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Research Article
Ekapa, a new genus of mygalomorph spiders (Araneae, Entypesidae) from South Africa
expand article infoDuniesky Ríos-Tamayo‡§, Robin Lyle|, Catherine L. Sole
‡ Fundación Miguel LIllo, Tucumán, Argentina
§ Unidad Ejecutora Lillo (CONICET–Fundación Miguel Lillo), Tucumán, Argentina
| Agricultural Research Council - Plant Health and Protection, Biosystematics, Pretoria, South Africa
¶ University of Pretoria, Pretoria, South Africa

Corresponding author: Duniesky Ríos-Tamayo ( riostamayo.dnk@gmail.com )

Academic editor: Bernhard A. Huber
© 2023 Duniesky Ríos-Tamayo, Robin Lyle, Catherine L. Sole.
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: Ríos-Tamayo D, Lyle R, Sole CL (2023) Ekapa, a new genus of mygalomorph spiders (Araneae, Entypesidae) from South Africa. African Invertebrates 64(1): 1-12. https://doi.org/10.3897/afrinvertebr.64.97018
ZooBank: urn:lsid:zoobank.org:pub:1BBCC36B-FB24-4D32-8332-F5B3528431B4
Open Access

Abstract

A new genus of mygalomorph spider Ekapa gen. nov., is described from South Africa. The new genus was formed to include the species Hermacha curvipes Purcell, 1902 and Hermacha nigra Tucker, 1917. These species are synonymized based on their somatic similarities, such as fovea shape, cheliceral teeth distribution, ocular pattern, presence of preening combs, and their close geographic distribution. The new genus proposed is distinguished by the presence of a projection in the retroventral side of the palpal tibia in males, together with a distinctive copulatory bulb shape; and in females by the shape of their spermathecae, with a high base and apical stalks that open into oval apical receptacles.

Keywords

new genus, new species, new synonymy, systematics, taxonomy

Introduction

The fauna of mygalomorph spiders from South Africa is poorly known and many of the current genera require revision (Dippenaar-Schoeman 2002; Ríos-Tamayo and Lyle 2020). Recently, taxonomic work on South African mygalomorph spiders has been focused on the newly described family Entypesidae Bond, Opatova & Hedin, 2020 (Opatova et al. 2020). Several regional revisions focused on the genera Lepthercus Purcell, 1902, Entypesa Simon, 1902 and Hermacha Simon, 1889. The genera Hermachola Hewitt, 1915 and Brachytheliscus Pocock, 1902 were revalidated. Lastly, a new genus Afropesa Zonstein & Ríos-Tamayo, 2021 (Ríos-Tamayo and Lyle 2020; Ríos-Tamayo et al. 2021; Zonstein and Ríos-Tamayo 2021) was described. Work relating to the genus Hermacha, evaluated the status of different species, re-describing many of them and describing three new species. Additionally, Hermacha curvipes Purcell, 1902 and Hermacha nigra Tucker, 1917 were categorized as species incertae sedis (Ríos-Tamayo et al. 2021).

In the present paper, we propose to synonymize the species H. curvipes and H. nigra, based on their morphological similarities observed through the examination of the type specimens and additional related material. Besides the similarities of the two species, the somatic characters and the morphology of the copulatory organs do not match the features of the genus Hermacha, and have not been observed in any other related genus. Thus, we propose the creation of a new genus, described here to accommodate these species.

Materials and methods

The material studied here is deposited in the following institutions (curators are noted in brackets): NM – Natal Museum Pietermaritzburg, KwaZulu-Natal, South Africa (Kirstin Williams); SAM – Iziko South African Museum, Cape Town, South Africa (Aisha Mayekiso) and NCA – National Collection of Arachnida, ARC – Plant Health Protection, Pretoria, South Africa (Petro Marais).

All measurements are given in millimeters and were taken from the left side of the specimen. Total length was measured without spinnerets. The photographs were taken using a high-resolution microscopy camera AxioCam MRc5 mounted on a Zeiss Axio Zoom V16 microscope. Extended focal range images were stacked using the ZEN module Z-stack software. Lengths of leg segments are always given in the same order: femur, patella, tibia, metatarsus, tarsus, and total. The male copulatory bulb and female genitalia were dissected using a microscalpel. The copulatory bulbs were photographed following the approaches of Goloboff (1995) and Ríos-Tamayo and Goloboff (2018). Spermathecae were clarified by immersion in clove oil, thus making the opaque glandular tissues that cover the structure transparent.

Abbreviation and notation for leg spines follow Goloboff and Platnick (1987) and Ríos-Tamayo and Goloboff (2018); describing the legs from I to IV and from proximal to distal (i.e., from femur to tarsus within each leg, from basal to apical within each segment); the sequence prolateral-dorsal-retrolateral-ventral (P-D-R-V) is always followed, using lowercase letters (p-d-r-v), when referring to spiniform setae instead of spines. The filiform trichobothria present in the metatarsi follow a pattern of alternating long and short trichobothria; the numbers of shorter trichobothria are given in parentheses, describing from proximal to distal (e.g. the notation (3)1(2)1 refers to three short trichobothria, followed by a long trichobothria, followed by 2 short ones, and a long one).

Abbreviations used in species descriptions are as follows: ALE, anterior lateral eyes; AME, anterior median eyes; D, dorsal; DB, dorsal basal; P, prolateral; PA, prolateral apical; PLE, posterior lateral eyes; PLS, posterior lateral spinnerets; PME, posterior median eyes; PMS, posterior median spinnerets; OQ, ocular quadrangle; R, retrolateral; RM retrolateral medial; RA, retrolateral apical; V, ventral; VB, ventral basal; VPOST, ventral posterior. In the notation of leg spines, an expression such as D–R, D–P indicates that the spines occupy the dorsal–posterior superior surfaces, or the dorsal–prolateral superior surfaces; 1:2 A, 3:4 B indicates that the spines or scopula referred to are on the apical half or basal three-fourths. In the description of the preening combs, II 2 or 3 PV indicates that the combs are on the second leg, composed by two or three setae and in prolateral-ventral position.

Taxonomy

Family Entypesidae Bond, Opatova & Hedin, 2020

Ekapa gen. nov.

https://zoobank.org/D93C9440-E3B3-4815-A4FC-C00F8DE56F22

Type species

Hermacha curvipes Purcell, 1902, by present designation.

Etymology

The new name “Ekapa” is taken from the type locality and means “Cape Town” in the Xhosa language. The gender is neutral.

Diagnosis

The new genus can be distinguished from all other related genera in the family Entypesidae by the presence of a retroventral projection in the male palpal tibia (absent in all other entypesid genera; Figs 2D, 4E). Males of Ekapa gen. nov. can also be distinguished from Hermachola and Brachytheliscus by the presence of retrolateral apical megaspines on tibia I (Figs 1F, 4H), similar to Afropesa and Hermacha. They can be differentiated from Afropesa by the male palpal bulb morphology which is piriform, with dorsal keels and strong flanges (Figs 2A–C, 4A–D) and from Hermacha by the presence of preening combs in both sexes. Ekapa gen. nov., differs from Lepthercus by the lack of a megaspine on a raised cuticular spur on tibia I of males. Females possess spermathecae with a high basal portion (fused or not) and stalks that open in oval apical receptacles (Fig. 5D, E).

Relationships

Given the morphological characters presented here, this new genus shows affinities with Afropesa, Hermacha and the Australian genus Stanwellia Rainbow & Pulleine, 1918 (currently belonging to the family Pycnothelidae), since they all present a retrolateral apical megaspine on tibia I. Nevertheless, Ekapa gen. nov. differs from them due to its characteristic retroventral projection on the male palpal tibia. Stanwellia spp, demonstrate a similar shape of the spermathecae (with a high base), but differ from Ekapa gen. nov. since they possess flexuous leg tarsi, and a straight thoracic fovea (Harvey et al. 2018).

Description

Cephalothorax and legs covered with short bristles; cephalothorax with dark setae on the margin, more abundant towards the posterior part. Thoracic fovea short, straight or slightly recurved. Eyes group broader than long, anterior row slightly procurved and posterior row recurved (Figs 1C, 3F, 5C); clypeus short. Abdomen covered with short black hairs, with a pattern of yellow chevrons (Fig. 3C). Sternum oval, covered with black hairs, with posterior sigilla well defined and separated from the sternal margin. Labium broad, with few cuspules (principally in females), labiosternal sigilla consisting of a pair of well-marked sigilla. Maxillae with strong cuspules on the posterior inner edge. Stout short bristles as a rastellum, chelicerae with bristles on dorsal and retrolateral face. Chelicerae with a continuous line of equidistant teeth on the promargin, with many small basal denticles (no equidistant line with a few basal denticles in Hermacha). Male intercheliceral tumescence well-marked and setose (Fig. 3E). Serrula absent in both sexes. Leg formula 4123; all legs sparsely covered with hairs. Legs with all tarsi aspinose. Tibia I of male with a megaspine, and with a series of spines, all of which are almost the same size (Figs 1F, 4H). Palpal tibia slightly swollen, with prolateral spines, ventrally with some rigid setae and a projection on the retroventral side (Figs 2D, 4E). Cymbium short without spines. Copulatory bulb piriform, flanged, with long dorsal keels and with a short slightly curved embolus (Figs 2A–C, 4A–D). Scopulae: on metatarsi I–II distributed in the entire segment; III sparse, apical; IV absent. Present on tarsi I–II, throughout each segment (dense in females), III–IV sparse. Trichobothria: filiform in two rows from basal to anterior; metatarsi with one straight line, tarsi with a narrow zigzag row along length. Metatarsal preening combs present in legs II–IV. Spermathecae: receptacles with a high base (fused or not), and with stalk that ends in the oval apical receptacle (Fig. 5D, E).

Figure 1. 

Ekapa curvipes (Purcell, 1902) comb. nov., male holotype (SAM-ENW-X006011) A cephalothorax, dorsal view B sternum, labium, maxillae, ventral view C ocular pattern, dorsal view D spinnerets, ventral view E–G tibia I E prolateral view F ventral view G retrolateral view.

Figure 2. 

Ekapa curvipes (Purcell, 1902) comb. nov., male holotype (SAM-ENW-X006011) A–C copulatory bulb A retrolateral view B ventral view C prolateral view D, E distal segments of right pedipalp D retrolateral view, arrow indicates the projection in the retroventral side E prolateral view.

Species included

Ekapa curvipes (Purcell, 1902), comb. nov.

Ekapa curvipes (Purcell, 1902), comb. nov.

Figs 1, 2, 3, 4, 5

Hermacha curvipes Purcell, 1902: 377; Tucker 1917: 106.

Hermacha nigra Tucker, 1917: 110; Hewitt 1919: 109. new synonymy.

Type material

Syntypes , 2 ♂♂: Hermacha curvipes (SAM-ENW-X006011, Fig. 1A–I): Western Cape, Simonstown (Waterfall) (34.18333°S, 18.43333°E), Jul 1899, Purcell, W. coll. Syntypes, 2 ♀♀: Hermacha nigra (SAM-ENW-X013899, Fig. 3A–F), Bergvliet Flats, Cape Peninsula (34.05°S, 18.45°E), Sept 1904, Purcell, W. coll.

Additional material examined

Western Cape, Cape Town, Aug 1965, collector not indicated, 1 ♂ (NM 16639); Bergvliet flats, Constantia, (34.05°S, 18.45°E), Oct 1902, collector not indicated, 1 ♀ (SAM-ENW-X012166), erroneously identified as Hermacha purcelli; Table Mountain, 1 Aug 2005, J. Pryke, C. coll., 3 ♂♂ (NCA 2008/770).

Synonymy

Purcell (1902), described Hermacha curvipes from Simonstown (Cape Town), based on two male specimens. Fifteen years later, Tucker (1917) described several female specimens from the same locality, as the unknown females of H. curvipes. Tucker (1917) gave no catalog number for these specimens, and it is therefore impossible to identify which specimens the author used in his description. In the same article, Tucker (1917: 110) described the species H. nigra Tucker, 1917, based on two females, from Bergvleit Flats, Cape Town which is 16 km from the type locality of H. curvipes. These two species have characters both similar and different to those found in the genus Hermacha (see Ríos-Tamayo et al. 2021) such as: fovea shape, cheliceral teeth distribution, ocular pattern, presence of preening combs and a relatively close geographical distance. The examination of additional male and female specimens (belonging to a new undescribed species) collected from Knysna Forest, with the same combination of characters, support our conclusion that the female of Hermacha nigra and the male of Hermacha curvipes are conspecific; with H. nigra as junior synonym.

Remarks

The holotype specimen (male, SAM-ENW-X006011) is not in the best condition; possesses a pallid coloration (seems to be discolored), with legs very soft (Figs 1A–G, 2A–E). To preserve the holotype specimen and prevent any damage resulting from manipulation of the specimen, descriptions were made based on the male specimen NCA 2008/770 from Table Mountain (about 25 km N from the type locality), 1 Aug 2005, J. Pryke, C. coll., 3 ♂♂, which shares all the characteristics observed in the holotype (Figs 3–F, 4A–I).

Figure 3. 

Ekapa curvipes, male (NCA 2008/770) A cephalothorax, dorsal view B sternum, labium, maxillae, ventral view C abdomen, dorsal view D spinnerets, lateral view E chelicerae, prolateral view, arrow indicate the intercheliceral tumescence F ocular pattern, dorsal view.

Diagnosis

As for the genus.

Description

Male (NCA 2008/770, Figs 3A–F, 4A–I): Total length: 14.74. Cephalothorax (Fig. 3A): length 7.09, width 5.36, with lateral dark marginal bristles, and dorsal pubescence. Cephalic region: length 4.27; clypeus short, with 6 bristles on the edge, seven bristles in front (the central bigger) and 4 on the back of the OQ. Fovea: width 0.69, recurved and divided. Eyes, diameters and interdistances (Fig. 3F): AME 1.08, ALE 2.09, PME 1.60, PLE 1.90, OQ length 3.83, 7.03 width; AME-ALE 0.57, PME-PLE 0.17, AME-PME 0.48, ALE-PLE 0.37, AME-AME 0.84, OMP-OMP 2.73. Chelicerae: length 3.38, width 1.94, with dark dorsal-anterior bristles, rastellum formed by setae; intercheliceral tumescence small, well-marked and with setae (Fig. 3E). Cheliceral furrow with 10 promarginal teeth and 20 mesobasal denticles. Labium: length 0.82, width 1.84. Maxillae: length 4.19, width 1.75, with 124 cuspules on inner corner, prolateral face curved, soft area small, with long uniformly distributed hairs. Sternum (Fig. 3B): length 6.96, maximum width 4.98. Abdomen (Fig. 3C): length 6.07, covered with small hairs and dark bristles. Spinnerets (Fig. 3D): PMS: length 2.28; PLS: length of basal:medial:apical segments 3.03:2.35:4.18. Lengths of legs and palp: I: 6.90, 4.09, 4.87, 4.97, 3.18, 24.01. II: 5.13, 2.96, 3.62, 3.74, 2.41, 17.86. III: 4.58, 2.53, 2.97, 4.08, 2.33, 16.49. IV: 5.73, 2.88, 4.13, 5.44, 2.59, 20.77; Palp: 7.39, 3.98, 5.04, —, 2.72, 19.13.

Figure 4. 

Ekapa curvipes, male (NCA 2008/770) A–D copulatory bulb A retrolateral view B ventral view C prolateral view D closeup of the tip, arrows indicate the two keels E, F distal segments of right pedipalp E retrolateral view, arrow indicates the projection in the retroventral side F prolateral view G–I tibia I G prolateral view H ventral view I retrolateral view.

Figure 5. 

Ekapa curvipes, female (SAM-ENW-X013899) A overall dorsal view B sternum, labium, maxillae, ventral view C ocular pattern, dorsal view D, E spermathecae D female from Bergvliet Flats, Western Cape (SAM-ENW-X013899) E female from same locality (SAM-ENW-X012166).

Chaetotaxy : Leg I: femur, 1-1-1 P, 1-1-1-1 D, 1-1 R; patella, 1-1 P; tibia, 1-1 P, 1 R (1/2 B), 2-1-2 V, and a strong apical spine (ventral posterior); metatarsus, 1-1 P, 1-2 V (1/2 A), strongly curved with a well-marked retrolateral elbow. Leg II: femur, 1-1-1 P, 1-1-1-1 D, 1-1-1-1 R; patella, 0; tibia, 1-1 P, 2-1-3 V; metatarsus, 1 P, 1-1 D, 1 RM, 2-2-2 V. Leg III: femur, 1-1-1 P, 1-1-1-1 D, 1-1-1 R; patella, 1-1 P, 1 R; tibia, 1-1-1 P, 1-1 D, 1-1 R, 2-2-3 V; metatarsus, 1-1-1 P, 1-1-1 D-P, 1-1-1 D-R, 2-2-3 V. Leg IV: femur, 1-1-1-1 P, 1-1-1-1 D, 1-1-1-1 R; patella, 1 R; tibia, 1-1 P, 1 DB, 1-1 R, 2-2-3 V; metatarsus, 1-1-1 P, 1-1-1 D-P, 1-0-1 D-R, 1-1-9-1 R, 2-1-1-3 V. Pedipalp: femur, 1 PA, 1-1-1 d; patella, 1 PA; tibia, 1-2-2 P, 1 D, 1 RA, developed retroventral projection (Fig. 4E); tarsus, 0. Copulatory bulb tapering, with two developed flags (center with a large spoon-like depression), with the distal portion truncate continuing with a short, slender and curved tip, with two strong posterior keels (Fig. 4A–D). Scopula: Metatarsi: I, dense, distributed throughout the segment (more abundant 1/2 A); II, dense, distributed throughout the segment; III, sparse apical, divided by setae; IV, 0. Tarsi: I—II, dense, uniformly distributed throughout the segment; III—IV, dense, uniformly distributed and divided by a wide band of setae (narrow on III). Trichobothria: Tibiae: I 11-12; II 11-10; III 10-11; IV 13-12. Metatarsi: I 1(5)1(3)1(3)1(4); II 1(5)2(4)1(2)1(5); III 1(5)1(3)1(2)1(1)1(4); IV 1(6)1(5)1(7). Tarsi: I-III 14; IV 15. Preening combs: III 3 RV-3 PV, IV 2 PV-3RV. Color: Entire body reddish-brown, first leg darker. Sternum and coxae lighter. All tarsi with ventral and laterals pallid, but not flexuous. Abdomen brown with dorsal chevrons (Fig. 3C); spinnerets with same color as abdomen.

Female (SAM-ENW-X013899, Fig. 5A–E): Total length: 17.61. Cephalothorax (Fig. 5A): length 5.15, width 3.88. Cephalic region: length 3.43, clypeus small (almost absent), with 4 marginal bristles; 12 bristles in front of the OQ, with a line of setae reaching as far as the fovea. Fovea: width 0.77, straight (slightly recurved). Eyes, diameters and interdistances (Fig. 5C): AME 0.10, ALE 0.26, PME 0.18, PLE 0.24, OQ length 0.49, width 0.89; AME-ALE 0.07, PME-PLE 0.02, AME-PME 0.11, ALE-PLE 0.05, AME-AME 0.17, PME-PME 0.42. Chelicerae: length 3.81, width 2.68; with dorsal-retrolateral dark bristles, rastellum with numerous rough bristles. Fang groove with 9 promarginal teeth and 16 mesobasal denticles. Labium: length 0.48, width 0.94, with 2 cuspules. Maxillae: length 2.16, width 0.92, with 133 cuspules on inner corner, prolateral face slightly curved, soft area small, with long uniformly distributed hairs, no serrula. Sternum (Fig. 5B): length 2.74, maximum width 2.29. Abdomen: length 8.74, covered with small hairs and setae. PMS: length 0.68; PLS: length of basal:medial:apical segments 1.26:0.79:0.86. Lengths of legs and palp: I: 3.93, 2.45, 2.68, 2.41, 1.62, 13.71. II: 3.71, 2.24, 2.32, 2.84, 1.57, 12.68. III: 2.97, 1.79, 1.68, 2.46, 1.44, 10.34. IV: 3.39, 2.22, 3.06, 3.55, 1.54, 14.30. Palp: 2.89, 1.59, 1.69, —, 2.00, 8.17.

Chaetotaxy : Leg I: femur, 1 PA; patella, 1 PA; tibia, 1-1 P, 1-1-3 V; metatarsus, 2-1-1 V. Leg II: femur, 1-1-1-1 d, 1 PA; patella, 1-1 P; tibia, 1-1 P, 1-1-2 V; metatarsus, 1-1 P (1/2 B), 2-2-2 V. Leg III: femur, 1 db, 1-1 P, 1 RA; patella, 1-1-1 P, 1 R; tibia, 1-1 P, 1-1 D, 1-1 R, 2-2-3 V; metatarsus, 1-1-1 P, 1-1-1 D-P, 1-1-1 D-R, 2-2-3 V. Leg IV: femur, 1-1-1-1 d, 1 PA; patella, 1 R; tibia, 1-1 R, 2-2-2 V; metatarsus, 1-1-1 P, 1-1 D-P (1/2 ANT), 1-1-1-1 D-R, 2-1-1-1-3 V. Pedipalp: femur, 1 PA; patella, 1-1 p; tibia, 2-1-1-3 P, 1-2-3 V; tarsus, 2 P, 1 VB. Spermathecae: complete, a high base with two lateral stalks, with an elongated receptacle (Fig. 5D, E). Scopula: Metatarsi: I, dense, uniformly distributed; II, dense, 3/4 A, divided by setae; III—IV, 0. Tarsi: I, dense, uniformly distributed; II, dense, uniformly distributed and divided by setae; III—IV, sparse, uniformly distributed (III divided by a wide band and IV by a very wide band of setae). Trichobothria: Tibiae: palp 8-9; I 9-9; II 6-5; III 8-8; IV 9-9. Metatarsi: I 1(5)1(4)1(2)1(3); II 1(4)1(3)1(1)1(3); III 1(4)1(3)1(1)1(4); IV 1(5)1(2)1(2)1(1)1(5). Tarsi: palp 10; I 14; II-III 12; IV 14. Preening combs: II 2 PV, III 3 PV, IV 3 RV-2 PV. Color: Entire body reddish-brown (chelicerae darker), legs same color as cephalothorax. Abdomen brown, with a mottled pattern dorsally; spinnerets same color as abdomen.

Distribution

Known from the Western Cape Province.

Acknowledgements

This research was made possible through scholarships awarded to the senior author: CONICET, Partial financial support for short stays of research; Res D N° 4367, 28 dec. 2017. It is part of the NRF (National Research Foundation, South Africa) grant No: 141937/Reference No: SRUG200417514326 to Catherine Sole and the authors. Special thanks are due to Petro Marais for providing access to the National Collection of Arachnida, ARC, Pretoria, where the senior author was hosted during his visit. Thanks to Ian Engelbrecht (SANBI, South African National Biodiversity Institute) for assistance provided during the earlier stages of the draft. To Sergei Zonstein (The Steinhardt Museum of Natural History Tel Aviv University, Israel) for useful comments. We are thankful to the second reviewer for their valuable comments and recommendations which helped to improve the original manuscript. Thanks to Ian Millar and Shannon Mitchel for the language review. We also thank all the curators who provided specimens of Entypesidae from South Africa on loan.

References

  • Dippenaar-Schoeman AS (2002) Baboon and Trapdoor Spiders of Southern Africa: An Identification Manual. Plant Protection Research Institute Handbook No. 13. Agricultural Research Council, Pretoria, 128 pp.
  • Goloboff PA (1995) A revision of the South American spiders of the family Nemesiidae (Araneae, Mygalomorphae). Part I: Species from Peru, Chile, Argentina, and Uruguay. Bulletin of the American Museum of Natural History 224: 1–189.
  • Goloboff PA, Platnick NI (1987) A review of the Chilean spiders of the superfamily Migoidea (Araneae, Mygalomorphae). American Museum Novitates 2888: 1–15.
  • Harvey MS, Hillyer MJ, Main BY, Moulds TA, Raven RJ, Rix MG, Vink C, Huey JA (2018) Phylogenetic relationships of the Australasian open-holed trapdoor spiders (Araneae: Mygalomorphae: Nemesiidae: Anaminae): multi-locus molecular analyses resolve the generic classification of a highly diverse fauna. Zoological Journal of the Linnean Society 184(2): 407–452. https://doi.org/10.1093/zoolinnean/zlx111
  • Hewitt J (1919) Descriptions of new South African Araneae and Solifugae. Annals of the Transvaal Museum 6: 63–111.
  • Opatova V, Hamilton CA, Hedin M, De Oca LM, Král J, Bond JE (2020) Phylogenetic systematics and evolution of the spider infraorder Mygalomorphae using genomic scale data. Systematic Biology 69(4): 671–707. https://doi.org/10.1093/sysbio/syz064
  • Purcell WF (1902) New South African trap-door spiders of the family Ctenizidae in the collection of the South African Museum. Transactions of the South African Philosophical Society 11(1): 348–382. https://doi.org/10.1080/21560382.1900.9525972
  • Ríos-Tamayo D, Goloboff PA (2018) Taxonomic revision and morphology of the trapdoor spider genus Actinopus (Mygalomorphae: Actinopodidae) in Argentina. Bulletin of the American Museum of Natural History 419: 1–83. https://doi.org/10.1206/0003-0090-419.1.1
  • Ríos-Tamayo D, Engelbrecht I, Goloboff PA (2021) A revision of the genus Hermacha Simon, 1889 (Mygalomorphae: Entypesidae) in Southern Africa with revalidation of Hermachola Hewitt, 1915 and Brachytheliscus Pocock, 1902. Argentina. American Museum Novitates 2021(3977): 1–80. https://doi.org/10.1206/3977.1
  • Tucker RWE (1917) On some South African Aviculariidae (Arachnida). Families Migidae, Ctenizidae, Diplotheleae and Dipluridae. Annals of the South African Museum 17: 79–138.
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