – ventral retrolateral tibial apophysis. Scale bars: 0.25 mm.
Digital microscope photographs of somatic morphology of Foordana flavipoda sp. nov., male (A, E) and female (B, I), F. kasouga sp. nov., male (C, F), F. distincta sp. nov., male (D) and female (G, H), and undescribed Foordana sp. from Zimbabwe, female (J). A–C habitus, dorsal view D–F palps, ventral view G, J uncleared epigynes, ventral view H, I cleared epigynes, ventral view. Scale bars: 1.0 mm (A–C), 0.5 mm (D–F), 0.25 mm (G–J).
The genus name is a patronym in honour of the late Stefan Foord, in recognition of his distinguished career and contribution to the development of African arachnology, with the suffix alluding to its superficial resemblance to Cetonana. Gender feminine.
Foordana distincta sp. nov., F. flavipoda sp. nov., F. kasouga sp. nov. and an undescribed species from Zimbabwe.
Holotype. South Africa • ♂; Free State Province; Clocolan, Amahelo-ho-Spitskop; 28°48.561'S, 27°39.255'E; 17 Mar. 2010; C. Haddad leg.; base of grass tussocks; NCA 2010/349.
Paratypes. South Africa • 4 ♀; together with holotype • Eastern Cape Province; 1 ♀; Queenstown/Komani district, Farm Arphen; 31°51.546'S, 26°34.112'E; 1160 m a.s.l.; 4 Apr. 2021; C. Haddad leg.; hand collecting; NCA 2021/1044 • 1 ♂ 2 ♀; Amatola Mountains, Hogsback, Amatole Forestry Company; 32°33.727'S, 26°54.924'E; 1460 m a.s.l.; 23 Mar. 2013; C. Haddad leg.; active search, grass tussocks and fynbos in grassland; NCA 2014/321 • Free State Province: 1 ♂ 2 ♀; Clocolan, Amahelo-ho-Spitskop; 28°48'S, 27°39'E; 9 Mar. 2007; C. Haddad leg.; Rhus litter; NCA 2007/1334 • 2 ♀; same collection data as for preceding; 8 Mar. 2007; dense grass; NCA 2007/1336 • 1 ♀; same collection data as for preceding; grassy litter near dam; NCA 2008/2897 • 1 ♀; same collection data as for preceding; Eucalyptus litter; NCA 2007/1335 • 1 ♀; Clocolan, Amahelo-ho-Spitskop; 28°48.561'S, 27°39.255'E; 2010; A. Jones leg.; in garden and around house; NCA 2010/328 • 3 ♀; Wepener district, Farm Dereham, 29°52.671'S, 27°04.456'E; 1520 m a.s.l.; 31 Mar. 2024; C. Haddad leg.; hand collecting in grassland; NMBA 19614 • KwaZulu-Natal Province; 3 ♀; Pietermaritzburg, Town Bush; 29°33'S, 30°21'E; 15 Apr. 1976; F. Wanless & A. Russell-Smith leg.; ground layer in grassland; BMNH • Western Cape Province: 1 ♀; Houw Hoek, Houw Hoek Inn; -34.2046, 19.1537; 275 m a.s.l.; 25 Nov. 2021; C. Haddad leg.; hand collecting in fynbos; NMBA 18796.
The male of this species shares with F. flavipoda sp. nov. a simple curved embolus in ventral view (Figs
Foordana flavipoda sp. nov., male (A, B) and female (C, D). A palp, ventral view B same, retrolateral view C epigyne, ventral view D same, dorsal view. Abbreviations: CD – connecting duct; Cd – copulatory duct; CO – copulatory opening; dRTA – dorsal retrolateral tibial apophysis; EM – embolus; FD – fertilization duct; SD – sperm duct; ST I – primary spermatheca; ST II – secondary spermatheca; vRTA – ventral retrolateral tibial apophysis. Scale bars: 0.25 mm.
Male (holotype, Amahelo-ho-Spitskop, NCA 2010/349): Measurements: CL 2.36, CW 2.03, AL 2.77, AW 1.87, TL 5.15, FL 0.14, SL 1.33, SW 1.15, AME-AME 0.07, AME-ALE 0.04, ALE-ALE 0.41, PME-PME 0.11, PME-PLE 0.19, PLE-PLE 0.68, MOQ: AW 0.35, PW 0.36, L 0.40. Length of leg segments: I 2.05 + 0.95 + 1.61 + 1.22 + 0.81 = 6.64; II 1.80 + 0.88 + 1.35 + 1.12 + 0.76 = 5.91; III 1.32 + 0.63 + 0.88 + 1.07 + 0.43 = 4.33; IV 1.92 + 0.85 + 1.60 + 1.78 + 0.63 = 6.78. Carapace deep red-brown, with four pairs of mediolateral mottled markings, corresponding to palps and first three pairs of legs (Fig.
Female (paratype, Amahelo-ho-Spitskop, NCA 2010/349): Measurements: CL 2.48, CW 2.05, AL 3.58, AW 2.35, TL 5.60, FL 0.24, SL 1.32, SW 1.05, AME-AME 0.06, AME-ALE 0.03, ALE-ALE 0.37, PME-PME 0.13, PME-PLE 0.14, PLE-PLE 0.65, MOQ: AW 0.31, PW 0.40, L 0.35. Length of leg segments: I 2.12 + 1.02 + 1.52 + 1.20 + 0.86 = 6.72; II 1.90 + 0.92 + 1.35 + 1.02 + 0.80 = 5.99; III 1.47 + 0.77 + 1.00 + 1.12 + 0.55 = 4.91; IV 2.25 + 0.98 + 1.80 + 1.95 + 0.70 = 7.68. Carapace deep orange-brown, with three paired mediolateral mottled grey markings and mottled patch in front of fovea (Fig.
The species name is the Latin for “distinct, clear”, referring to the well-defined abdominal markings of this species.
Widespread across the southern half of South Africa (Fig.
Distribution of Foordana distincta sp. nov. (black circles), F. flavipoda sp. nov. (white circles), F. kasouga (black triangles), Foordana sp. Zimbabwe (white triangle), Mushimane tswibilinki sp. nov. (black square), Namaquella arida sp. nov. (white square), N. samanthae sp. nov. (black star), Rukuluk gramineus sp. nov. (white star) and Rukuluk sp. Tembe (white pentagon) in southern Africa.
Holotype. South Africa • ♂; Free State Province; Bloemfontein district, Farm Hopefield; 28°51.683'S, 26°09.788'E; 1275 m a.s.l.; 28 Oct. 2001; C. Haddad leg.; Eucalyptus bark; NMBA 19613.
Paratypes. 2 ♀; together with holotype.
South Africa • 1 ♀; Free State Province; Harrismith, Platberg Nature Reserve, Platberg, near cross; 28°15.136'S, 29°09.958'E; 2215 m a.s.l.; 11 Mar. 2006; C. Haddad leg.; under rocks, mountainside; NCA 2006/863.
The male of this species can be recognised from congeners by the lack of ventral leg cusps on the anterior legs and the short broad dRTA (Fig.
Male (holotype, Hopefield, NMBA 19613): Measurements: CL 2.27, CW 1.91, AL 2.45, AW 1.77, TL 4.72, FL 0.18, SL 1.20, SW 1.03, AME-AME 0.07, AME-ALE 0.04, ALE-ALE 0.38, PME-PME 0.14, PME-PLE 0.13, PLE-PLE 0.61, MOQ: AW 0.30, PW 0.37, L 0.33. Length of leg segments: I 1.90 + 1.02 + 1.53 + 1.20 + 0.62 = 6.27; II 1.57 + 0.78 + 1.56 + 1.03 + 0.60 = 5.54; III 1.21 + 0.64 + 0.88 + 0.98 + 0.45 = 4.16; IV 1.65 + 0.85 + 1.43 + 1.55 + 0.60 = 6.08. Carapace orange-brown, sightly paler on posterior slope (Fig.
Female (paratype, Hopefield, NMBA 19613): Measurements: CL 1.80, CW 1.58, AL 2.10, AW 1.52, TL 4.05, FL 0.13, SL 1.12, SW 0.98, AME-AME 0.06, AME-ALE 0.02, ALE-ALE 0.30, PME-PME 0.11, PME-PLE 0.11, PLE-PLE 0.51 MOQ: AW 0.28, PW 0.32, L 0.29. Length of leg segments: I 1.50 + 0.84 + 1.04 + 0.98 + 0.58 = 4.94; II 1.25 + 0.71 + 0.91 + 0.80 + 0.47 = 4.14; III 0.92 + 0.54 + 0.60 + 0.80 + 0.48 = 3.34; IV 1.75 + 0.80 + 1.42 + 1.52 + 0.59 = 6.08. Carapace yellow-brown, paler on posterior slope (Fig.
This species is a contraction of the Latin flavus (yellow) and poda (legs), referring to the colouration of the legs of both sexes.
Only known from two localities in central South African grasslands (Fig.
The placement of this species in Foordana gen. nov. is tentative, as the lack of leg cusps in males, position of the sperm duct in the male palpal tegulum and the presence of a small hood in the female epigyne differ from the type species. Furthermore, the condition of the type material is not ideal, as the specimens are somewhat damaged and may have lost some of their colouration. As such, finding fresh material and incorporating sequence data into future analyses will be an essential step in resolving its placement.
Holotype. South Africa • ♂; Eastern Cape Province; Kasouga, 16 km WSW of Port Alfred; 33°39'S, 25°45'E; Jan. 1940; J. Omer-Cooper leg.;
Paratype. 1 ♂ ; South Africa • KwaZulu-Natal Province; 75 km WSW of Estcourt, Cathedral Peak Forest Station, Meteorology Station, Little Berg; 28°59'S, 29°11'E; 1860 m a.s.l.; 13–31 Dec. 1979; S. & J. Peck leg.; pan trap;
South Africa • 8 ♂ 1 ♀; KwaZulu-Natal Province; 75 km WSW of Estcourt, Cathedral Peak Forest Station, Meteorology Station, Little Berg; 28°59'S, 29°11'E; 1860 m a.s.l.; 21–31 Dec. 1979; S. & J. Peck leg.; veld, Malaise trap;
The male of this species can be recognised by the distinctive loop of the embolus and the far longer dRTA, which extends to more than a quarter of the tegulum length. In comparison, the emboli of F. distincta sp. nov. and F. flavipoda sp. nov. are only slightly curved, and the dRTA is clearly larger than the vRTA in lateral view (compare Figs
Male (holotype, Kasouga,
The species is named after the type locality; noun in apposition.
Only known from single localities in the Eastern Cape and KwaZulu-Natal provinces, South Africa (Fig.
At the proof stage I received a loan of Trachelidae from the
Zimbabwe • 1 ♀; Matabeleland; Bulawayo, Hillside; 20°10'S, 28°38'E; 28 Feb. 1999; M. FitzPatrick leg.; pitfall traps;
A clearly different female to the two species described above was discovered in the
Mushimane tswibilinki sp. nov.
This genus includes possibly the smallest known trachelids, with adults only ~1.8 mm in total length. Both sexes superficially resemble Falcaranea Haddad & Lyle, 2024 but are smaller, lighter in colour and have very different genitalic morphology. Males can be distinguished by from all other trachelids by the distinctive spiralling loop in the distal section of the sperm duct of the palp, close to the base of the embolus, which is short and spike-like (Fig.
Small spiders, 1.79–1.84 mm in length; carapace creamy-yellow, yellow-brown in eye region (Fig.
Digital microscope photographs of somatic morphology of Mushimane tswibilinki sp. nov. male (A–E) and female (F–J). A, F habitus, dorsal view B, G same, lateral view C, H same, ventral view D, I leg I, prolateral view E, J metatarsus and tarsus IV. Scale bars: 1.0 mm (A–C, F–H); 0.5 mm (D, E, I, J).
Scanning electron micrographs of Mushimane tswibilinki sp. nov. female A carapace, dorsolateral view B eye region, anterodorsal view C tarsus I, dorsolateral view, arrow indicating position of tarsal organ D same, enlargement of claws, dorsolateral view, arrow indicating tarsal organ E same, paired dorsal trichobothria F same, enlargement of tarsal organ G anterior lateral spinneret H posterior median spinnerets I posterior lateral spinneret. Abbreviations: Ac: aciniform gland spigot(s); Cy: cylindrical gland spigot(s); mAmp: minor ampullate gland spigot; MAmp: major ampullate gland spigot; n: nubbin; Pi: piriform gland spigot(s).
Mushimane tswibilinki sp. nov., male (A, B) and female (C, D). A palp, ventral view B same, retrolateral view C epigyne, ventral view D same, dorsal view. Abbreviations: CD – connecting duct; Cd – copulatory duct; CO – copulatory opening; EM – embolus; FD – fertilization duct; RTA – retrolateral tibial apophysis; SD – sperm duct; ST I – primary spermatheca; ST II – secondary spermatheca. Scale bars: 0.1 mm.
The name is a noun in the indigenous African Sesotho language for “boy” or “little man”, referring to the diminutive size of the type species. Gender masculine.
Monotypic.
Holotype. South Africa • ♂; KwaZulu-Natal Province; Ndumo Game Reserve, Staff housing; 26°54.660'S, 32°17.930'E; 130 m a.s.l.; 5 Dec. 2018; C. Haddad, R. Booysen & J. Neethling leg.; canopy fogging, Commiphora neglecta; NCA 2019/757.
Paratype. 1 ♀; together with holotype.
1 ♀; Same data as types; S.E.M. preparations, epigyne cleared and retained preserved with types.
As for the genus diagnosis.
Male (holotype, Ndumo, NCA 2019/757): Measurements: CL 0.97, CW 0.75, AL 0.98, AW 0.59, TL 1.79, SL 0.56, SW 0.41, AME-AME 0.03, AME-ALE 0.01, ALE-ALE 0.16, PME-PME 0.06, PME-PLE 0.05, PLE-PLE 0.27, MOQ: AW 0.14, PW 0.18, L 0.16. Length of leg segments: I 0.67 + 0.35 + 0.48 + 0.47 + 0.27 = 2.24; II 0.54 + 0.30 + 0.39 + 0.40 + 0.25 = 1.88; III 0.40 + 0.24 + 0.21 + 0.29 + 0.16 = 1.30; IV 0.60 + 0.27 + 0.46 + 0.40 + 0.18 = 1.91. Carapace creamy-yellow, slightly darker in eye region (Fig.
Female (paratype, Ndumo, NCA 2019/757): Measurements: CL 0.83, CW 0.63, AL 1.27, AW 0.75, TL 1.84, SL 0.52, SW 0.37, AME-AME 0.02, AME-ALE < 0.01, ALE-ALE 0.14, PME-PME 0.05, PME-PLE 0.05, PLE-PLE 0.25, MOQ: AW 0.13, PW 0.17, L 0.16. Length of leg segments: I 0.54 + 0.30 + 0.33 + 0.34 + 0.25 = 1.76; II 0.46 + 0.26 + 0.31 + 0.30 + 0.24 = 1.57; III 0.38 + 0.21 + 0.25 + 0.27 + 0.16 = 1.27; IV 0.54 + 0.25 + 0.44 + 0.38 + 0.17 = 1.78. Carapace yellow-brown (Fig.
This species name is a derogatory slang term in the indigenous African Sesotho, mocking a man with a small penis, in reference to the short embolus of the type species.
Only known from the type locality in northeastern KwaZulu-Natal, South Africa (Fig.
Due to the limited number of specimens available, one female was sacrificed for scanning electron microscopy; its epigyne was cleared prior to preparation of the material for S.E.M., used for the illustrations, and has been retained in the same vials as the types. In addition, this female had an even more strongly ventrally curved abdomen than the paratype.
Namaquella arida sp. nov.
Both sexes of Namaquella gen. nov. resemble Poachelas Haddad & Lyle, 2008 and Rukuluk gen. nov., with a pale body to support a lifestyle associated with grasses, although the body proportions are less elongate than in these genera. Males of Namaquella gen. nov. can be recognized from these and other trachelid genera by the oval tegulum with a simple, slightly curved distal embolus directed retrodistally and the simple sperm duct and RTA (Figs
Small spiders, 2.18–3.52 mm in length; carapace creamy-yellow in N. arida sp. nov. to bright yellow-orange in N. samanthae sp. nov.; carapace oval, broadest near posterior of coxae II, gradually narrowed towards eye region (Figs
Digital microscope photographs of somatic morphology of Namaquella arida sp. nov., male (A–E) and female (F–J). A, F habitus, dorsal view B, G same, lateral view C, H same, ventral view D, I leg I, prolateral view, arrowheads in D indicating ventral cusps E, J metatarsus and tarsus IV. Scale bars: 1.0 mm (A–C, F–H); 0.5 mm (D, E, I, J).
Scanning electron micrographs of Namaquella arida sp. nov., subadult male A carapace, dorsal view B same, surface texture and setae C eye region, dorsal view D chelicerae, arrows indicating promarginal teeth E mouthparts F modified setae on posterior surface of paturon G detail of serrula H patellar indentation, leg IV I same, detail of lyriform organ.
Scanning electron micrographs of Namaquella arida sp. nov., subadult male A tibia I, lateral view B metatarsus I, lateral view C same, metatarsal stopper D metatarsus IV, metatarsal stopper and preening comb E tarsus I, lateral view, arrow indicating position of tarsal organ F same, detail of tarsal organ G same, detail of claw tuft and claws, ventrolateral view H same, distolateral view I same, base of dorsal trichobothrium.
Namaquella arida sp. nov., male (A–C) and female (D, E). A schematic representation of cusp arrangement on legs I and II B palp, ventral view C same, retrolateral view D epigyne, ventral view E same, dorsal view. Abbreviations: CD – connecting duct; Cd – copulatory duct; CO – copulatory opening; EM – embolus; FD – fertilization duct; RTA – retrolateral tibial apophysis; SD – sperm duct; ST I – primary spermatheca; ST II – secondary spermatheca. Scale bars: 0.1 mm.
The name is derived from the Namaqualand region in arid western South Africa, from which the type species originates. Gender feminine.
Two species, Namaquella arida sp. nov. and N. samanthae sp. nov.
Holotype. South Africa • ♂ ; Northern Cape Province; Calvinia, Akkerendam Nature Reserve; 31°24.425'S, 19°46.823'E; 1170 m a.s.l.; 18 Jan. 2021; C. Haddad & R. Booysen leg.; hand collecting, grass tussocks; NCA 2021/229.
Paratype. 2 ♀; together with holotype.
1 subadult ♂ 2 subadult ♀; same data as types; S.E.M. preparations.
The male of this species can be recognised by the orientation of the embolus, at approximately 45° to the longitudinal axis of the palp (approximately 30° in N. samanthae sp. nov.) and the shape of the RTA, which is rounded and more strongly bent distally, with the tip directed dorsally, whereas the tip is directed dorso-distally in N. samanthae sp. nov. (compare Figs
Male (holotype, Akkerendam, NCA 2021/229): Measurements: CL 1.45, CW 1.22, AL 1.60, AW 1.07, TL 3.02, SL 0.87, SW 0.64, AME-AME 0.03, AME-ALE 0.03, ALE-ALE 0.25, PME-PME 0.08, PME-PLE 0.10, PLE-PLE 0.44, MOQ: AW 0.20, PW 0.25, L 0.21. Length of leg segments: I 1.25 + 0.65 + 1.06 + 0.85 + 0.49 = 4.30; II 0.90 + 0.54 + 0.73 + 0.61 + 0.41 = 3.19; III 0.68 + 0.40 + 0.44 + 0.51 + 0.30 = 2.33; IV 0.98 + 0.48 + 0.81 + 0.82 + 0.37 = 3.46. Carapace creamy-yellow (Fig.
Female (paratype, Akkerendam, NCA 2021/229): Measurements: CL 1.67, CW 1.18, AL 1.96, AW 3.53, TL 2.18, SL 1.05, SW 0.76, AME-AME 0.06, AME-ALE 0.03, ALE-ALE 0.29, PME-PME 0.10, PME-PLE 0.13, PLE-PLE 0.52, MOQ: AW 0.21, PW 0.28, L 0.24. Length of leg segments: I 1.30 + 0.70 + 1.05 + 0.87 + 0.48 = 4.40; II 1.03 + 0.61 + 0.80 + 0.69 + 0.44 = 3.57; III 0.78 + 0.50 + 0.52 + 0.60 + 0.30 = 2.70; IV 1.25 + 0.65 + 1.08 + 1.10 + 0.42 = 4.50. Carapace creamy-yellow (Fig.
This species name refers to the arid Succulent Karoo environments that it was collected from; adjective.
Only known from the type locality (Fig.
Holotype. South Africa • ♂; South Africa; Western Cape Province; Fisherhaven, nr Hermanus; 34°21.430'S, 19°07.557'E; 24 Mar. 2008; S. Oliver leg.; walking on pathway during day; NCA 2008/573.
For detailed diagnosis, see N. arida sp. nov. above. Female unknown.
Male (holotype, Fisherhaven, NCA 2008/573): Measurements: CL 1.70, CW 1.43, AL 1.73, AW 1.17, TL 3.52, FL 0.12, SL 0.98, SW 0.79, AME-AME 0.06, AME-ALE 0.03, ALE-ALE ~0.27, PME-PME 0.11, PME-PLE 0.09, PLE-PLE 0.56 MOQ: AW 0.26, PW ~0.32, L 0.27. Length of leg segments: I 1.38 + 0.73 + 1.12 + 0.92 + 0.50 = 4.65; II 1.08 + 0.59 + 0.79 + 0.73 + 0.45 = 3.64; III 0.72 + 0.47 + 0.50 + 0.61 + 0.30 = 2.60; IV 1.15 + 0.57 + 0.91 + 0.98 + 0.40 = 4.01. Carapace deep yellow-brown, with faint grey mottling; surface finely wrinkled; fovea short, distinct, at ⅔ CL. AER slightly procurved, almost straight; clypeus height equal to approximately ½ AME diameter; AME and ALE subequal in size; AME separated by distance approximately ½ their diameter; AME separated from ALE by distance slightly less than ⅓ AME diameter; PER recurved, PME and PLE subequal; PME separated by distance approximately equal to their diameter; PME separated from PLE by distance equal to their diameter. Chelicerae deep yellow-brown, anterior surface covered with quite dense long, fine setae; promargin with three slightly separated teeth, retromargin with three adjacent teeth on common base; endites and labium orange. Sternum shield-shaped, yellow-brown, with orange-brown borders; surface finely wrinkled, covered with scattered short, fine setae. Abdomen elongate-oval, broadest in posterior half; creamy-yellow dorsally, with yellow dorsal scutum from ¼ AL to posterior; broad grey mottled stripe along midline, sides of abdomen also mottled grey, markings separated by pair of cream mediolateral strips; two pairs of distinct round sigilla, first and second pairs respectively just behind ¼ and ½ AL. Legs I to IV with yellow femora, remaining segments progressively darker to yellow-brown tarsi; cusps only present on metatarsi and tarsi I (Fig.
This species is named for my wife Samantha, who collected the holotype.
Only known from the type locality.
Rukuluk gramineus sp. nov.
Members of Rukuluk gen. nov. share with Namaquella gen. nov. and Poachelas the pale body with adaptations to grass-living but can be distinguished from the former by the very different genitalic morphology and more elongate body, and from the latter by the lack of paired spines ventrally on the anterior legs. Males of Rukuluk gen. nov. can be recognized from all other trachelid genera by the distinctive palpal morphology, particularly the presence of retrolateral grooves in the cymbium that correspond to the RTA and distal section of the embolus (Figs
Small spiders, 3.06–3.20 mm in length; carapace bright creamy-yellow to yellow-brown; carapace oval, broadest at posterior of coxae II, gradually narrowed towards eye region (Fig.
Digital microscope photographs of somatic morphology of Rukuluk gramineus sp. nov., male (A–E, K–N) and female (F–J, O). A, F habitus, dorsal view B, G same, lateral view C, H same, ventral view D, I leg I, prolateral view E, J metatarsus and tarsus IV K–N left male palp in prolateral (K), ventral (L), retrolateral (M) and dorsal (N) views; O epigyne, ventral view. Scale bars: 1.0 mm (A–C, F–H); 0.5 mm (D, E, I, J); 0.25 mm (K–O).
Scanning electron micrographs of Rukuluk gramineus sp. nov., female (A, C, D, F–I) and male (B, E). A carapace, dorsal view B eye region, anterior view C same, dorsal view D chelicerae, anterior view E distal end of chelicerae, ventral view F mouthparts, ventral view G detail of serrula H sternum, arrow indicating slit sensilla I same, enlarged.
Scanning electron micrographs of Rukuluk gramineus sp. nov., female A leg I, prolateral view B patellar indentation and lyriform organ, leg I C same, detail of lyriform organ D tibia I, prolateral view E same, detail of scopulate setae F metatarsus I, prolateral view G same, detail of scopulate setae H same, detail of metatarsal stopper I tarsus I, prolateral view J same, retrodistal view, black arrow indicating position of tarsal organ K same, enlargement of tarsal claws L same, tarsal claws in distal view.
Scanning electron micrographs of Rukuluk gramineus sp. nov., male (A–H) and female (I). A tibia I, ventral view B same, prolateral view C metatarsus I, ventral view D same, prolateral view, arrow indicating distal slit sensillium E tarsus I, ventral view F same, prolateral view G tibia II, lateral view, black arrows indicating ovoid sensory pores (H) I metatarsus IV, stopper and distal comb/brush.
Scanning electron micrographs of Rukuluk gramineus sp. nov., male (A, B, G, H) and female (C–F, I–L). A, B metatarsi I (A) and II (B), ventral cusps, arrows indicating triad of pores at base of cusp on proximal side C distal half of tarsus IV, seemingly lacking a tarsal organ D tarsus I, lateral trichobothrium base E tarsus IV, dorsal trichobothrium F, G tarsus I, tarsal organ H abdominal dorsum I ventral abdominal setae J anterior lateral spinneret K posterior median spinneret L posterior lateral spinneret. Abbreviations: Ac: aciniform gland spigot(s); Cy: cylindrical gland spigot(s); mAmp: minor ampullate gland spigot(s); MAmp: major ampullate gland spigot; Pi: piriform gland spigot(s).
Scanning electron micrographs of Rukuluk gramineus sp. nov., male (A–D) and female (E). A right palp, retrolateral view B left palp, ventral view C detail of dorsal tibial trichobothrium D palpal patella, enlargement of retrolateral lyriform organ E epigyne, ventral view. Abbreviations: LO – lyriform organ; RB – retrobasal cymbial groove; RD – retrodistal cymbial groove; Tr – trichobothrium.
Rukuluk gramineus male (A–C) and female (D, E). A schematic representation of cusp arrangement on legs I and II B palp, ventral view C same, retrolateral view D epigyne, ventral view E same, dorsal view F diagrammatic course of the insemination ducts. Abbreviations: AG – accessory glands; AT – atrium; Cd – copulatory duct; CO – copulatory opening; dRTA – dorsal retrolateral tibial apophysis; EM – embolus; FD – fertilization duct; SD – sperm duct; ST I – primary spermatheca; ST II – secondary spermatheca; vRTA – ventral retrolateral tibial apophysis. Scale bars: 0.25 mm.
The name is an arbitrary combination of letters. Gender feminine.
Holotype. South Africa • ♂; Northern Cape Province; Witsand Nature Reserve, Brulsand; 28°34.688'S, 22°27.769'E; 1210 m a.s.l.; 23 Mar. 2023; C. Haddad & R. Booysen leg.; grass tussocks on sand dunes; NMBA 18616.
Paratypes. South Africa • 2 ♀; together with holotype NMBA 18616 • 1 ♂ 1 ♀; Northern Cape Province; Witsand Nature Reserve, Viewing Point; 28°33.584'S, 22°29.568'E; 1225 m a.s.l.; 25 Mar. 2023; C. Haddad & R. Booysen leg; grass tussocks on sand dunes; NMBA 18636 • 1 ♀; Witsand Nature Reserve, Rest Camp; 28°33.773'S, 22°29.095'E; 1195 m a.s.l.; 25 Mar. 2023; C. Haddad & R. Booysen leg.; grass tussocks, woodland; NMBA 18661 • 1 ♀; same collection data as for preceding; MACN.
South Africa • 1 ♂ 2 ♀; Northern Cape Province; Witsand Nature Reserve, Rest Camp; 28°33.773'S, 22°29.095'E; 1195 m a.s.l.; 25 Mar. 2023; C. Haddad & R. Booysen leg.; grass tussocks, woodland; S.E.M. preparations.
As for the genus diagnosis.
Male (holotype, Witsand, NMBA 18616): Measurements: CL 1.53, CW 1.20, AL 1.72, AW 0.96, TL 3.06, FL 0.10, SL 1.36, SW 1.05, AME-AME 0.03, AME-ALE 0.02, ALE-ALE 0.23, PME-PME 0.08, PME-PLE 0.05, PLE-PLE 0.38, MOQ: AW 0.24, PW 0.26, L 0.25. Length of leg segments: I 1.30 + 0.62 + 1.13 + 0.90 + 0.45 = 4.40; II 1.07 + 0.51 + 0.88 + 0.77 + 0.42 = 3.65; III 0.70 + 0.41 + 0.50 + 0.63 + 0.33 = 2.57; IV 1.15 + 0.52 + 1.02 + 1.05 + 0.40 = 4.14. Carapace bright yellow-orange, with black mottled pentagonal marking in front of fovea (Fig.
Female (paratype, Witsand, NMBA 18661): Measurements: CL 1.27, CW 1.06, AL 1.93, AW 1.15, TL 3.20, FL 0.11, SL 1.40, SW 1.05, AME-AME 0.03, AME-ALE 0.02, ALE-ALE 0.21, PME-PME 0.08, PME-PLE 0.05, PLE-PLE 0.37, MOQ: AW 0.22, PW 0.27, L 0.23. Length of leg segments: I 1.12 + 0.57 + 0.92 + 0.70 + 0.40 = 3.71; II 0.95 + 0.50 + 0.70 + 0.61 + 0.36 = 3.12; III 0.62 + 0.38 + 0.48 + 0.55 + 0.29 = 2.32; IV 1.10 + 0.51 + 0.97 + 0.98 + 0.36 = 3.92. Carapace yellow, with faint black mottled marking in front of fovea (Fig.
This species name is the Latin word meaning “belonging to grass”, referring to the microhabitat that the species was collected from.
Only known from the type locality (Fig.
South Africa • 2 imm.; KwaZulu-Natal; Tembe Elephant Park, Viewing Tower; 26°02.227'S, 32°24.907'E; 115 m a.s.l.; 5 Dec. 2018; C. Haddad & R. Booysen leg.; base of grass tussocks; NCA 2019/760.
Aside from the type species, another species of Rukuluk only known from two juvenile specimens in eastern South Africa was collected. Based on DNA barcode data, both specimens were placed in a monophyletic group with R. gramineus sp. nov. (Fig.
The current contribution takes the diversity of Afrotropical trachelid genera and species to 20 and 107, respectively, making it the biogeographical realm with the richest diversity. It is not entirely surprising that most of these taxa have been described during the last two decades, as this family has become the focus of intensive taxonomic study during this period. Despite these advances, the phylogenetic relationships of the family remain poorly understood, although recent analyses based on the COI gene have provided some preliminary indications (
The analysis conducted in the present study restricted the terminals to a single representative of the described species in the SPIZA project on BOLD, with three terminals (two Afroceto Lyle & Haddad, 2010 and one Spinotrachelas Haddad, 2006) represented by undescribed species. In performing the analysis, one Foordana distincta sp. nov. individual (a paratype female from Houw Hoek in the Western Cape, SPIZA1583-23, NMBA 18796) consistently grouped together with Capobula infima (Simon, 1897), so was assumed to be a contaminated sequence and was removed from the final analysis. The holotype male of Namaquella samanthae sp. nov. (SPIZA461-19, NCA 2008/573) was submitted for sequencing but this was unsuccessful, so this species could not be included.
As in the previous analyses, Planochelas haddadi Khoza & Lyle, 2011 was placed as sister to all other Trachelidae (Fig.
This paper pays tribute to the life and career of the late Stefan Foord, who made invaluable contributions to the study of African spider biodiversity. Robin Lyle is thanked for the use of her unpublished illustrations of the male palp of Foordana kasouga and female epigyne of F. distincta. The collection curators are thanked for the loans of material that made this study possible. The reviewers, Chi Jin and Jan Bosselaers, and editor, Galina Azarkina, are thanked for their valuable inputs that helped improve the manuscript.
The author has declared that no competing interests exist.
No ethical statement was reported.
This work was funded through various grants from the National Research Foundation of South Africa (SABI # FA2005040700016, TTK #2008050500003, CFRR #95569, CFRR #112127, IFRR #95902 and FBIP #129108).
The author solely contributed to this work.
Charles R. Haddad https://orcid.org/0000-0002-2317-7760
All of the data that support the findings of this study are available in the main text or Supplementary Information.
The collecting data and specimen accessions of four new genera of Trachelidae from southern Africa
Data type: xlsx
Explanation note: table S1. Details of collecting data of four new genera of Trachelidae spiders from southern Africa.