A revision of Pachyballus Simon, 1900 and Peplometus Simon, 1900 (Araneae, Salticidae, Ballini) with descriptions of new species

Abstract Two genera from the tribe Ballini (Araneae, Salticidae), Pachyballus Simon, 1900 and Peplometus Simon, 1900, are remarkable for their resemblance to beetles. Their biology is, however, poorly known and taxonomy has hitherto been rarely analysed. Thirteen species are included in this taxonomic revision of the two genera. Six of them are new to the science: Pachyballus caelestissp. nov. (♂♀, Congo D.R.), Pachyballus miniscutulussp. nov. (♂♀, South Africa), Pachyballus mombasensissp. nov. (♂♀, Kenya), Pachyballus ornatussp. nov. (♂♀, Congo D.R. and Tanzania), Peplometus congoensissp. nov. (♂♀, Congo and Congo D.R.), and Peplometus nimbasp. nov. (♂, Guinea). One species (Pachyballus cordiformis Berland et Millot, 1941) and a subspecies (P. flavipes aurantius Caporiacco, 1949) are recognised as synonyms of Pachyballus flavipes Simon, 1910. One new combination is proposed: Peplometus oyo (Wesołowska et Russell-Smith, 2011) comb. nov. (ex Pachyballus). The previously unknown females of Pachyballus transversus Simon, 1900 and Peplometus chlorophthalmus Simon, 1900, along with the males of Pachyballus castaneus Simon, 1900 and Peplometus biscutellatus (Simon, 1887) are newly diagnosed and described. Neotypes for Pachyballus castaneus and P. flavipes are designated. Numerous new data on the distribution are provided here and a key to Pachyballus females and to the males of Peplometus is presented. Identity of one species remains doubtful, Pachyballus gambeyi (Simon, 1880).


MNHN
National The specimens were examined in 70% ethanol. The epigynes were macerated in cold 5% KOH for 24 hours, dehydrated with absolute ethanol, cleared in xylene and put in clove oil in a temporary microscope slides. A reticular eyepiece attached to a stereomicroscope was used for drawing. After examination and reverting the above described sequence, the female genitalia were placed to microvials and stored with specimens. Specimens were measured as in Metzner (1999); all dimensions are given in millimetres. A Nikon Coolpix 8400 or Canon EOS 550D mounted on the stereomicroscope was used to take digital photos, which were stacked using Helicon Focus image stacking software. Male-female matching was based both on the co-occurrence of specimens and morphological similarities between the sexes (e.g., shape and colouration of body). SEM microphotographs were taken with SEM Hitachi TM-1000. The photographed parts were dried, and then mounted on an adhesive specimen stub. The maps were prepared using DIVA-GIS.

Distinguishing genera
Due to numerous similarities of Pachyballus and Peplometus, many collectors have failed to distinguish the two genera and labelled the specimens simply as Pachyballus. All previously undetermined specimens analysed in this work had been assigned to Pachyballus before and mostly determined only to the genus level. Peplometus after the work by Simon (1901) was only found by Berland and Millot (1941) and mentioned by Bodner and Maddison (2012; for specimen from Ghana used in molecular phylogenetic analysis). Simon (1901) differentiated the two genera by the shape of their carapace -significantly wider than long in Pachyballus and elongate in Peplometus (Figs 7,8). However, Simon depicted this feature on Pachyballus transversus, which has the widest carapace among its congeners. In most species that can be assigned to this genus the ratio of carapace width and length is approximately 1:1 and it may overlap with the proportions seen in Peplometus. A better character allowing the separation of these genera is the shape of abdomen. It is clearly elongated in Peplometes, whereas in Pachyballus the length of abdomen is equal to its width. The other reliable feature for telling apart the two genera, which may be applied only to males though, is the form of the tibia I. These tibiae in Pachyballus are not modified (Fig. 129), but in Peplometus are always conspicuously altered. They are strongly thickened and often flattened , or considerably elongated, but in this case the metatarsus has a basal process (Figs 134,135). Tibia I in Pachyballus bears dense and long hairs on ventral surface (Fig. 129), whereas in Peplometus it has a "brush" of long leaf-like setae . The structure of tibia I of males may even be used in distinguishing Peplometus species. Identifying females in this genus is very difficult in general and the results may be doubtful. Conversely, species recognition by Pachyballus is relatively straightforward in females, where a combination of morphological characters and genitalia conformation is used. Determination of the males is sometimes impossible, because the structure of their palpal organs is extremely similar and there often lack reliable distinguishing morphological features. As a consequence of this, we were able to construct the key only for Pachyballus females and males of Peplometus. Most probably, many taxonomic intricacies will be possible to solve only using molecular methods. Fresh material for analysis of the two genera will, however, be difficult to obtain.

Taxonomic account
Pachyballus Simon, 1900Pachyballus Simon 19001901: 486. Type species. Pachyballus transversus Simon, 1900. Diagnosis. Pachyballus is closely related to Peplometus. From the latter genus it can easily be separated by the rounded abdomen (elongated in Peplometus), by the form of leg I in males (not modified) and by the absence of the leaf-like setae on tibia I.
Description. Small to medium-sized spiders (ca. 3.0-5.0 mm length), with very flat body, covered with tough highly sclerotised integument. Body colouration in the majority of species dark brown or black with metallic lustre, only legs (especially in females) lighter. Carapace rounded, its width usually slightly larger than the length, eye field clearly trapezoid. Posterior part of carapace covered with abdomen. Chelicerae with three (exceptionally two) teeth on promargin and two to four teeth on retromargin, basally fused together. Abdomen short and wide, heart-shaped or rounded, its length to width ratio is 0.8-1.1, clearly wider than carapace, with straight anterior border. Abdominal dorsum totally covered with strongly sclerotised scutum, its edges sloping so that the abdomen has shape of a shallow bowl. Venter with a narrow scutum along anterior margin, its lobes extending laterally as lobes. A second, trapezoid scutum in posterior part, numerous very small sclerotised bumps on both sides of abdomen (Fig. 1). Legs short, first pair slightly bigger in males, with enlarged femora. Tibia I not modified, setae on its ventral side not so dense as in Peplometus. Tarsus of the female palp usually black. Male palp with oval bulb, embolus spirally coiled around its tip (Fig. 3). Tibial apophysis thin and straight. Epigyne with large semi-circular depression (atrium), usually divided by epigynal septum, copulatory ducts usually long, forming a spiral in their initial parts and with several complex loops distally.

Key to Pachyballus (females only)
1 Ventral scutum in posterior half of the abdomen present (Fig. 2)  Diagnosis. This species is covered very densely with short hairs, which is the best feature to distinguish it from congeners. Male palpal organ has a characteristic embolus that forms a considerably high and narrow coil similar to that of P. castaneus. The coil comprises four loops (two and a half in the latter species). The female is distinctive in having the epigyne with broad, short ridge on posterior edge of the epigynal depression; the copulatory ducts are relatively straight (they do not form any loops).
Etymology. The specific name is from Latin, meaning "soaring" and refers to this species living high in a forest canopy.
General appearance as in Fig. 9. Small, very flat spider with strongly sclerotised, pitted integument. Carapace black, clothed in dense short light hairs. Eye field trapezoid, distance between anterior lateral eyes shorter than between posterior laterals. Eyes encircled by white scales. Clypeus low, with a few white hairs. Chelicera with two teeth on promarginal edge and four retromarginal teeth. Endites and labium light brown with whitish tips. Abdomen heart-shaped, wider than long, blackish with white scales on sides and posterior part. Anterior margin of abdomen covers posterior part of carapace. Venter brown, with typical scuta. Spinnerets black. Legs I the stoutest, femur and patella brown, tibia slightly thickened, black with long dark setae ventrally, metatarsus and tarsus yellowish. Tibia with one short ventral spine distally, metatarsus with two pairs of ventral spines. Legs II the same in colour as the first pair. Legs III and IV brown with yellow distal segments. Pedipalp yellowish, its structure similar to that in other species, embolus forms a considerably high and narrow coil that comprises four loops as in Figs 12-15.
General appearance as in Figs 10, 11. Slightly larger than male, shape of body similar. Carapace covered with white hairs. Colouration of abdomen a little lighter than in male, brown, blackish in the mid part and along edge. Palps light brown. Epigyne oval, central depression divided posteriorly by short, wide ridge (Fig. 16). Copulatory ducts wide, spermathecae slightly smaller than in other species, small accessory glands opening into copulatory ducts (Fig. 17).
Distribution. Known only from the type locality (Fig. 193).
Remarks. All specimens were collected by fogging. Probably, this species lives high in the forest canopy.  Simon, 1900 Figures 6, 18-28, 129, 193 Pachyballus castaneus Simon 1900: 400 (♀). Diagnosis. The male is indistinguishable from the males of P. flavipes and P. mombasensis by body shape and colouration, but its bulb is slightly narrower than in these species and the embolic spiral is tightly convoluted; width of the basal embolic loop equals only a half of tegulum width, whereas in both other species it is as wide as tegulum. The female can be separated from congeners in having copulatory ducts compactly arranged and not forming loose loops (see Fig. 28).
General appearance as in Figs 18, 19. Small spider with flattened body covered with strongly sclerotised and pitted integument (Fig. 6). Carapace dark brown, eye field on more or less half of carapace, vicinity of eyes black. A few long bristles at anterior eyes. Posterior edge of carapace covered by abdomen. Chelicerae with three teeth on both margins (Fig. 22). Mouth parts brown, sternum oval, brown, clypeus extremely low. Abdomen dark brown, heart-shaped, as wide as long, dorsal scutum turned back via margins to venter (Fig. 19), ventral scuta typical; the anterior one narrow with lateral extensions, posterior trapezoid. Numerous small bumps on sides. Colouration dark brown to black, iridescent, integument clearly pitted, short hairs on edges of carapace, dense bristles near eyes. Spinnerets brown. Legs I the thickest (Fig. 19), brown (except yellow tarsi and metatarsi), femur, patella and tibia slightly thickened, tibia slightly flattened dorsally, covered with dense setae ventrally. Tibiae and metatarsi with two pairs of short stout spines ventrally. Other legs yellow, with brownish femora. Pedipalp  brown, its structure as in Figs 23-26, slightly narrower than in congeners. Palpal tibia short with single thin straight apophysis, bulb oval, embolus thin, long, spirally coiled on bulb tip.
General appearance as in Figs 20, 21. Similar to male, abdomen almost round or oval, but its anterior margin almost straight. Abdominal venter with two scuta as in male (Fig. 21). All legs and palps dark yellow. Epigyne typical, with horseshoe-shaped anterior depression (Fig. 27). Internal structures relatively simple, copulatory ducts shorter than in congeners, compactly arranged, spermathecae strongly sclerotised (Fig. 28).
Immature specimens. Similar to adults, abdomen covered dorsally with one large scutum.
Distribution. Known from South Africa and Zimbabwe (Fig. 193). Designation of neotype. Pachyballus castaneus was originally described from Natal (South Africa) on the basis of a single female. The type specimen was lost (the collection manager informed us that the type could not be found in Simon's collection in MNHN). The original description is very superficial (only body size and colouration of legs are mentioned) and insufficient for identification of the species. Colouration of legs varies within different Pachyballus species (see Figs 32, 33, 104-106), so it cannot be used as a taxonomic character. The neotype, a female that originates from the same province as the type, is herein designated to stabilise the nomenclature.
Remarks. The male of this species is described here for the first time. Simon, 1910 Figures 1, 3, 5, 29-57, 194 Pachyballus flavipes Diagnosis. The male is almost indistinguishable from that of P. mombasensis, though it differs a little by having a protruding tibial apophysis while in the latter species tibial apophysis is adpressed to cymbium. The female resembles females of P. castaneus and P. mombasensis, but has very long copulatory ducts, forming several loops, whereas in two other species these ducts are relatively short (cf. Fig. 51 with Fig. 73 and Fig. 28).
General appearance as in Figs 29-31. Shape of body typical for Pachyballus; small, flat, with heart-shaped abdomen. Body covered with strongly sclerotised and clearly pitted integument. Colouration of body brown to black dorsally and dark brown ventrally, dorsum iridescent, almost hairless. Some light hairs near anterior eyes and below anterior median eyes. Anterior margin of abdomen covers distal part of carapace. Venter with typical scuta and small sclerotised bumps (Fig. 1). Cheliceral dentition variable, with three teeth on promargin and two, three or (exceptionally) four on retromargin . Tips of mouth parts whitish. First pair of legs the stoutest, femora and patellae dark brown, tibiae black with dense long dark setae ventrally and two pairs of very short thick spines (Fig. 31), dorsal part of tibia slightly flattened, metatarsi and tarsi creamy. Other legs with clear contrasts: brown femora, other segments light, only sometimes dark ring on tibiae proximally (Fig. 30). Palps dark, tip of cymbium light. Structure of palpal organ as in Figs 3, 37, 44-46.
General appearance as in Figs 32-36. Similar to male, abdomen more oval. Anterior median eyes surrounded by short light hairs. All legs yellowish (Fig. 35 Distribution. Species widely distributed in Africa (Fig. 194). Some of the records from Kenya were already mentioned in Dawidowicz and Wesołowska (2016).
Synonymisation. Caporiacco (1949) described P. flavipes aurantiacus from Kenya on the basis of a single female. According to him, the epigyne of this subspecies was the same as in P. flavipes and the only difference consisted of colouration. However, this feature is variable. Moreover, according to Berdondini and Whitman (2003) type specimen of P. flavipes aurantiacus kept in the Natural History Museum of Florence collection is juvenile. Thus, we recognise P. f. aurantiacus as a synonym of the name P. flavipes. Type of P. cordiformis was destroyed and only two palps and three legs persisted in the vial. Although Berland and Millot (1941) reported that they had collected only a single male by the city Man, the two palps in the sample are left palps, so they must have been taken from two males. Structure of palpal organ and the figure in Berland and Millot (1941) suggest that this species is identical with P. flavipes. Therefore, we recognise the name P. cordiformis as the synonym of P. flavipes.
Designation of neotype. Simon (1910) described this species from Bioko (Fernando Po) on the basis of a single female. Wanless and Clark (1975) compared a female from Ivory Coast with the type and concluded their conspecificity. Unfortunately, the type specimen was lost (we were informed that the type could not be found in Simon's collection in MNHN) and the female collected by the latter authors lacked epigyne. Original description was insufficient to recognise the species; only the shape of body and colouration of legs were given. Taking this under consideration it is justified to propose the neotype to stabilise the nomenclature. The neotype, a female collected in Cameroon, the nearest continental country to Bioko (ca. 40 km), is herein designated. Remarks. This species probably lives both in the forest canopy and in the understory. Given this fact, i.e. the variety of preferred microhabitats, the large geographical range and high variation of cheliceral dentition, it is possible that it consists of several cryptic species. The sole morphology may be insufficient to solve this taxonomic problem and there is a need to support further analysis with molecular methods.

Pachyballus gambeyi (Simon, 1880)
Homalattus gambeyi Simon 1880: 166 (♂). Pachyballus gambeyi Simon 1901: 485;Prószyński 1987: fig. on  Remarks. The holotype was examined by Prószyński (1987). His drawings show that this specimen has palpal organ typical for Pachyballus, but morphological characters do not allow any specific identification. Simon described this species from Canala (21°32'S, 165°57'E), central New Caledonia. All species of Pachyballus are restricted to Africa, which is thousands kilometers from New Caledonia. This distribution pattern is highly improbable, and the situation might have resulted simply by mislabelling the sample, which is not an uncommon problem in collections. As the provenience of the specimen is doubtful we conclude that the status of P. gambeyi remains unclear.  Diagnosis. This species is distinctive in having a unique size of ventral posterior scutum (Figs 59, 61) that is clearly smaller than in other species. Its width is equal to spinnerets area (2-3 times larger in the congeners). The female has the epigyne similar to that in Pachyballus mombasensis, but the copulatory ducts are longer (cf. Fig. 74 with Figs 66, 67).
Etymology. The specific name is derived from the Latin words "mini-" and "scutum", meaning "small" and "shield" correspondingly, and refers to the small size of ventral posterior scutum.
General appearance as in Figs 58, 59. Colouration of carapace dark brown, with black rings around eyes, some long bristles at first row of eyes. Chelicerae dark brown. Clypeus and cheeks dark brown, covered with sparse white hairs. Labium and endites yellowish brown, paler apically. Sternum yellowish brown. Abdomen heart-shaped, dark brown dorsally. Venter brownish grey, with a small posterior scutum, ranging at one fifth of abdomen length (Fig. 59). Book-lung covers yellow. Spinnerets yellowish brown. First pair of legs brown with yellow tarsi. Legs II-IV light brown. Leg hairs brown. Structure of palpal organ as in Figs 62, 63, embolic coil wide, comprises 2.5 loops, palpal tibia with protruding apophysis.
Body flattened, integument strongly sclerotised, clearly pitted. Colouration of carapace dark brown, some bristles by the first row of eyes. Mouth parts brownish with lighter tips. Chelicera with three retromarginal teeth. Abdomen very flat, heartshaped, dark brown, venter with typical scuta. First pair of legs brown, tibia short, slightly thickened, its dorsal part a little flattened, two pairs of ventral spines and dense dark setae. Other legs yellowish, only femora brownish. Palpal organ as in Figs 70, 71, embolic coil wide and low, tibial apophysis adpressed to cymbium.  Female. Measurements. Cephalothorax: length 1.3, width 1.5, height 0.5. Eye field: length 0.7, anterior width 1.1, posterior width 1.5. Abdomen: length 2.1, width 2.1.
General appearance as in Figs 68, 69. Similar to male, first pair of legs not stouter than others, palps dark. Posterior ventral scutum large. Epigyne with semicircular depression (Fig. 72). Internal structure as in Figs 73, 74, copulatory ducts clearly shorter than in congeners, spermathecae comparatively large, strongly sclerotised. Immature specimens. As adults, dorsum of abdomen covered with one large scutum.
Distribution. Known only from the type locality (Fig. 195). Diagnosis. A characteristic body pattern allows an easy recognition of this species; the abdomen is light with dark margins and a dark streak or central patch (Figs 78,81). Some specimens, probably young adult females (soon after moulting), can be uniformly black and resemble females of P. flavipes, but they are distinguishable in having dark palps (that are yellow in the latter species).

Pachyballus ornatus
Etymology. The specific name is Latin and means decorated, which refers to the characteristic colour pattern.
General appearance as in Figs 75-80. Body flat, integument strongly sclerotised, clearly pitted. Carapace wider than long, eye field trapezoid (distance between anterior lateral eyes shorter than between posterior laterals), occupies about half of carapace.
Colouration of carapace brown, black around eyes, in some specimens with two large yellowish patches in eye field (Figs 76,78,80). Dense bristles at eyes of first row and at lateral edges of carapace. Chelicera with three short teeth on promargin and sawshaped tooth with four tips on retromargin. Labium and endites brownish with lighter tips, sternum rounded, brown. Abdomen very flat, triangular to heart-shaped, wider than long, its anterior margin almost straight. Colouration of abdomen yellow, brown on the edges, with brown longitudinal median streak (Fig. 77), which is reduced to round patch positioned centrally in some specimens (Fig. 79). Variation of pattern as presented in Figs 76-80. Abdomen with typical ventral scuta (Figs 2, 75). Spinnerets yellow. First pair of legs slightly stouter than other legs, femora enlarged, tibiae slightly thickened, brown, only tarsi light. Other legs yellowish to light brown, femora darker. Palps as in Figs 86, 87, embolic coil low, but basal loop high.
General appearance as in Figs 81-84. Body shape and colouration similar to that in male. First legs not enlarged. Palps dark. Epigyne with anterior semi-circular de- Distribution. Tanzania and Congo D.R. (Fig. 195).
Remarks. Some specimens, probably young females (soon after the moulting), were dark, almost black (Fig. 84). They appear to become brighter with age due to guanine accumulation (light spots are formed by guanine crystals stored under integument). Black females were collected together with light specimens. Internal structure of epigynes similar in light, Figs 92-94, and black, Figs 85, 95-97 specimens. Colouration seems to turn brighter in the course of life also in males. Simon, 1900 Figures 98-122, 191, 192, 196 Pachyballus transversus Diagnosis. The body proportions of this species are different than in other Pachyballus spp., namely width of carapace and width of abdomen are clearly greater than their length. Sigilla are strongly marked. Shape of the eye field is more trapezoid than in congeners, its width at posterior eye row is a quarter larger than anterior width.
General appearance as in Figs 98-102. Slightly larger than P. flavipes, body flattened, covered with hard integument, pitted. Carapace wide, dark brown to blackish, vicinity of eyes black. Eye field strongly trapezoid, more than in other Pachyballus spp. Its width at last row of eyes is a quarter larger than its anterior width. A few hairs and long bristles at anterior eyes. Clypeus low. Chelicerae with short fang, three small teeth on promargin and four teeth with fused base on retromargin (Fig. 110). Mouth parts brown, sternum oval, dark. Abdomen heart-shaped, wide, clearly wider than long. Colouration of abdomen usually light, orange brownish with blackish edge, sometimes yellowish or brown. Sigilla clearly visible. Venter brown, with typical scuta (Fig. 98). First pair of legs brown, slightly larger than other pairs, with femora and tibiae thickened, tibiae slightly flattened dorsally. All legs brownish with lighter tarsi. Palps dark, structure as in Figs 103, 111, 112. Female. Measurements. Cephalothorax: length 1.4-1.6, width 1.6-2.0, height 0.7. Eye field: length 0.7-0.8, anterior width 1.2-1.5, posterior width 1.8-2.0. Abdomen: length 2.4-2.9, width 2.8-3.2.
Immature specimens. Shape of body as in adults, abdomen with two oval scuta, close to each other on dorsum (Figs 191,192). Ventral scuta absent.  (Fig. 196). Remarks. Simon (1900) described only the female of P. transversus. He recorded this species from Congo, Transvaal (South Africa) and Zanzibar (Simon 1900(Simon , 1901. He found it also (Simon 1910) later in Guinea Bissau and noted in this publication that this species had been described from Congo. This suggests that the Congolese specimens constitute the "type material". However, the two specimens from Congo, personally labelled by Simon and kept in the MNHN collection, are males. The fact was also mentioned by Berland and Millot (1941). Simon most probably described these specimens and apparently published the wrong data on sex of specimens. This description is very superficial and the structure of copulatory organs is not depicted. There are also some other males from Zanzibar in Simon's collection in MNHN (but no female). Simon's specimens from Natal, kept in MZC (examined) are immature and probably misidentified. Thus, it should be concluded that Simon's description concerned the male and the female of this species is described here for the first time. The figure in Simon (1901) shows characteristic body proportions of P. transversus (width of carapace greatly exceeds its length, eye field is clearly trapezoid with long distance between eyes in the posterior row, and abdomen is rounded), which allows the proper species recognition.  Lessert, 1925Figures 123-128, 193 Pachyballus variegatus Lessert 1925.
Diagnosis. This species is the only one in the genus that does not have the posterior ventral scutum on abdomen. Its other diagnostic feature is the pattern on abdomen; the anterior one-fourth of the abdomen is bright, the posterior part dark with a wide, light median patch (Figs 123, 127).
General appearance as in Fig. 123. Body flattened, integument strongly sclerotised, clearly pitted. Eye field light, yellowish orange, blackish rings around eyes, thoracic part light brown anteriorly, posterior slope yellowish. A few delicate hairs at anterior eyes. Chelicerae brown, with three teeth on promargin and a tooth with two tips on retromargin (Fig. 124). Labium and endites basally brownish, with light tips. Sternum light brown. Abdomen heart-shaped, widest anteriorly and tapering, anteriorly orange, sides brown with wide orange area in the middle (Fig. 123). Venter yellowish, without scutum. Spinnerets short, yellow. Legs relatively short, dark yellow (first pair absent in the syntype, only coxae extant, brown). Pedipalp (only right present) brown. Palpal organ as in Figs 125, 126, embolic coil wide, with three loops.
Remarks. Lessert (1925) wrote in the original description that the basic colour of body was black. The specimens must have bleached heavily, however the outline of lighter patches has been preserved.
Distribution. Known only from the type locality (Fig. 193).

Diagnosis.
Peplometus is closely related to Pachyballus. From the latter genus it can easily be separated by the elongated abdomen (rounded in Pachyballus). It differs also in the form of leg I in males having the leaf-like setae on tibia I (absent in the other genus).
Description. Small spiders (ca. 3.0-4.0 mm length), with very flat body, covered with hard strongly sclerotised integument. Carapace almost square-shaped, pitted dorsally, its length only slightly exceeds its width. Chelicerae with two (exceptionally three) small teeth on promargin and on retromargin a saw-shaped tooth with four to five denticles. Abdomen slightly elongated, ratio of its length to width 1.3-1.5 (only in female of P. biscutellatus 1.2). In males, width of abdomen almost equal to width of carapace, abdomen of females wider and rather oval. Anterior margin of abdomen straight. Hard scutum covers dorsum of abdomen, venter also strongly armoured, with scuta as in Pachyballus, namely the narrow scutum with backwards extending "horns" at anterior edge, and a large trapezoid scutum in posterior half of abdomen. Males with a thinner sclerotised plate in front of epigastric furrow, its thick posterior margin forms narrow wedge-shaped bar (Figs 138, 185). Numerous sclerotised bumps on sides of the abdominal venter (Fig. 138). Legs short, in males the first pair slightly thicker, with enlarged femora and modified tibiae. In some species conspicuously thickened, usually flattened, in other species elongated (in this case the metatarsus I with a dorsal process). Tibiae I black, with ventral brush of long dense flattened setae, contrasting with other segments. Copulatory organs similar to those in Pachyballus. Tibia I with a distal process, metatarsus shorter than tibia (Fig. 134) ...P. oyo -Tibia I without process, length of metatarsus equal to length of tibia (Fig.  135)  3 Tibia I not flattened (Fig. 133)  Tibia I flattened dorsally (Fig. 131) , 130, 136-153, 189, 190, 197 Homalattus biscutellatus Simon 1887: 263 (♀). Peplometus biscutellatus Simon 1901: 486;Berland and Millot 1941: 398. General appearance as in Figs 136, 137. Body flattened, completely covered with strongly sclerotised and pitted integument. Dorsum of body dark brown, clothed in dense short white hairs (lost in some of the studied specimens). Eye field on more than half of carapace length, black rings around eyes, the first row of eyes encircled with white hairs. Long white bristles between anterior eyes. Clypeus extremely low, brown with a few white hairs. Chelicerae with three diminutive teeth on promargin and five retromarginal teeth fused basally (Fig. 139). Mouthparts brown. Sternum oval, brown. Abdomen elongated, widest anteriorly and tapering rearward, anterior edge almost straight. Dorsum of abdomen totally covered by very strongly sclerotised and pitted scutum, laterally slightly turned back to venter. Venter with typical scuta; anterior scutum narrow, extending backwards laterally, posterior ventral scutum largest, cordiform. In front of epigastric furrow a thin sclerotised plate with thick posterior margin forms narrow wedge-shaped bar. Numerous bumps on sides ventrally; armour-plate pattern is shown in Fig. 138. Spinnerets brown, obscured by dorsal scutum. Legs I the biggest, femora enlarged, brownish with darker prolateral side, three short spines on dorsum. Tibia thickened, clearly flattened prolaterally, blackish with pale dorsal streak, ventrally black brash of dense long feathershaped setae (Figs 130, 140), metatarsi and tarsi short, yellowish, two pairs of ventral spines on metatarsi. Other legs pale yellowish with dark marks at bases of segments, only femora IV brown (or yellow with brown streak on prolateral side). Palps light, their structure as in Fig. 142. Palpal tibia short with single thin straight apophysis, bulb oval, embolus thin, long, spirally coiled, forming three loops on bulb tip.
Immature specimens. Abdomen not elongated, heart-shaped, with two oval dorsal scuta on abdomen, close to each other (Figs 189,190).
Remarks. The first description of the male is given here. Simon described P. biscutellatus based on a female from Ivory Coast, however in his samples from Sierra Leone the two sexes were present (labelled by Simon himself ). This fact has already been mentioned by Berland and Millot (1941), but they have not described the missing male. Material collected by Simon is in a very poor condition. Numerous specimens have lately been collected by fogging, this species typically inhabits canopy.
Distribution. West equatorial Africa (Fig. 197). Simon, 1900 Figures 131, 132, 154-171, 197 Peplometus chlorophthalmus  Diagnosis. The tibia of the leg I in male is strongly flattened dorsally (Figs 131,158). The female is difficult to distinguish from P. biscutellatus, but it has a narrower abdomen (see Diagnosis of the latter species).
General appearance as in Figs 154-156. Small spider with flattened body. Carapace dark brown to blackish, pitted. White bristles around anterior median eyes and between all eyes of first row. Clypeus low, black, with a few white hairs. Fang of chelicerae short, two small teeth on promargin, retromarginal tooth with four tips (Fig.  160). Sternum oval, dark brown. Abdomen narrow, covered with strongly sclerotised, pitted, dark integument. Venter with typical scuta (same as by P. biscutellatus). Legs yellowish with dark lines along femora and tibiae III and IV on prolateral surface. Legs I the stoutest, tip of prolateral side of femur with small patch, tibia strongly thickened, black with yellow line along dorsum and large light patch on retrolateral side, long flattened black setae ventrally (Figs 157, 159). Tibia I characteristic for having a large flattened part on dorsal side (Figs 131, 158). Pedipalp light, its structure as in Figs 161-164, embolic coil wide. Female. Measurements. Cephalothorax: length 1.0-1.4, width 1.1-1.3, height 0.5-0.6. Eye field: length 0.6-0.7, anterior width 1.0-1.2, posterior width 1.2-1.3. Abdomen: length 2.0-2.4, width 1.5-1.8. Similar to male, general appearance as in Figs 166, 167. White hairs at anterior eyes and on posterior carapace slope. Abdomen wider than in male, but relatively narrow, narrower than in P. biscutellatus. Legs darker than in male. Femora of I-III legs brown with yellow ventral surface, femora IV completely brown. Patellae IV with pro-and retrolateral brown stripes. First leg not larger, its tibia black with black long setae ventrally, in some specimens with narrow light streak along dorsum. Tibiae II with prolateral brown stripe, tibiae IV brown with thin yellow longitudinal stripes dorsally and ventrally. Other leg segments light yellow. Palps blackish. Epigyne as in Figs 168, 169, rectangular with shallow depression. Ventral structure of epigyne similar to other species, copulatory ducts long, weakly sclerotised in initial part, forming several loops (Figs 170, 171). Immature specimens. Abdomen not elongated, heart-shaped, with two oval scuta on dorsum, close to each other, not covering whole dorsum of abdomen (Fig. 165).
Remarks. The first description of the female is given here. Simon described only the male of P. chlorophthalmus, although the vial with a type specimen contains also an undescribed female and a few immature specimens. This material is however in a very poor condition.
Length of the apical part of embolus varies. It is very long in South African specimen, extending beyond the retrolateral edge of apical part of cymbium (Figs 161,162), and short in specimen from Congo D. R. (Figs 163, 164).
Distribution. Known from Congo and South Africa (Fig. 197).  Diagnosis. The most characteristic feature of this species is colouration and shape of the first pair of legs. Tibia I in male is totally black, it is not flattened (Fig. 172), whereas in congeners tibia is flattened dorsally or laterally and has a light streak or patch. Tibia I of the female is light retrolaterally, with large dark patch at its basis (Figs 179,180). The abdomen of female is narrow, similar as in male, while females of other species have abdomen wider than males. Etymology. The species is named after its terra typica (Congo). Description. Male. Measurements. Cephalothorax: length 1.7, width 1.8, height 0.6. Eye field: length 0.9, anterior width 1.3, posterior width 1.7. Abdomen: length 3.0, width 2.2.
General appearance as in Figs 172, 173. Body flattened, integument strongly sclerotised and clearly pitted. Carapace trapezoid, widest posteriorly, dark brown, black around eyes, a few bristles at eyes of first row. Chelicera with two small teeth on promargin and four-tip retromarginal tooth. Mouth parts and sternum light brown. Abdomen elongated, only slightly wider than carapace, venter also covered by scuta, posterior scutum large (Fig. 173). Legs I stout, brown, lateral side of femur slightly darker, tip of patella dark, tibia strongly thickened, black, with long dense black feather-shaped setae ventrally (Figs 133,172). Other legs yellowish brown, with distal small dark brown patches on patellae and tibiae II-III and proximal brown rings on patellae, tibiae and metatarsi IV, femora IV brown. Palpal organ as in Figs 177, 178, apical part of embolus long.
General appearance as in Figs 174-176. Carapace rounded, brown to black, strongly sclerotised and clearly pitted, eye field on half of its length. Some dark bristles at eyes of first row. Chelicerae yellowish brown, with short fang, two small teeth on promargin and three-tip tooth on retromargin. Mouth parts creamy. Sternum oval, blackish. Abdomen relatively narrow, elongated, widest anteriorly, anterior edge almost straight. Dorsum of abdomen totally covered by very strongly sclerotised, pitted, black scutum, turned back on its margin. Abdomen ventrally with hard scuta; anterior scutum narrow along anterior margin of abdomen, extending backwards, posterior scutum large, trapezoid. Numerous bumps on sides of abdomen venter (Fig. 175). Spinnerets dark. Legs generally light, creamy with blackish stains. Legs I the biggest (not so markedly as in males); femora with black patch distally on prolateral side, tibia thickened, prolaterally black, basal half of retrolateral part black (Figs 179,180). Dense long black setae on ventral side of tibia, two pairs of ventral spines. Dark line along dorsum of femora IV. Palps blackish. Epigyne large, rectangular, with small depression divided by median ridge, hardly visible horizontal crevices laterally (Fig. 181). Internal structure as in Figs 182, 183.
Distribution. Known only from Congo and DR Congo (Fig. 198). Diagnosis. Male of this species is very similar to that of Pachyballus oyo, it can be distinguished by the lack of a conspicuous, sharp process on tibia I, which is very characteristic for the latter species (compare Fig. 135 with Fig. 134). In P. nimba metatarsus and tibia of leg I are of equal length, while in P. oyo metatarsus I is clearly shorter, about half of tibia I length.
Etymology. The specific name is a noun in apposition referring to the Nimba Mts, type locality of this species.
General appearance as in Fig. 184. Body very flat, covered with strongly sclerotised integument, clearly pitted, brown with blackish area around eyes. Carapace slightly trap-  ezoid, white bristles near eyes of first row. Clypeus very low. Chelicerae with two teeth on promargin and five on retromargin (apical tooth very small), fang short. Endites, labium and sternum yellow. Abdomen elongated, shield shaped (its anterior margin almost straight), dorsum covered with large strongly sclerotised scutum. Ventrally abdomen with typical large scuta (Fig. 185), as in other species. In front of epigastric furrow clearly visible narrow wedge-shaped sclerotised swelling. Spinnerets yellow. First pair the stoutest, femora basically yellow, slightly darker on sides; patella light yellow, tibia slightly thickened, brown (prolateral surface darker), with light streak dorsally, dense long whitish (probably bleached) setae ventrally; metatarsus long, creamy with black dorsal hump and black line along prolateral side (Fig. 187). Legs II-IV yellow, with thin dark line along prolateral sides of patellae and tibiae. Legs without spines, except two short ventral spines on metatarsus I. Palps yellow, tibial apophysis very thin, bulb triangular, embolus spirally coiled on bulb tip (Fig. 188). Female unknown. Distribution. Known only from the type locality, Nimba Mts in western Africa (Fig. 198). Russell-Smith, 2011), comb. nov. Figures 134,198 Pachyballus oyo Wesołowska and Russell-Smith 2011: 588, f. 126-135, 232-234 (♂♀). Diagnosis. The male of this species is similar to that of Peplometus nimba, but it can be identified by the presence of a big apical process on the first pair of legs and the metatarsus of this pair shorter than tibia (Fig. 134).

Mimicry
The mimicry is a common phenomenon among spiders (Pocock 1909). Jumping spiders imitate various models, mainly ants (overview in Cushing 1997Cushing , 2012, but also pseudoscorpions (Platnick 1984), flies (Morrison 1981), wasps (Reiskind 1976(Reiskind , Żabka 1992, velvet wasps (Edwards 1984, Wesołowska 2009  Thompson 1995) e.g. Cylistella Simon, 1901, Coccorchestes Thorell, 1881, Planiemen Wesołowska and van Harten, 2007. The Ballini genera Pachyballus Simon, 1901 andPeplometus Simon, 1901 also resemble beetles, most probably from the family Chrysomelidae, which is expressed in their body shape and a very strongly sclerotised integument. The microsculpture of the integument and bright or iridescent colouring emphasise this resemblance. In juveniles of all Peplometus species and Pachyballus transversus the two dorsal scuta on the abdomen are arranged in a pattern similar to elytra of beetles (Figs 165,(189)(190)(191)(192), it is the unique feature among spiders. This type of resemblance in spiders usually suggests a Batesian type of mimicry. On the other hand, the two presented genera live in the canopy and their body type (small, flattened, colours often matching their preferred habitats) may suggest that it is a type of camouflage. The resemblance to other tree-living arthropods could be just a result of a shared cryptic body pattern.