Smeringopus Simon, 1890

HUBER, BERNHARD A., 2012, Revision and cladistic analysis of the Afrotropical endemic genus Smeringopus Simon, 1890 (Araneae: Pholcidae), Zootaxa 3461 (1), pp. 1-138 : 8-13

Smeringopus Simon, 1890

Smeringopus Simon 1890: 94 ; type species by original designation: Pholcus elongatus Vinson, 1863 = Pholcus pallidus Blackwall, 1858 . Simon 1893: 476. Kraus 1957: 217. Timm 1976: 70–72.

Diagnosis. Relatively large pholcids (body length usually about 5–8 mm) with elongate abdomen ( Figs. 2–13 ), usually with vivid dark pattern, deep thoracic pit ( Figs. 82 , 414 ), male palpal femur usually with deep retrolateral furrow with distinct proximal rim ( Figs. 344 , 411 ; absent in S. mpanga , S. ruhiza , and chogoria group), cymbium with macrotrichia ( Figs. 158 , 797 ), legs usually with curved hairs on tibiae and metatarsi (absent in thomensis group and cylindrogaster group), without spines on male femora (present in S. saruanle ). Distinguished from Smeringopina by absence of proximal lateral apophyses on male chelicerae, by barely modified male palpal trochanter, and by low number of modified hairs on male chelicerae (usually one on each side, rarely zero). Distinguished from other Smeringopinae genera by male gonopore with only two epiandrous spigots ( Figs. 159 , 347 ) and by absence of stridulatory ridges on chelicerae.

Description. Male: Total body length ~3.5–10 (usually ~5–8); carapace width 1.2–3.3 (usually 1.5–2.5). Carapace with deep pit; ocular area weakly raised, eye triads relatively close together (distance PME-PME usually about same as PME diameter), each secondary eye accompanied by more or less distinct elevation ( Figs. 215 , 539 ; ‘pseudo-lenses’; cf. Huber 2009), AME relatively large, in low position. Clypeus high, never modified, usually with pair of dark stripes ( Figs. 179 , 570 ). Chelicerae never with stridulatory ridges, usually with pair of small apophyses near fang-joints, each provided with one modified hair ( Figs. 213 , 417 ); representatives of the rubrotinctus group with larger apophyses and without modified hairs ( Figs. 87 , 106 ). Palpal coxa with or without retrolateral apophysis, trochanter barely modified, femur usually with deep retrolateral furrow with distinct proximal rim ( Figs. 344 , 411 ; absent in S. mpanga , S. ruhiza , and chogoria group), cymbium always with macrotrichia ( Figs. 158 , 797 ), sometimes with process near palpal tarsal organ ( Figs. 357 , 367 ; very long in S. chogoria and S. bujongolo : Figs. 131, 132 ), palpal, tarsal organ capsulate ( Figs. 395 , 421 ), procursus never with hinge, tip usually with spine-like process and membranous structures, bulb with usually rather complex embolus and one or two processes arising from embolus or fused proximally to embolus.

Legs long and thin, leg 1 length ~25–80 (usually ~35–60), tibia 1 ~6–20 ( Fig. 14 ; usually ~7–15), tibia 2 usually shorter than tibia 4 ( Fig. 16 ), especially in small species ( S. saruanle , S. oromia ). Tibia 1 L/d usually ~40–70, higher only in leaf-dwelling species ( cylindrogaster group: ~80–95). Legs usually without spines on femora (present in S. saruanle ), with curved hairs on tibiae and metatarsi (absent in thomensis group and cylindrogaster group), retrolateral trichobothrium very proximal (at 1.5–3.0%), prolateral trichobothrium always present (also on tibiae 1; Fig. 644 ). Tarsal pseudosegments very indistinct, apparently never regular rings but rather irregular platelets ( Figs. 89 , 346 ).

Abdomen elongate, posteriorly rather pointed, never elevated above spinnerets, usually with distinct dark pattern dorsally, oblique lines or marks laterally, and distinctive ventral pattern usually consisting of dark epigastric area, two or three black lines in median part and two lines in posterior part ( Figs. 559, 567 ). Male gonopore always with two spigots ( Figs. 159 , 347 ), each ALS with large widened spigot, pointed spigot, and 5–6 cylindrically shaped spigots ( Figs. 419 , 665 ; S. thomensis with only two small cylindrically shaped spigots: Fig. 737 ; other species of the thomensis group also with reduced number but not studied with SEM).

Female usually very similar to male, no sexual dimorphism in PME-PME distance, chelicerae unmodified, legs usually slightly shorter than in males, only in S. natalensis on average longer ( Fig. 15 ). Epigynum either a simple plate ( Figs. 168 , 399 ) or provided with pair of pockets ( Figs. 666, 672 ), very derived in S. isangi (with additional pair of pockets on lateral membranous processes; Fig. 542 ). Internal genitalia with frontal valve that is sometimes widened and divided medially (e.g. Figs. 616 , 790 ); pores of pore plates either homogeneously distributed ( Figs. 99 , 114 ) or in groups ( Figs. 349 , 397 ); rarely with internal pockets in female genitalia ( Figs. 342 , 361 ).

Monophyly. In this monograph, I follow a conservative approach, keeping S. rubrotinctus and close relatives (the rubrotinctus species group) in Smeringopus even though preliminary molecular data (Dimitrov, Astrin & Huber, in press) suggest that these taxa (together with the two species comprising the chogoria group) may be more closely related with Smeringopina than with Smeringopus . Following this conservative delimitation of Smeringopus , all cladistic analyses above agree on two synapomorphies for the genus: (1) the presence of curved hairs on the legs (char. 20); and (2) the presence of macrotrichia on the male palpal cymbium (char. 24). The analyses using equal and implied character weighting at K=6 identify one or two further synapomorphies, none of them convincing (char. 25: presence of process on cymbium near palpal tarsal organ; char. 35: epigynum with large posterior indentation). A re-delimited Smeringopus (as suggested by molecular data, i.e. excluding the rubrotinctus group and the chogoria group) would also be supported by two morphological characters: (1) male palpal femur with retrolateral furrow (char. 23); and (2) pore plates in female internal genitalia with pores arranged in groups (char. 36). Further analyses (including a more complete sample of Smeringopina and DNA sequences of more species) are needed to decide if the rubrotinctus and chogoria groups belong either to Smeringopus or to Smeringopina .

Generic relationships. All analyses, under all weighting regimes used, agree on a sister group relationship between Smeringopus and Smeringopina . Three morphological characters support this relationship: (1) an elongated abdomen (char. 3); (2) the unique (among Pholcidae ) reduction of epiandrous spigots from 4 to 2 (char. 8); (3) the loss of cheliceral stridulation (char. 9). Preliminary molecular data support this close relationship (Dimitrov, Astrin & Huber, in press), but the details need further investigation (especially regarding the positions of the rubrotinctus and chogoria groups; see Monophyly above).

The outgroup sample is too small to allow well-founded conclusions about relationships among the other Smeringopinae , but it is noteworthy that Cenemus (which is endemic to the Seychelles) consistently groups with these other genera (which are geographically restricted to northern Africa, the Mediterranean, and the Middle East) rather than with Smeringopus and Smeringopina (the two sub-Saharan genera). Cenemus shares with these northern genera the reduction of ALS spigots from 7–8 to 2, but it also shares a derived character with the two sub-Saharan genera (the elongated abdomen, char. 3).

Specific relationships. Based on the cladistic analyses above and partly also on superficial similarity and geographic closeness, Smeringopus is here divided into twelve operational species groups, three of them monospecific. Some of these groups are likely monophyletic (e.g., rubrotinctus group, chogoria group, cylindrogaster group, thomensis group), one may be monophyletic even though the cladograms suggest otherwise ( hypocrita group), and at least one is very probably not monophyletic ( peregrinus group). This grouping, even though preliminary and not strictly cladistic, structures the existing diversity and reflects biogeographic patterns. In the descriptive section below, species are ordered according to species groups in the order used here (which in turn is derived from the cladogram in Fig. 1 ).

1. rubrotinctus group. This group includes five species ( S. rubrotinctus , S. mgahinga , S. bwindi , S. ruhiza , S. mpanga ) with an epigynum with posterior indentation ( Figs. 38–57 ; char. 35). In the analyses above, the group is always sister to all other Smeringopus , but molecular data (Dimitrov, Astrin & Huber, in press) suggest a closer relationship with the chogoria group than with other groups. These two groups also share a rather dark, almost purplish coloration ( Figs. 18–23 , 115–118 ) and the geographic distribution ( Fig. 58 ).

2. chogoria group. This group includes two species that are extremely similar in most respects and share a very long process on the male palpal cymbium ( Figs. 131 , 157 ; char. 26).

3. S. ngangao . This species appears very isolated morphologically, especially by the absence of curved hairs on legs and by the absence of a distal apophysis on the procursus.

4. arambourgi group. Three species ( S. arambourgi , S. oromia , S. turkana ) of this largely Ethiopian group share a transversal dark band ventrally on the abdomen ( Figs. 170, 172, 174 ; char. 6). In two species ( S. lineiventris , S. saruanle ) the ventral abdominal pigment is largely or entirely reduced and the transversal band is thus barely visible or absent. Two further species are very likely part of this group but the types are lost and no new material is known to me ( S. affinitatus , S. zonatus ).

5. natalensis group. Most species of this large southern African group ( S. natalensis , S. koppies , S. badplaas , S. florisbad , S. lesnei , S. harare , S. blyde , S. hanglip , S. lydenberg ) are characterized by three (rather than one or two) processes arising from the genital bulb ( Figs. 302, 307 ; char. 32). Two species ( S. mlilwane , S. ndumo ) have only two processes arising from the genital bulb but are tentatively assigned to this group because of other specific similarities (e.g., ventrally strongly curved procursus; female genitalia) and geographic closeness.

6. S. pallidus . The type species of the genus appears very similar to representatives of the arambourgi group above but shares with the following groups a distinct retrolateral apophysis on the male palpal coxa ( Figs. 413 , 732 , 802 ; char. 21).

7. S. lesserti . This species shares with all the following groups the presence of epigynal pockets ( Figs. 423 , 666, 672 ; char. 33) but otherwise (bulbal apophyses, procursus tip) it appears very isolated.

8. hypocrita group. This group is restricted to southern Africa but in contrast to the natalensis group rather to the western than to the eastern side ( Fig. 475 ). The cladistic analyses never resolved this group as monophyletic, but apart from their geographic distribution, the species share a procursus tip that is strongly bent towards prolateral ( Figs. 477, 484 ; char. 18). The cladistic analyses separate the group into a more southern clade ( S. hypocrita , S. sederberg , S. dehoop ; probably also S. lotzi and S. ubicki that were not included in the matrix) and a more northern clade ( S. atomarius , S. uisib , S. tombua ).

9. cylindrogaster group. This group is unique (among Smeringopus ) in its pale coloration, resulting from its unique biology (high in the vegetation rather than near the ground; see Natural history below). The three species ( S. luki , S. isangi , S. cylindrogaster ) share a distinctive color pattern (abdomen dorsally monochromous, ventrally with black spots), an elongated cymbium, and a prominent proximal ventral process of the procursus. Two species occur in central Africa; the third is the only Smeringopus (other than S. pallidus ) that also occurs in western Africa.

10. peregrinus group. Representatives of this large group share with the two following groups a distinctive structure in the female internal genitalia (part of the valve appears medially widened and divided; Figs. 616 , 790 ; char. 37) but the group itself is probably not monophyletic. It is widely distributed in central and eastern Africa ( S. peregrinus , S. peregrinoides , S. katanga , S. butare , S. dundo ) but reaches further south until Namibia ( S. similis ), Zimbabwe, and Mozambique ( S. kalomo , S. chibububo ). S. moxico is tentatively assigned to this group even though the cladistic analysis suggests otherwise.

11. thomensis group. The three species of this group ( S. thomensis , S. principe , S. mayombe ) share the loss of curved hairs on the legs (char. 20) and the reduction of ALS spigots (char. 7; in S. thomensis two tiny cylindrically shaped spigots are still present: Fig. 737 ; in the other two species the spinnerets were studied in the light microscope only and it remains unknown if the spigots are entirely reduced or if two tiny spigots are still present too). They also share a distinctive pattern dorsally on the abdomen ( Figs. 699, 701, 703 ; not coded) and the geographic distribution ( São Tomé and Príncipe Islands and western Congo D.R.).

12. roeweri group. The four species of this group ( S. roeweri , S. lubondai , S. carli , S. sambesicus ) share rounded light projections proximally on the male chelicerae ( Figs. 771 , 779 ; char. 10). The group is widely distributed in central and eastern Africa.

Natural history. Even though S. pallidus is a pantropical species, very few studies have been dedicated to exploring its biology in any detail. Jackson (1992) and Jackson et al. (1992) studied whirling behavior as a defense mechanism against predators such as web-invading jumping spiders. It is remarkable that in many places S. pallidus seems to have been largely replaced by other synanthropic species such a Physocyclus globosus . For example, Sánchez Roig (1911) cites S. pallidus (under Pholcus tipuloides ) as “una de las especies más vulgares de Cuba ”, but this does no longer seem to be the case (A. Pérez G. & B. A. Huber, unpubl. obs.); also, Mello-Leitão (1918, 1946) reports the species (under S. geniculatus ) to occur in “todo o Brazil, no littoral” and to be “muito comum no interior das habitações” in Rio de Janeiro, but the species does no longer seem to be common along the Brazilian coast (B.A. Huber, unpubl. obs.).

For most other species, label data and observations by the author in South Africa, Kenya, Uganda, Cameroon, Gabon, and Guinea are the only sources of information. Most species prefer the same type of shady, protected habitat that is typical for pholcid spiders: holes and caverns, undersides of overhangs, dark spaces under logs and rocks and between buttresses. Here the spiders build their sheet webs that are more or less domed, with the animal hanging from the apex of the dome. Unlike most other pholcids but like many other Smeringopinae , several species of Smeringopus seem to be fairly tolerant against aridity. This may explain to some degree the fact that several species have invaded human constructions and it is thus remarkable that only one species has spread all over the world.

Smeringopus occurs from sea level to over 3700 m, but only representatives of the rubrotinctus species group have been found beyond 2300 m. Most species of the rubrotinctus group appear restricted to high altitudes, and S. bujongolo is currently the pholcid spider with the highest known record in Africa (at 3780 m).

Most exceptional both in its morphology and behavior is S. cylindrogaster (its two close relatives share the morphology but their behavior has not been studied). Smeringopus cylindrogaster has shifted its microhabitat to the undersides of alive (green) leaves where it rests in an unusual lamp-shade web in an inverted position ( Fig. 13 ; Huber 2009). The entire spider has changed to a pale whitish coloration with black spots that break the contours. Only the dorsal side of the abdomen (that is pressed against the leaf) is monochromous (cf. Figs. 530, 534 ). Unlike other Smeringopus species that whirl or vibrate their bodies when disturbed, S. cylindrogaster remains tightly pressed against the leaf.

The facultative construction of silk balls that are attached to the domed webs appears to be plesiomorphic for Smeringopus (see Cladistic analysis, char. 40), but only in a few species have such silk balls actually been observed: in S. pallidus ( Japyassú & Macagnan 2004) , S. cylindrogaster ( Huber 2009) , S. peregrinus ( Figs. 8 , 623–625 ) and S. ngangao ( Fig. 7 ). Silk balls also occur in other genera of Smeringopinae ( Hoplopholcus , Holocnemus ; Wiehle 1933; Sedey & Jakob 1998; J. &. F. Murphy, unpubl. obs.), but they have been studied in more detail in only two species ( Hajer & Řeháková 2003; Japyassú & Macagnan 2004). The conditions under which such structures are incorporated into the web seem to vary among taxa.

Distribution. With the exception of the pantropical S. pallidus , Smeringopus is largely restricted to central, eastern, and southern Africa ( Fig. 17 ). Other than S. pallidus , only two species occur outside Africa: (1) S. natalensis (originally from southern Africa) has been able to establish stable populations in Western Australia and New South Wales ( Huber 2001); (2) S. lineiventris is only known from Yemen. Only two species occur in western Africa ( S. pallidus and S. cylindrogaster ), where Smeringopus is largely replaced by Smeringopina . The two genera have a wide range of overlap in central Africa. The same occurs with Crossopriza in the Sahel and in north-eastern Africa. At least four species occur on Madagascar ( S. pallidus , S. carli , S. peregrinus , S. kalomo ), but none of them is endemic to the island and it is likely that at least three of them are recent (possibly human) introductions.

Composition. As delimited here, Smeringopus now includes 55 described species, 36 of which are newly described below. The collections seen include about 20 further undescribed species that are not treated for various reasons: some are very similar to species treated herein; some are only represented by poorly preserved specimens; most are represented by only one sex. Considering the patchiness of collecting efforts and known distribution patterns in the genus, it appears likely that 50% of the actual species may remain undescribed.

Figures

FIGURE 1. One of the four most parsimonious cladograms resulting from the analysis of the matrix in Appendix 1 using successive weighting in NONA. See Cladistic analysis for discussion.

FIGURE 17. Known distribution of Smeringopus. Excluded is Smeringopus pallidus which has a worldwide distribution and whose distribution in Africa is shown in Fig 386.

FIGURE 58. Known distributions of the rubrotinctus group, the chogoria group, and S. ngangao. Squares: further undescribed species.

FIGURE 475. Known distribution of the hypocrita group. Squares: further undescribed species.

FIGURES 2–13. Smeringopus, alive specimens in their natural habitats. 2. S. bwindi, female with eggsac (Uganda, Bwindi). 3–4. S. chogoria, females with eggsac and spiderlings (Kenya, Mau Mau and Chogoria). 5. S. mpanga, female with spiderlings (Uganda, Kalinzu). 6–7. S. ngangao, male and web with silk-balls (Kenya, Ngangao). 8–10. S. peregrinus, web with silk-balls (Kenya, Gonkonyi), female (Kenya, Hell’s Gate), female with eggsac (Gonkonyi). 11. S. pallidus, female with eggsac (Uganda, Semuliki). 12. S. lesserti, male (Gabon, Mayebout). 13. S. cylindrogaster (Guinea, Dieke).

FIGURES 81–93. Smeringopus mgahinga. 81–82. Male prosoma, frontal and dorsal views. 83–84. Right palp, retrolateral and retrolatero-dorsal views. 85. Left procursus and embolus, prolateral view. 86. Tip of right procursus. 87. Tip of male cheliceral apophysis. 88. Trichobothria on male tibia 1. 89. Male tarsus 1 pseudosegmentation. 90. Male tarsus 1 claws. 91. Male gonopore. 92. Epigynum. 93. Female ALS. Scale lines: 20 µm (86), 30 µm (87, 93), 40 µm (89), 50 µm (90), 100 µm (85, 88, 91), 200 µm (83, 84, 92), 500 µm (82), 600 µm (81).

FIGURES 411–423. Smeringopus lesserti. 411. Left male palpal femur, retrolateral view. 412–413. Left procursus tip, distal and retrolateral views. 414. Female prosoma. 415. Male bulbal processes, prolatero-dorsal view. 416–417. Male cheliceral apophyses. 418. Male gonopore. 419. Male ALS. 420. Male PMS. 421. Male palpal tarsal organ. 422. Female ocular area. 423. Epigynum. Scale lines: 10 µm (416–417), 20 µm (419–421), 80 µm (412–413, 418), 100 µm (411, 415), 200 µm (422–423), 500 µm (414).

FIGURES 343–356. Smeringopus natalensis (343–349) and S. hanglip (350–356). 343. Left procursus tip, retrolateral view. 344. Left palpal femur, ventral view. 345. Male cheliceral apophysis. 346. Tarsal pseudosegments. 347. Male gonopore. 348. Cleared female genitalia, dorsal view. 349. Detail of pore plate. 350. Left procursus, dorsal view. 351. Male cheliceral apophysis. 352. Bulbal processes. 353. Male ALS and PMS. 354. Male gonopore. 355. Epigynum. 356. Female ALS. Scale lines: 20 µm (345, 351), 30 µm (346, 356), 40 µm (353), 50 µm (349), 60 µm (343), 80 µm (347), 100 µm (344, 352), 200 µm (348, 350, 354), 300 µm (355).

FIGURES 155–168. Smeringopus chogoria (155–164) and S. ngangao (165–168). 155–156. Right palp, retrolatero-dorsal and prolatero-dorsal views. 157–158. Left palp, dorsal and prolatero-dorsal views. 159. Male gonopore. 160–161. Male chelicerae and cheliceral apophysis (arrow points to modified hair). 162–163. Male and female ALS. 164. Epigynum. 165. Male gonopore. 166. Modified hair on male cheliceral apophysis. 167. Female ALS. 168. Epigynum. Scale lines: 10 µm (166), 20 µm (161, 167), 30 µm (162, 163), 50 µm (165), 80 µm (159), 100 µm (160), 200 µm (155–158, 168), 300 µm (164).

FIGURES 791–804. Smeringopus roeweri (791–800) and S. carli (801–804). 791. Male prosoma, frontal view. 792–793. Left procursus, prolateral and dorsal views. 794. Left embolus. 795. Male gonopore. 796. Male ocular area. 797. Cymbial macrotrichia. 798. Epigynum. 799. Female ALS. 800. Female PMS. 801. Epigynum. 802. Left palp, retrolateral view. 803–804. Male and female ALS. Scale lines: 20 µm (799–800, 803–804), 50 µm (797), 80 µm (792–793), 100 µm (794–795), 200 µm (796, 798, 801–802), 500 µm (791).

FIGURES 210–218. Smeringopus arambourgi. 210. Left procursus tip, retrolatero-distal view. 211. Left bulbal processes, prolaterodistal view. 212–213. Male chelicerae and cheliceral apophysis. 214. Male ALS. 215. Female ocular area (arrows point to ‘pseudolenses’). 216. Female ALS. 217. Cleared female genitalia, dorsal view. 218. Detail of pore plate. Scale lines: 10 µm (213), 20 µm (214, 216), 30 µm (218), 80 µm (212), 100 µm (210, 211, 215), 200 µm (217).

FIGURES 530–544. Smeringopus cylindrogaster group. S. luki (530–533, 538–541) and S. isangi (534–537, 542–544). 530–531, 534–535. Males, dorsal and ventral views. 532–533, 536–537. Left male palps, prolateral and retrolateral views. 538. Male prosoma, frontal oblique view. 539. Male ocular area. 540–544. Female epigyna and cleared female genitalia, dorsal (541, 544) and ventral (543) views (arrows point to median and lateral pockets).

FIGURES 169–183. Smeringopus arambourgi group, habitus and male prosomata, oblique frontal views. 169–170. S. arambourgi, male, dorsal view and female, ventral view. 171–172. S. turkana, male, dorsal view and female, ventral view. 173–174. S. oromia, male, dorsal and ventral views. 175–176. S. saruanle, male, dorsal view and female, ventral view. 177–178. S. lineiventris, male, dorsal view and female, ventral view. 179. S. arambourgi. 180. S. turkana. 181. S. oromia. 182. S. saruanle. 183. S. lineiventris.

FIGURES 558–571. Smeringopus peregrinus group, habitus and male prosomata, oblique frontal views. 558–559. S. peregrinus, male, dorsal and ventral views. 560–561. S. peregrinoides, female, dorsal view and male, ventral view. 562. S. butare, male dorsal view. 563. S. similis, male, dorsal view. 564–565. S. kalomo, male and female, dorsal views. 566–567. S. chibububo, male, dorsal view and female, ventral view. 568. S. katanga, male, dorsal view. 569. S. moxico, male, dorsal view. 570. S. chibububo. 571. S. kalomo.

FIGURES 106–114. Smeringopus mpanga. 106. Male cheliceral apophyses. 107. Left male palpal femur. 108. Tip of left procursus. 109. Male gonopore. 110–111. Epigynum, ‘closed’ and ‘open’. 112. Female ALS and PMS. 113. Female internal genitalia, dorsal view. 114. Detail of pore plate. Scale lines: 20 µm (112), 30 µm (114), 60 µm (106, 108), 80 µm (109), 200 µm (107, 113), 300 µm (110, 111).

FIGURES 357–366. Smeringopus lydenberg (357–361) and S. mlilwane (362–366). 357, 362. Left cymbia and procursi, retrolateral views. 358, 363. Left procursi, dorsal views. 359–360, 364–365. Left bulbal processes, prolateral and dorsal views. 361, 366. Cleared female genitalia, dorsal views (arrows point to internal pockets). Scale lines: 0.3 mm.

FIGURES 367–371. Smeringopus ndumo. 367. Left cymbium and procursus, retrolateral view. 368. Left procursus, dorsal view. 369–370. Left bulbal processes, prolateral and dorsal views. 371. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm.

FIGURES 131–137. Smeringopus chogoria (131–136) and S. bujongolo (137). 131. Left cymbium and procursus, retrolateral view. 132. Left procursus, dorsal view. 133–134. Left embolus, prolateral and dorsal views. 135. Male chelicerae, frontal view. 136–137. Cleared female genitalia, dorsal views. Scale lines: 0.3 mm (133–134), 0.5 mm (131–132, 135–137).

FIGURES 394–400. Smeringopus pallidus. 394. Left procursus tip, retrolatero-dorsal view. 395. Male palpal tarsal organ. 396. Female ALS. 397. Detail of pore plate. 398. Male gonopore. 399. Epigynum. 400. Cleared female genitalia, dorsal view. Scale lines: 10 µm (395), 20 µm (396–397), 60 µm (394), 100 µm (398, 400), 200 µm (399).

FIGURES 14–16. Scatter diagram and histograms of specific measurements in Smeringopus. 14. Representatives of the cylindrogaster group have relatively slender legs in relation to their length. 15. Of the 26 species with at least 5 males and 5 females measured, only S. natalensis has shorter male than female legs. 16. In most species, tibia 2 is shorter than tibia 4, which is unusual in long-legged pholcids; rather usual values are found in representatives of the cylindrogaster group (c) and the thomensis group (t).

FIGURES 634–646. Smeringopus peregrinoides. 634. Male prosoma, frontal view. 635. Left male palp, dorsal view. 636. Left procursus tip, retrolatero-dorsal view. 637. Left bulb, prolateral view. 638. Male ocular area. 639. Left embolus, prolatero-distal view. 640. Male gonopore. 641–642. Male chelicerae and cheliceral apophysis. 643. Male ALS. 644. Male tibia 1, trichobothria. 645. Epigynum. 646. Female ALS. Scale lines: 10 µm (642), 20 µm (643, 646), 50 µm (636), 80 µm (639–641), 100 µm (637, 638, 644), 200 µm (635, 645), 400 µm (634).

FIGURES 663–672. Smeringopus butare (663–666) and S. kalomo (667–672). 663. Male cheliceral apophysis. 664. Male gonopore. 665. Male ALS. 666. Epigynum. 667. Male cheliceral apophysis. 668–669. Left procursus, retrolatero-dorsal and dorsal views. 670. Male gonopore. 671. Male ALS. 672. Epigynum. Scale lines: 10 µm (663, 667), 20 µm (671), 30 µm (665), 60 µm (668), 80 µm (664, 669), 100 µm (670), 200 µm (666, 672).

FIGURES 732–744. Smeringopus thomensis (732–740) and S. principe (741–744). 732. Left male palpal femur, retrolateral view. 733. Left procursus tip, prolateral view. 734. Male cheliceral apophysis. 735. Left embolus, distal view. 736. Male gonopore. 737. Female ALS. 738. Epigynum. 739. Cleared female genitalia, dorsal view. 740. Detail of pore plate. 741. Left embolus, distal view. 742. Left procursus tip, prolateral view. 743. Cleared female genitalia, dorsal view. 744. Detail of pore plate. Scale lines: 10 µm (734, 737), 20 µm (740), 30 µm (744), 80 µm (733, 735–736), 100 µm (739, 741–742), 200 µm (732, 738, 743).

FIGURES 609–616. Smeringopus peregrinus. 609. Left cymbium and procursus, retrolateral view. 610. Left procursus, prolateral view. 611–613. Left embolus, prolateral, dorsal, and ventral (slightly prolateral) views (arrow points to variable structure; see text). 614–615. Male chelicerae, frontal and lateral views. 616. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm.

FIGURES 782–790. Smeringopus carli. 782. Left cymbium and procursus, retrolateral view. 783–784. Left procursus, prolateral and dorsal views. 785–787. Left embolus, prolateral, prolatero-ventral, and retrolatero-ventral views. 788–789. Male chelicerae, frontal and lateral views. 790. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm (782–787, 790), 0.5 mm (788–789).

FIGURES 94–99. Smeringopus ruhiza. 94. Left cymbium and procursus, retrolateral view (arrow points to variable structure, see text). 95. Left procursus, dorsal view. 96–97. Left embolus, prolateral and dorsal views. 98. Male chelicerae, frontal view. 99. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm (96–97), 0.5 mm (94–95, 98–99).

FIGURES 336–342. Smeringopus hanglip. 336. Left cymbium and procursus, retrolateral view. 337. Left procursus, dorsal view. 338–339. Left bulbal processes, prolateral and dorsal views. 340–341. Male chelicerae, frontal and lateral views. 342. Cleared female genitalia, dorsal view (arrow points to internal pocket). Scale lines: 0.3 mm (336–339), 0.5 mm (340–342).

FIGURES 38–57. Smeringopus rubrotinctus group, epigyna, ventral views (47: lateral view) and cleared female genitalia, ventral and dorsal views. 38–41. S. rubrotinctus (38: syntype, 39: Karisimbi, 40–41: Rwegura). 42–45. S. bwindi (Buhoma). 46–49. S. mgahinga (46–47: Ruhiza, 48–49: Mgahinga). 50–53. S. ruhiza (50: Kitahurira, 51–53: Buhoma). 54–57. S. mpanga (Kanyanchu).

FIGURES 18–27. Smeringopus rubrotinctus group, habitus and male prosomata, oblique frontal views. 18. S. rubrotinctus, female, ventral view. 19. S. bwindi, male, dorsal view. 20–21. S. ruhiza, male and female, dorsal views. 22–23. S. mpanga, male and female, dorsal views. 24. S. rubrotinctus. 25. S. mgahinga. 26. S. ruhiza. 27. S. mpanga.

FIGURES 115–130. Smeringopus chogoria group, habitus, left male palps (prolateral and retrolateral views), male prosomata (oblique frontal views), epigyna (ventral views), and cleared female genitalia (ventral and dorsal views). 115–116, 119–120, 123–126. S. chogoria. 117–118, 121–122, 127–130. S. bujongolo.

FIGURES 300–309. Smeringopus natalensis (300–304) and S. koppies (305–309). 300, 305. Left cymbia and procursi, retrolateral views. 301, 306. Left procursi, prolateral views. 302–303, 307–308. Left bulbal processes, prolateral and dorsal views. 304, 309. Cleared female genitalia, dorsal views. Scale lines: 0.3 mm.

FIGURES 476–486. Smeringopus hypocrita (476–482) and S. sederberg (483–486). 476, 483. Left cymbia and procursi, retrolateral views. 477, 484. Left procursi, dorsal views (arrows point to variable structures; see text). 478–479. Male chelicerae, frontal and lateral views. 480–481, 485–486. Left emboli, prolateral and dorsal views (arrow points to variable structure; see text). 482. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm.

FIGURES 699–708. Smeringopus thomensis group. S. thomensis (699–700, 705–706), S. principe (702–704, 707–708), and S. mayombe (701). 699, 701, 703. Males, dorsal views. 700, 702. Male prosomata, oblique frontal views. 704. Female, ventral view. 705–708. Left male palps, prolateral and retrolateral views.

FIGURES 765–773. Smeringopus roeweri. 765. Left cymbium and procursus, retrolateral view. 766–767. Left procursus, prolateral and dorsal views. 768–770. Left embolus, prolateral, prolatero-ventral, and prolatero-dorsal views. 771–772. Male chelicerae, frontal and lateral views. 773. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm.

FIGURES 774–781. Smeringopus lubondai. 774. Left cymbium and procursus, retrolateral view (arrow points to variable structure; see text). 775–776. Left procursus, prolateral and dorsal views. 777–778. Left embolus, prolateral and dorsal views. 779–780. Male chelicerae, frontal and lateral views. 781. Cleared female genitalia, dorsal view. Scale lines: 0.3 mm (777–778), 0.5 mm (774–776, 779–781).

FIGURES 617–625. Smeringopus peregrinus. 617. Left embolus, ventral (slightly prolateral) view. 618. Male gonopore. 619. Male ALS. 620. Epigynum. 621. Cleared female genitalia, dorsal view. 622. Detail of pore plate. 623. Silk ball (from web shown in Fig. 8). 624–625. Details of silk ball. Scale lines: 8 µm (625), 10 µm (624), 30 µm (619), 40 µm (622), 60 µm (618), 80 µm (623), 100 µm (617), 200 µm (620–621).