MRACRoyal Museum for Central Africa, Tervuren, Belgium;

NCA National Collection of Arachnida, ARC-Plant Protection Research Institute, Pretoria, South Africa;

NHMLNatural History Museum, London, UK;

NRSNaturhistoriska riksmuseet, Stockholm, Sweden;

SMFSenckenberg Museum, Frankfurt am Main, Germany;

TAU Steinhardt Museum of Natural History, Tel-Aviv University, Israel;

ZMUT Zoological Museum, University of Turku, Finland.

Boagrius incisus Tullgren, 1910: TANZANIA: 1♂ syntype Kibonoto (Kilimanjaro), viii–ix.1905, Y. Sjöstedt (NRS).

Boagrius pumilus Simon, 1893: MALAYSIA: Pahang Province: 1♂ Bukit Fraser, 18.xi.1984, P. Lehtinen (ZMUT).

Chedima purpurea Simon, 1873: MOROCCO: 1♂ Taza, 25.ii.2004, D.W. Wrase (SMF); 1♀ Had Msila, 21.ii.2004, D.W. Wrase (SMF).

Diaphorocellus biplagiatus Simon, 1893: SOUTH AFRICA: Western Cape: 1♂ Beaufort-West, Farm Katdoornkuil, 3–6.xii.2007, D.H. Jacobs (NCA 2008/4672); 1♀ Beaufort-West, Farm Kantkraal, 3–6.xii.2007, D.H. Jacobs (NCA 2008/2607).

Hybosida dauban Platnick, 1979: SEYCHELLES: 1♂ Silhouette Isl., Mon Plaisir, 20.xii.1993, J. Gerlach (ZMUT); 1♀ same island, Gratte Fesse, 12–24.i.1999, J. Gerlach and M. Saaristo (ZMUT).

Levymanus gershomi Zonstein & Marusik, 2013: ISRAEL: ♂ holotype, 3♂ 2♀ paratypes Qetura, 8.v.2003, E. Topel (TAU).

Palpimanus sogdianus Charitonov, 1946: TAJIKISTAN: 1♂ 1♀ Beshkent, 17.iv.1989, S. Zonstein (TAU).

Palpimanus transvaalicus Simon, 1893: SOUTH AFRICA: Limpopo: 4♂ 1♀ Tuinplaas, 17.ix.2002, M. van Jaarsveld (NCA 2003/379).

Sarascelis junquai Jézéquel, 1964: Côte d’Ivoire: 2♂ 1♀ Titekro, 29.ii.1984, R. Schouten and J. Buysen (MRAC 166073).

Scelidocteus schoutedeni Benoit, 1974: D.R. CONGO: Équateur Province: ♂ holotype Kumungu, iv.1921, H. Schouteden (MRAC 15551).

Scelidomachus socotranus Pocock, 1899: YEMEN: Socotra: ♂ holotype Dahamis, xii.1898, W.R.O. Grant (NHML); 1♂ paratype Thluteen, xii.1898, W.R.O. Grant (NHML).

Photographs were taken in dishes of different sizes with a paraffin or cotton layer on the bottom. Specimens were photographed using a Canon EOS 7D camera attached to an Olympus SZX16 stereomicroscope and with a SEM JEOL JSM-5200 scanning electron microscope at the Zoological Museum, University of Turku. Digital images were processed using the “CombineZP” image stacking software (http://www.hadleyweb.pwp.blueyonder.co.uk/).

The width of the sternum was measured between the bases of the coxae of legs II. Illustrations of the female copulative organs were made after maceration in 20% potassium hydroxide aqueous solution and exposure for a few minutes in an alcohol/water solution of Chlorazol Black. Lengths of the leg segments were measured on the dorsal side. All measurements are given in millimetres.

Abbreviations (except encoded in the text):

ALE anterior lateral eyes

ALS anterior lateral spinnerets

AME anterior median eyes

CH carapace height

CL carapace length

CW carapace width

CyH clypeus height

PLE posterior lateral eyes

PLS posterior lateral spinnerets

PMS posterior median spinnerets

SL sternum length

SW sternum width

TL total length of body in dorsal view

Simon (1893) divided all the palpimanids known to him into two tribes, according to the eye pattern. The tribe Chedimeae was established for the spiders with anterior and posterior lateral eyes almost touching each other. Simon assigned the rest of the palpimanids (with widely spaced lateral eyes) into the nominative tribe Palpimaneae. Platnick (1975) elevated both the above-mentioned groups to subfamily rank. He also separated these taxa from another subfamily concurrently described in his work, the Otiothopinae, by possessing accessory terminal sclerites in the male bulb (present in the former groups, but absent in the latter). Additionally, the Otiothopinae have been found to be an exclusively Neotropical group (Platnick et al. 1999), while the subfamily Chediminae is a chiefly Paleotropical taxon, with only some representatives distributed also in a few adjoining far southern Palearctic areas (Simon 1873; Hu and Li 1987; Zonstein and Marusik 2013; El-Hennawy 2014; Zonstein et al. in press). The subfamily Palpimaninae is widely distributed in Africa, the Mediterranean Region, Middle East and Central Asia, but is very poorly represented in the Oriental Region (WSC 2016). The puzzling occurrence of Palpimanus argentinus Mello-Leitão, 1927 in South America is still based exclusively on the old collection data, and since description has not been confirmed by field studies. The only representative, collected by F. Silvestri, is kept in the Simon’s collection in Paris. Conforming to Mello-Leitão (1927), it originates from Salta (Argentina). However, on the labels (even those based on the received material), E. Simon rarely noted the collection country name, but mostly only locality. Thus, the specimen interpreted as South American could actually have been collected by F. Silvestri from Salto (Italy). In view of the complete absence of later records, this suggestion seems to be more reasonable than assuming that the species was occasionally introduced (see Platnick 1975).

For the above-mentioned reasons (structure of the male palp and the geographical confinement), the following discussion will deal with only two Old World subfamilies, Palpimaninae and Chediminae.

The nominative subfamily Palpimaninae currently comprises 2 genera and 37 very uniformly looking species: the widespread Palpimanus Dufour, 1820 (35) and South African Ikuma Lawrence, 1938 (2). Prior to this study, the more diverse subfamily Chediminae was considered to include 9 genera and 31 species: Boagrius Simon, 1893 (2), Chedima Simon, 1873 (1), Hybosida Simon, 1898 (4), Diaphorocellus Simon, 1893 (4), Levymanus Zonstein & Marusik, 2013 (1), Sarascelis Simon, 1887 (7), Scelidocteus Simon, 1907 (7), Scelidomachus Pocock, 1899 (1) and Steriphopus Simon, 1887 (4). The distribution of almost two-thirds of these species (23) is confined to tropical Africa (counted from WSC 2016). Most probably, the monotypic Badia Roewer, 1962 was misplaced in the Palpimanidae, and in any case it should be certainly excluded from the chedimine genera because, unlike the Chediminae, B. rugosa Roewer, 1962 was shown to have widely spaced lateral eyes (see Zonstein and Marusik 2013).

Since the representatives of Chedimanops gen. n. and Hybosidella gen. n. completely lack the lateral eyes, they cannot be assigned to any of the two Old World subfamilies by using such a simple and traditional criterion as the distance between ALE and PLE. Therefore, any reliable suggestion concerning the taxonomic position of the two newly-described genera should be based on the available characters. These potentially diagnostic features are surveyed below.

Body size: The Palpimanidae of the Old World can be divided into two informal groups. The first includes the tiny palpimanids 2–4 mm long, belonging to the genera Hybosida, Levymanus and Steriphopus (see Platnick 1979; Hu 2001; Zonstein and Marusik 2013). Small (approximately 3 mm) south Asian Boagrius pumilus Simon, 1893 also belongs to the latter group. The considerably larger African B. incisus Tullgren, 1910, with body length 7–8 mm (♂♀ syntypes in NRS, examined) is in our opinion misplaced in this genus. The latter group includes the rest of the chedimine genera and all representatives of the Palpimaninae. In members of Chedima, Diaphorocellus, Sarascelis, Scelidocteus, and Scelidomachus the body is 4.5–8.5 mm long (Simon 1873, 1907; Pocock 1899; Jézéquel 1964; Benoit 1974; Zonstein et al. 2016). In some smaller members of Scelidocteus it varies from 4 to 5 mm long (Simon 1907). According to Nentwig et al. (2016), in the Mediterranean species of Palpimanus, occurring in southern Europe, body length ranges from 4.4 to 7.6 mm. African representatives of Palpimanus and adult members of Ikuma are 4.5–13.0 mm long (see Pocock 1898, Simon 1907, 1910; Lawrence 1927, 1937, 1938, 1947, 1952, 1962; Lessert 1936; Benoit 1974).

Thoracic fovea: In most chedimine genera and in all representatives of the Palpimaninae, the fovea is represented by a narrow, short and deep, and bilaterally constricted longitudinal groove (as shown in Marusik and Zonstein 2013: figs 3–5, 7). In Scelidocteus, the fovea is anchor-like and widened posteriorly (Forster and Platnick 1984: fig. 283). A modified variant of this type can be observed in Hybosida and Steriphopus, possessing the angular and opened posteriorly triangular or Λ-shaped (lambda-shaped) foveal groove (Platnick 1979: fig. 1; Saaristo 2010: fig. 24.1). The posteriorly opening fovea is also characteristic for Boagrius pumilus (see Murphy and Murphy 2000: fam. 45: fig. 1). In Diaphorocellus, the right and left parts of the fovea show a certain tendency for further separation from one another (Zonstein et al. 2016: fig. 6). Finally, in Levymanus the thoracic fovea is represented by two separate sulci located side by side (Zonstein and Marusik 2013: figs. 10, 12; Zonstein et al. in press). It should be noted that all the mentioned modifications are characteristic for genera of the subfamily Chediminae, but not for those of the Palpimaninae (see Cerveira and Jackson 2005: fig. 1).

Abdominal scuta: Within the Palpimaninae, the anterior part of the abdomen overhangs the posterior part of the carapace, making a dorsal portion of the scutum de facto frontal and thus invisible from above in both males and females (see Cerveira and Jackson 2005: figs 1–3; Lecigne 2016: fig. 9G). In contrast, in the Chediminae the abdomen is more evenly expanded by the middle, and the dorsal portion of the scutum in a dorsal view becomes visible, almost always in males and very often in females (Platnick 1979: figs 1, 3; Hu 2001: figs 8–15; Deeleman-Reinhold 2001: fig. 76; Zonstein and Marusik 2013: figs 1–9, 12, 14, 15; Zonstein et al. 2016: figs 1, 3, 5).

Male copulatory organs: In the nominative subfamily, the embolus is represented with a fairly robust and rigid heavy-sclerotised process which seems to be quite commensurable with the “conductor” (or the tegular appendage, sensuJocqué and Dippenaar-Schoeman 2006: fig. 75i– j). This is true at least for those representatives of the Palpimaninae in which construction of the palpal organ has been examined and illustrated – in some Mediterranean species of Palpimanus Dufour, 1820 (Platnick 1981: figs 1–9), African P. crudeni Lessert, 1936, P. giltayi Lessert, 1936, P. pseudarmatus Lawrence, 1952, P. tuberculatus Lawrence, 1952, P. stridulator Lawrence, 1962, and P. lualabanus Benoit, 1974 (Lessert 1936: figs 15–18; Lawrence 1952: figs 19, 20; 1962: fig. 1e–f; Benoit 1974: figs 16–17), and Asian P. sogdianus Charitonov, 1946 (Marusik and Guseinov 2003: figs 29–32). Within the Chediminae, the embolus is thinner, weaker and more fragile and it is certainly smaller than the surrounding outgrowths. Most often, this difference may be very sharply expressed, e.g. in Hybosida (Simon and Fage 1922: fig. V2; Platnick 1979: figs 5–7, 10–12), Sarascelis (Jézéquel 1964: figs 7b, 9a–b, 11a), Steriphopus (Hu 2001: figs 8–15), Levymanus (Zonstein and Marusik 2013: figs 41–46), and Diaphorocellus (Zonstein et al. 2016: figs 11–15, 17, 18). The same holds true for Chedima and Scelidomachus (Zonstein and Marusik, in prep.). In some species of Scelidocteus the embolus appears stronger; however, even in this case it conforms to the general trend, being noticeably thinner than the “conductor” (Lessert 1930: fig. 4; Jézéquel 1964: fig. 2a–b; Benoit 1974: figs 8, 9, 12, 13; Dippenaar-Schoeman 2006: fig. 75i–j).

Female copulatory organs: The ventral surface of the epigastric scutum in females, carrying a pair of the fairly well expressed transversal S-shaped or V-shaped sclerotized marks, is very characteristic for the Palpimaninae (Lawrence 1927: fig. 53; 1938: fig. 3; 1962: fig. 1d; Kritscher 1996: fig. 1; Marusik and Guseinov 2003: figs 33, 35, marked as cl), but is rarely observed within the Chediminae (e.g. in Diaphorocellus – Lawrence 1927: fig. 54; Zonstein et al. 2016: fig. 2). Most chedimine genera lack such marks (see Platnick 1979: figs 8, 14; Saaristo 2010: fig. 3; Zonstein and Marusik 2013: fig. 13). In those species of Palpimanus in which the structure of the endogyne has been studied (the Mediterranean congeners, Asian P. sogdianus, African P. transvalicus), the receptacles are fairly large, extended and weakly sclerotized (as shown in Platnick 1981: figs 10–18; Marusik and Guseinov 2003: fig. 35). In contrast, in representatives of the Chediminae subjected to a special examination, the endogyne is shown to be more compact, with mostly round or oval, and more sclerotized receptacles (Jézéquel 1964: figs 1, 3, 6, 8, 10; Platnick 1979: figs 9, 15; Zonstein and Marusik 2013: fig. 49–50; Zonstein et al. 2016: figs. 20–26).

As has been shown above, Chedimanops gen. n. and Hybosidella gen. n. include very minute spiders, with males having the upper part of the abdominal scutum well-visible in the dorsal projection (Figs 3, 6, 7, 41, 42), and a fragile embolus surrounded by stouter conducting outgrowths (Figs 49–54, 56–60, 62–66). All these features are fairly characteristic for taxa within the Chediminae, while none of them have been found in the Palpimaninae (see above). Thus, regardless of the full absence of the lateral eyes, there is nothing to prevent these genera from being placed in the Chediminae.

Despite the fact that, unlike all other palpimanids, Chedimanops gen. n. and Hybosidella gen. n. comprise only two-eyed species, these two genera cannot be considered as closely related. Strictly speaking, there is nothing else in common, other than their similar diminutiveness, similar eye reduction, and the characters shared within the entire subfamily Chediminae (see above). In both species of Chedimanops gen. n., the carapace with a Λ-shaped and posteriorly opened thoracic fovea is combined with a spotted dorsal abdominal pattern (Figs 1, 3, 4, 6, 12, 15). The similar shape of the thoracic fovea is documented for Hybosida and Steriphopus (cf.Platnick 1979: fig. 1; Saaristo 2010: fig. 24.1), whereas a spotted abdominal pattern is characteristic for Chedima (see the corresponding description by Simon 1873). In contrast, like the majority of the chedimine palpimanids, Hybosidella etinde gen. et sp. n. has the carapace with a narrowly-closed thoracic groove and a uniformly pale-coloured abdomen behind the sclerotised shields (Figs 41, 42, 48). Within the Chediminae, the male palpal organ provided with a long and bent claw-shaped outgrowth, like that in Chedimanops gen. n., is characteristic for Chedima (Zonstein and Marusik, in prep.). The structure of the palpal organ in Hybosidella etinde gen. et sp. n. resembles that in Scelidomachus socotranus Pocock, 1899, with the amendment that the apically dilated conductor is bifurcate in the former, but tridentate in the latter taxon (cf. Figs 53, 54, 64–66 and Pocock 1903: p. 194, text-figure).