On Levymanus, a remarkable new spider genus from Israel, with notes on the Chediminae (Araneae, Palpimanidae)

Abstract Levymanus gershomi gen. n. et sp. n., is described from southern Israel. The eye arrangement and structure of the male palp indicate that this genus belongs to Chediminae Simon, 1893. Levymanus gen. n. differs from other chedimine genera by its unusually long and slender legs, an elongate body, a unique shape of the bipartite thoracic fovea, reduced leg scopulae, smaller spinnerets, and other characters, which are presumably apomorphic. We propose two taxonomic changes: 1) based on widely spaced lateral eyes the Western African genus Badia Roewer, 1961 is transferred from Chediminae to Palpimaninae, and 2) Fernandezina gyirongensis Hu & Li, 1987 from China, based on palpal morphology, is transferred to the Asian genus Steriphopus Simon, 1887 for a new combination Steriphopus gyirongensis (Hu & Li, 1987) comb. n.


Introduction
The Palpimanidae is a relatively small family of araneophagous spiders consisting of 131 species in 15 genera (Platnick 2013), distributed in tropical and sub-tropical zones worldwide and absent only in the Nearctic and Australia. The family was divided by Platnick (1975) into three subfamilies: Palpimaninae Thorell, 1870 (Africa and Eurasia), Otiothopinae Platnick, 1975 (almost entirely Neotropical), and Chediminae Simon, 1893 (mainly Paleotropical). The Chediminae includes taxa with closely spaced or touching lateral eyes (Palpimaninae have widely spaced lateral eyes) and with tegular sclerites (lacking in Otiothopinae). Currently, the Chediminae includes 30 species in 9 genera, three of which are monotypic (Platnick 2013). While studying spiders in Israel we found one species that seems to belong to Chediminae, but which had a peculiar carapace shape (especially the fovea) and strongly reduced scopulae. In order to allocate this taxon, the senior author examined representatives of all available genera referred to Chediminae -Boagrius Simon, 1893, Chedima Simon, 1873, Diaphorocellus Simon, 1893, Hybosida Simon, 1898, Sarascelis Simon, 1887, Scelidocteus Simon, 1907, Scelidomachus Pocock, 1899and Steriphopus Simon, 1893 (including the holotypes), as well as some specimens belonging to Colopaea Simon, 1893 (Stenochilidae). As a result we concluded that the specimens from southern Israel belonged to a new genus and species. The main goals of this paper are: 1) to diagnose and describe the new genus and species; 2) to discuss the relationships of the new genus; 3) to discuss the questionable position of some taxa within the subfamily Chediminae.

Material and methods
Specimens of the following spider taxa were studied.
Photographs were taken using a Zeiss Discovery V20 stereomicroscope with a Canon PowerShot G9 camera, and an Olympus SZX16 stereomicroscope with an Olympus E-520 camera, and prepared using the CombineZP software. Scanning electron micrographs were made using the SEM JEOL JSM-5200 scanning microscope at the Zoological Museum, University of Turku, Finland. Photographs of landscapes showing the surroundings of the type locality were taken by Vasiliy Kravchenko. Background maps were obtained from the public internet source http://www.maps-for-free.com. Measurements were made to an accuracy of 0.01 mm. Lengths of leg and palp segments were measured on the dorsal side, from the midpoint of the anterior margin to the midpoint of the posterior margin. All measurements are given in millimetres.
Etymology. Both the generic name and the specific epithet are given in honour and memory of Gershom Levy (1937Levy ( -2009, the prominent Israeli arachnologist, for his immense contribution to Israeli and Near East arachnological research. The gender is masculine. Diagnosis. In general appearance, especially by the elongate body and the extended dorsal abdominal scutum, Levymanus gen. n. resembles the otiothopine genus Fernandezina Birabén, 1951(cf. Platnick 1975Grismado 2002, fig. 1; Grismado and Ramírez 2008, fig. 4;Piacentini et al. 2013, fig. 5a-f), but can be easily distinguished from it by the presence of the accessory structures in the male palp, accompanying the embolus (Figs 38-48), characteristic for the Chediminae, but absent in the Otiothopinae. Both males and females belonging to the new genus are easily distinguishable from other palpimanids due to the characteristic bipartite thoracic fovea (Figs 10, 12).
Description. Small bicolored chedimine palpimanids with body length 2.0-2.5 in males and 2.5-3.0 in females; legs and abdomen without pattern. Carapace with corrugated cuticle, diamond-oval in dorsal view, narrowed anteriorly and posteriorly. Cephalic part somewhat raised behind eye area -slightly in males, and more noticeably in females. Thoracic fovea short and bipartite, with two separate sulci located side by side; posterior edge of carapace slightly raised (Figs 14-15). Eight eyes. ALE largest, about four to five times larger than other eyes, which are subaequal in size. ALE and PLE almost touching each other. PME widely spaced from each other, as well as from AMEs and from PLEs. Clypeus about two times higher than AME diameter. Chilum inconspicuous. Chelicerae small, equal in length with clypeus; stridulatory ridges absent; cheliceral furrow without true or peg teeth. Sternum shield-like with fine reticulation; labium about as broad at base as it is long. Prosoma posteriorly with short paired triangular extentions and narrow tubular structure (Figs 11, 13) entering pedicel tube of abdomen (= scutopetiolar apparatus sensu Saaristo and Marusik (2008)). Palps relatively short; legs I-IV long and slender. Leg formula: 1=4,2,3. Femora with well-developed scale-like microsculpture on the cuticle; scales weakly developed on patella and other segments. Femur I rather long and moderately swollen; patella very long (longer that tibia). Tibia and metatarsus I with weakly developed prolateral scopula. Leg tarsi slightly curved and ascopulate. Claw tufts weakly developed. Leg tarsi with two narrow and dentate claws. Abdominal scuta conforming a rather long and corrugated pedicel tube; dorsal portion of scutum with irregular posterior margin. Scutum in male larger than in female, its dorsal part longer than the ventral in the male (Fig. 15) and ventral and dorsal parts subequal in the female (Fig. 14). Although the dorsal and ventral parts of the scutum are fused, the seams are clearly visible in females ( Fig. 12) and the dorsal part is rather narrow. Spinneret group very small (Fig.  34). Sclerotised ring encircling spinnerets present but weakly raised. AMS tiny and domed (Fig. 32); PMS and PLS reduced to a few sessile spigots in females, absent in males. Male palp: patella very small without dorsal process; tibia enlarged (swollen); both articles sub-globular. Cymbium moderately long lacking processes and with clusters of setae: a bunch of setae near prolateral base (Figs 43,48), and larger setae on retrolateral side (Figs 44,48 Description. Male (holotype): Total length 2.55; carapace, sternum and labium intensive carmine-red; chelicerae, palps (entirely), coxa and femur I light reddish orange, other segments of leg I and entire legs II-IV pale yellowish red; abdomen milk-white with intensive reddish orange dorsal scutum. Carapace (Fig. 10): 1.10 long, 0.76 wide. Diameters of AME, ALE, PLE, PME: 0.10, 0.02, 0.02, 0.02. Interdistances: AME-AME 0.08, ALE-AME 0.05, ALE-PLE <0.01, PLE-PME 0.11, PME-PME 0.14. Chelicerae as shown in Figs 17-18. Sternum (Fig. 11)  Palp : femur short and swollen, 2 times longer than wide, subequal in length to tibia and slightly shorter than cymbium. Patella globular. Tibia without apophyses, strongly swollen, 1.5 times longer than wide, 1.3 times wider than femur. Cymbium narrow, shorter than bulb, without outgrowths, with two clusters of hairs on prolateral side (Figs 39,46). Bulb as wide (in widest part) as long (not counting embolic division and tegular process), tegulum with strong and long retrolateral tegular process (Tp); base of process with conical outgrowth (Co) and deep furrow (Tf); embolus fused with other sclerites of the embolic division (Ed). Embolic division, at- tached to tegulum by a flexible membrane, and bears embolus (Em) with retrolateral outgrowth (Eo), and lamella (La) located between embolus and cymbium. Female (paratype): coloration as in male. Total length 2.93. Carapace (Fig. 12): 1.34 long, 0.92 wide. Diameters of AME, ALE, PLE, PME: 0.10, 0.03, 0.02, 0.02. Interdistances: AME-AME 0.08, ALE-AME 0.05, ALE-PLE <0.01, PLE-PME 0.10, PME-PME 0.12. Sternum (Fig. 13)  wide at base. Measurements of palp and leg segments as shown in Table 1. Spermathecae weakly sclerotised, round, touching each other, each spermatheca with a pair of accessory glands (Figs 49, 50). Due to the weak sclerotisation the outline of the spermathecae and their ducts are not very clear.
Variability. Carapace length in males 1.01-1.12, in females 1.22-1.34. Coloration varies very slightly, recently moulted specimens are lighter. Distribution and habitat. The species is known only from the type locality (Qetura) represented by an extra-arid stony desert at 200-500 m above sea level. All specimens were collected with pitfall traps.

The composition and distribution of the Chediminae
Of the three recognized palpimanid subfamilies, the Palpimaninae was reviewed by  and the Otiothopinae was revised by Platnick (1975) and Platnick et al. (1999). On the contrary, the Chediminae was briefly surveyed only once, by Simon (1893).
The holotype of B. rugosa was not found. Only a single microslide containing a separate leg of this specimen is deposited in SMF (Julia Altmann, personal communication). We thus consider the holotype of B. rugosa to be lost as is the case in several other types from the same study (see Sierwald 1997, Lotz 2007. According to the original description, Badia possesses lateral eyes, ALE and PLE, widely spaced from each other (cf. Roewer 1961, fig . 3a), like in the members of the  Palpimaninae. Meanwhile, all chedimine genera recognised by Simon (1893Simon ( , 1898Simon ( , 1907 have these eyes (near) contiguous. This feature is considered as one of the key characters of the group and a reliable criterion to distinguish representatives of both subfamilies (see Forster and Platnick 1984, p. 76, fig. 282;Dippenaar-Schoeman and Jocqué 1997, pp. 239, 240, figs 100a, b, d-f). Hence, we conclude that Badia should be excluded from the Chediminae and transferred to the Palpimaninae.
Within three Oriental genera of the Chediminae, Boagrius, Sarascelis and Steriphopus, the two former genera possess much larger anterior median eyes (see Hu 2001, figs 8-15, 1;Jézéquel 1964, figs 5a-c;Deeleman-Reinold 2001, fig. 76); additionally, in males belonging to these two genera the additional palpal structures are either consid-erably longer (Boagrius) or look more massive (Sarascelis) than in Fernandezina gyirongensis. Furthermore, the palpal patella in males of Sarascelis is more or less hooked and the cymbium is either asymmetrical or with a well-developed lateral process. All these characters are absent in F. gyirongensis.
The third genus, Steriphopus (described originally under Pachypus Pickard-Cambridge, 1873 nom. praeocc.), possesses smaller AMEs (like in F. gyirongensis). Strictly speaking, at the first view other characters noted and figured by Pickard-Cambridge (1873) make Steriphopus dissimilar not only to F. gyirongensis, but also to any other palpimanids. According to the original description, the holotype male of S. macleayi (Pickard-Cambridge, 1873), the generotype, has a developed ventral scutum that extends almost to the spinnerets (Pickard-Cambridge 1873, pl. 16, figs 2b, 2c) and an unusually long cymbium which is figured as a structure about three times longer then the palpal tibia (Pickard-Cambridge 1873, pl. 16, fig. 2m). But it should be noted that these described features and the corresponding figures do not reflect the actual characters of the holotype we examined. Contrary to the description, the holotype possesses a moderately sclerotised abdomen and a large sub-globular palpal tibia that appears to be even slightly longer then the cymbium (Figs 8,9). Other characters of the holotype, including the broad-oval shape of the carapace and configuration of the bulb bear a certain resemblance to F. gyirongensis.
Therefore, the given species is provisionally placed in Steriphopus and the new combination is proposed: Steriphopus gyirongensis (Hu & Li, 1987), comb. n., with reservation and assumption, that this species may represent a separate chedimine genus, as yet undescribed (since we could not study the holotype, which is lost -Shuqiang Li personal communication).

Characters and relationships of Levymanus gershomi sp. n.
The distinctive characters of the new taxon are listed and discussed below. It should be noted that within the Chediminae some characters, such as the structure of the spermathecae and fine structure of the male palp are known only for a few species described or surveyed after the 1960s (Jézéquel 1964;Platnick 1979;Forster and Platnick 1984). Since we have to exclude these parameters from the comparison, our conclusions are thus preliminary (the putative apomorphies of the new taxon are marked A1-A7).
First and foremost, Levymanus gershomi sp. n. differs from all other chedimine palpimanids by having long slender legs (A1) and an elongate body (A2) -as shown in Figs 1-2. All examined members of Chediminae may be referred to the "standard" palpimanid type with a more or less compact or robust body and considerably shorter and thicker legs, as in Figs 3-9. Among other palpimanids, some species of Fernandezina (Otiothopinae) also have somewhat longer and thinner legs and a more elongate body (Platnick 1975, figs 80, 85;Grismado 2002, fig. 1). However, the modifications are considerably less strong than in L. gershomi sp. n. In addition, species of Fernandezina possess a much shorter pedicel tube (cf. Platnick 1975, fig . 88).
A dense prolateral brush of scopular hairs on the tibia, metatarsus and tarsus of leg I is very characteristic for the whole family Palpimanidae (Forster and Platnick 1984;Jocqué and Dippenaar-Schoeman 2006), though we found that in Steriphopus macleayi it seems to be less developed than in other examined palpimanids (cf. Figs 3-9). However, in L. gershomi sp. n. the scopula is even weaker and represented only by sparsely distributed spatulate hairs .
The structure of the palp in the new taxon does not differ strongly from that in other members of the subfamily. The most significant distinction is the presence of a tegular furrow (A7) (Figs 38, 40, 41, 43), a detail that has not been found in any other examined genera of the chedimine palpimanids.
We thus conclude that Levymanus gen. n. is distinct from all other genera within the subfamily. Moreover, the above-noted characters contrast with all other genera of Chediminae taken as a whole. Currently it is not certain whether this new taxon represents a separate subfamily or whether it should be considered only as a specialized chedimine palpimanid. This question might be answered in the course of a taxonomic revision and phylogenetic study of the Chediminae.