Ballomma, a new Afrotropical genus in the Cryptothelinae (Araneae, Zodariidae): eyes on the run

Ballomma gen. nov. is described in the zodariid subfamily cryptothelinae. It is characterized by the pairs of adjacent anterior median eyes and anterior lateral eyes and triangular palpal tarsus in the female, the concave tegulum with thin embolus in the male. Apart from the type species B. erasmus gen. et sp. nov. (♂♀), the genus contains B. haddadi gen. et sp. nov. (♂♀), B. neethlingi gen. et sp. nov. (♂♀), and B. legala gen. et sp. nov. (♀). A key to the species is provided. The restricted distribution of the genus is discussed.


Introduction
The Cryptothelinae remain a subfamily within the Zodariidae in which numerous novelties are to be expected. The recent description of Palindroma Jocqué & Henrard, 2015 with the discovery of a novel synapomorphy for the family, is symptomatic for this lack of knowledge. The subfamily has its distribution centre in southern Africa. Although South Africa has been fairly well inventoried in the last few years by the SANSA initiative (Dippenaar-Schoeman et al. 2015), many new taxa remain to be discovered such as in Limpopo, the northernmost province of the country. The area is characterized by a complex mosaic of biomes, with a high degree of endemism. Some of the mountain areas are among the oldest on the African continent and harbour species that belong to old evolutionary lineages (Jocqué et al. 2013). The species Australutica africana Jocqué, 2008 is one of these (Jocqué 2008). It was found in a thoroughly inventoried area in the Soutpansberg (Foord et al. 2008). Many of the other less important mountain ranges have only been inventoried superfi cially, but the few collections that have been made there, reveal a highly diverse fauna, often containing species with surprising affi nities.
Here we describe a new genus with four new species, all collected at medium altitude in the Limpopo Province. One of the most remarkable characters of the genus is the position of the ALE which are very close together, an eye arrangement which is particularly rare. These eyes would not be recognized as ALE without the observation of intermediate positions between lateral and central in related genera.

Material and methods
Specimens were observed, drawn and measured with a WILD M 10 stereo microscope. Details of the female genitalia and male palps were observed with a Zeiss Stemi 2000 stereo microscope. Measurements and photographs of the habitus, details of mouthparts, detached male palps and female genitalia were taken with a Leica MZ16 using the LAS automontage software (ver. 3.8). The female genitalia were dissected and digested with pancreatin then immersed in 75% ethanol.
For scanning electron micrographs (SEM) photos, specimens were dried in Hexamethyldisilazane (36h), gold coated and examined and photographed with a JEOL 6480 LV scanning electron microscope. Types are deposited in the National Collection of Arachnids, Pretoria (NCA) and the Royal Museum for Central Africa, Tervuren, Belgium (MRAC).
All measurements are in mm. All palp illustrations are from left palps.

Diagnosis
Ballomma gen. nov. is a member of the Cryptothelinae characterized by the eye position with touching ALE and AME, the shield-shaped sternum without or with tiny precoxal sclerites, the female palp with triangular fl attened palpal tarsus, the male palp with concave tegulum accommodating the thin, slightly curved embolus directed diagonally over the tegulum and the epigyne with cul-de-sac ducts.

Etymology
The genus name is derived from the combined Greek terms βαλλω (= to touch) and ομμα (=eye), referring to the touching anterior lateral eyes. The gender is neuter.

Affi nities
Ballomma gen. nov. clearly belongs to the Cryptothelinae, a subfamily of the Zodariidae that has been characterized by nomenclatorial problems: poor defi nitions and misplacements (Jocqué 1991;Jocqué & Henrard 2015). It has the modifi ed endites and an eye position of a type that only occurs in that subfamily. The present genus has some superfi cial resemblance with Capheris Simon, 1887 with which it shares the triangular female palpal tarsus and the closely set AME and ALE. However, in Ballomma gen. nov. the eyes are even more closely set and characters of sternum, carapace profi le and structure of the genitalia are all different. As for the latter character there is a superfi cial resemblance with Aschema Jocqué, 1991 from Madagascar in which the ALE and AME are also very closely set. The shape of a number of undescribed Aschema species have somatic and genitalic characters that come close to those of the Ballomma gen. nov. species in the present article. However, the characteristics of the sternum and carapace and of the genitalia, and the modifi ed leg IV, clearly separate them from the genus here described.

Description
HABITUS. Medium size to small spiders (4.5-7.5) with smooth teguments. Carapace longer than wide (L / W < 1.4-1.6), without setae apart from a few longer hairs on clypeus; widest at level of coxae II-III, narrowed to about 0.43-0.66 times maximum width in eye region (cephalic width measured on posterior tangent of PME). Cervical grooves poorly indicated. Profi le: fl at; fovea in shallow depression.
COLOURATION. Carapace medium brown; chelicerae, legs, mouthparts and sternum medium to orange brown; abdomen dorsum with complex dark pattern on pale background; in some cases with four apodemes, sides grey, venter pale sometimes with dark pattern; in males sclerotized in front of epigastric fold and slightly sclerotized in front of spinnerets.
EYES. In three rows; ALE in front of AME and touching each other; AME close together sometimes touching; posterior row recurved, eyes apart. All eyes pale and subequal. Clypeus retreating, height 2.5 to 3.6 times diameter of AME, with some dispersed setae.
PROSOMA. Chilum double, sometimes poorly delimited, slightly more than twice as wide as high; no setae. Chelicerae conical with many evenly dispersed setae; fangs shorter than wide at base. Labium triangular, narrowed base. Endites roughly triangular, converging, with basolateral extension accommodating palpal coxae. Sternum shield-shaped, as wide as long or slightly longer than wide, without triangular extensions or precoxal sclerites; anterior margin straight, lateral margins slightly sinuous.
LEGS. Slender; formula 4123. Spination reduced on legs I and II, well developed on III and IV with spines numerous on T and Mt III and IV. Spines slender. With few or without hinged hairs. FEMALE PALP. With tarsus triangular and fl attened, with row of prolateral thorns, retrolaterally with some thin spines; palpal claw with some small teeth at base; turned inward over less than 35°; without distal patch of chemosensitive setae.
ABDOMEN. Oval; tracheal spiracle small, somewhat advanced. Females and juveniles with six spinnerets. Female ALS large, conical, biarticulate. PLS without, PMS with one cylindrical gland spigot. Colulus represented by fi eld with few setae. MALE PALP. Patella sometimes with swelling; RTA variable; cymbium with well developed basal fl ange and with group of truncated subapical thorns; subtegulum strongly developed, with transverse ridges; tegulum with prolateral concavity, distally with dorsal and ventral extensions; accommodating thin, slightly curved embolus, directed diagonally over tegulum. Epigyne simple, with internal structure characterized by cul-de-sac ducts, visible in transparency.

Distribution
Ballomma gen. nov. is so far only found in the mountain forests of the Limpopo Province in northern South Africa.

Diagnosis
The male of B. erasmus gen. et sp. nov. is easily recognized by the fi eld of spines in front of the epigastric fold and the RTA with three short elements. The female is characterized by the S-shaped ducts in the anterior half of the epigyne.

Distribution
Only known from the type locality in the Limpopo Province, South Africa (Fig. 14).

Diagnosis
The male of B. haddadi gen. et sp. nov. is easily recognized by the peculiar palp with the long, thin, sinuous embolus provided at its base with a large, sharp process pointing forward. The female has a characteristic epigyne with dark central area from where procurved cul-de-sac tubes originate, followed by well-separated spermathecae.
LEGS. Without hinged hairs. ABDOMEN. Epiandrum well developed with oval membranous patch with two comma shaped marks along posterior margin and two tiny darker spots along frontal margin. MALE PALP (Figs 7A-B, 8A-B). Patella with pro-and retrolateral hair tufts. tibia with dorsolateral apophysis, roughly triangular and pointed, retrolaterally concave; cymbium with basolateral slightly indented protrusion, seven distolateral spines; tegulum high, in the middle with retrolateral protrusion with rounded tip; distally with membranous part ending in long, sickle-shaped apophysis (could be homologous to MA); embolus originating on posterior part of tegulum, thin, slender, curved over central part of tegulum, with large sickle-shaped sclerite at base. Female (largest of additional material) MESUREMENTS. Total length 7.46; carapace 3.34 long, 2.13 wide, narrowed to 0.92 in eye region.

Distribution
Known from the Limpopo Province in South Africa (Fig. 14).

Etymology
The species name is a noun in apposition taken from the type locality.

Remark
The tube contains four juveniles and three subadult males, most likely conspecifi c with the holotype. with complex pale pattern on dark grey background; venter pale with three longitudinal bands of darker mottling; spinnerets yellow. CARAPACE. Finely reticulate. Clypeus with dispersed setae. Endites with dense fi eld of small spinules in distal half (Fig. 9E). With small crescent-shaped precoxal sclerite at coxae II and III.
LEGS. Hinged hair on T I and II; preening brush on II-IV poorly developed. PALP. Tarsus triangular, slightly fl attened ventrally, with row of prolateral short spines; claw with three teeth, not turned inward. EPIGYNE (Fig. 10A-D). Roughly quadrangular; anterior ducts clearly visible, forming a V-shaped pattern, followed by transverse row of globular structures.

Distribution
Only known from the type locality in the Limpopo Province, South Africa (Fig. 14).

Diagnosis
The male of D. neethlingi gen. et sp. nov. is easily recognized by the fi eld of spines in front of the epigastric fold and the palp with a retrolateral patellar swelling and a broad triangular transparent RTA. The female has a characteristic epigyne in which the ducts in transparency are in the shape of an X with a slight extension where upper and lower parts of X meet.

Etymology
The species name is a patronym for Jan Neethling a pseudoscorpion specialist and the collector of the holotype.
Other material examined SOUTH AFRICA: 5 juveniles, same data as holotype.
LEGS. Hinged hair on TI and TII. Palp with tarsus triangular and ventrally fl at; provided with many prolateral and few retrolateral spines; pectinated claw turned inward over > 30°. EPIGYNE (Figs 12C, 13C). Roughly rounded area with ducts visible in transparency forming X-shaped pattern, slight extension where top and bottom parts meet.

Distribution
Known from the Limpopo Province in the northern part of South Africa (Fig.14).

Discussion
Most genera in the Zodariidae have a fairly large distribution area. This is partly due to the fact that the family has escaped far-reaching splitting. Genera do indeed remain largely subjective systematic levels and whether the species-groups are deeply split or not, makes an evident difference for their distribution. In general, genera are far too small (see Jocqué et al. 2013) thus obscuring patterns of evolution and distribution. But in the Zodariidae, most genera are fairly well delimited on the base of somatic characters which makes the grouping plausible and has prevented unnecessary splitting on the base of genitalic divergence (Jocqué 1990(Jocqué , 2009Nzigidahera et al. 2011). Only genera with an apparent relictual distribution, like Suffrica Henrard & Jocqué, 2015, tend to have smaller distribution areas. This probably also applies to Ballomma gen. nov., which turns out to sit near the base of genealogic tree of the Zodariidae (unpublished). A prediction of the genus' distribution ( Fig. 15) on the base of climatic data (DIVA-GIS 2011) expects a much larger distribution area than presently known in South Africa, overfl owing in Botswana, Zimbabwe and Mozambique. Predictions on the base of climatic data for other zodariid genera have shown that areas are sometimes largely overestimated because the calculation cannot take geographical barriers into account. Mallinella Strand, 1906, for instance, is expected to have a large area on Madagascar (unpublished data), but is not present there probably because it was unable to reach the island. The infl uence of geographical barriers is expected to be particularly important for those genera that occur in habitats that are likely to be disrupted like mountain forest which is apparently preferred by the species of Ballomma gen. nov.
Species with the anterior lateral eyes touching have so far only been found in Ballomma gen. nov. and in Aschema, endemic to Madagascar. The apparent endemicity of Ballomma gen. nov. to a few highlands in the Limpopo Province and their affi nities with endemics from Madagascar emphasizes the special character of these habitats and the fact that they are inhabited by taxa from old evolution lines. Further inventories of these montane forest patches might reveal the presence of more taxa representing old evolution lines and makes the study of these mountains a priority for the SANSA project.