A revision of the continental species of Copa Simon, 1885 (Araneae, Corinnidae) in the Afrotropical Region

Abstract The cryptic ground-dwelling castianeirine genus Copa Simon, 1885 (Araneae: Corinnidae) is revised in the continental Afrotropical Region. The type species of the genus, Copa flavoplumosa Simon, 1885, is redescribed and considered a senior synonym of Copa benina Strand, 1916 syn. n. and Copa benina nigra Lessert, 1933 syn. n. It is widespread throughout the Afrotropical Region but has not been introduced to any of the associated regional islands. A new species, Copa kei sp. n., is described from South Africa. Copa agelenina Simon, 1910, originally described from a subadult female from southern Botswana, is considered a nomen dubium. Copa flavoplumosa is a characteristic species of leaf litter spider assemblages and is particularly prevalent in savanna habitats on the continent, but also occurs in various forest types, grasslands, fynbos and semi-arid Nama Karoo habitats. In contrast, Copa kei sp. n. has only been recorded from Afromontane and coastal forests in south-eastern South Africa.


introduction
The spider genus Copa Simon, 1885 (Araneae: Corinnidae) is only known from the Afrotropical and Palaearctic Regions, and some of the species from the latter have recently been studied by Deeleman-Reinhold (1995. Deeleman-Reinhold (2001) described the genus Echinax to include three species of Copa from South-East Asia that she had earlier described in 1995, and an additional new species. Yang et al. (2004) subsequently described a fifth species of Echinax from China. Thus, only two Asian species of Copa remain, both described from Sri Lanka (Simon 1896). Prior to this revision, seven species and one subspecies of Copa were known from the Afrotropical Region (Dippenaar-Schoeman and Jocqué 1997), of which one described by Simon (1909) has been transferred to Echinax (Haddad 2012a). Another species, described by Strand (1916), will be transferred to a new castianeirine genus (Haddad 2012b in prep.).
Although most genera in this subfamily resemble ants, Copa species have cryptic colouration and closely resemble wolf spiders of the family Lycosidae (Figs 1-6), a characteristic shared with Echinax and two undescribed lycosiform castianeirine genera (Haddad 2012a, b, in prep.). Copa are very common spiders in the leaf litter of various habitats and are predominantly ground-living, occurring widely in savanna woodlands but also occasionally in forests, where they are well camouflaged. They usually share litter microhabitats with species of several other castianeirine genera, including Cambalida Simon, 1909, Merenius Simon, 1909and Castianeira Keyserling, 1879(Haddad 2012b, 2012c. Copa flavoplumosa Simon, 1885 is also recorded here from drier habitats, including fynbos, grassland and Nama Karoo in South Africa and the arid savannas of north-western South Africa, Botswana and Angola, thereby showing considerable ecological flexibility and adaptability. In contrast, the species of the other three cryptic lycosiform genera are primarily arboreal and are rarely collected in leaf litter. The current study presents the first revision of the continental Afrotropical species of the genus, and the Madagascan fauna, including two species described by Simon (1903) and Strand (1907) and nearly 30 new species, will be treated at a later stage in a separate paper.

Material and methods
All specimens examined during this study were observed in 70% ethanol using a Nikon SMZ800 stereomicroscope for descriptions, digital photographs and measurements. A series of digital photographs of the male emboli of each species were taken using a Nikon Coolpix 8400 mounted on a Nikon SMZ800 stereomicroscope. The photographs were then stacked using Combine ZM software (http://www.hadleyweb. pwp.blueyonder.co.uk) to increase depth of field. Photographs of live C. flavoplumosa and C. kei sp. n. were taken in the field using a Canon EOS 40D digital camera with 50mm or 100mm macro lenses. Material of both aforementioned species was prepared for scanning electron microscopy by dehydrating the specimens in a graded ethanol series and critical-point drying them in an argon chamber, after which they were fixed to aluminium stubs and sputter-coated with gold three times for three minutes. The material was then studied in a JEOL 6400 WinSEM and digital micrographs were taken.
All measurements are given in millimetres (mm). Total body length measurements are given for the smallest and largest specimens of each sex to give an indication of size variation, and body, eye and leg measurements are given for the specimens indicated intercoxal sclerites only present between coxae I and II (Figs 9,12). Leg formula 4132 in females, 4312 or 4123 in males, legs I and III nearly equal in length; legs strongly spined, femora, patellae, tibiae and metatarsi covered in short straight black setae and black and white feathery setae (31)(32)(33)(34)(35)(36)(37)(38)(39), feathery setae sparse on tarsi; retrocoxal window absent; femora with several scattered erect ventral setae (Figs 24,31); anterior patellae with proximal and distal long fine dorsal setae (Figs 24,34); posterior patellae with fine long proximal dorsal seta and distal spine, clearly shorter than patella (Fig. 35); patellar indentation narrow, broad at proximal end (Figs 25,26,32,33); metatarsi III longer than metatarsi I and II; metatarsi distally scopulate (Fig. 39); tibiae, metatarsi and tarsi with several dorsal and lateral trichobothria with sunken distal plate (Fig. 29), also with several short erect setae (Fig. 38); tarsal organ oval, slightly elevated from integument, surface finely wrinkled, opening oval (Figs 30,42); paired tarsal claws short, with very dense claw tufts in between (Figs 28,41); metatarsi III and IV without terminal preening brush or comb. Abdomen oval, Figures 7-12. Digital microscope photographs of Copa flavoplumosa Simon, 1885 from D.R. Congo (7-9) and C. kei sp. n. from South Africa (10-12): 7, 10 female, dorsal habitus 8, 11 male, dorsal habitus 9, 12 sternum of female in ventral view. Scale bars = 1.0 mm. either yellow-orange with black markings or black with white markings; three pairs of fine straight setae on anterior margin above pedicel; dorsal scutum small, strongly sclerotized, extending less than ⅛ abdomen length in females and slightly more than ½ abdomen length in males; two pairs of distinct sclerotised dorsal sigilla present in both sexes (Figs 7,8,11,12); epigastric region moderately sclerotised, venter without post-epigastric sclerites and ventral sclerite, inframamillary sclerite present, distinct, densely covered in short setae; two paired rows of tiny sclerites from epigastric furrow to spinnerets, outer row weakly sclerotised and indistinct. Spinnerets: ALS of female (Figs 43,49) with two major ampullate gland spigots and many piriform gland spigots; ALS of male (Figs 44,50) with single major ampullate gland spigot, single large adjacent nubbin and many piriform gland spigots; PMS of female (Figs 45,51) with three large cylindrical gland spigots, one small minor ampullate gland spigot and one or two aciniform gland spigots; PMS of male (Figs 46,52) with one large minor ampullate gland spigot, one tartipore and one nubbin, with zero aciniform gland spigots in C. flavoplumosa and two in C. kei sp. n.; PLS of female (Figs 47,53) with two large cylindrical gland spigots and zero (in C. flavoplumosa) or several (in C. kei sp. n.) aciniform gland spigots; PLS of C. flavoplumosa male with two reduced aciniform gland spigots and several tiny nubbins present (Fig. 48); PLS of C. kei sp. n. with three aciniform gland spigots only (Fig. 54). Female epigyne with 6-shaped or curved sclerotized epigynal ridges leading to lateral copulatory openings (Figs 55, 57); copulatory ducts directed anteriorly or transversely before entering ST II posteriorly; ST II usually oval, sometimes expanded posterolaterally, connected broadly to somewhat elongated posterior ST I. Male palpal segments without apophyses; cymbium with spines prolaterally and ventrally, dorsal surface covered in curved finely plumose setae with round tip and thicker straight setae with sharp tips (Fig. 59); distal end of cymbium in C. kei sp. n. with shallow depression, densely covered in setae (Fig. 60); embolus with variable width and angle of base, and length and curvature of distal coil (Figs 56,58,(61)(62)(63)(64)(65)(66).       nated as a paralectotype. A lectotype female and paralectotype female are designated here, of which the larger of the two in the type series is the lectotype.

Key to the continental Afrotropical species of Copa
The specimens of Copa benina nigra Lessert, 1933 available in the MNHG are not specifically labelled as types and their status is thus uncertain, although the labels indicate localities consistent with that in Lessert's (1933) description, i.e. Chimporo and Rio Mbale. Neither of these localities could be traced on modern maps or electronic resources. Some maps from the early 20 th century indicate that the Rio Mbale runs northwards between 16°20'E and 16°40'E with its source at approximately 12°00'S in central Angola. Chimporo has been cited by Mansell (1996) as being located at 17°20'S, 17°17'E in southern Angola. From Lessert's (1933) figure of the male embolus it is clear that C. b. nigra is a junior synonym of C. flavoplumosa.
Female (Parc National Albert, MRAC 234182). Measurements: CL 3.60, CW 2.69, AL 4.85, AW 3.10, TL 8.20 (6.35-9.30), FL 0.40, SL 1.70, SW 1.58, AME-AME 0.10, AME-ALE 0.02, ALE-ALE 0.44, PME-PME 0.12, PME-PLE 0. General appearance as in Fig. 7. Carapace bright yellow-orange, eye region black; broad median black line comprising black feathery setae from PER to posterior slope of carapace, broken up by narrow asetose line from between PME to midpoint and Y-shaped asetose area from fovea towards anterior coxae and posterior of carapace; striae absent; lateral margins with narrow fringe of black feathery setae; areas between markings covered in orange feathery setae. All eyes with black rings; AER procurved, medians much larger than laterals; AME separated by distance equal to ½ their diameter; AME separated from ALE by distance approximately 1 ∕ 10 AME diameter; clypeus height slightly less than 1½ AME diameter; PER strongly procurved, eyes subequal in size; PME separated by distance equal to ¾ their diameter; PME separated from PLE by distance equal to 4 ∕ 5 PME diameter; CW:PERW = 3.45:1. Chelicerae yelloworange, with pectinate curved setae on promargin; promargin with two teeth separated by basal width of proximal tooth, distal tooth much larger; retromargin with two teeth separated by ½ their basal width, distal tooth slightly larger than proximal tooth, close to fang base. Endites yellow, cream prolaterally and distally, with small black prolateral proximal markings; labium yellow-orange, cream distally, without markings; sternum pale orange, without markings (Fig. 9). Legs yellow-brown, posteriors slightly darker, femora slightly darker dorsally than ventrally; femora with broad dorsal line between proximal and distal spines and incomplete dorsal rings at ⅔ their length and distally, each composed of black feathery setae; patellae with dense black feathery setae laterally; tibiae I & II without markings, with scattered black feathery setae, III & IV with rings proximally and medially corresponding to ventral spines, distal ends with black ring, all covered in black feathery setae with white feathery setae between them; metatarsi I & II without markings, with scattered black and white feathery setae, III & IV with proximal, medial and distal rings corresponding to paired leg spines, covered in black feathery setae with white feathery setae between them; tarsi uniform yellow; palp yellow, spines without spots. Leg spination: femora: I pl 2 do 3 rl 2, II pl 2 do 3 rl 2, III pl 2 do 3 rl 1-2, IV pl 2 do 3 rl 1; all femora with scattered erect ventral setae; patellae: I & II with fine proximal and distal do setae, III & IV with proximal and distal do spines, proximal spine finer and shorter than distal; tibiae: I do 1 pl 2 long fine setae, plv 2 rlv 2 spines, II do 1 long fine seta, pl 2 plv 2 rlv 1 spines, 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 rl 1, with scattered erect ventral setae, mainly retrolaterally; patellae: pl 1 do 2; tibiae: pl 1 do 2 plv 1-2; tarsi: pl 1 rl 1 plv 2 rlv 1. Abdomen with very small orange-brown anterior dorsal scutum; dorsum cream, densely covered in black straight and feathery setae, interspersed with white feathery setae forming small spots in anterior two-thirds and fine transverse chevrons posteriorly; sides of abdomen cream, densely covered in white feathery setae; venter cream, covered in short straight black setae, with broad densely setose subrectangular marking medially from epigastric furrow to spinnerets, comprising black and white feathery setae and short straight black setae. Epigyne longer than broad, with large 6-shaped ridges laterally at midpoint of epigyne, separated by approximately 1½ times their width, with copulatory openings distinct (Figs 55, 67); copulatory ducts directed anteriorly, slightly obliquely, with anterior bend and characteristic loop before entering anterior ST II; broad ducts connecting ST II to elongate posterior ST I; ST I clearly narrower than ST II (Fig. 68).
Colour variation. Throughout the geographical range of C. flavoplumosa there is considerable variation in the colouration of specimens, particularly with regard to the intensity of yellow/orange and white/cream markings on the body. Three main generalised colour forms can be found. The most widespread variation has a yellow-brown to bright orange carapace and abdomen, with black markings (Fig. 1), and is found throughout the region except in the rainforests of central Africa. This colour form is mainly associated with populations in savanna and grassland habitats.
The second colour form (corresponding to the redescriptions above) has a yellow to orange carapace with black markings and a black abdomen with white markings, similar in pattern and arrangement to the previous form (Figs 2, 3, 7, 8). This form is found in moist savannas and forests across tropical Africa. While most South African populations of this species have colouration corresponding to the first form described here, the populations in the fynbos and grasslands along the southern coast of the country also have a black abdomen with white markings, but the carapace is even darker, nearly dark red-brown in colour.
The third form, corresponding to the description of C. benina nigra, is a nigrito form restricted to central and western Africa but only occurring in isolated populations. This form has an entirely black body with white markings corresponding to those described for the other two types above (Fig. 4).
The distribution of the three forms can partly be explained by the habitats they occupy, although some populations (e.g. Faro Game Reserve in Cameroon and Mankono in Ivory Coast) have representatives of all three colour forms but in varying proportions. The colouration of the first form is clearly an adaptation for camouflage in the litter layer of savanna and other habitats that are exposed to sunlight for a considerable portion of the day. The second form apparently occurs in closed canopy forests and dense woodlands that are shaded for most of the day or the entire day. The black abdominal colouration with strongly contrasting markings enables these spiders to blend into dark patches with low light levels in these habitats. The nigrito form can exploit such microhabitats in a similar way, but this does not explain the occurrence of this colour form at some Miombo woodland localities (e.g. Wildlives Game Farm in Zambia). A possible explanation for this case could be the occurrence of natural fires in these habitats. The burned trees, logs and grasses in such disturbed environments may provide sites where these spiders may optimally exploit their colouration for camouflage. Whether individuals have the capability for colour change in response to changing environmental conditions (e.g. following fire) through the use of chromatophores or ommochromes, or whether individual phenotypes are stable, has yet to be determined and should be the subject of future research.
Distribution. Widespread throughout the continental Afrotropical Region (Fig. 71). Biology. Specimens were mainly collected from the leaf litter layer of all of the main biome types in Africa except for true deserts and karoo habitats, although records from semi-arid temperate grasslands and dry savannas are scarce. The greatest density of records is in moist savannas and closed canopy forests, although the species seems largely absent from rainforests; considering the extensive sampling in the D.R. Congo, especially by the MRAC, only three records from rainforests in this country are reported here. Specimens were most regularly collected by pitfall traps, litter sifting and by hand from the ground surface.
Etymology. The specific name is a noun in apposition taken from the type locality, the town Kei Mouth, located at the estuary of the Great Kei River in the Eastern Cape Province.
Female (holotype, Kei Mouth, NCA 2007/3843). Measurements: CL 3.84, CW 2.75, AL 6.00, AW 4.55, TL 9.65 (6.40-9.80), FL 0.40, SL 1.75, SW 1.60, AME-AME 0.10, AME-ALE 0.01, ALE-ALE 0.46, PME-PME 0.20, PME-PLE 0. General appearance as in Fig. 10. Carapace bright yellow-orange, eye region black except between PME; broad median black line covered in black feathery setae from PER to posterior slope of carapace, broken up by asetose line from PME to midpoint and paired oblique asetose line from fovea towards anterior coxae; black striae present, falling within broad median band; lateral margins black from chelicerae to posterior marking, markings expanded from coxae I and from coxae I-IV, densely covered in black feathery setae; areas between markings covered in white feathery setae. All eyes with black rings; AER procurved, medians much larger than laterals; AME separated by distance equal to ½ their diameter; AME separated from ALE by distance approximately 1 ∕ 10 AME diameter; clypeus height approximately 1½ AME diameter; PER strongly procurved, medians very slightly larger than laterals; PME separated by distance equal to 1¼ their diameter; PME separated from PLE by distance equal to 4 ∕ 5 PME diameter; CW:PERW = 3.31:1. Chelicerae yellow-orange, with pectinate curved setae on promargin; three closely spaced teeth on promargin, distal tooth smallest, median tooth largest; median tooth closer to distal tooth than to proximal tooth; retromargin with two teeth separated by their basal width, distal tooth slightly smaller than proximal tooth, close to fang base. Endites yellow, cream prolaterally; labium yellow-brown, cream distally, with broad transverse black marking along proximal margin; sternum bright yellow, with broad black marking along margins, expanded at coxae (Fig. 12). Legs yellow-brown, with faint black mottling; spine bases with distinct black spot; trochanters with distal margins black laterally; femora all with black lateral and distal mottling, ventrally with faint distal ring; patellae with fine dorsal proximal line and lateral and distal mottling; tibiae with faint rings proximally and medially corresponding to ventral spines, distal ends with black ring; metatarsi with proximal, medial and distal rings, corresponding to paired leg spines; tarsi yellow; palp yellow, spines with black spots. Leg spination: femora: I pl 2 do 3 rl 1, II pl 2 do 3 rl 1, III pl 2 do 3 rl 2, IV pl 2 do 3 rl 2; all femora with scattered erect ventral setae; patellae: I & II with fine proximal and distal do setae, III & IV with proximal and distal do spines, proximal spine finer and shorter than distal; tibiae: I do 1 long fine seta, plv 2 rlv 2, II do 1 long fine seta, plv 1 rlv 2 spines, III pl 2 do 1 rl 2 plv 2 rlv 1-2 vt 2, IV pl 2 do 1 rl 2 plv 2 rlv 2 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, with scattered erect ventral setae, mainly retrolaterally; patellae: pl 1 do 2; tibiae: pl 1 do 2 plv 1; tarsi: pl 1 rl 1 plv 3 rlv 1. Abdomen with very small red-brown anterior dorsal scutum beneath marking; dorsum mottled grey, with large black spot anteriorly, dark grey median stripe from anterior spot to midpoint, and small cream chevrons posteriorly; short straight black setae and white feathery setae on markings dorsally and laterally; sides of abdomen cream; venter cream, covered in short straight black setae, with dark marking medially on epigastric plate covering epigyne, broadened from epigastric furrow, extending to and surrounding spinnerets. Epigyne small, with strongly curved ridges laterally at midpoint of epigyne, separated by approximately three times their width, with copulatory openings distinct (Figs 57, 72); copulatory ducts almost straight, nearly transverse, slightly oblique, entering rounded anterior ST II; broad ducts connecting ST II to subrectangular posterior ST I; ST I slightly narrower than ST II (Fig. 73).
Distribution. Known from the south-eastern parts of South Africa (Fig. 76); endemic to the Maputaland-Pondoland-Albany Centre of Endemism (Driver et al. 2005).
Biology. Specimens were mainly collected from the leaf litter layer of closed canopy Afromontane and coastal forest habitats. Remarks. The holotype is a subadult female specimen (Figs 77-79), clearly with a pre-epigynum and lack of epigynal sclerotisation typical of adults, and is not an adult female as described by Simon (1910) and listed on Platnick (2012). The species is definitely different to C. flavoplumosa specimens collected in north-western South Africa (Sunnyside and Hermitage) and has a clearly broader PER (CW:PERW = 2.84:1 as opposed to 3.45:1 in C. flavoplumosa females) that is less strongly procurved than in C. flavoplumosa. Since no adult Copa material is available from the arid savanna of southern Botswana, it is impossible to match this specimen to either of the two conti- nental species or to recognise it as a distinct species. I would therefore propose that C. agelenina be considered a species nomen dubium.
The eye arrangement and measurements (Fig. 80) suggest that this species may belong to one of the new cryptic lycosiform castianeirine genera (Haddad 2012b, in prep.), but adults will have to be collected before the generic placement can be confirmed and the species be revalidated and redescribed.

Discussion
The current study treated the continental species of Copa in the Afrotropical Region, reducing the number of species from four to two, of which one species is newly described. The type species of the genus, C. flavoplumosa, is widespread throughout the region and includes two synonyms newly proposed here. It is distributed from Guineé in the west to Tanzania in the east, and from Nigeria in the north to South Africa in the south. The new species, C. kei, is endemic to south-eastern South Africa. While C. flavoplumosa provides a useful example of extreme habitat flexibility, occupying habitats from forests to semi-deserts, C. kei is very closely associated with Afromontane and coastal forests in South Africa. The latter species has a distribution falling entirely within the Maputaland-Pondoland-Albany Centre of Endemism in South Africa (Driver et al. 2005). Surprisingly, very few C. flavoplumosa records exist from the tropical rainforests of the D.R. Congo and Congo Republic, despite the former being one of the best sampled countries on the continent (Fig. 71). The two species therefore represent extremes regarding both vagility and ecological flexibility. Both species are clearly ground-dwelling leaf litter specialists and were mainly collected by pitfall trapping, litter sifting and hand collecting. Copa flavoplumosa may be very abundant in some habitats (e.g. forests in Ivory Coast), but they tend to be considerably less common in savannas and other habitat types (Modiba et al. 2005;Dippenaar-Schoeman and Wassenaar 2006;Foord et al. 2008;Haddad et al. 2010;Muelelwa et al. 2010). They have occasionally been collected in agroecosystems, specifically from the canopies of orchard crops in South Africa (avocadoes, macadamias and pistachios), but never exceed 2% of the total spider fauna (Dippenaar-Schoeman et al. 2001Haddad et al. 2005). Their arboreal habits in agroecosystems are in stark contrast to their almost exclusive ground-dwelling habits in natural habitats, and the reasons for this ecological divergence are unknown.