Stenaelurillus Simon, 1886

Logunov, Dmitri V. & Azarkina, Galina N., 2018, Redefinition and partial revision of the genus Stenaelurillus Simon, 1886 (Arachnida, Araneae, Salticidae), European Journal of Taxonomy 430, pp. 1-126 : 4-15

Genus Stenaelurillus Simon, 1886

Stenaelurillus Simon, 1886: 351 . Type species: Stenaelurillus nigricaudus Simon, 1886 ; by subsequent designation by Simon (1903: 669).

Mashonarus Wesołowska & Cumming, 2002: 165 . Type species: Mashonarus guttatus Wesołowska & Cumming, 2002 ; by original designation. Syn. nov.

Microheros Wesołowska & Cumming, 1999: 204 . Type species: Microheros termitophagus Wesołowska & Cumming, 1999 ; by monotypy. Syn. nov.

Definition

Small to medium spiders, ranging in body length from 3.30 to 6.40 mm in males (4.69 ± 0.72, n = 23) and 4.00 to 8.20 mm in females (5.74± 1.16, n = 19). Sexes similar in general body shape. Sexual dimorphism is poorly marked and can be seen in the following characters: males are usually smaller (on average, by some 20% of their body length) and brighter coloured; their Tb, Mt and Tr of legs I can be entirely dark brown or black in some species (e.g., S. jocquei sp. nov., S. fuscatus ; Figs 150 , 300 ) or with dark brown ventral sides (e.g., S. nigricaudus ; Figs 391, 393 ), or all their legs are darker (dark brown to black; e.g., S. modestus Wesołowska, 2014 ; Figs 361–363 ); males of some species have their carapaces covered with dense bunches of long (black, white and iridescent) hairs (e.g., S. hirsutus , S. jocquei sp. nov.; Figs 252 , 299 ); in other species (e.g., S. albopunctatus ) the male endites differ in having visible distal ectal projections (sensu Ramírez 2014; Fig. 46 ) which are absent in females.

CARAPACE. Rather high, with the posterior half of the thorax abruptly declining, almost vertical ( Figs 61 , 118 , 133 , etc.); densely covered with elongated recumbent scales making its colour pattern of two wide, white longitudinal stripes running along the ALE–PLE lines and two white marginal stripes ( Figs 55 , 102 , 115 , etc.); fovea present and situated between PLEs, but sometimes poorly seen because of the dense scale cover; in many species (e.g., in S. bandama sp. nov.) the anterior part of the eye field is covered with short erect bristles, similar to what was described in the aelurilline genus Asianellus Logunov & Heçiak, 1996 as ‘rod-hairs’ (see Logunov & Heçiak 1996: 105, figs 14–16), or in some species (especially in the males) carapace bears dense, mane-like bunches of hairs, sometimes also occupying cheeks ( Figs 164 , 252–255 , 298–300 ).

EYES. In three rows, with large black areas surrounding eyes ( Figs 85, 87 , 118 , etc.); in most species, the anterior eye row is slightly wider in both sexes (by 3–9% in males and 2–4% in females), but females of at least five species (viz., S. albopunctatus , S. jocquei sp. nov., S. darwini Wesołowska & Russell-Smith, 2000 , S. kronestedti and S. modestus ) have the anterior and posterior eye rows equally wide, and yet the anterior eye row is narrower by 3–5% than the posterior one in S. guttiger (in both sexes); the second row midway between ALEs and PLEs; quadrangle length 32–50% of carapace length.

CLYPEUS ( Figs 53 , 89 , 117 , etc.). Vertical, medium to high, ranging between 45–79% of the AME diameter in males and 42–60% in females; clypeal colourful coloration in many species of Stenaelurillus is diagnostic at the species level (e.g., in S. hirsutus , S. lesserti Reimoser, 1934 , S. pilosus , S. striolatus Wesołowska & Russell-Smith, 2011 , etc.; Figs 255 , 413 , 468 ).

CHELICERAE. Medium, vertical and of usual shape, with no projections or other modifications ( Figs 10– 13 ); promargin with two small teeth, usually fused together and sometimes forming a low blade-shaped ridge (e.g., in S. nigricaudus ; Fig. 10 ), retromargin with a small tooth ( Figs 10–12 ).

ENDITES. Subparallel, of usual shape, but in the males of some species (e.g., S. albopunctatus ) with poorly marked but visible distal ectal projections (arrowed in Fig. 13 ); in all species of Stenaelurillus endites with pale yellow to white apices.

LABIUM. Wide-triangular, with the obtuse tip directed forward.

STERNUM. Ovoid, longer than wide ( Figs 56 , 78 , 131 , etc.).

PEDICEL. Short, in live specimens not visible in dorsal view.

ABDOMEN. Elongate ( Figs 55 , 130 , etc.); males with the large dorsal scutum occupying the proximal half/two-thirds of the dorsum ( Figs 45 , 62 , 112 , etc.), the scutum may be invisible under the dense scale cover; colour markings on dorsum simple, either consisting of paired and/or singular white spots ( Figs 176 , 215 , 254 ) or white longitudinal stripes ( Figs 102 , 130 , 350 , 482 ), sometimes with no white colour pattern at all ( Figs 147 , 462 ).

BOOK- LUNG COVERS. Not sclerotized.

SPINNERETS. Medium, 21–36% of the abdominal length; the posterior pair slightly longer than the anterior one ( Figs 134–135 , 462 , 484 ), in some species (e.g., S. albopunctatus , S. furcatus , S. mirabilis , S. strandi ; Figs 45 , 347 , 385 ) the tips of posterior spinnerets or entire posterior spinnerets are contrastingly dark brown, especially in males.

LEGS. Subequally developed ( Figs 49–56 , 111–118 ); male legs I are usually darker than those of females, being entirely dark brown/black or with dark brown/black Tb, Mt and Tr ( Figs 204, 210 , 298–303 ); tarsal claws narrow, with well-developed and numerous teeth ( Figs 14–23 ): 7–20 teeth prolaterally and 5–19 retrolaterally. Leg formula: IV/III,II/I in both sexes, the third and fourth pairs of legs are always longer than the first and second pairs.

TRICHOBOTHRIA. Tr with 2–3 trichobothria in a single row, Mt with 2–4 trichobothria in a single row, Tb with a group of 5–6 trichobothria on its dorso-proximal end (other leg segments were not examined in this regard); trichobothrial proximal and distal plates present ( Figs 1–3, 5–8 ); the proximal plate (= trichobothrial ‘hood’) is finely striated (= fingerprint-like cuticular sculpture), with a medial differentiation and with one–two clearly marked transverse ridges on its distal margin; the margin of the alveolus smooth; the distal plate smooth, with or without a pair of transverse ridges in front of it; bothrial base thin.

TARSAL ORGAN. Of the capsulate type, with an ovoid pore being situated at the top of a low, round, domeshaped elevation and slightly disposed to one end, distally ( Figs 4, 9 ).

LEG SPINATION. All legs with numerous spines, especially on Tb and Mt of legs III–IV; the spine patterns of legs I–II are similar/identical, as are those of legs III–IV; the common spine patterns for most species are as follows: Fm I–II d 0-1-1/2-3/5; Fm III–IV d 0-1-1/2-4/5; Pt I–II pr 0-1-0; Pt III–IV pr and rt 0-1-0; Tb I–II pr 1-1, v 0/1-0/1-2ap; Tb III–IV d 1-0-0, pr and rt 1-1-1-1 or 1-1-1/2, v 1-0-2ap; Mt I–II pr 0/1-1ap, v 2-2ap; Mt III–IV pr and rt 1-0/1-1/2ap; some species have spineless Pt I–II, for instance, S. belihuloya sp. nov., S. brandbergensis comb. nov., S. guttatus comb. nov., and S. pseudoguttatus

sp. nov., of which two ( S. brandbergensis comb. nov. and S. guttatus comb. nov.) were earlier considered in the separate genus Mashonarus .

FEMALE PALP. General form; without an apical claw ( Figs 89 , 128 , 233 ).

MALE PALP. Femora of usual shape but in some species can be densely covered with long hairs and bristles (e.g., in S. lesserti ; see Sebastian et al., 2015: fig. 5A–C) or bears a low but clearly developed laminar crest (e.g., in S. darwini ; see Wesołowska & Russell-Smith, 2000: fig. 273); tibia short and wide, with two/three processes, ventral (VTA), retrolateral (RTA) and dorsal (DTA; e.g., in S. albopunctatus , S. kronestedti ); VTA usually looks like a ventral bulge of the tibia ( Figs 90 , 156 , 165 ) or like the short process (e.g., in S. abramovi Logunov, 2008 or S. fuscus Cao & Li, 2016 ; see Logunov 2008: fig. 1l; Cao et al. 2016: fig. 40A–B), whereas the well-developed DTA can bear a sharp apical spur or bunches/ rows of thick bristles (iridescent, dark brown or black; Figs 41 , 106 , 137 , 318 : DTA); the retrolateral and

dorsal sides of the tibia can be densely covered with thick and long hairs and bristles ( Figs 282 , 293 ); the cymbium is oval, with no apical spines and usually with no processes or projections, except for a few species having the proximo-lateral corner of the cymbium developed into a marked cymbial lateral projection (e.g., S. guttatus comb. nov., S. strandi , S. termitophagus comb. nov.; Figs 166 , 455 , 474 : CLP); the cymbial apex is slightly extended beyond the alveolus and in many species can bear a poorlymarked ventral groove (sensu Ramírez, 2014; arrowed in Figs 193 , 246 ); the basal haematodocha is welldeveloped ( Figs 30, 32–33 : BH); the subtegulum is simple, bean-shaped and visible in an expanded palp only ( Fig. 30 : ST); the tegulum is narrow and elongated, with an obtuse shoulder ( Figs 31 , 39 , 58 , 91 , 377 : T), which can bear a finger-shaped tegular process directed distad (e.g., in S. guttiger , S. nigricaudus , S. strandi , etc.; Figs 64 , 192 , 455 : TP); the distal haematodocha is well-developed ( Figs 30, 32–33 : DH), clearly separating the true tegulum from the salticid radix; the functional tegulum consists of the salticid radix (sensu Logunov 1999; Logunov & Cutler 1999; Logunov & Marusik 2003; Figs 31–32 , 37 : SR), which seems to be fused with a large distal sclerite on unknown origin that in some species (e.g., S. termitophagus comb. nov.) is visible and delimited from the salticid radix by a membranous area (arrowed in Figs 472 ); the salticid radix usually has a pointed or obtuse proximal projection ( Figs 31 , 58 , 166 : PP) and also bears a finger-shaped, bifurcated or pointed distal projection ( Figs 40 , 105 , 120 , 246 : DP), which in some species can be decorated with crest-forming stiff bristles (e.g., in S. lesserti ; Wesołowska 2014a: fig. 2A–D), in other species (e.g., S. abramovi , S. fuscus ; Figs 36–39 , see also Logunov 2008: figs 1–3; Cao et al. 2016: fig. 41B–C) the DP is strong, distinct and movable (see below for further discussion); the embolic division consists of the embolus (E) with the heavy embolic base (EB), which is either fused by its median edge with the distal edge of the salticid radix ( Figs 31 , 376 ) or connected to it by a visible membrane (arrowed in Fig. 40 ); the embolic conformation varies from whipshaped, with the poorly-developed embolic base ( Fig. 457 , 475 ), to needle-, hook- or ribbon-shaped, with a well-developed base ( Figs 33 , 93 , 160 , 378 ).

FEMALE COPULATORY ORGANS. The epigyne is usually flat, without epyginal depression or fossae, with openly displayed copulatory openings ( Figs 47 , 99 , 126 : CO), but in a few species ( S. darwini and S. termitophagus comb. nov.; Figs 109 , 476 ) they are hidden under a pair of bulge-shaped, chitinous flaps; the copulatory openings are either as rounded pores ( Figs 126 , 262 ), which sometimes are poorly seen on the epigynal plate ( Figs 99 , 171 ), or as deep furrows that could be subparallel (e.g., S. jocquei sp. nov., S. pilosus ; Figs 296 , 412 ; Wesołowska & Russell-Smith 2011: figs 165–166) or positioned transversely, forming a single transverse line ( S. bandama sp. nov., S. iubatus Wesołowska & Russell- Smith, 2011, S. sudanicus Wesołowska, 2014 ; Figs 70 , 278 , 452 ; Wesołowska 2014b: fig. 12B–C; Wesołowska & Russell-Smith, 2011: figs 159–160); in most species, the copulatory openings are widely separated, but in some (e.g., S. lesserti , S. abramovi , S. pecten Wesołowska, 2014 ; Logunov 2008: fig. 4; Wesołowska 2014 b: fig. 10F–G) they are close to each other, being separated by a space that is equal to or smaller than the diameter of each; epigynal pocket present, usually narrow and deep, situated at the edge of the epigastric furrow ( Figs 262 , 278 : EP) or in between the copulatory openings ( Figs 126– 227 , 288 : EP), but in some species it can be absent or poorly-marked (e.g., in S. guttatus comb. nov., S. latibulbus Wesołowska, 2014 ; Figs 171 , 334 ) or in some species (viz., S. abramovi ; see Logunov 2008: fig. 4) be displaced forward and look like a round depression; the insemination ducts usually short and wide ( Figs 335 , 399 : ID), except for S. termitophagus comb. nov. ( Figs 477–478 ), and lead to the bean-shaped, round or ovoid, medium/large primary spermathecae (sensu Ramírez 2014; Figs 71 , 127 , 172 : PS); the fertilization ducts are pronounced and well-visible ( Figs 127 , 145 , 228 : FD).

Morphological notes

The terms ‘functional tegulum’ and ‘salticid radix’ are used here in the same sense as in Logunov (1998, 1999), Logunov & Cutler (1999) and Logunov & Marusik (2003) to emphasise the composite nature of what is usually described in the advanced salticids as the tegulum; the cited works can be consulted for more details and evidence for this terminology.

The present analysis has shown that the sclerite composition of the male palp in Stenaelurillus is similar to that of Habrocestoides Prószyński, 1992 (see Logunov 1999): viz., in both genera the true tegulum is small and hidden behind the large salticid radix forming the main part of the functional tegulum ( Figs 31–33 , 39 , 64 , 90 ). Compared to Habrocestoides , the functional tegulum of Stenaelurillus also includes the second large sclerite, which in the majority of species is firmly fused with the salticid radix but in some is visible, being delimited from the latter sclerite by a membranous area, for instance, in S. albus Sebastian, Sankaran, Malamel & Joseph, 2015 ( Sebastian et al. 2015: fig. 2E), S. arambagensis (Biswas & Biswas, 1992) (see Prajapati et al. 2016: fig. 6B, sub. S. digitus Prajapati, Murthappa, Sankaran & Sebastian, 2016 ; Caleb et al. 2017), or S. termitophagus comb. nov. (arrowed in Fig. 472 ). It is difficult to homologise this sclerite, as the detailed structure of male copulatory organs yet remains poorly studied in the majority of salticid genera. Based on its position between the salticid radix and the embolic division, it is possible to conclude that this sclerite seems to be the same one as the ‘sclerite-1’ described in the genus Pseudeuophrys Dahl, 1912 (see Logunov 1998 for further discussion). The present finding also allows us to argue that the composite nature of the functional tegulum in Salticidae seems to be the rule rather than an exception. In many salticid genera, particularly in the Dendryphantinae and the Chrysillini (sensu Maddison 2015), the true tegulum is seen as a sclerotized, prolateral outgrowth (= shoulder of tegulum, sensu Maddison 1996) separated from the functional tegulum (= the salticid radix) by a visible tegular ledge (e.g., Maddison 1996: fig. 3). Nevertheless, in the alpha-taxonomic literature, both structures (the true tegulum and the salticid radix) are usually described together as ‘the tegulum’, though the latter is better called ‘the functional tegulum’ or ‘the salticid tegulum’.

This clarification of what is the true tegulum in Stenaelurillus has allowed us to correctly code its proximal finger-shaped process, which is well-developed in many species (e.g., in S. nigricaudus , S. strandi , etc.). This structure is coded here as ‘the tegular process’ ( Figs 33 , 64 , 192 , 241 : TP), because it clearly represents a distal prolongation of the true tegulum. It seems to be homologous to the distal tegular projection of Pisauridae (sensu Sierwald 1990) or the suprategulum of Linyphiidae (sensu Saaristo 1977; = the median apophysis in Merret 1963). By mistake, the TP in Stenaelurillus was coded by some earlier authors as the ‘terminal apophysis’ (e.g., Logunov 2008; Prajapati et al. 2016; Sebastian et al. 2015), although the true terminal apophysis is known to be part of the embolic division (see Merrett 1963: figs 1–3; Saaristo 1977).

In a few species (e.g., S. abramovi , S. fuscus ), the distal projection (DP) of the functional tegulum is strong, distinct and even movable, being separated from the functional tegulum by a visible membrane ( Figs 36, 39 ; see also Logunov 2008: figs 1–3; Cao et al. 2016: fig. 41B–C). The latter authors coded this sclerite as the ‘sclerotized apophysis’. Based on its position at the top of the salticid radix and the fact that it is separated from it by a membrane, it is possible to hypothesize that this sclerite could be either the terminal apophysis or the lamella described, for instance, in the Linyphiidae ( Merrett 1963: fig. 3). This assumption needs verification when more salticid taxa have been studied with regards to the detailed structure of their male copulatory organs.

It is worth noticing that in some Asian species (e.g., S. abramovi , S. albus , S. fuscus ) the proximal projection of the bulbus seems to have been formed by the proximal extention of the true tegulum ( Figs 38–39 ; see also Sebastian et al. 2015: fig. 2E) rather than the saltcid radix, as in most other species. In these species the functional tegulum has a narrow membranous area running across it, likely marking the border between the true tegulum and the salticid radix. Due to the limited/no material of these species available for the present study, we have been unable to expand their male palps and to verify this observation. The matter needs further attention when more fresh material of Asian species of Stenaelurillus have been made available.

Diagnosis and affinities

According to Maddison (2015) and his molecular data, Stenaelurillus belongs to the subtribe Aelurillina Simon, 1901 of the tribe Aelurillini Simon, 1901 in the subfamily Salticinae Blackwall, 1841 . It is a compact group of ground-dwelling jumping spiders containing some 281 species in nine genera ( Table 2 ; p. 119), which was formely known as the subfamily Aelurillinae Prószyński, 1976 (e.g., Metzner 1999). From all other aelurillines Stenaelurillus can be distinguished by the following characters: (1) the longest, conspicuous spinnerets ( Figs 45 , 134 , 205 , 462 ; 21–36% of the abdominal length), in other genera spinnerets are usually small, inconspicuous and poorly visible in dorsal view; (2) well-developed and numerous teeth of tarsal claws ( Figs 14–23; 7–20 teeth prolaterally and 5–19 retrolaterally), in other genera 3–5 teeth ( Figs 24–29 ) (this character is diagnostic even in immature specimens); (3) the least developed cymbial pocket (sensu Logunov 1996) as compared to other aelurillines, usually the embolic division is situated openly, in a shallow cavity at the top of the bulbus ( Figs 34 , 40 , 64 , etc.); (4) the true tegulum is marked and visible as a heavily sclerotized sclerite hidden behind (retro-laterad of) the salticid radix ( Figs 39 , 377 ; see above for further details); and (5) the bright body coloration consisting of numerous stripes and paired/singular spots ( Figs 102 , 130 , 178 , 215 , etc.), in other genera the body colour pattern is usually limited to longitudinal stripes (e.g., in Phlegra Simon, 1876 or Aelurillus Simon, 1884 ).

Based on the refined generic diagnosis, it is safe to conclude that the monotypic genus Microheros and the small genus Mashonarus with three species ( WSC 2017), both from Africa, are to be synonymized with Stenaelurillus . Both the conformation of the copulatory organs ( Figs 90–94 , 165–172 , 472–478 ) and the somatic morphology (long spinnerets, trichobothria and dentation of tarsal claws; Figs 5–9 , 16–19 , 176 , 484 ) of their generotypes are identical or in full agreement with those of Stenaelurillus nigricaudus ( Figs 1–2 , 14–15 , 369–381 , 395–399 ), the generotype of Stenaelurillus , and of other species included in the latter genus. All the species of Microheros and Mashonarus , except for the Indian Mashonarus davidi Caleb, Mungkung & Mathai, 2015 ( Figs 504–506 ), the holotype of which we have been unable to borrow, are redescribed and illustrated in detail below.

Composition

At present, Stenaelurillus consists of 45 valid species ( Table 1 ), of which 33 are known from both sexes, eight species from the males and four species from the females ( Table 2 ; p. 119). Details on the distribution of and references to all these species are provided in Table 1 .

Distribution

The geographical distribution of Stenaelurillus is clearly palaeotropical ( Fig. 507 ), covering almost the entire African continent, except for the western half of the equatorial zone, and then extending to southern Iran, south and SE Asia, including the southern and south-eastern territories of China. There are no common species between the Afrotropical and Oriental regions. As evidenced from the data at hand, there are three main centres of the present day species diversity (= chorological centres): western Africa (7 species), the south-eastern part of central Africa (21 species) and South Asia (5 species). The revealed distributional disjunction between the African and Asian parts of the generic range and the absence of species in Indonesia and Polynesia-Micronesia seem to be due to the incompleteness of our knowledge.

The outlined distribution of Stenaelurillus is virtually restricted to the Palaeotropical Zoogeographic Kingdom (sensu Kryzhanovsky 2002: inset 1), with a few records originating from the neighbouring regions of the Ancient Mediterranean (e.g., S. nigricaudus from N Algeria, S. marusiki Logunov, 2001 from SW Iran and S. triguttatus Simon, 1886 from Nepal and Tibet; Figs 509–510 ). Genera with such distributional patterns are classified as either “palaeotropical” (sensu Kryzhanovsky 2002: 58) or “multiregional of the eastern hemisphere” (sensu Pravdin & Mishchenko 1980: 28) and are commonly considered to be of Neogene origin (i.e., 23 Ma or later; sensu Hilgen et al. 2012). Numerous examples of the palaeotropical distribution in tribes and genera are known from Coleoptera, Orthoptera, Diptera, Mantodea and other orders ( Pravdin & Mishchenko 1980; Nartschuk 1994; Kryzhanovsky 2002; Davies et al. 2002; etc.).

It is worth noticing that so far no species of Stenaelurillus have been recorded or described from Madagascar. This fact could be explained either by the current incompleteness of our knowledge of Stenaelurillus , or, more likely, by the later origin of the genus, after Madagascar had already separated from Africa (183–158 Ma) and India (96–65 Ma) during the late Jurassic to late Cretaceous fragmentation of Gondwana (see Vences et al. 2009); the latter assumption seems to be in agreement with a possible Neogene origin of Stenaelurillus (see above). To date, the only described Malagasy aelurilline species is Aelurillus madagascariensis Azarkina, 2009 (see Table 3 , p. 120), a true member of the genus Aelurillus Simon, 1885 that is related to some species from the Levant ( Azarkina 2009).

The genus Stenaelurillus is not unique in the order Araneae , as far as its distributional pattern is concerned. A similar palaeotropical distribution is also known in a number of better studied spider genera, for instance: Borboropactus Simon, 1884 (Thomisidae) recorded from tropical Africa, S and SE Asia, but compared to Stenaelurillus also including Malaysia, Indonesia, the Phillipines and Papua New Guinea ( Benjamin 2011; WSC 2017); Calommata Lucas, 1837 (Atypidae) recorded from Africa, the Near East and SE Asia ( Fourie et al. 2011); Mallinella Strand, 1906 (Zodariidae) , a recently revised genus occurring in central and west Africa, S and SE Asia, southeastward to the northern tip of Australia ( Dankittipakul et al. 2012 a: fig. 1415; 2012 b); Sarascelis Simon, 1887 and the entire subfamily Chediminae (Palpimanidae) known from tropical Africa, India and Indonesia, but compared to Stenaelurillus with a lesser number of records ( Zonstein & Marusik 2013: fig. 51); Synaphosus Platnick & Shadab, 1980 (Gnaphosidae) known from central Africa, the Near East, Central Asia and SE Asia (yet two species were also introduced to the USA; see Ovtsharenko et al. 1994; WSC 2017); and other spider genera. Two spider genera display an almost identical distributional pattern as that of Stenaelurillus , except for some singular records from the Neotropics, namely: Nesticella Lehtinen & Saaristo, 1980 (Nesticidae) is known from tropical Africa, but not as widespread as Stenaelurillus , S and SE Asia, including the south-easternmost areas of the Palearcrtic Region and Papua New Guinea ( Lin et al. 2016; WSC 2017), plus a single record from Brazil ( Rodrigues & Buckup 2007); Bianor Peckham & Peckham, 1885 (Salticidae) is known from the Afrotropical, Oriental and Palaearctic (southern parts) Regions, with a single species – B. biocellosus Simon, 1902 – with its type locality in Brazil ( Logunov 2001 a). Of other jumping spider genera ( Salticidae ), most of which still remain rather poorly studied with regards to their composition and general distribution, the following genera seem to also demonstrate a palaeotropical distribution: Hyllus C.L. Koch, 1846 is widespread in tropical Africa and SE Asia (including the generotype; WSC 2017), but the congenerity of many species included in this genus requires revision; Harmochirus Simon, 1885 , a revised genus the distribution of which is almost identical to that of Stenaelurillus but with a lesser number of species and records ( Logunov 2001 a; WSC 2017); and other salticid genera. A further comparative analysis of the distributional pattern of Stenaelurillus with that of other spider genera, including the Salticidae , lies outside the goals of the present study and seems to be premature at the currently incomplete state of knowledge.

Figures

Figs 146–151. Stenaelurillus fuscatus Wesołowska & Russell-Smith, 2000, holotype, ♂ (150–151) and ♂ from Kenya (Taita Discovery Centre; MRAC 213.133) (146–149), general appearance. Scale bars: 1 mm.

Figs 298–308. Stenaelurillus jocquei sp. nov., light form, general appearance, ♂♂ (298–300, 302–303, 305–306) and ♀♀ (301, 304, 307–308). 298–301, 303–304, 307–308. Paratypes from Cameroon (Faro GR). 302, 305–306. Holotype. Scale bars: 1 mm.

Figs 382–394. Stenaelurillus nigricaudus Simon, 1886, ♂♂. 382–390. General appearance. 391, 393. Legs 1, prolaterally. 392, 394. Legs 1, retrolaterally. Specimens: 382, 385–386, 388–389, 391– 392 from Burkina Faso; 383–384, 387, 390, 393–394 from Senegal (Richard Toll). Scale bars: 1 mm.

Figs 361–368. Stenaelurillus modestus Wesołowska, 2014, from South Africa (uMkhuze GR), general appearance. 361–363, 365. ♂. 364, 366–368. ♀. Scale bars: 1 mm.

Figs 252–257. Stenaelurillus hirsutus Lessert, 1927 (holotype of Stenaelurillus cristatus Wesołowska & Russell-Smith, 2000) from Tanzania (Mkomazi GR), general appearance. Scale bars: 1 mm

Figs 40–48. Stenaelurillus albopunctatus Caporiacco, 1949, lectotype, ♂ and paralectotype, ♀. 40. Male palp, ventral view. 41. Male palp, retrolateral view. 42. Embolic division, dorsal view. 43. Embolic division, retrolateral view. 44. Embolic division, median view. 45. ♂, dorsal view. 46. ♂, ventral view. 47. Epigyne, ventral view. 48. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 40–44, 47–48 = 0.1 mm; 45–46 = 1 mm.

Figs 57–62. Stenaelurillus arambagensis (Biswas & Biswas, 1992), ♂ from Pakistan (Jehlum). 57. Male palp, ventral view. 58. Male palp, retrolateral view. 59. VTA, median view. 60. Clypeus, front view. 61. Body, lateral view. 62. Body, dorsal view. Abbreviations: see Material and methods. Scale bars: 57–59 = 0.1 mm; 60 = 0.5 mm; 61–62 = 1 mm.

Figs 111–118. Stenaelurillus darwini Wesołowska & Russell-Smith, 2000 from Tanzania (Mkomazi GR; MMUE G7571.48, 50), general appearance. 111–114. ♂. 115–118. ♀. Scale bars: 1 mm.

Figs 128–135. Stenaelurillus furcatus Wesołowska, 2014 from Namibia (Etosha National Park; MRAC 215.642), general appearance. 128–131. ♀. 132–135. ♂. Scale bars: 1 mm.

Figs 49–56. Stenaelurillus albopunctatus Caporiacco, 1949 from Kenya (Nairobi), general appearance. 49–52. ♂ (Langata, MMUE G7583.1). 53–56. ♀ (grounds of National Museum, MMUE G7583.2). Scale bars: 1 mm.

Figs 99–104. Stenaelurillus brandbergensis (Wesołowska, 2006), paratype, ♀ from Namibia (MRAC 216.021). 99. Epigyne, ventral view. 100. Spermathecae, dorsal view. 101–104. General appearance. Scale bars: 99–100 = 0.1 mm; 101–104 = 1 mm.

Figs 156–164. Stenaelurillus glaber Wesołowska & Russell-Smith, 2011, holotype, ♂ (156–160) and paratype, ³ from Nigeria (Mokwa) (161–164). 156. Male palp, ventral view. 157. Male palp, retrolateral view. 158. Embolic division, dorsal view. 159. Embolic division, median view. 160. Embolic division, retrolateral view. 161–164. General appearance. Abbreviations: see Material and methods. Scale bars: 156–160 = 0.1 mm; 161–164 = 1 mm.

Figs 84–89. General appearance of the holotype of Stenaelurillus belihuloya sp. nov. (84–86) and a female of S. lesserti Reimoser, 1934 from Sri Lanka (Pollonaruwa) (87–89). Scale bars: 1 mm.

Figs 413–416. Stenaelurillus pilosus Wesołowska & Russell-Smith, 2011, holotype, ♂, general appearance. Scale bars: 1 mm.

Figs 463–471. Stenaelurillus striolatus Wesołowska & Russell-Smith, 2011, holotype, ♂. 463. Palp, ventral view. 464. Palp, retrolateral view. 465. Embolic division, dorsal view. 466. Embolic division, median view. 467. Embolic division, retrolateral view. 468–471. General appearance. Scale bars: 463– 467 = 0.1 mm; 468–471 = 0.5 mm.

Figs 1–13. Trichobothrial bases (1–3, 5–8), tarsal organs (4, 9), left chelicerae (10–12) and right endite (13) of Stenaelurillus Simon, 1885. 1–2. S. nigricaudus Simon, 1886, ♀. 1. Tb 1. 2. Tr 1. 3. S. hirsutus Lessert, 1927, ♀, Tr 2. 4–7. S. termitophagus (Wesołowska & Cumming, 1999) comb. nov., ♀. 4. Tr 1. 5. Mt 1. 6. Tr 2. 7. Tr 1. 8–9. S. guttatus (Wesołowska & Cumming, 2002) comb. nov., ♀. 8. Tr 1. 9. Tr 1. 10. S. nigricaudus Simon, 1886, ♀. 11. S. termitophagus comb. nov., ♀. 12. S. darwini Wesołowska & Russell-Smith, 2000, ♀. 13. S. albopunctatus Caporiacco, 1949, ♂, right endite. Abbreviations: see Material and methods. Scale bars: 1–9 = 5 µm; 10–12 = 0.25 mm; 13 = 0.1 mm.

Figs 73–78. Stenaelurillus bandama sp. nov., holotype, ♂, general appearance. Scale bars: 1 mm.

Figs 173–179. Stenaelurillus guttatus (Wesołowska & Cumming, 2002) comb. nov., ♂, from Zambia (Namwala Kafue NP; NCA 2007/573), general appearance. Scale bars: 1 mm.

Figs 213–226. Stenaelurillus guttiger (Simon, 1901), ♂♂, general appearance. Specimens: 213–217, 220–221 from Botswana (Francistown; NCA 2009/3744); 218, 222–223 from South Africa (Ophathe GR; NCA 2007/2973); 219 from South Africa (Kruger NP, Letaba; NCA 2007/2284); 224–226 from South Africa (Kruger NP, Shingwedzi Camp), photos ©Galina Azarkina, juvenile (224) and two adult ♂♂ (225–226). Scale bars: 1 mm.

Figs 344–351. Stenaelurillus mirabilis Wesołowska & Russell-Smith, 2000, from Tanzania (Mkomazi GR, Ibaya camp), general appearance. 344–347. ♂. 348–351. ♀. Scale bars: 1 mm.

Figs 480–487. Stenaelurillus termitophagus (Wesołowska & Cumming, 1999) comb. nov. from South Africa (Doompoort, NCA 2010/4945), general appearance. 480–483. ♀. 484–487. ♂. Scale bars: 1 mm.

Figs 454–462. Stenaelurillus strandi Caporiacco, 1939, holotype, ♂ (459–462) and paratype (454–458). 454, 459. Male palp, ventral view. 455, 460. Male palp, retrolateral view. 456–458. Embolic division. 456. Retrolateral view. 457. Apical view. 458. Median view. 461–462. General appearance. Scale bars: 454–460 = 0.1 mm; 461–462 = 1 mm.

Figs 204–212. Stenaelurillus guttiger (Simon, 1901), ♂♂, general appearance. Specimens: 204–206, 208, 210 paratype of S. natalensis Haddad & Wesołowska, 2006 from South Africa (Ndumo GR; NCA 2006/678); 207, 209, 211–212 from South Africa (Polokwane GR; NCA 2008/2067). Scale bars: 1 mm.

Figs 14–29. Tarsal claws of legs 1 in females of Stenaelurillus Simon, 1886, scanning electron micrographs. 14–15. S. nigricaudus Simon, 1886. 14. Retrolateral. 15. Prolateral. 16–17. S. guttatus (Wesołowska & Cumming, 2002) comb. nov. 16. Retrolateral. 17. Prolateral. 18–19. S. termitophagus (Wesołowska & Cumming, 1999) comb. nov. 18. Retrolateral. 19. Prolateral. 20–21. S. fuscatus Wesołowska & Russell-Smith, 2000. 20. Retrolateral. 21. Prolateral. 22–23. S. guttiger (Simon, 1901). 22. Retrolateral. 23. Prolateral. 24–25. Aelurillus v-insignitus (Clerck, 1757). 24. Retrolateral. 25. Prolateral. 26–27. Phlegra fasciata (Hahn, 1826). 26. Retrolateral. 27. Prolateral. 28–29. Langona bristowei Berland & Millot, 1941. 28. Retrolateral. 29. Prolateral. Scale bars: 0.1 mm.

Figs 227–238. Stenaelurillus guttiger (Simon, 1901), ♀♀. 227, 229. Epigyne, ventral view. 228, 230. Spermathecae, dorsal view. 231–238. General appearnce. Specimens: 227–228, 235, 238 from South Africa (Ndumo GR; NCA 2006/1275); 229–230, 232 from South Africa (Kruger NP, Satara; NCA 2008/2001); 231 from South Africa (Kruger NP, Pafuri; NCA 2007/2474); 233–234, 236–237 from South Africa (Kruger NP, Oliphants; NCA 2009/3738). Scale bars: 227–230 = 0.1 mm; 231–238 = 1 mm.

Figs 90–98. Stenaelurillus brandbergensis (Wesołowska, 2006), paratype, ♂ from Namibia (MRAC 216.021). 90. Palp, ventral view. 91. Palp, retrolateral view. 92. Embolic division, retrolateral view. 93. Embolic division, dorsal view. 94. Embolic division, median view. 95–98. General appearance. Abbreviations: see Material and methods. Scale bars: 90–94 = 0.1 mm; 95–98 = 1 mm.

Figs 165–172. Stenaelurillus guttatus (Wesołowska & Cumming, 2002) comb. nov., holotype, ♂ (165–170) and paratype, ♀ (171–172). 165. Male palp, ventral view. 166. Male palp, retrolateral view. 167. Male palp, dorsal view. 168. Embolic division, dorsal view. 169. Embolic division, median view. 170. Embolic division, retrolateral view. 171. Epigyne, ventral view. 172. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 105–110. Stenaelurillus darwini Wesołowska & Russell-Smith, 2000 from Tanzania (Mkomazi GR; MMUE G7571.48,50). 105. Male palp, ventral view. 106. Male palp, retrolateral view. 107. Embolic division, dorsalview. 108. Embolic division, median view.109. Epigyne, ventral view.110. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 0.1 mm

Figs 136–145. Stenaelurillus fuscatus Wesołowska & Russell-Smith, 2000, ♂ (136–142), and ♀ from Kenya (Taita Discovery Centre; MRAC 213.133) (143–145). 136. Male palp, ventral view. 137. Male palp, retrolateral view. 138. DTA, dorsal view. 139–142. Embolic division. 139. Ventral view. 140. Retrolateral view. 141. Median view. 142. Dorsal view. 143–144. Epigyne, ventral view. 145. Spermathecae, dorsal view. Scale bars: 0.1 mm. Abbreviations: see Material and methods.

Figs 317–323. Stenaelurillus kronestedti Próchniewicz & Heçiak, 1994, holotype, ♂ (317–322) and paratype ♀ (323). 317. Bulbus, ventral view. 318. Male palp, retrolateral view. 319. Embolic division, dorsal view. 320. Embolic division, retrolateral view. 321. Embolic division, median view. 322. General appearance. 323. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 317– 321, 323 = 0.1 mm; 322 = 1 mm.

Figs 280–287. Stenaelurillus jocquei sp. nov., holotype (280–283) and paratype (284–287), ♂♂ from Cameroon (Faro GR), light form. 280–282. Male palp. 280. Ventral view. 281. Retrolateral view. 282. Dorso-retrolateral view. 283–287. Embolic division. 283. Dorsal view. 284. Ventral view. 285. Median view. 286. Retrolateral view. 287. Apical view. Scale bars: 0.1 mm.

Figs 292–297. Stenaelurillus jocquei sp. nov., paratypes from Cameroon (Rumsiki; MRAC 168.689), dark form. 292. Male palp, ventral view. 293. Male palp, retrolateral view. 294. Embolic division, retrolateral view. 295. Embolic division, median view. 296. Epigyne, ventral view. 297. Spermathecae, dorsal view. Scale bars: 0.1 mm.

Figs 472–479. Stenaelurillus termitophagus (Wesołowska & Cumming, 1999) comb. nov., ♂ from Botswana, (Mopane Woodland) and ♀ from Botswana (island in Moanachira River). 472–474. Male palp. 472. Ventral view. 473. Ventro-retrolateral view. 474. Retrolateral view. 475. Embolic division, apical view. 476. Epigyne, ventral view. 477. Spermathecae, dorsal view. 478. Spermathecae, ventral view. 479. Diagrammatic course of the insemination ducts. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 193–203. Male palp of Stenaelurillus guttiger (Simon, 1901). 193. Ventral view. 194. Retrolateral view. 195, 197, 203. Embolic division, dorsal view. 196, 200. Embolic division, ventral view. 198, 201. Embolic division, retrolateral view. 199, 202. Embolic division, median view. Specimens: 193– 194, 196–199 from Botswana (Francistown; NCA 2009/3744); 195 from South Africa (Ophathe GR; NCA 2007/2973); 200–203 from South Africa (Ndumo GR; NCA 2006/1275). Scale bars: 0.1 mm.

Figs 246–251. Stenaelurillus hirsutus Lessert, 1927 (holotype of Stenaelurillus cristatus Wesołowska & Russell-Smith, 2000). 246. Male palp, ventral view. 247. Male palp, retrolateral view. 248. Male palp, dorsal view. 249. Embolic division, ventral view. 250. Embolic division, median view. 251. Embolic division, retrolateral view. Scale bars: 0.1 mm.

Figs 30–33. Expanded female palp of Stenaelurillus nigricaudus Simon, 1886 from Burkina Faso (Sissamba). 30. Median view. 31. Ventral view. 32. Retrolateral-dorsal view. 33. Retrolateral view. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 34–39. Male palp of Stenaelurillus abramovi Logunov, 2008 from Thailand (Chai Chet Cape), scanning electron micrographs. 34. Ventral view. 35, 39. Retrolateral view. 36. Embolic division, ventral view. 37. Median view. 38. Dorsal view. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 376–381. Stenaelurillus nigricaudus Simon, 1886, from Senegal (Richard Toll), scanning electron micrographs. 376. Bulbus of male palp, ventral view. 377. Male palp, retrolateral view. 378. Male palp, dorsal view. 379. Embolic division, ventral view. 380. Embolic division, retrolateral view. 381. Embolic division, dorsal view. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 63–72. Stenaelurillus bandama sp. nov., holotype, ♂ (63–65), paratype, ♂ (66–69) and paratype, ♀. 63. Male palp, retrolateral view. 64. Male palp, ventral view. 65. Male palp, dorsal view. 66–69. Embolic division. 66. Ventral view. 67. Median view. 68. Dorsal view. 69. Retrolateral view. 70. Epigyne, ventral view. 71. Spermathecae, dorsal view. 72. Diagrammatic course of the insemination ducts. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 184–192. Male palp of Stenaelurillus guttiger (Simon, 1901). 184. Male palp, ventral view. 185– 186. Male palp, retrolateral view. 187–192. Embolic division. 187, 189. Ventral view. 188, 192. Dorsal view. 190. Median view. 191. Retrolateral view. Specimens: 186, 189–192 = paratype of S. natalensis Haddad & Wesołowska, 2006 from South Africa (Ndumo GR; NCA 2006/678); 184–185, 187–188 from South Africa (Polokwane GR, NCA 2008/2067). Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 119–127. Stenaelurillus furcatus Wesołowska, 2014 from Namibia (Etosha National Park; MRAC 215.642). 119. Male palp, retrolateral view. 120. Male palp, ventral view. 121–124. Embolic division. 121. Retrolateral view. 122. Dorsal view. 123. Ventral view. 124. Median view. 125. RTA, retrolateral view. 126. Epigyne, ventral view. 127. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 0.1 mm.

Figs 258–263. Stenaelurillus hirsutus Lessert, 1927, ♀ (paratype of Stenaelurillus cristatus Wesołowska & Russell-Smith, 2000) from Tanzania (Mkomazi GR). 258–261. General appearance. 262. Epigyne, ventral view. 263. Spermathecae, dorsal view. Scale bars: 258–261 = 1 mm; 262–263 = 0.1 mm.

Figs 406–412. Stenaelurillus pilosus Wesołowska & Russell-Smith, 2011, holotype, ♂, and paratype, ♀. 406. Male palp, ventral view. 407. Male palp, retrolateral view. 408–410. Embolic division. 408. Dorsal view. 409. Median view. 410. Retrolateral view. 411. Spermathecae, dorsal view. 412. Epigyne, ventral view. Scale bars: 0.1 mm.

Figs 274–279. Stenaelurillus iubatus Wesołowska & Russell-Smith, 2011, ♀ (paratype of Stenaelurillus pilosus Wesołowska & Russell-Smith, 2011) from Nigeria (Ibadan). 274–277. General appearance. 278. Epigyne, ventral view. 279. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 258–261 = 1 mm; 262–263 = 0.1 mm.

Figs 448–453. Stenaelurillus sudanicus Wesołowska, 2014, holotype, ♀. 448–451. General appearance. 452. Epigyne, ventral view. 453. Spermathecae, dorsal view. Scale bars: 448–451 = 1 mm; 452–453 = 0.1 mm.

Figs 288–291. Stenaelurillus jocquei sp. nov., paratype ♀ from Cameroon (Faro GR; MRAC 221.132), light form. 288–289. Epigyne, ventral view. 290. Spermathecae, dorsal view. 291. Diagrammatic course of the insemination ducts of the spermathecae. Scale bars: 0.1 mm.

Figs 330–335. Stenaelurillus latibulbis Wesołowska, 2014, ♀ from Zambia (Luambe). 330–333. General appearance. 334. Epigyne, ventral view. 335. Spermathecae, dorsal view. Abbreviations: see Material and methods. Scale bars: 330–333 = 1 mm; 334–335 = 0.1 mm.

Figs 395–399. Stenaelurillus nigricaudus Simon, 1886. 395–398. Epigyne, ventral view. 399. Spermathecae, dorsal view. Specimens: 395–396, 399 from Senegal (Richard Toll); 397–398 from Burkina Faso. Abbreviations: see Material and methods. Scale bars: 1 mm.

Figs 239–245. Stenaelurillus hirsutus Lessert, 1927, holotype. 239. Male palp, ventral view. 240. Embolic division, retrolateral view. 241. Embolic division, dorsal view. 242. Embolic division, median view. 243–245. General appearance. Abbreviations: see Material and methods. Scale bars: 239–242 = 0.1 mm; 243–245 = 1 mm.

Figs 369–375. Stenaelurillus nigricaudus Simon, 1886, from Senegal (Richard Toll). 369–371. Male palp. 369. Ventral view. 370. Retrolateral view. 371. Dorsal view. 372–375. Embolic division. 372. Ventroapical view. 373. Dorsal view. 374. Median view. 375. Apical view. Scale bars: 0.1 mm.

Figs 504–506. Phlegra davidi (Caleb, Mungkung & Mathai, 2015) comb. nov., ♂ from India (Tamil Nadu). 504. Expanded palp, median view. 505. Ditto, retrolateral view. 506. Chelicerae, endites and labium, ventral view. Scale bars: 0.1 mm.

Fig. 507. General distribution and modern centres of species diversity of Stenaelurillus Simon, 1886 .

Fig. 509. Collecting localities of ten Asian species of Stenaelurillus Simon, 1885.

Fig. 510. Collecting localities of seven African species of Stenaelurillus Simon, 1885.

Tables

Table 1 (continued on next pages). Species composition and distribution of Stenaelurillus Simon, 1885.

Species name	Known	Zoogeographical regions		References
	sexes	Afrotropical	Palaearctic Oriental
S. abramovi Logunov, 2008	♂♀		S Vietnam, Thailand (Fig. 508)	Logunov (2008), Logunov & Jäger (2015), present data
S. albopunctatus Caporiacco, 1949	♂♀	Kenya (Fig. 511)		Caporiacco (1949), Wesołowska (2014b), present data
S. albus Sebastian, Sankaran, Malamel & Joseph, 2015	♂♀		S India (Fig. 509)	Sebastian et al. (2015), Prajapati et al. (2016)
S. arambagensis (Biswas & Biswas, 1992)	♂♀		India (Andhra Pradesh, Gujarat, Maharashtra, West Bengal), Pakistan (Punjab) (Fig. 509)	Prajapati et al. (2016: sub. S. digitus ), Caleb et al. (2017), present data
S.bandama sp. nov.	♂♀	Cote d’Ivoire (Fig. 510)		Wesołowska (2014b: sub. S. hirsutus , the records from Cote d’Ivoire), Present data
S. belihuloya sp. nov.	♂		Sri Lanka (Fig. 509)	Present data
S. brandbergensis ( Wesołowska, 2006) comb. nov.	♂♀	Namibia (Fig. 511)		Wesołowska (2006: sub. Mashonarus b.), present data
S. darwini Wesołowska & Russell-Smith, 2000	♂♀	Tanzania, Kenya (Fig. 511)		Prószyński (1984: 140, sub. Stenaelurillus sp. nov.2), Wesołowska & Russell-Smith (2000), Wesołowska (2014b), present data
S. furcatus Wesołowska, 2014	♂♀	Namibia (Fig. 511)		Wesołowska (2014b), present data
S. fuscatus Wesołowska & Russell-Smith, 2000	♂♀	Tanzania, Kenya (Fig. 512)		Wesołowska & Russell-Smith (2000), present data
S. fuscus Cao & Li, 2016	♂		S China (Yunnan) (Fig. 508)	Cao et al. (2016)
S. gabrieli Prajapati, Murthappa, Sankaran & Sebastian, 2016	♂♀		NW India (Gujarat) (Fig. 509)	Prajapati et al. (2016)
S. glaber Wesołowska & Russell-Smith, 2011	♂	Ghana, Nigeria, Uganda (Fig. 511)		Wesołowska & Russell-Smith (2011), Wesołowska (2014b), present data
S. guttatus ( Wesołowska & Cumming, 2002) comb. nov.	♂♀	Botswana, Zambia, Zimbabwe (Fig. 511)		Wesołowska & Cumming (2002: sub Mashonarus g.), present data
S. guttiger ( Simon, 1901)	♂♀	Botswana, Kenya, Mozambique, Zimbabwe, South Africa (Fig. 512)		Haddad & Wesołowska (2006: sub. both S. g. and S. natalensis ), Wesołowska & Cumming (2011), present data