A revision of the genus Cicynethus Simon, 1910 (Araneae, Zodariidae), a tale of colour patterns

Abstract. The genus Cicynethus Simon, 1910 is revised. Apart from the type species C. peringueyi Simon, 1893, only known from a juvenile, it contains five species, all of which are here described or redescribed based on adults: C. acer sp. nov. (♂♀), C. decoratus (Lawrence, 1952) comb. nov (♂), C. floriumfontis Jocqué, 1991 (♂♀), C. mossambicus sp. nov. (♂♀) and C. subtropicalis (Lawrence, 1952) comb. nov. (♂♀). The species are characterized by the genitalia, but also by the colour pattern. The distribution of the genus is extended to northern Mozambique. Cicynethus hongfuchui is transferred to Storenomorpha hongfuchui (Barrion, Barrion-Dupo & Heong, 2013) comb. nov. Cicynethus acanthopus Simon, 1910 is considered a species incertae sedis. A key to the species is provided.


Introduction
The faunistic approach of taxonomy, that prevailed into the mid sixties of the previous century, led to many misplaced species and synonymies as explained in Platnick & Raven (2013). Consequently, the spider family Zodariidae was also riddled with nomenclatorial problems and insufficiently described genera before the revision by Jocqué (1991). Many genera were only known from juveniles (e.g., Systenoplacis Simon, 1907, Hetaerica Rainbow, 1916, sometimes leading to unnecessary synonymies (see Jocqué 2009).
A perfect example in this respect is the genus Cicynethus Simon, 1910 that, at the time of the family revision (Jocqué 1991), was only known from two species, both exclusively represented by juveniles. The redescription (Jocqué 1991) was based on a single male only and females have still not been described for these species. The present paper aims to redefine the genus, this time based on both sexes, correct the misplacement of a few species, including the species from China, and describe two new species, thereby considerably expanding the distribution of the genus in Africa.

R e s e a r c h a r t i c l e
urn:lsid:zoobank.org:pub:D73665B3-6043-4528-9A89-CCCBBC90D6B9

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. Female genitalia were digested using half a tablet of Total Care Enzima product (protein removal system originally for cleaning contact lenses and containing subtilisin A, 0.4 mg per tablet; Abbott Medical Optics, Santa Ana, CA) in a few millilitres of distilled water overnight and then immersed in 75% ethanol. Photographs of these female genitalia and male palps, as well as measurements and photographs of the habitus and details of mouthparts were taken with a Leica MZ16 A using the Leica Application Suite (LAS) automontage software (ver. 3.8). The habitus of some specimens was photographed by the focus stacking technique developed at the Royal Museum for Central Africa (part of the Digit 03 project) and explained in Brecko et al. (2014). For SEM photos, specimens were air dried (36 h), gold coated, examined and photographed with a JEOL 6480 LV scanning electron microscope. Illustrations were assembled and edited in Photoshop CS5 (white balance and colour contrast adjusted). Maps were created with the online tool SimpleMappr (Shorthouse 2010).
Types of the new species are deposited in the National Collection of Arachnids (Pretoria, South Africa) (NCA), the Ditsong National Museum of Natural History (TMSA, former Transvaal Museum) and the Royal Museum for Central Africa, Tervuren, Belgium (RMCA).
All palp illustrations are from left palps unless stated otherwise. All measurements are in mm unless otherwise indicated.

Diagnosis
Cicynethus are medium-sized, elongated spiders with eyes in two procurved rows and spination on legs reduced to a few small distal spines on the ventral side of the metatarsi. It differs from Chariobas Simon, 1893, which has the eyes in a close group, and from Thaumastochilus Simon, 1897, which has strong spines on the anterior leg pair.

Description
Body. Medium-sized to large spiders (9.2-14.4 mm) with smooth to finely granulate teguments. Carapace longer than wide (L / W < 1.5-2.2) and fairly flat, protruding anteriorly (Figs 12B, 16D), widest at level of coxae II-III, not much narrowed in front to about 0.6-0.65 times maximum width (cephalic width measured on strong frontal bend), with few hairs apart from a few longer hairs in front of fovea and on clypeus. Cervical grooves faint. Profile highest just behind eyes and with slight dip at level of fovea.
Colour. Carapace medium to dark brown; chelicerae, mouthparts and sternum medium to orange brown; leg colour complex, frontal legs often darker and more uniform than posteriors; abdomen dorsum grey with species-specific colour pattern. Remark: colour pattern often darker in vivo ( Fig. 15A-B), much more contrasted in alcohol.
EyEs. Eyes in two procurved rows (Fig. 5C). All eyes pale and subequal, AME usually the largest. MOQ quadrangular or slightly longer than wide. Clypeus straight, height 0.8 to 2 times diameter of ALE, with dispersed setae.
Prosoma. Chilum double, poorly sclerotized, about or slightly more than twice as wide as high, lateral margin poorly defined, with a few setae. Chelicerae without teeth, broad from base to tip with evenly dispersed setae; fangs shorter than wide at base. Labium diamond-shaped, widest in distal half, narrowed base. Endites roughly triangular, converging, with basolateral extension accommodating palpal coxa; with field of thick setae in distal half. Sternum elongated, roughly oval (Figs 8C, 9B, E, 12D), longer than wide, widest at middle, without triangular extensions; sometimes with a few precoxal sclerites (Fig. 16B); anterior margin straight or slightly concave.

Diagnosis
The male of Cicynethus acer sp. nov. is recognized by the sharp excrescence at the tip of the RTA and the retrolateral knob on the tegular swelling. The epigyne is similar to that of C. floriumfontis, but the median septum is provided with two conspicuous anterior arches in the latter.

Etymology
The species name 'acer' is an adjective meaning 'sharp' and referring to the sharp excrescence of the RTA. Colour ( Fig. 3A-C). Carapace medium reddish brown, darker on sides, with paler median spot near posterior margin and central dark band in front of fovea, widened behind and reaching ocular area; chelicerae medium brown with pale mediodistal patch; sternum medium brown, paler towards posterior, pale yellow tip, darker along margins; endites and labium pale yellow, darker along margins; legs pale yellow with all P medium brown on sides; leg I with distal half of T, Mt and t medium brown; leg II with Mt and t brownish yellow, legs III and IV with distal part of Mt and t brownish yellow; abdomen: dorsum with dark grey mottling provided with two median triangles and white median stripe in front of spinnerets; sides with dark grey mottling; venter pale with poorly defined dark stripes in posterior half. stErnum (Fig. 3B). Elongated oval, 1.42 wide, 2.06 long. No precoxal sclerites.

JOCQUÉ R. & HENRARD A., Revision of the genus Cicynethus Simon, 1910
EyEs. AME larger than other eyes: ALE 0.20; AME 0.16; PLE 0.15; PME 0.15; ALE-AME 0.10; AME-AME 0.07; PLE-PME 0.15; PME-PME 0.08. Clypeus 0.28 high.  Colour ( Fig. 4A-D). Carapace uniform reddish brown, with paler triangle behind fovea reaching posterior margin. Chelicerae reddish brown, with pale patch near fangs. Endites reddish brown at base, fading to yellowish in front; labium reddish brown, paler in front. Sternum uniform dark reddish brown, paler near posterior tip. Legs I with F yellow, distal part and remainder of legs orange; legs II yellow, with distal orange rings on F and T orange, P, Mt and t orange; legs III-IV yellow, with narrow distal ring on F, T and Mt; entire P and distal part of t orange. Abdomen pale with three longitudinal dark stripes, the central one with two faint triangles in front of white longitudinal bar; sides dark mottled grey; venter with two ill-defined longitudinal stripes. stErnum (Fig. 4C). Elongated oval, 1.49 wide, 2.41 long. Precoxal sclerites in front of coxae II poorly delimited.

Variation
In the juveniles (Fig. 5A-C) the colour pattern is much more contrasted than in the adults. The carapace has three well-defined dark bands and in the median stripe of the dorsal abdominal pattern, the two triangles in front of the white bar are much more clearly marked than in the adults.

Diagnosis
The female of Cicynethus decoratus comb. nov., is recognized by the epigyne with broad plate deeply indented in front and by the dark rings on the ventral part of all Fe and TII-IV. Colour (Fig. 7A). Carapace reddish brown, with paler median spot near posterior margin. Chelicerae reddish brown with pale patch near fangs. Endites reddish brown fading to yellowish in front; labium dark reddish brown, paler in front. Sternum uniform dark reddish brown, darkened along anterior margins. Legs yellowish orange; anterior legs darker reddish brown from P onwards; F with dark ventral distal spot; d F II with dark distal ring interrupted dorsally; F II-IV with dark ring on distal third; all P darkened on sides; T II with basal dark ring; T II-IV with dark ring on distal third. Abdomen cream, dorsum with dark dorsal pattern: lateral bands with three ill-defined triangles and median band consisting of diamond, stripe, 2 chevrons and 2-segmented spot in front of spinnerets; sides with one large frontal spot and two small posterior ones; venter with two longitudinal dark lines.
Colour (Fig. 7B). Carapace reddish brown, slightly suffused with black in cephalic region, with narrow long dark 'V' in front of fovea and four dark marginal triangles on either side. Chelicerae reddish brown with pale patch near fangs. Endites yellowish orange, darker at base. Labium reddish brown. Sternum medium brown, darkened along margin and with faint dark spots in posterior half; legs yellowish orange, darkened to medium brown on M tI, t I and t II, and extremity of t III and t IV; F and T II-IV with median ring on ventral half. Abdomen cream, dorsum with complex sepia dorsal pattern: diamond-shaped and 3-segmented black spot in front of spinnerets; sides with one large frontal spot and two small posterior ones; venter with two longitudinal dark lines. stErnum (Fig. 8C). Elongated oval, 1.14 wide, 1.63 long. Precoxal sclerites in front of coxae II.
EPigynE (Fig. 7E). With broad plate, longer in front; with deep median indentation; produced into median septum; frontal corners of plate surrounded by dark circular area.

Male
Unknown. Subadult male (Fig. 8E) with contrasted pattern on carapace and abdomen.

Distribution
Known from the vicinity of Port Edward (Fig. 21).

Variation
Total length: 9.23-12.57. Carapace: length 4.12-5.11, width 2.34-2.91. The lateral dark triangles on the carapace may be very vague (Fig. 7C). The median septum in the epigyne is sometimes strongly developed, as in the female from Ingogo Forest (RMCA_ARA_131806) (Fig. 7F, H). Juveniles (n = 16, including subadult males and females from RMCA_ARA_131806) have a yellow carapace with a medium brown pattern consisting of a median band widening from the fovea towards the front, including the eye region and margins with a row of dark coalescing triangular spots (Fig. 8E); sternum with broad dark margins; legs and abdomen as in the adult female. Jocqué, 1991 Figs 2A-C, 9-11, 21 Cicynethus floriumfontis Jocqué, 1991: 104, figs 236-241 (descr. ♂).

Diagnosis
Both male and female of Cicynethus floriumfontis are recognized by the colour pattern, with numerous small dark spots and the broad dark bands on the carapace, the central one not reaching the PME. Colour (Fig. 9A-C). Carapace brownish yellow with many tiny dark dots, brown median band widening towards the front, not reaching eye region; broad dark marginal bands with irregularly serrated median margin; chelicerae medium brown with darker condyle and dark spot near fang insertion and pale mediodistal spot; endites yellow; labium medium brown; sternum with yellow centre, darker margins  (2018) and medium brown anterior part (Fig. 9B); legs brownish yellow with many dark spots at setae sockets; spots on F coalescing into dark dorsal median stripe (Fig. 9A); all P, T I and II, Mt I with dark proand retrolateral stripes; distal dorsal margin of trochanters and coxae brown; abdomen cream, sparsely dotted with small darker spots; epiandrum orange; spinnerets yellow ( Fig. 9A-B). stErnum (Fig. 9B). Elongated oval, 1.63 wide, 2.20 long. No precoxal sclerites.

Variation
The colour pattern of the males, mainly that of the legs and the spots on the abdomen, may vary in intensity. The number of distal cymbial spines varies between two and five.

Distribution
Known from two localities in the Eastern Cape Province, South Africa (Fig. 21).

Natural history
No details are known on the habitat in which the specimens were collected. One specimen was 'caught in association with termites' without further explanation.

Diagnosis
Both male and female Cicynethus mossambicus sp. nov. are recognized by the uniform dark chestnut carapace and the hairy legs; males are recognized by the shape of the RTA, the simple, not strongly swollen tegulum and the distal tegular protrusion (DTP); females by the epigyne provided with broad procurved scape and the wide but short median septum separating the small copulatory openings.

Etymology
The species name is an adjective referring to the country where it is found. Colour (Fig. 12A-B). Carapace dark chestnut brown with small pale brown patch at posterior margin. Chelicerae dark brown; endites brown, with distal half white; labium brown, with distal margin white; legs: tarsi brown with pale ring at base; metatarsi dark brown; tibia I with broad dark ring at base and distal extremity, orange in between, tibia II similar to I but orange part longer; tibiae III and IV: yellow with dark rings at base and distal extremity; trochanters orange with brown parts; coxae dark brown, with large ventral yellow patch; abdomen cream with dark median stripe ending in two triangles followed by interrupted chevrons; sides cream with longitudinal stripe all along; venter pale, with two faint longitudinal stripes; spinnerets greyish brown.

Distribution
Known from mountain area in northwest Mozambique (Fig. 22). EPigynE (Fig. 15A). With posterior, slightly procurved plate, in the middle produced into median septum, delimiting large atria; large dark area on either side.

Note
Since the holotype was collected more than 110 years ago, it is not surprising that is bleached. We therefore provide the description of a more recent female in addition to the description of a male. Colour (Fig. 15A). Carapace reddish brown with darker median band in front of fovea, widening towards the front, reaching eye region; dark marginal bands with irregularly serrated median margin; chelicerae reddish brown with distal median part white; endites yellowish orange; labium reddish brown; sternum reddish brown with darker margins in anterior half; legs: all F yellow with distal dorsal extremity dark brown, remainder of first leg pair reddish brown; second pair with P brown, T yellow, Mt and t brown; third and fourth pairs with P yellow with brown sides, remainder yellow to slightly orange on Mt and t; abdomen base colour cream; dorsum with three dark stripes, the median one divided by thin white line anteriorly and by large white bar in posterior third; venter with white mottling; spinnerets yellow.
EPigynE (Fig. 17B). With wide posterior plate slightly procurved, in the middle produced into faint median septum delimiting large atria; large dark area on each side.

Variation
The colour of the live specimen ( Fig. 15A-B) is quite aberrant from what is observed in specimens kept in ethanol (Fig. 15C-G). The carapace of the live specimen is almost uniform dark brown except for a small paler patch at the posterior margin, whereas in all the preserved specimens there is a clear pattern composed of three dark bands interrupted by stretches of medium brownish orange. Also, the abdominal pattern is somewhat different: the dorsolateral dark bands are less pronounced than in the collection specimens. Considering the locality where the live specimen was photographed, we assume that its colour pattern falls within the variation observed in the species, with the characteristic contrasting white posterior stretch of the median abdominal band (Fig. 15A-G). The epigyne is also subject to quite some variation, as shown in the pictures (  The female specimens from Empangeni (2000/438) have an epigyne in which the contours of the dark internal structure (Fig. 17E-F) look slightly different from those of the other specimens ( Fig. 17A-D). However, since the colour pattern of all these specimens is similar and in the absence of males with different palps, we have refrained from describing another new species based on only two specimens.

Distribution
Known from several localities near the type locality in Kwazulu-Natal Province, South Africa (Fig. 21).

Systematic notes
Cicynethus acanthopus Simon, 1910, only known from a subadult female (Fig. 20E-F) from Namibia, is aberrant in many respects (see Jocqué 1991): the shape and the setation of the carapace, the shape of the labium, the height of the clypeus and the leg spination. All these characters point in the direction of another genus, not yet identified. We therefore consider C. acanthopus as 'species incertae sedis'.
Cicynethus hongfuchui Barrion, Barrion-Dupo & Heong, 2013 is not a member of Cicynethus and most probably an elongated species of Storenomorpha Simon, 1884. In that genus, the posterior eye row is strongly recurved, whereas it is clearly procurved in Cicynethus. The colour pattern and number European Journal of Taxonomy 465: 1-35 (2018) of spines on the legs, very few in Cicynethus, also point to Storenomorpha. One character mentioned by Barrion et al. (2013: 45), "maxillae yellow except short black serrula", is puzzling because one of the synapomorphies of the family Zodariidae is the absence of a serrula. We here present the new combination Storenomorpha hongfuchui (Barrion, Barrion-Dupo & Heong, 2013) comb. nov.

Discussion
The only specimen of C. peringueyi is a juvenile (Fig. 20C-D) from Cape Town and it remains questionable whether it belongs to one of the species described on adults. Its pattern resembles that of C. floriumfontis and it might thus be either a senior synonym of that species or a species that has never been collected again. The juvenile specimen from Port Elizabeth (MNHN AR 10010), on the other hand, has a pattern ( Fig. 20A-B) that is reminiscent of that of C. subtropicalis comb. nov., but the locality falls completely outside its presently known distribution area and is even separated from it by that of C. decoratus comb. nov. Considering the collecting efforts in the framework of the SANSA inventory (Dippenaar-Schoeman et al. 2015), it is surprising that no specimens of Cicynethus have been found in the vicinity of the two localities of these juveniles or along the coast in between these.
The microhabitat preference of Cicynethus is far from clear and puzzling. It is remarkable that most samples are small and there is not a single sample in which adults of both sexes are present. The majority of the specimens have been found at ground level, either in pitfall traps or by sifting leaf litter. But, as could be expected, other specimens were found in Malaise traps, or by sweeping the shrub layer. Representatives of the Storenomorphinae are, indeed, expected to live in tube-shaped retreats above ground level (Jocqué & Bosmans 1989;Leroy & Jocqué 1993;Jocqué 1994). However, the frequency with which Cicynethus is found in pitfalls might indicate that they have a mixed life style and spend part of their life cycle at ground level and part in the shrub layer. According to C. Haddad (pers. comm.), juveniles are primarily litter-dwellers and adults tend to move to foliage. The experience with C. subtropicalis comb. nov. has been to find adults in low shrubs (< 1.5m high) only, where they construct a tubular retreat, sticking several broad leaves together with silk. This observation is in accordance with what can be deduced from some of the samples listed above.
This brings us to the colour pattern. There is one large sample (n = 17) of C. decoratus comb. nov. in which both juveniles and one adult female are present (RMCA_ARA_131806). The colour pattern of the carapace of the juveniles is clearly different from that of the adult, which might point in the direction of a different life style. Yet, the colour pattern of the adults is apparently stable and clearly different between the species. This is quite unusual. Colour patterns are hardly ever used to a large extent to differentiate spiders living at ground level, and in the many revisions of Zodariidae, this character has been used only once, more precisely to differentiate some species in the key to females of Suffascar Henrard & Jocqué, 2017(Henrard & Jocqué 2017). Yet, in Cicynethus it is perfectly possible to combine the keys for males and females to identify the species almost solely on the base of their colour pattern. It was thus possible to identify the females from the Oribi Gorge Nature Reserve and Ingogo Forest Reserve as C. decoratus comb. nov. Surprisingly, the terminology used in both descriptions (the present one was carried out independently from the one by Lawrence 1952), is fairly different. This illustrates the phenomenon that there is hardly a standardized terminology to describe colour patterns.
NCA (Petro Marais, Ansie Dippenaar-Schoeman, Robin Lyle) and NMBA (Leon Lotz) for the loan of material. Alain Reygel is thanked for the drawings and Bennie Kruger for the picture of a live C. subtropicalis comb. nov. Charles Hadded made an important comment on the lifestyle of Cicynethus and made corrections to an earlier draft. Photographs of some spiders are available in the digital collections of the RMCA at http://digit03.africamuseum.be/home. This is publication number BRC 423 of the Biodiversity Research Center, Université Catholique de Louvain, Belgium.