Spermophora, HENTZ, 1841

Huber, Bernhard A., 2003, Southern African pholcid spiders: revision and cladistic analysis of Quamtana gen. nov. and Spermophora Hentz (Araneae: Pholcidae), with notes on male-female covariation, Zoological Journal of the Linnean Society 139 (4), pp. 477-527 : 515-522

SPERMOPHORA HENTZ, 1841

The results of the cladistic analysis provide support, albeit weak, for at least some of the following species being more closely related to ‘real’ Spermophora than to any other African genus, or to species assigned tentatively to Spermophora . The character supporting this position is the ‘ Spermophora flap’, a distinctive ventral projection distally on the procursus (e.g. Figs 227 , 233 ). Note that in two of the species below - S. pembai and S. suurbraak - this flap is absent, and their assignment is tentative. This phylogenetic position is surprising, as the closest relatives of the type species (the synanthropic S. senoculata (Dugès) with worldwide distribution) occur in East Asia and Australia, and other African representatives were assigned to Spermophora tentatively for want of a better hypothesis ( Huber, 2003a,b).

South African representatives of Spermophora are long-legged, six-eyed pholcids with globular or oval opisthosoma, varying in total size from 2.0 to 3.5 mm. They are distinguished from other southern African genera as follows: from Quamtana by the dorsal attachment of the bulb (e.g. Figs 226 , 232 ), hinged process on the procursus (e.g. Figs 226 , 232 ), and single or missing modified hair on the male cheliceral apophysis (e.g. Figs 229 , 235 ); from Smeringopus , Crossopriza and Artema by the lateral cheliceral apophyses (e.g. Figs 228 , 234 ); from

Figures 215–225. Quamtana embuleni (215–218), Q. oku (219–223) and Q. biena (224, 225) spp. nov. 215, 219, 220, Modified hairs on male cheliceral apophyses (sc: sclerotized cone). 216, Female ALS and PMS. 217, Tip of female pedipalp with tarsal organ. 218, Epigynum with pair of pockets (p). 221, Male ALS. 222, Male AME and lateral triad. 223, 224, Male gonopores with epiandrous spigots. 225, Male ALS and PMS.

Leptopholcus and Pholcus by the short opisthosoma (vermiform in Leptopholcus , long cylindrical in Pholcus ).

In South African Spermophora species , the clypeus is always unmodified; chelicerae never with stridulatory ridges; legs without spines, with few vertical hairs, without curved hairs; prolateral trichobothrium missing on tibiae 1, present in all others. For other characters, see descriptions below.

Most species are from Western Cape; only S. pembai occurs in Eastern Cape ( Fig. 264 ).

SPERMOPHORA SCHOEMANAE SP. NOV.

( FIGS 26 , 48 , 226–231 , 244–247 )

Spermophora sp. 3 : Huber, 2003a,b

Type. Male holotype from forest in Grootvadersbosch near Heidelberg (~34∞00¢S, 21∞00¢E), Western Cape ,

South Africa; February 1, 1989 (A. S. Dippenaar- Schoeman), in NCP (89/887).

Etymology. Named after the collector, Ansie Dippenaar-Schoeman from the Plant Protection Research Institute, Pretoria.

Diagnosis. Medium size species, distinguished from congeners by the pointed projection of the bulb and the shape of the bulbal apophysis ( Fig. 227 ).

Male (holotype). Total length 1.90 (2.1 with clypeus), carapace width 0.90. Leg 1: 18.9 (4.7 + 0.4 + 5.2 + 6.8 + 1.8), tibia 2: 3.1, tibia 3: 2.2, tibia 4: 2.7; tibia 1 L/ d: 58. Habitus as in Figure 26 . Carapace orange-ochre with some darker marks; sternum brown with many tiny light spots; legs orange-ochre; opisthosoma grey with blackish pattern; ventral pattern as in female (cf. Fig. 48 ) but with triangular black mark in genital area. Ocular area slightly elevated; distance PME–PME 120 Mm; diameter PME 80 Mm; no trace of AME. Clypeus unmodified. Sternum wider than long (0.70/ 0.55). Chelicerae as in Figure 228 , with one coneshaped hair on each apophysis ( Figs 229 , 246 ), tips 175 Mm apart. Palps as in Figures 226, 227 ; trochanter with wide retrolateral apophysis; femur proximodorsally with distinct apophysis, ventrally with small cone; procursus prolaterally with proximal projection and distal hinged process; bulb with hook-shaped apophysis, membranous embolus, and distinctive pointed projection ( Fig. 227 ). Retrolateral trichobothrium of tibia 1 at 13%; tarsus 1 with ~15 pseudosegments, distally very distinct. Gonopore with four epiandrous spigots ( Fig. 244 ). ALS with several piriform gland spigots ( Fig. 245 ). Palpal tarsal organ capsulate ( Fig. 247 ).

Variation. Tibia 1 in five other males: 4.3–5.2 (x = 4.93).

Female. In general similar to male. Tibia 1 in two females: 3.7, 4.1. Epigynum simple externally ( Figs 48 , 230 ) but strongly protruding, with pair of pockets 195 Mm apart. Dorsal view as in Figure 231 .

Distribution. Known from type locality only ( Fig. 264 ).

Material examined. SOUTH AFRICA: WESTERN CAPE: Heidelberg : type above, together with 7♂ 2♀ ( NCP 89 /887) .

SPERMOPHORA GORDIMERAE SP. NOV.

( FIGS 27 , 49 , 232–237 )

Type. Male holotype from Kirstenbosch Botanical Gardens, Cape Town (~34∞00¢S, 18∞30¢E), Western

Cape, South Africa; August 12, 1978 (A. Russell Smith), in ZFMK.

Etymology. Named after Nadine Gordimer, South African writer and Nobel laureate.

Diagnosis. Medium size species, distinguished from the very similar S. schoemanae by the shape of the bulb (no pointed projection, Fig. 232 ), from other southern African congeners also by the shape and position of the male cheliceral apophyses ( Fig. 234 ).

Male (holotype). Total length 2.4 (2.6 with clypeus), carapace width 1.05. Leg 1: 5.6 + 0.4 + 5.7 + 8.1, tarsus missing, tibia 2: 3.5, tibia 3: 2.5, tibia 4: 3.3; tibia 1 L/d: 53. Habitus as in Figure 27 . Carapace ochreyellow with darker median line and posterior mark, sternum dark brown, clypeus slightly darker than carapace; legs ochre-yellow; opisthosoma pale ochregrey with black spots dorso-laterally and ventrally. Ocular area barely elevated; distance PME–PME 140 Mm; diameter PME 100 Mm; no trace of AME. Clypeus unmodified. Sternum wider than long (0.75/0.65). Chelicerae as in Figure 234 , with one cone-shaped hair on each apophysis ( Fig. 235 ), tips 205 Mm apart. Palps as in Figures 232, 233 ; trochanter with wide retrolateral apophysis (distinct in dorsal view); femur proximo-dorsally with distinct apophysis; procursus with distal hinged process prolaterally; bulb with distinctive apophysis and membranous embolus ( Figs 232, 233 ). Retrolateral trichobothrium of tibia 1 at 13%.

Variation. Tibia 1 in other male: 5.3.

Female. In general similar to male, but dark mark on carapace more distinct, palps dark brown; ventral pattern on opisthosoma confined to area behind epigynum. Tibia 1 in three females: 4.4, 4.6, 4.9. Epigynum simple externally ( Figs 49 , 236 ), with pair of pockets 175 Mm apart. Dorsal view as in Figure 237 .

Distribution. Known only from Cape Town area ( Fig. 264 ).

Material examined. SOUTH AFRICA: WESTERN CAPE: Cape Town: type above, together with 1♀ ( ZFMK); Cape Town, Table Mountain (~34∞00¢S, 18∞35¢E), Newlands forest , river valley, March 7, 1993 (R. Jocqué), 1♂ 2♀ ( MRAC 174.693 ) .

SPERMOPHORA PENINSULAE LAWRENCE, 1964

( FIGS 29 , 50 , 238–243 , 248–251 )

Spermophora peninsulae Lawrence, 1964: 65–67 ; figs 9–14. Huber, 2003a,b

Types. Male holotype and 1♀ paratype from Kalk Bay Caves , Cape Peninsula (34∞15¢S, 18∞30¢E), Western Cape, South Africa; September 1932 (R. F. Lawrence), in SAM (B7897); not seen (see Notes below) .

Notes. I have not seen the holotype, but Lawrence’s (1964) illustrations of the palp and male chelicerae (his figs 10, 11) leave no doubt as to the correct identification of the specimens below. Moreover, I have seen some of the nontype material described by Lawrence (see below) all of which is identical to the material described below. There are several inconsistencies between the information on labels and the data presented by Lawrence (1964), but these mostly concern collection dates, and do not seem relevant.

Diagnosis. Large species, distinguished from other southern African congeners by the two pairs of apophyses frontally on the male chelicerae ( Figs 240 , 250 ). Also by the shapes of bulbal apophysis ( Fig. 238 ) and epigynum (pockets close together, Figs 50 , 242 ).

Male (CAS). Total length 3.1 (3.3 with clypeus), carapace width 1.4. Leg 1: 33.2 (8.3 + 0.6 + 8.4 + 13.1, 2.8), tibia 2: 5.4, tibia 3: 3.9, tibia 4: 5.0; tibia 1 L/d: 63. Habitus as in Figure 29 . Carapace ochre, ocular area and clypeus light to dark brown, sternum dark brown; legs light brown; opisthosoma grey with blackish pattern, ventrally like female (cf. Fig. 50 ) but with black genital area. Ocular area slightly elevated; distance PME– PME 160 Mm; diameter PME 100 Mm; no trace of AME. Clypeus unmodified. Sternum wider than long (1.0/ 0.8). Chelicerae with distinctive pair of apophyses ( Fig. 250 ), with one modified hair on each of the distal apophyses ( Figs 241 , 250 ), tips 40 Mm apart. Palps as in Figures 238, 239 ; trochanter with large retrolateral apophysis; femur proximo-dorsally with small apophysis; procursus with distal hinged process prolaterally; bulb with distinctive apophysis and membranous embolus ( Figs 238, 239 ). Retrolateral trichobothrium of tibia 1 at 9%. Tarsus 1 with>30 pseudosegments, ~20 quite distinct. Gonopore with four epiandrous spigots ( Fig. 248 ). ALS with several piriform gland spigots ( Fig. 249 ). Palpal tarsal organ capsulate ( Fig. 251 ).

Variation. Tibia 1 in nine other males: 7.25–8.25 (x = 7.70). Lawrence (1964) noted considerable variation in the degree of coloration. It was not clear, however, whether the paleness of some specimens was an artefact of preservation or whether cave populations are indeed paler than forest populations. I have not seen newly collected cave specimens, but Lawrence himself described several specimens from caves with distinct patterns.

Female. In general similar to male. Tibia 1 in seven females: 5.25–6.33 (x = 5.82), in one exceptional female from Wynberg Cave entrance: 8.1. Epigynum simple externally ( Figs 50 , 242 ), with pair of pockets 30 Mm apart. Dorsal view as in Figure 243 .

Distribution. Known only from Cape Town area ( Fig. 264 ).

Material examined. SOUTH AFRICA: WESTERN CAPE: Table Mountain, Wynberg Caves, Oread Hall (34∞07¢S, 18∞27¢E), June 13, 1954 (J. R. Grindley), 1♂ 2 juveniles (not adult females as stated in Lawrence, 1964) ( SAM B10021) ; Wynberg Caves (33∞59¢S, 18∞24¢E), March 1931 (collector not given, probably R. F. Lawrence), 4 juveniles (not adult females as stated in Lawrence, 1964) ( SAM B7896 ) ; Wynberg Caves, Kalk Bay Caves (33∞59¢S, 18∞24¢E), July 1932 (collector not given, probably R. F. Lawrence), 1♀ (not ♂ as on label) ( SAM B7895 ) ; Wynberg Caves, Table Mountain (33∞59¢S, 18∞24¢E), August 5, 1956 (J. R. Grindley), 1 penultimate ♂ and 1♀ (not 2♀ as stated in Lawrence, 1964) ( SAM B10022 and B10024) ; Table Mountain, Powder Room Cave (33∞58¢S, 18∞25¢E), March 4, 1956 (Speleological Assoc.), 1♀ ( SAM B10023) ; Table Mountain, Wynberg Cave entrance (~34∞00¢S, 18∞30¢E), February 13, 1991 (V. D. & B. Roth), 3♀ 1 juvenile ( CAS) ; Cape Town, Table Mountain, Fernwood Gully indigenous forest (33∞58¢S, 18∞27¢E), 150 m a.s.l., December 18, 1996 (C. E. Griswold), 7♂ 5♀ ( CAS) ; Table Mountain, Newlands Ravine indigenous forest (33∞58¢S, 18∞27¢E), 120 m a.s.l., December 18, 1996 (C. E. Griswold), 5♂ 4♀ ( CAS) .

SPERMOPHORA PEMBAI SP. NOV.

( FIGS 28 , 252–255 )

Type. Male holotype from Great Fish River Reserve (33∞08¢S, 26∞39¢E), Eastern Cape, South Africa; at boundary of Farm Ulster, on soil, pittrap, December 3, 1993 (M. Burger), in NCP (96/113) .

Etymology. Named after George Milwa Mnyaluza Pemba (1912–2001), one of South Africa’s greatest pioneering artists.

Diagnosis. Large species with elongated and pointed opisthosoma, distinguished from southern African congeners by the shape of the bulb (no pointed projection, shape of apophysis; Fig. 252 ), and by the wide distance between the long male cheliceral apophyses ( Fig. 254 ).

Male (holotype). Total length 3.2 (3.3 with clypeus), carapace width 0.9. Leg 1: 4.9 + 0.3 + 4.7, metatarsus and tarsus missing, tibia 2: 2.8, tibia 3: 2.0, tibia 4: 2.9; tibia 1 L/d: 53. Habitus as in Figure 28 . Carapace ochre-yellow with darker median band, ocular area medially light, clypeus dark brown under triads, sternum dark brown with slightly lighter spots at bases of coxae and medially. Legs ochre-yellow; opisthosoma grey with few blackish spots dorsally and prominent brown pattern ventrally. Ocular area slightly elevated; distance PME–PME 115 Mm; diameter PME 90 Mm; no trace of AME. Clypeus unmodified. Sternum wider than long (0.70/0.55). Chelicerae as in Figures 254, 255 , with pair of long apophyses, tips 360 Mm apart, without modified hairs on tips. Palps as in Figures 252, 253 ; trochanter with retrolateral apophysis (prominent in dorsal view); femur proximodorsally without apophysis; procursus distally very complex, with up to three hinged structures ( Fig. 252 ). Retrolateral trichobothrium of tibia 1 at 12%.

Variation. One of the males is much paler and the pattern on the carapace very indistinct. Tibia 1 missing in other males.

Female. Unknown.

Distribution. Known from type locality only ( Fig. 264 ).

Material examined. SOUTH AFRICA: EASTERN CAPE: Great Fish River Reserve : type above, together with 2♂ ( NCP 96 /113) .

SPERMOPHORA SUURBRAAK SP. NOV.

( FIGS 30 , 256–258 )

Type. Male holotype from 10 km E of Suurbraak (34∞01¢S, 20∞46¢E), Western Cape, South Africa; tall fynbos, pitfall, January 12–18, 1989 (R. Jocqué), in MRAC (169.700) .

Etymology. Named after the type locality.

Diagnosis. Medium size species with oval opisthosoma, distinguished from southern African congeners by the shapes of the bulb (single projection, Fig. 257 ), the procursus ( Figs 257, 258 ), and the male cheliceral apophyses ( Fig. 256 ). The ZFMK has a very close relative from Signal Hill, Cape Town, with minimally different procursus but with the male cheliceral apophyses much wider apart (distance between tips: 150 Mm).

Male (holotype). Total length 2.1 (2.3 with clypeus), carapace width 0.9. Leg 1: 16.15 (4.2 + 0.35 + 4.45 + 5.4 + 1.75), tibia 2: 2.9, tibia 3: 2.1, tibia 4 missing; tibia 1 L/d: 50. Habitus as in Figure 30 . Carapace pale ochre with median and lateral blackish lines, sternum brown with dark brown margins and many light speckles. Legs ochre-yellow; opisthosoma ochre-grey with blackish pattern also ventrally. Ocular area slightly elevated; distance PME–PME 140 Mm; diameter PME 70 Mm; no trace of AME. Clypeus unmodified. Sternum wider than long (0.65/0.55). Chelicerae as in Figure 256 , with pair of apophyses curved backwards at tips; tips 45 Mm apart. Palps as in Figures 257, 258 (the angle between femur and tibia may not represent the position at rest, but the two segments are flattened at the contact area suggesting that this position is natural at some time); trochanter with short ventral and distinct retrolateral apophysis; patella ventrally almost completely reduced; procursus distally very complex, entire distal part appears hinged; bulb with distinctive embolar division, sclerotized in middle part, but membranous proximally and distally ( Figs 257, 258 ). Retrolateral trichobothrium of tibia 1 at 15%. Tarsus 1 with ~15 pseudosegments, quite distinct distally.

Female. Unknown. Distribution. Known from type locality only ( Fig. 264 ).

Material examined. SOUTH AFRICA: WESTERN CAPE: 10 km E Suurbraak : type above .

Figures

Figure 264. Known distribution of Quamtana gen. nov. (contd.) and Spermophora Hentz within South Africa.

Figures 226–231. Spermophora schoemanae sp. nov. Left male palp in prolateral (226) and retrolateral (227) views, male chelicerae in frontal view (228), modified hair on male cheliceral apophysis (229), and cleared epigynum in ventral (230) and dorsal (231) views. ba: bulbal apophysis, e: embolus, f: Spermophora-flap, hp: hinged process, pp: pointed projection of bulb. Arrowed: femur-apophysis. Scale bars = 30 Mm (229), 0.3 mm (228, 230, 231), 0.4 mm (226, 227).

Figures 232–237. Spermophora gordimerae sp. nov. Left male palp in prolateral (232) and retrolateral (233) views, male chelicerae in frontal view (234), modified hair on male cheliceral apophysis (235), and cleared epigynum in ventral (236) and dorsal (237) views. ba: bulbal apophysis, e: embolus, f: Spermophora-flap, hp: hinged process. Arrowed: femur-apophysis. Scale bars = 30 Mm (235), 0.2 mm (234), 0.3 mm (236, 237), 0.4 mm (232, 233).

Figures 16–30. Quamtana and Spermophora habitus, males in dorsal and lateral views. 16. Q. tsui. 17. Q. merwei. 18. Q. bonamanzi. 19. Q. embuleni. 20. Q. filmeri. 21. Q. knysna. 22. Q. kabale. 23. Q. oku. 24. Q. kitahurira. 25. Q. biena. 26. S. schoemanae. 27. S. gordimerae. 28. S. pembai. 29. S. peninsulae. 30. S. suurbraak.

Figures 31–50. Quamtana and Spermophora, female opisthosomata, ventral views with epigynum. 31. Q. entabeni. 32. Q. mbaba. 33. Q. umzinto. 34. Q. meyeri. 35. Q. nylsvley. 36. Q. molimo. 37. Q. leptopholcica. 38. Q. hectori. 39. Q. mabusai. 40. Q. nandi. 41. Q. merwei. 42. Q. bonamanzi. 43. Q. embuleni. 44. Q. filmeri. 45. Q. kabale. 46. Q. oku. 47. Q. biena. 48. S. schoemanae. 49. S. gordimerae. 50. S. peninsulae.

Figures 244–251. Spermophora schoemanae sp. nov. (244–247) and S. peninsulae (248–251). 244, 248, Male gonopores with epiandrous spigots. 245, 249, Male ALS. 246, 250, Male cheliceral apophyses with modified hairs. 247, 251, Male palpal tarsal organs.

Figures 238–243. Spermophora peninsulae. Left male palp in prolateral (238) and retrolateral (239) views, male chelicerae in frontal view (240), modified hair on male cheliceral apophysis (241), and cleared epigynum in ventral (242) and dorsal (243) views. ba: bulbal apophysis, e: embolus, f: Spermophora-flap, hp: hinged process. Arrowed: femur-apophysis (239) and distinctive small cone on male chelicera (240). Scale bars = 30 Mm (241), 0.3 mm (240, 242, 243), 0.5 mm (238, 239).

Figures 252–255. Spermophora pembai sp. nov. Left male palp in prolateral (252) and retrolateral (253) views, and male chelicerae in frontal (254) and lateral (255) views. ba: bulbal apophysis, e: embolus. Scale bars = 0.2 mm (254, 255), 0.5 mm (252, 253).

Figures 256–258. Spermophora suurbraak sp. nov. Male chelicerae in frontal view (256), and left male palp in prolateral (257) and retrolateral (258) views. Scale bars = 0.2 mm (256), 0.5 mm (257, 258).