A revision of the Afrotropical spider genus Cambalida Simon, 1909 (Araneae, Corinnidae)

Abstract The non-mimetic Afrotropical spider genus Cambalida Simon, 1909, placed within a subfamily of predominantly ant-mimicking spiders (Araneae: Corinnidae: Castianeirinae), is revised. Three species are transferred from Castianeira Keyserling, 1879 to Cambalida: Cambalida deminuta (Simon, 1909), comb. n., Cambalida fulvipes (Simon, 1896), comb. n. and Cambalida loricifera (Simon, 1885), comb. n.. A fourth species, Cambalida fagei (Caporiacco, 1939), comb. n., is transferred from Brachyphaea Simon, 1895 to Cambalida. Two species, Castianeira depygata Strand, 1916, syn. n. and Cambalida mestrali Lessert, 1921, syn. n., are considered junior synonyms of Cambalida fulvipes. The males of Cambalida deminuta and Cambalida loricifera are redescribed and their unknown females are described for the first time. The female and male of Cambalida fulvipes and Cambalida coriacea Simon, 1909 are also redescribed. The type material of the type species of the genus, Cambalida insulana Simon, 1909 from Pagalu (Annobon) Island, is lost, and only immature specimens have been subsequently collected from a nearby island. The species is regarded as a nomen dubium until fresh adult material can be collected. A replacement name, Cambalida simoni nom. n. is proposed for Cambalida fulvipes Simon, 1909, the latter being a secondary junior homonym of Cambalida fulvipes (Simon, 1896). The type material of this species is also lost and it is too considered nomen dubium. The following new species are described: Cambalida compressa sp. n. from West Africa, Cambalida dippenaarae sp. n. from southern Africa, Cambalida griswoldi sp. n. and Cambalida lineata sp. n. from Madagascar, and Cambalida unica sp. n. from Cameroon. Notes are provided on the biology of each species and the distribution of the genus in the Afrotropical Region.


introduction
The spider genus Cambalida Simon, 1909, endemic to the Afrotropical Region, was initially described in the subfamily Micariinae of the Clubionidae by Simon (1909). Reiskind (1969) also listed this genus in the Clubionidae in his revision of North American Castianeirinae, but was uncertain of its subfamily placement. Brignoli (1983) listed the genus in the Gnaphosidae: Micariinae, while Platnick (1989) subsequently placed Cambalida and many of the other micariine genera in the Liocranidae. Dippenaar-Schoeman and Jocqué (1997) already listed Cambalida in the Corinnidae: Castianeirinae, but did not formally transfer this genus. Bosselaers and Jocqué (2000) only recently transferred Cambalida from the Liocranidae to the Corinnidae based on characters it shares with other members of the Castianeirinae, particularly regarding genitalic structure, a placement confirmed through subsequent phylogenetic analyses Jocqué 2002, Haddad et al. 2009).
During this study it became apparent that this small genus posed a large number of taxonomic problems that needed resolution. For example, Simon (1909) described the three species of Cambalida from females only, and in the same paper described four species in the genus Castianeira Keyserling, 1879 from males only, raising the possibility that some of these sexes could possibly be matched, which turned out not to be the case. Unfortunately, several of these types are lost, including those of the type species of Cambalida (C. insulana Simon, 1909), raising problems in clarifying the taxonomic status of these species. Bosselaers and Jocqué (2000) recently redescribed C. coriacea Simon, 1909, a species that Simon (1909) considered close to the type species but with less sclerotisation, and based on this redescription all castianeirines considered congeneric with C. coriacea are considered in this revision to be true Cambalida.
The validity of C. coriacea was recently put under threat by the discovery that Castianeira fulvipes Simon, 1896 may be a senior synonym. This would have resulted in a nomenclatorial change Cambalida fulvipes (Simon, 1896), which is a senior homonym of Cambalida fulvipes Simon, 1909. I proposed to the International Commission on Zoological Nomenclature that C. coriacea have priority over its secondary senior homonym, which has been an unused name since its description (Haddad 2006). However, this proposal was rejected by Kraus (2006), and consequently Cambalida fulvipes (Simon, 1896) retains priority (ICZN 2007) and a new name (C. simoni nom. n.) is proposed for the secondary junior homonym in the current study. Incidentally, the type material of this species is lost and thus C. simoni nom. n. is considered a nomen dubium. Detailed examination of the genitalic morphology indicates that C. coriacea is, in fact, a good species and that the proposed synonymy with C. fulvipes would be incorrect, as was my earlier proposed conservation of the junior name (Haddad 2006). The present study uncovered a rich diversity of species in the Afrotropical Region, many of which have very similar male embolic structures .
Cambalida are castianeirines with relatively unspecialised colouration (Figs 1-4), which contrasts with many genera in this subfamily that mimic ants in both colouration and behaviour. It should be noted that Cambalida immatures display behaviour similar to mimetic castianeirines, moving the front legs up and down to resemble antennal movements of ants. This behaviour was only rarely observed in adults. Cambalida are entirely ground-dwelling and are mainly associated with savanna and forest habitats on the continent, although two species occurring in southern Africa are also found in drier grassland, Nama Karoo and/or fynbos habitats.
The genus is revised here for the first time and ten species are recognised, of which five are described as new. Based on current data, Cambalida is considered endemic to the Afrotropical Region.

Material and methods
Material used in this study was observed in 70% ethanol using a Nikon SMZ800 stereomicroscope for descriptions, digital photographs and measurements. The epigynes and male palps of representative specimens were dissected and cleaned in a Branson 3200 ultrasonic bath for 10 minutes in 70% ethanol, after which they were drawn. Digital photographs of the male emboli of each species, as well as the dorsal habitus of C. fulvipes and C. lineata sp. n., were taken using a Nikon Coolpix 8400 mounted on a Nikon SMZ800 stereomicroscope. Additional photographs were taken of carapace and abdominal structures of C. fulvipes and the holotype female of C. fagei (Caporiacco 1939). The photographs were then stacked using Combine ZM software (http://www. hadleyweb.pwp.blueyonder.co.uk) to increase depth of field. Material for scanning electron microscopy was dehydrated through a graded ethanol series and then critical-point dried in an argon chamber. Specimens were then glued to aluminium stubs and sputter-coated three times with gold for 2 minutes, and subsequently studied in a JEOL WinSEM at 10kV. Digitized micrographs were taken of the morphological structures examined.
All measurements are given in millimetres (mm). Total body length measurements were determined for the smallest and largest specimens of each sex to indicate size variation, and body, eye and leg measurements are given for the specific specimens indicated. Descriptions of the eye arrangements are given for the anterior view of the anterior eye row and dorsal view of the posterior eye row.
Remark. The majority of the specimens examined have a much lighter colouration than their congeners, as described below, i.e. a yellow carapace and legs and yellow abdomen with black mottling. The remaining specimens had a brown body and legs with similar markings to the majority of congeners. Since the genitalic morphology is very stable in all of the specimens examined it is clear that this species is a case of colour polymorphism.
Distribution. Widespread in West Africa but only known from a few scattered localities (Fig. 65).
Biology. Several records come from agroecosystems (fallow, sorghum and rice); the rest are from riparian forest and savanna habitats. Diagnosis. Females can be recognised by the relatively long entrance ducts that are initially directed anterolaterally before looping posteriorly, medially and then anteriorly before entering triangular ST II (Figs 61, 62). Males have an embolus with a triangular basal section, and distal section of the coil narrow and tapering to a sharp point (Figs 51,63,64).
Distribution. Widespread in central and western Africa (Fig. 65).
Biology. This species has been collected from a variety of tropical forest types, mainly by pitfall trapping, litter sifting and by hand.  Remarks. The lectotype is the larger of the two C. deminuta males in the type series. Fernand Vaz is a river in the D.R. Congo and no specific locality is mentioned in the original description or on the label accompanying the types.
Distribution. Widespread in central and western Africa (Fig. 79).
Biology. This species has been collected from a variety of tropical forest types, mainly by pitfall trapping, litter sifting and by hand.
Etymology. The species name is a patronym in honour of Ansie Dippenaar-Schoeman, in recognition of her contributions to the study and promotion of research on African arachnids.

Cambalida fagei
Diagnosis. Females of this species share with C. deminuta the very large ST II and compact ST I, but can be distinguished from that species by the much larger copulatory openings (Figs 77, 78).
Remarks. The holotype female is in very poor condition but the eye arrangement (Fig. 74) and abdominal sclerotisation (Figs 75,76) are consistent with the placement of this species in Cambalida. No material other than the holotype could be found and it will not be redescribed here. The epigyne of the species is distinct, notably the very large ST II and compact ST I (Figs 77, 78), and it is clearly different to C. fulvipes, which also occurs in Ethiopia.
Diagnosis. Females of this species can be recognised by the broad curved epigynal ridges and the nearly round ST II of the female epigyne (Fig. 80). Males have a curved basal section of the embolus and a distal section that is gently curved towards the tip of the embolus (Figs 54, 82).
Remarks. The larger of the two females in the type series is designated here as the lectotype.
Female ( Carapace dark orange-brown with black mottling, clypeus slightly paler, eye region darker; black striae radiating from fovea towards palps and leg coxae; surface finely granulate, sparsely covered in white plumose setae. All eyes with black rings; AER procurved, ALE much larger than AME; AME separated by distance approximately 3 ∕ 5 their diameter, AME separated from ALE by 1 ∕ 6 AME diameter; clypeus height slightly less than 1½ AME diameter; PER procurved, PLE slightly larger than PME; PME separated by distance slightly less than their diameter, PME separated from PLE by distance slightly more than ½ PME diameter; CW:PERW = 2.39:1. Chelicerae deep orange-brown with black mottling on anterior surface, pale orange-brown proximally and along prolateral distal margin; three teeth on promargin, median tooth largest, proximal and distal teeth smaller and subequal, distal tooth situated closest to median tooth; two slightly separated subequal teeth on retromargin; endites mottled dark brown, fading to yellow and cream prolaterally; labium mottled dark brown, cream distally; sternum deep orange-brown with dark brown mottling, except at setal bases, giving speckled appearance. Legs finely granulate; femora I brown with black mottling, except along dorsal midline, yellow dorsally at distal end; femora II and III yellow with broad black mottled ring at ¾ their length; femora IV yellow, with dark brown ring with black mottling from ½ their length, yellow at distal end; patellae I-IV yellow with faint black mottling, dark around patellar indentation; tibiae, metatarsi and tarsi I-III yellow with faint lateral mottling; tibiae IV yellow-orange with broad incomplete mottled black ring medially, mottling absent dorsally, proximally and distally; metatarsi IV yellow with faint black lateral mottling; tarsi IV yellow. Leg spination: femora: I pl 1 do 3, II do 3, III pl 2 do 3 rl 1, IV pl 1 do 3 rl 1; patellae with do 1 long distal seta; tibiae: I plv 1 rlv 1, II rlv 1-2, III pl 2 rl 2 plv 2 vt 2, IV pl 2 rl 2 plv 2 vt 2; metatarsi: I plv 2 rlv 2, II plv 2 rlv 2, III pl 2 rl 2 plv 2 rlv 2 vt 3, IV pl 3 rl 3 plv 2 rlv 2 vt 3. Palpal spination: femora do 2, patellae pl 1 spine and do 2 short setae, one proximally and one distally, tibiae pl 1 do 1 plv 1, tarsi pl 1 plv 3 rlv 1. Abdomen mottled dark grey, with cream spots around sigilla, fine cream chevrons in posterior ⅔ of abdomen and small white spot above spinnerets; dorsal scutum mottled dark brown, extending 1 ∕ 5 abdomen length; venter mottled pale grey, darker towards spinnerets, epigastric sclerite brown and inframamillary sclerite yellow-brown. Epigyne with broad curved epigynal ridges with lateral copulatory openings (Fig. 80); copulatory ducts initially directed laterally, looping sharply towards posterior then transversely towards midline, bending at nearly a right angle before entering ST II posteromedially; ST II somewhat round, joined broadly to large kidney-shaped posterior ST I (Fig. 81).
Biology. This is the Cambalida species that occupies the greatest range of habitats, from tropical and temperate forests, to savannas, grasslands, karoo and fynbos. Although generally scarce in agroecosystems, this species has been caught in the ground cover layer of pistachio nuts in the Northern Cape Province of South Africa (Haddad and Dippenaar-Schoeman 2006), maize fields in Kenya and South Africa, and rice paddies in West Africa.
Feeding in C. fulvipes follows a similar pattern to that observed for other corinnids such as Graptartia (Haddad 2004). Prey is grasped using the first two pairs of legs, which form a basket in which the prey is subdued. Following the bite, prey may die within 1 minute (e.g. vinegar flies Drosophila melanogaster Meigen), after which feeding commences. Once complete, only a small ball of macerated prey remains is left. Other material examined. None. Diagnosis. Females have similar spermathecal proportions to the continental C. fulvipes but can be recognised by the narrower, distinctly coiled epigynal ridges (Fig. 84), which are broad and curved in C. fulvipes (Fig. 80). The distal section of the male embolus is short and slightly curved towards the tip (Fig. 86).

Etymology.
A patronym in honour of Charles Griswold, one of the collectors of the types, in recognition of his vast and significant collections of spiders on Madagascar.
Remark. The palp of the paratype male is greatly expanded and only the tegulum is illustrated here (Fig. 86). The cymbium has the distal setae characteristic of the genus, confirming the placement of this species and the occurrence of Cambalida on Madagascar.
Male ( Carapace dark brown, eye region darker; faint black striae radiating from fovea towards palps and leg coxae; surface finely granulate, densely covered in white plumose setae. Rings around eyes faded to orange-brown; AER procurved, ALE larger than AME; AME separated by distance equal to ½ their diameter, AME separated from ALE by 1 ∕ 5 AME diameter; clypeus height slightly less than 1½ AME diameter; PER procurved, PLE slightly larger than PME; PME separated by distance slightly less than ¾ their diameter, PME separated from PLE by distance slightly less than ⅓ PME diameter; CW:PERW = 2.42:1. Chelicerae brown with faint black mottling on anterior surface, yellow prolaterally in distal half; three teeth on promargin, median tooth largest, distal tooth smallest, median and distal teeth closest; two slightly separated teeth on retromargin, distal tooth slightly smaller than proximal tooth, closer to fang base than promarginal teeth; endites pale brown with dark brown mottling, cream prolaterally; labium pale brown with dark brown mottling, cream distally; sternum red-brown with dark brown mottling. Legs finely granulate; femora I-IV dark brown, slightly paler dorsally, yellow dorsally at distal end; patellae I-IV yellow with black mottling laterally and ventrally distally, darker on posterior legs; tibiae I and II yellow with black mottling; tibiae III and IV orange-brown with dense black mottling, paler along dorsal midline; metatarsi I and II yellow-brown, with sparse dark brown mottling laterally; metatarsi III and IV dark brown, yellow at proximal and distal ends; tarsi I-IV yellow, tarsus IV with black mottling medially. Leg spination: femora: I pl 1 do 3, II do 3, III pl 1 do 3 rl 1, IV pl 1 do 3 rl 1; patellae with do 1 long distal seta; tibiae: I and II spineless, III pl 2 rl 2 plv 2, IV pl 2 rl 2 plv 2 vt 2; metatarsi: I plv 2 rlv 1, II plv 1 rlv 1, III pl 2 rl 1 plv 1 vt 3, IV pl 3 rl 3 plv 2 vt 3. Palpal spination: femora pl 1 do 2, patellae pl 1, tibiae pl 1 plv 1, tarsi pl 1 plv 3. Abdomen with dark red-brown, nearly black scutum covering entire dorsum; venter mottled pale grey between sclerites; epigastric sclerite, post-epigastic sclerites and ventral sclerite deep red-brown with black mottling, inframamillary sclerite orange-brown. Palps orange-brown with dense black mottling; embolus short, basal coil slightly curved, distal section gently curved towards tip (Fig. 86).
Distribution. Known from isolated localities in northern and southern Madagascar (Fig. 95).
Other material examined. None.

Diagnosis.
The species is unique amongst Cambalida in the cream median stripe on the abdomen, which extends to the carapace in the holotype (Fig. 7) and one of the paratypes, but is very indistinct in two of the paratypes. The ST II are very large and round, and nearly twice as broad as the ST I (Figs 87, 88 Carapace orange-brown, clypeus slightly paler laterally, eye region slightly darker, with slightly paler yellow-orange line from fovea to posterior margin; black striae radiating from fovea towards palps and coxae and faint black mottling on slopes; surface finely wrinkled, sparsely covered in white plumose and short straight setae. All eyes with black rings; AER procurved, laterals larger than medians; AME separated by distance slightly less than ½ their diameter, AME separated from ALE by distance equal to 1 ∕ 5 AME diameter; clypeus height slightly larger than 1½ times AME diameter; PER procurved, PME and PLE equal in diameter; PME separated by distance slightly less than ⅔ their diameter, PME separated from PLE by distance slightly less than ⅓ PME diameter; CW:PERW = 2.45:1. Chelicerae deep orange-brown with black mottling on anterior surface, yellow along prolateral distal margin; bent setae on promargin pectinate; three teeth on promargin, median tooth largest, proximal tooth smallest, median tooth closer to distal tooth than to proximal tooth; two teeth on retromargin, distal tooth slightly larger than proximal tooth, closer to fang base than promarginal teeth; endites pale orange with faint black mottling, cream prolaterally; labium orange, paler distally; sternum bright orange with faint black mottling. Legs finely granulate; femora I-IV mottled brown, with paler dorsal lines proximally and distally and club-shaped retrolateral paler line; patellae I-IV brown, yellow-brown dorsally; tibiae I yellow, mottled proximally and laterally at distal end; tibiae II-IV yellow dorsally and ventrally, mottled brown laterally, tibiae IV yellow at distal end; metatarsi and tarsi I-IV yellow with faint lateral brown mottling proximally. Leg spination: femora: I pl 1 do 2, II do 2, III pl 1 do 3, IV pl 1 do 3-4 rl 1; patellae with do 1 long distal seta; tibiae: I plv 3 rlv 2, II plv 2 rlv 2, III pl 2 do 1 rl 2 plv 2 rlv 2 vt 2, IV pl 2 do 1 rl 2 plv 2 rlv 1 vt 2; metatarsi: I plv 2 rlv 2, II plv 2 rlv 2, III pl 3 rl 3 plv 2 rlv 2 vt 3, IV pl 3 rl 3 plv 2 rlv 2 vt 3. Palpal spination: femora pl 1 do 2, patellae pl 1, tibiae pl 1 plv 1, tarsi pl 2 plv 2 rlv 1. Abdomen mottled dark grey dorsally, with narrow triangular median cream line, broadest anteriorly and extending past middle of abdomen, cream spots surrounding sigilla, and small white spot above spinnerets (Fig. 7); dorsum with yellow scutum extending ¼ abdomen length; sides of abdomen mottled dark grey, with cream line extending ⅔ the distance from epigastric furrow to spinnerets; venter mottled pale grey, darker towards spinnerets, epigastric sclerite and inframamillary sclerite yellow.
Epigyne with small lateral copulatory openings situated within hemispherical epigynal ridges (Fig. 87); copulatory ducts initially directed posteriorly, bending at nearly right angle before running transversely towards midline, entering ST II posteromedially; ST II large and round, joined broadly to narrower kidney-shaped posterior ST I (Fig. 88).
Male. Unknown. Distribution. Known from three isolated localities in northern and central Madagascar (Fig. 95).
Biology. A ground-dwelling species collected in rainforest. ( Diagnosis. This species is recognised by the distinctly coiled epigynal ridges and the oblique curved entrance ducts of the females (Figs 89, 90). Males can be recognised by the broad, somewhat flattened embolus (Figs 43, 56).

Cambalida loricifera
Remarks. The right leg I and right palp of the male holotype are missing. The redescription of the male is provided for the holotype, although more recently collected specimens are darker brown in colour.
Etymology. The species name is Latin for "unique". Remark. The holotype is the smaller of the two females in the vial containing the types and has the epigyne dissected.

Cambalida insulana Simon, 1909
Cambalida insulana Simon, 1909: 369. Remarks. The type material could not be traced in MNHN and is probably lost (Christine Rollard, pers. comm.), which was confirmed by Bosselaers and Jocqué (2000), who also could not successfully locate the type. The type locality given by Simon (1909) is "Ile Annobon", also known as Pagalu Island, which is situated 160km south-west of Sao Tomé in the Atlantic Ocean. The only Cambalida material available from these islands is a series of immature and subadult specimens from Parc Nacional Ôhó, Sao Tomé, collected in 2001 and deposited in CAS. It is thus likely that C. insulana populations may still be viable and not extinct. Additional adult material from the type locality is needed before this species can be validated, redescribed and neotypes designated.

Cambalida simoni nom. n.
Cambalida fulvipes Simon, 1909: 369 [preoccupied by senior homonym] Remarks. A replacement name is here proposed for Cambalida fulvipes Simon, 1909, a name occupied by its secondary senior homonym, C. fulvipes (Simon, 1896), proposed earlier in this paper. The type material of this species could not be traced in MNHN and is probably lost, and the original description is inadequate for its successful identification. The occurrence of three different species in West Africa, and the lack of any comparable fresh material from Guinee-Bissau, from which this species was described, means that this species should remain nomen dubium until fresh material can be collected from the vicinity of the type locality (Bolama, 11°34'N, 15°29'W).

Discussion
The current revision of the Afrotropical genus Cambalida increased the species compliment from three to ten species, with a further two species (including the type species of the genus) being considered nomina dubia. The biogeographical patterns of Cambalida are quite different to most of the other Afrotropical genera. Apochinomma Pavesi, 1881 and Copa Simon, 1885(Haddad 2012 each have a single widespread Afrotropical species and Echinax Deeleman-Reinhold, 2001 (Haddad in press) has two widespread species, with the other species in each genus generally being restricted to a single biogeographical region (e.g. West Africa) or a single country. In contrast, Cambalida has one species widespread throughout the region (C. fulvipes), two additional species widespread through tropical West and Central Africa (C. coriacea and C. deminuta), and one widespread in southern Africa (C. dippenaarae). The remaining six species are all comparatively range restricted. While Copa has radiated considerably on Madagascar, with more than 30 new species to be described from the island (Haddad 2012), only two new Madagascan species of Cambalida have been described in this study. However, there is a large quantity of unidentified material from the island, and it is plausible that several new Cambalida have yet to be discovered. Until such time as the fauna has been more thoroughly studied, no discussion or hypotheses of the biogeographical relationships of the Madagascan and continental faunas should be made. Cambalida appear to be exclusively ground-dwelling leaf litter spiders occurring mainly in savanna and forest habitats. In savannas they are generally uncommon but are similar in abundance to Copa flavoplumosa Simon, 1885 and Merenius spp. (Foord et al. 2008;Haddad et al. 2010;Muelelwa et al. 2010). In contrast, they contribute a more significant proportion of spider assemblages in the leaf litter of shrubs in the South African Grassland Biome, where they are the most abundant corinnids (Butler and Haddad 2011). They are only occasionally collected from agroecosystems in South Africa (Haddad and Dippenaar-Schoeman 2006), but several species are quite common in rice and fallow habitats in West Africa.
The current study has significantly increased the species compliment of Cambalida to ten, and it is likely that further new species will be sampled and need to be described in the future. This is supported by the reasonably small distribution ranges of several species (e.g. C. griswoldi, C. lineata and C. loricifera) and others that are only known from the type locality (C. fagei and C. unica). It is probable that many historically poorly sampled biodiversity hotspots may yield considerable additions to the fauna, particularly in East and West Africa. The possible occurrence of Cambalida on the Indian Ocean islands (other than Madagascar) also requires further investigation.