Actinopus Perty 1833

Actinopus Perty, 1833

Figs 1–225, Maps 1–19

Actinopus Perty, 1833: 198 (type species by monotypy, Actinopus tarsalis Perty, 1833); C. L. Koch, 1842: 98 [revalidation]; Ausserer, 1871: 138; Simon, 1892: 80; Mello-Leit„o, 1923: 17; Raven, 1985: 145, 147; World Spider Catalog, 2020.

Closterochilus Ausserer, 1871: 141 (type species: Pachyloscelis nigripes Lucas, 1834, by original designation).

Theragretes Ausserer, 1871: 142 (type species: Actinopus walckenaerii Lucas, 1837, by original designation).

Diagnosis. Within Actinopodidae, species of Actinopus are recognized by the procurved anterior row of eyes, with the lateral eyes aligned antero-posteriorly (Fig. 4 A, F); labium elongated antero-posteriorly, without cuspules on males and with an average of 15–30 cuspules on females (Fig. 4 B–C); sternal sigilla close together, positioned in a deep median common depression (Fig. 4 D). Additionally, males and females of Actinopus differ from those of Missulena by the pronounced rastellum (Fig. 4 E), sternum not rebordered (Fig. 10 C) and superior tarsal claws with a tooth and one or two denticles (Fig. 4 G– H). They differ from those of Plesiolena by the flattened cheliceral apex (Fig. 10 E–F) and by the crown of robust spines on the dorsal, apical surface of the patella and tibia III (Fig. 11 A, C). Males of Actinopus differ from those of Plesiolena by the ventral pseudoscopula in metatarsi and tarsi (Fig. 5 A) and from those of Missulena by the cymbium without ventral incision. They differ from males of both genera by the copulatory bulb with paraembolic apophysis (Fig. 5 B) and with twisted keels in prolateral view of embolus (Fig. 5 C). Females of Actinopus differ from those of Missulena by tarsi I and II with strong and sharp spines and spermathecae with robust receptacles; differ from those of both genera by the short spermathecal receptacles (Fig. 5 D).

Description. Total length (males and females) from 7.0 to 35.9 mm.

Carapace. Anterior prolateral angle of carapace tapered (Fig. 172 B) or subquadrate (Fig. 5 F). Carapace subquadrate in dorsal view, smooth with strongly demarcated radial grooves and wider at coxae III, anterior median ocular tubercle absent, with delimited cephalic area and strong inclination at fovea level, anterior margin rounded; fovea U-shaped, strongly procurved, deep, sharply demarcated (Figs 5 E); in lateral view, thoracic area with abrupt anterior slope and gradual posterior slope. Clypeus, anteriorly less sclerotized than posteriorly, height from 1.0–1.5 times diameter of ALE. Anterior eyes row slightly procurved; posterior eyes row recurved d in dorsal view (Fig. 5 F). No or few bristles between AME-clypeus; no or few bristles between PME–PME and PLE–PLE. Ocular quadrangle wider than long; ALE diameter bigger than that of other eyes. Interdistances: Males: PME–PME separated by more than four diameters of PME, ALE–AME separated by ca. twice diameter of AME, ALE–PLE separated by less than twice diameter of ALE,AME–PME separated by ca. twice diameter of AME (Fig. 4 A). Females: PME–PME separated by ca. three times interdistance of PME–PLE, PLE–PME separated by half diameter of PME, AME–AME separated by half the diameter of ALE, ALE–AME separated by four times diameter of AME, ALE–PLE separated by twice diameter of PLE, AME–PME separated by four times diameter of AME (Fig. 5 F). Chelicerae not geniculate, longer than half carapace length, with apical rastellum. Rastellum smooth, with bristles (Fig. 13 C) or spines (Fig. 108 F). Females with very conspicuous crown of thorns bordering rastellum (Fig. 210 F). Cheliceral teeth, in ventral view, arranged in two rows, prolateral and retrolateral. Prolateral teeth row bordered by denticles (Fig. 5 G). Labium fused to sternum. Labium and maxillae longer than wide. Males without labial and maxillary cuspules (Figs 4 B, 6 A); females with numerous cuspules and bristles (Fig. 5 H). Sternum longer than wide. Sternum with 6–10 hardly distinguishable sigilla, strongly fused at center of sternum. Pre-coxal triangles absent (Fig. 24 C).

Legs. Males: Long, smooth (Fig. 6 B) or covered with few simple and sparse hairs (Figs 14 D, 31 A–D); Many prolateral spines on the sides of the patellae III and IV, forming crown spines on dorsal surface of Patella and tibia III (Fig. 14 A, C); tibiae, metatarsi and tarsi with filiform dorsal trichobothria, with high bases (Fig. 6 C, D); clavate tarsal trichobothria absent. Tarsi and metatarsi III and IV with compact pseudoscopula (Fig. 6 E–F), subapical chemosensitive fissures present (Fig. 6 E–H) or absent. Inferior tarsal claws without teeth; superior tarsal claws with one tooth, one tooth and one denticle or one tooth and two denticles (Fig. 4 G–H). Subinguinal tufts absent; all trochanters with a fissure. Females: Legs short, robust and covered by several short and strong spines, with many long bristles (Fig. 7 A), rows of strong spines on prolateral and retrolateral surfaces of distal articles (tibiae, metatarsi and tarsi) in all legs (Fig. 7 B), except leg IV, with rows only on prolateral surface; many spines on prolateral patellae III and IV, forming a crown of spines on dorsal surface of patella and tibia III; ventrally, spines replacing pseudoscopula on tarsi III and IV; tibiae, metatarsi and tarsi with filiform dorsal trichobothria with a high bases (Fig. 7 C), tricoma with large fixation base (Fig. 7 D); Clavate tarsal trichobothria absent. Inferior and superior tarsal claws without teeth. Subinguinal tufs absent. All trochanters with fissure.

Abdomen. Long, oval, homogeneously covered with simple setae and dorsally with long simple setae (Fig. 10 A); male and female dorsal scutum, when present, restricted to anterior half of abdomen, whole or divided; scutal bristles with more sclerotized bases that those of ordinary tegument bristles (Fig. 13 A). Four spinnerets (Fig. 7 E). Posterior median spinnerets (PMS) small or medium-sized, with one segment; posterior lateral spinnerets (PLE) with three bigger segments; spigot types: principal ampullary fusule and secondary ampullary fusule (Fig. 7 D–E), tartipores (Fig. 7 D) as remnants of small ampular in some segments; apical digitiform article of PLS.

Male Palp. Femur, patella and tibia without or with few spines. Tibia without retrolateral tibial apophysis, more thickened at base than apically (Fig. 7 H); cymbium rounded, without spines and scopula; subtegulum subrectangular, sclerotized, turned laterally (Fig. 8 A); tegulum large, sclerotized, reservoir with two or three loops in prolateral view; distal region of tegulum with prolateral paraembolic apophysis; apical tegulum with prolateral keels; embolus inserted in angle relative to tegulum, sclerotized, with visible duct (Figs 9 A–C and 12 A–C). Generally, keels represented by three paraembolic apophyses in prolateral view of embolus (Actinopus paranensis, A. pinhao, A. ducke, A. hirsutus, A. jaboticatubas, A. confusus, A. pampulha, A. candango, A. paraitinga, A. cornelli, A. nattereri, A. rufipes, A. vilhena, A. harveyi, A. itapitocai, A. ipioca, A. itaqui, A. xingu, A. mesa, A. caxiuana, A. utinga, A. crassipes, A. dubiomaculatus, A. pusillus, A. emas, A. bocaina, A. laventana, A. osbournei, A. dioi, A. longipalpis, A. tutu, A. azaghal, A. itacolomi, A. tarsalis, A. castelo, A. parafundulus, A. urucui and A. panguana). In A. panguana (Fig. 206 A–C), a dorsal tegular process (DTP); in A. cucutaensis (Fig. 176 A–C), A. robustus, A. lomalinda, A. palmar and A. guajara, an apical tegular process (ATP); basal tegular apophysis (BTA) absent in A. tarsalis and A. castelo (Fig. 12 A–C).

Spermathecae. Spermatic ducts robust, short (Fig. 217 B–D), well separated from base by a membranous region or strongly fused to it (Figs 26 D, 181 D); each receptacle without or with one or three lobes (Fig. 220 A), lobes quite sclerotized, apically covered by pores (Fig. 8 B).

Composition. Eighty species, 42 of which are newly described. Ten old species names are herein considered species inquirendae: A. rojasi (Simon, 1889) and A. valencianus (Simon, 1889) were described from juveniles. The type of A. pertyi Lucas, 1843, originally deposited at Muséum National d´Histoire Naturelle (MNHN), Paris, France, is lost. The holotype female of A. rufibarbis Mello-Leit „o, 1930, deposited at Museu Nacional do Rio de Janeiro, Rio de Janeiro, Brazil, is without spermathecae and in poor condition. The types of A. nigripes (Lucas, 1834), A. piceus (Ausserer, 1871) and A. liodon (Ausserer, 1875), all originally deposited at Naturhistorisches Museum, Wien, Austria, and A. harti Pocock, 1895 originally deposited at The Natural History Museum, London, England, were not found and are considered lost. Specimens identified as A. caraiba (Simon, 1889) and A. scalops (Simon, 1889) were examined and photographed by J. P. Guadanucci at Muséum National d´Histoire Naturelle (MNHN), Paris, France, but the localities on the labels are slightly different from the respective published type localities reported in the original descriptions and the specimens were not labelled as types. These specimens were not considered as types in the present paper by all prior knowledge about Simon’s specimens deposited at MNHN and the totally absence of any other type evidence on labels of other specimens deposited in the Paris Museum.

Key to species groups

The following key is based on males only and, besides the 12 groups here defined, includes also the three species (known from males) which were not included in any group.

1. Discontinuous thoracic fovea that is interrupted in middle (Ríos-Tamayo & Goloboff, 2018, fig. 42 A) and sternum with a deep central depression (Ríos-Tamayo & Goloboff, 2018, fig. 42 B)................................................gr. goloboffi

Continuous thoracic fovea (Figs 5 A, H) and sternum without a deep central depression (Fig. 4 D).................... 2

2(1). Tibia III with three basal, dorsal spines (Fig. 208 B)..................................................................................... A. clavero

Tibia III with a single basal, dorsal spine (Fig. 158 B).... 3

3(2). ATP present (Fig. 176 A).................................................. 4

ATP absent (Fig. 18 A)..................................................... 5

4(3). ATP oblique in relation to embolus (Fig. 195 A–B)................................................................................... A. concinnus

ATP parallel in relation to embolus (Fig. 176 A–B)................................................................................gr. cucutaensis

5(3). Embolus strongly curved prolaterally (distal third not visible in dorsal view); DTP present (Fig. 206 A–C)................................................................................. A. panguana

Embolus, at most, gently curved prolaterally, DTP absent (Fig. 18 A–C).................................................................... 6

6(5). Base of embolus with extremely developed serrated area, composed of three rows of large cusps (Figs 198 A–C)................................................................................ A. apiacas

Serrated area not so developed, or absent (Figs 18 A–C, 67 A–C)................................................................................. 7

7(6). PA inserted apically on bulb, flattened and apically folded, oblique in relation to embolus (Fig. 201 A–C).... A. jamari

PA inserted on the prolateral bulb, or absent (Fig. 75 A–C).......................................................................................... 8

8(7). Keels short, restricted to median sector of embolus, represented by short prominent lamellae in prolateral view (Figs 162 A, 165 A)..................................gr. parafundulus

Keels otherwise (Fig. 62 A).............................................. 9

9(8). Two embolar keels, PAc absent (Fig. 168 A–C)............. 10

Three embolar keels, PAc present (Fig. 35 A–C)........... 12

10(9). Tegulum and embolus thin and elongated along longitudinal line; embolar insertion ample (Fig. 168 A–C)....gr. reznori

Tegulum notably wider than embolar insertion (Fig. 75 A–C)............................................................................... 11

11(10). Serrated area on embolar prolateral surface absent or represented by at most two small cusps (Fig. 15 A–C)...................................................................................gr. apalai

Serrated area represented by three or more cusps (Fig. 32 A–C)................................................................... gr. pinhao

12(9). Embolus, as seen from dorsal view, inserted on tegulum at a 90 o angle (Fig. 9 B)......................................... gr. tarsalis

Embolar insertion, as seen from dorsal view, otherwise (Fig. 153 A–C)................................................................ 13

13(12). Sternal sigilla fused medially of sternum (Fig. 133 C)............................................................................. gr. osbournei

Sternal sigilla converging to the middle but separated (Fig. 196 C)............................................................................. 15

14(13). Small spiders, total length less than 13 mm................... 15

Large spiders, total length more than 17 mm and less than 36 mm............................................................................. 16

15(14). Serrated area on prolateral embolus small (Fig. 143 A–C) or, when large, represented by sparse small denticles (Fig. 153 A–D).................................................................gr. tutu

Serrated area on prolateral embolus large, densely covered by relatively large denticles (Fig. 124 A–C)... gr. crassipes

16(14). Serrated area on prolateral embolus reduced or absent (Fig. 67 A)................................................................ gr. nattereri

Serrated area on prolateral embolus well developed (Fig. 32 A)........................................................................ gr. pinhao