Vytfutiini, Griswold, 1990

Griswold, Charles E., 2022, The Lace Web Spider Genus Vytfutia Deeleman-Reinhold (Araneae, Phyxelididae) in the Indo-Pacific Region, Proceedings of the California Academy of Sciences 67 (14), pp. 329-355 : 333-335

Vytfutiini , Griswold, 1990

Type genus Vytfutia Deeleman-Reinhold, 1986: 34 . Comprise a single genus, Vytfutia .

Diagnosis, Synapomorphies and Description See below under Vytfutia .

Vytfutia Deeleman-Reinhold, 1986: 34 (type species by monotypy Vytfutia bedel Deeleman-Reinhold , holotype male in RMNH, examined). V. Deeleman-Reinhold, 1986: 34, type V. bedel Deeleman-Reinhold, 1986 ; N.B.: transferred from Agelenidae to Amaurobiidae by Griswold, 1990: 186, to Phyxelididae by Griswold et al., 1999: 59.

Diagnosis: Vytfutia may be distinguished from other Phyxelididae by having the epigynum ( Figs. 24 , 39 , 41 , 48 ) with the posterior median lobe fused to the lateral lobes so that no suture is visible, by the presence of a retrolateral process ( RL) on the male palpal tarsus and a sclerotized dorsal spur on male metatarsus I ( Figs. 16 , 37 , 46 ), by the entire chilum ( Fig. 4 ) and by the PLS lacking the stout, curved, apicolateral seta ( Figs. 12, 13 ).

Synapomorphies: Among the Phyxelididae , the dorsal spur on male metatarsus I ( Figs. 56–61 ), RTA on the male palpal tibia, and fusion of median and lateral sectors of the epigynum into a single plate ( Figs. 24 , 39 , 41 , 48 ) comprise Vytfutia synapomorphies.

Notes: Vytfutia halandrefana and V. pallens are the most morphologically divergent species of Vytfutia , and also very different to each other. Vytfutia halandrefana is small, with a broad thoracic fovea, few leg spines and complex female genitalia with convoluted ducts. Unfortunately, the male of V. halandrefana remains unknown. Vulval morphology suggests that the V. halandrefana male palp has an elongate embolus. Vytfutia pallens is much larger than other Vytfutia , with longer limbs and pale coloration, as befits a cave-dwelling organism.

Description: ( Vytfutia ) Total length 2.12–7.70. Markings differ significantly among species and are described under each species description. Carapace length 1.19–1.43 times width, height 0.39–0.53 width; thoracic fovea typically linear, length 0.11–0.18 carapace length, narrow oval in V. halandrefana , length 0.07 carapace length. PER straight to slightly recurved, AER straight, ocular area width 2.25–3.33 times length; clypeal height typically 1.20-1.70 times AM diameter, but lower in V. halandrefana , 0.60–0.80 times AM diameter; chelicerae with small boss ( Figs. 1 , 35 ), length 6.50–14.00 times clypeal height, pro- and retromargins of fang furrow with 3–5 heterogeneous teeth, retromargin with 3-4 escort setae at fang base, promargin with large group of fluffy whisker setae, rake setae not found (Ramirez 2014); cheliceral gland on flat cuticle, with numerous pores; labrum with flattened, anteriad pointing labral tongue, tongue apex free, deeply concave, with plumose setae dorsally and laterally on labrum, minute, bristle-like setae occur distad of tongue apex; sternum length 1.17–1.39 times width, apex a blunt point ( Figs. 29 , 33 ). Legs long, femur I 1.01–1.48 times carapace length; leg I always longest, III shortest, leg formula 1423, in some species 1243 or 1 [2=4] 3; male metatarsi I ( Figs. 27, 28 , 56–61 ) with retrolateral, concavity at midsegment, concavity lacks spinules, at base of concavity a short, prolateral process surmounted by stout clasping spine, concavity arising at 0.21-0.47 length of metatarsus, concavity length 0.21–0.27 metatarsus, with short, dorsal spur at apex of concavity, spur height 0.06–0.95 of metatarsus length; legs otherwise unmodified; palpal claw and STC with numerous (up to 11) teeth ( Figs. 52, 55 ), ITC with 2-3 teeth or smooth ( V. pallens ); calamistrum subapical, origin at 0.22-0.29 distance from metatarsus base, length 0.33–0.46 metatarsus length ( Figs. 8 , 50 ). Spination (following summary is based on males and females of Vytfutia labalaba ; V. bedel and V. pallens are very similar; the female of V. halandrefana is very different, which is noted in that species’ description). Male: palp: femur d0-1-1, p0-1-0; leg I: femur d1-1-1, p0-0-0-1, r0-0-1, tibia d0-0-1, p0-1-1-0, r1-1-1-0, metatarsus p0-0-1, v2-0-0-2, r0-1(clasper)-0; leg II: femur d1-1-1, p0-0-1-0, r0-0-1-0, tibia p0-1-1-0, r0-1-2-0, metatarsus p0-1-1-2, r0-0-2; leg III: femur d1-1-1, p0-0-1-0, r0-0-1-0, patella d0-1, p1, r1, tibia d1-0-0, p0-1-1-0, v0-0-1-0, r0-1-1-0, metatarsus p1-1-0-2, v2-0-0-2, r1-1-0-1; leg IV: femur d1-1-1, p0-0-1-0, r0-0-1-0, patella d0-1-0, tibia d1-0-0, p0-1-1-0, v0-1-1-0, r0-1-1-0, metatarsus p1-1-0-2, v2-0-0-2, r1-1-0-1. Female: palp: femur d0-0-1-1, r0-0-1; leg I: femur d0-1-1-0, p0-0-0-1, tibia p0-1-1-0, r0-1-1-0, metatarsus p0-0-1, v2-0-0, r0-0-1; leg II: femur d1-1-1, p0-0-1, r0-0-1, tibia d1-1-0-2, v2-0-0-2, r1-1-0-2, p0-1-1-0, metatarsus d0-0-1, p0-1-1-0, v2-0-0-2, r0-0-1-0; leg III: femur d1-1-0-1, p0-0-0-1, r0-0-0-1, patella p1, r1, tibia p0-1-1-0, v1-1-0-0, r0-1-1-0, metatarsus p1-1-0-2, v2-0-2-1, r1-1-0-2; leg IV: femur d1-0-0-1, p0-0-1, r0-0-1, patella d1, tibia p0-1-1-0, v1-1-0-0, r0-1-1-0, metatarsus d2-0-01, p1-1-0-2, v0-1-0-2, r0-0-0-1. Male epiandrum lacking spigots. Tracheae comprise four simple tubes that are limited to the abdomen ( Deeleman-Reinhold, 1986: 34). Spinnerets (based on scanning electron microscope [SEM] examination of three species: males and females of V. pallens Deeleman-Reinhold ( Griswold et al., 2005, figs. 46D, 47, 48) and of V. labalaba new species and a female of V. halandrefana , new species). Vytfutia are typical phyxelidids in cribellum and paracribellar morphology, and universally differ from Vidoleini and Phyxelidini in lacking the large, stout, curved seta (black when viewed with light microscopy) at ectal margin of the PLS (compare Figs. 14 and 12 ). Cribellum width 0.43–0.67 spinnerets width ( Fig. 12 ). Female ALS may have one mesal MAP spigot with a posterior MAP nubbin; V. halandrefana differs in retaining two MAP spigots. Piriform gland ( PI) spigots in females range from as few as 18 ( V. halandrefana ) to more than 50 ( V. pallens ); males have fewer; small tartipores are scattered throughout PI spigot field; female PMS with as few as 10 ( V. halandrefana ) to more than 25 ( V. labalaba ) PC spigots; replaced by nubbins in the male; the PMS of both sexes have one anteromedian mAP spigot and at least two ( V. halandrefana ) to six ( V. labalaba ) AC spigots and one ( V. halandrefana ) to four ( V. pallens ) CY spigots; female PLS with large anterior MS spigot with cylindrical base and shaft, accompanied by a flanking AC, MS and flanking AC are replaced by nubbins in male; female with one ( V. halandrefana ) to three ( V. labalaba ) CY spigots and fewer than eight AC spigots. Palpal femur with anterobasal row of four to eight stout setae set in enlarged bases, these setae shortened and enlarged as thorns ( Figs. 9 , 53, 54 ); male palpal tibia ( Figs. 16 , 46, 47 ) with short to long, convex, simple DTA extending apically, RTA developed as a trapezoidal flange; cymbium with retrolateral paracribellar projection ( Figs. 15 , 38 , 46, 47 ); palpal bulb with a small, oval petiole, anneli of subtegulum weakly developed, tegulum central, convex, course of reservoir a simple curve within bulb, without switchbacks, median apophysis present, fleshy, arising at mid-bulb near embolic base; embolus a slender, narrow, tapering blade to a slender spine, pars pendula fused basally with truncus of embolus, embolus flexibly attached to tegulum; conductor apical, fleshy, grooved to receive apex of embolus ( Figs. 15 , 36, 37 , 45, 46 ). Female genitalia with epigynum ( Figs. 18 , 24 , 39 , 41 ) a simple, undivided plate, copulatory openings slit like, transverse, oblique to longitudinal; vulva with capsulate structures(s) with internal chambers, with 2-3 lobes, at least V. labalaba with sessile HS and posterdorsal Bennett’s gland pore ( Fig. 42 ), may have convoluted ducts ( V. halandrefana , Fig. 25 ).

Composition: Four species, all in the genus Vytfutia .

Distribution: The Indo-Pacific islands of Madagascar, Sumatra and Borneo ( Fig. 62 ).

Key to the species of Vytfutia

1 Males .................................................................... 5

— Females.................................................................. 2

2 (1) Epigynum deep, quadrangle less than 1.00, copulatory opening rims oblique to nearly longitudinal, posterior margin of epigynum convex or concave ( Figs. 18 , 39 , 48 ), vulva with 2-3 simple lobes, without slender ducts, vulva width greater than twice length ( Figs. 19 , 40 , 49 ) ......................................................................... 3

— Epigynum broad, quadrangle greater than 2.50, copulatory opening rims transverse, posterior margin of epigynum concave ( Figs. 23 , 24 ), vulva complex with large anterior lobe and slender ducts, vulva width less than 1.50 times length ( Fig. 25 ), Madagascar............................................................... V. halandrefana , new species

3 (2) Epigynum with copulatory opening rims oblique ( Figs. 39 , 48 ), vulva with integrated lobes anteriad of copulatory openings ( Figs. 40 , 49 ).................................... 4

— Epigynum with copulatory opening rims longitudinal, epigynum posterior margin convex, elongated posteriorly in center ( Fig. 18 ), vulva with spherical lobe laterad of copulatory openings ( Fig. 19 )................................... V. bedel Deeleman-Reinhold, 1986

4 (3) Epigynum with copulatory opening rims converging anteriorly, posterior margin slightly concave ( Figs. 39 , 41 ), vulva with paired lobes laterad of copulatory openings ( Figs. 40 , 42 )......................................................... V. labalaba , new species

— Epigynum with copulatory opening rims diverging anteriorly, posterior margin straight to slightly convex ( Fig. 48 ), vulva with paired lobes far anteriad of copulatory openings ( Fig. 49 )...................................... V. pallens Deeleman-Reinhold, 1989

5 (1) Male palpal tibia with DTA short, less than 0.25 length tibia ( Figs. 37, 38 , 46, 49 ), tibia with subapical spine or cuspule, embolus short, arising near base of bulb (past 6 o’clock) and making turn of less than 180º, median apophysis long, length greater than 5.00 times width ( Figs. 36, 37 , 45, 46 )........................................................ 6

— Male palpal tibia with DTA elongate, greater than 0.50 length tibia ( Figs. 16, 17 ), tibia lacking subapical enlarged seta, embolus long, arising on retrolateral side of bulb (4 o’clock) and making turn of more than 225º, median apophysis short, length less than 4.00 times width ( Figs. 15, 17 )................................... V. bedel Deeleman-Reinhold, 1986

6 (5) Palpal tibia with elongate apical spine ( Figs. 46, 47 ), embolus short, straight, arises on prolateral side of bulb (near 8 o’clock), median apophysis hooked at apex ( Figs. 45, 46 ); metatarsus I long, base of concavity originates at more than 0.45 metatarsus length, spur high, height great than 0.09 metatarsus length ( Figs. 60, 61 )...... V. pallens Deeleman-Reinhold, 1989

— Palpal tibia with short subapical cuspule ( Figs. 37, 38 ), embolus long, curved, arises at base of bulb (near 6 o’clock), median apophysis flattened, spoon shaped at apex ( Figs. 36, 37 ); metatarsus I shorter, base of concavity originates at less than 0.35 metatarsus length, spur low, height greater than 0.08 metatarsus length ( Figs. 58, 59 )......... V. labalaba , new species

Figures

FIGURES 24, 25. Vytfutia halandrefana new species, female genitalia drawings, Tsingy de Bemaraha, Madagascar, CASENT9008848. 24. Epigynum, ventral. 25. Vulva, left half, dorsal.

FIGURES 36–40. Vytfutia labalaba new species, genitalia-drawings, Holotype male and paratype female collected on the wall of a building at 1550 m elevation on Mt. Kinabalu, Sabah-Borneo, Malaysia, CASENT9023842. 36–38. Male left palp, tibia and tarsus. 36. Ventral. 37. Retrolateral. 38. Dorsal. 39, 40. Female genitalia. 39. Epigynum, ventral. 40. Vulva, dorsal.

FIGURES 41–44. Vytfutia labalaba new species, Mt. Kinabalu, Borneo, CASENT9023842, female genitalia, scanning electron micrographs (SEM). 41. Epigynum, ventral. 42. Vulva, dorsal. 43. Bennet’s gland pore (BG), left. 44. Pores of spermathecal head (HS), left. Scale bars, 41 = 100µm, 42 = 30µm, 43, 44 = 10µm.

FIGURES 45–49. Vytfutia pallens Deeleman-Reinhold, 1989, genitalia-drawings, male and female paratypes from Niah Cave, Sarawak, Malaysia. 45–47. Male left palp, tibia and tarsus. 45. Ventral. 46. Retrolateral. 47. Dorsal. 48. 49. Female genitalia. 48. Epigynum, ventral. 49. Vulva, dorsal. C: conductor, D: dorsal tibial apophysis, MA: median apophysis, PC: paracymbium, RL: retrolateral lobe of tibial apex. Grey arrow to tibial apical spine.

FIGURES 15–19. Vytfutia bedel Deeleman-Reinhold, 1986, genitalia-drawings, Holotype male and paratype female collected from Gunung Leuser, border of National Park at Bohorok in primary rainforest, N. Sumatra, Indonesia. 15–17. Male left palp. 15. Tibia and tarsus, ventral. 16. Tibia, dorsal. 17. Tibia and tarsus, retrolateral. 18, 19. Female genitalia. 18. Epigynum, ventral. 19. Vulva, dorsal. C = conductor; D = Dorsal tibial apophysis; E = embolus; MA = median apophysis; RL = retrolateral tibial apophysis.

FIGURES 1–11. Vytfutia labalaba new species, Mt. Kinabalu, Borneo, CASENT9023842, female, morphology photos. 1. Cephalothorax, anterior, showing chelicerae and palps. 2. PME, dorsal, showing canoe-shaped tapeta. 3. ALE and PLE, lateral, showing canoe-shaped tapeta. 4. Cephalothorax, anterior, showing AER, clypeus and chilum. 5. Trochanter I. 6. Trochanter IV. 7. Posterior margin of carapace and pedicel, dorsal. 8. Left metatarsus IV, retrolateral, showing calamistrum. 9. Left palp, prolateral, arrows to probasal thorns. 10. Left leg I, prolateral. 11. Left leg IV, retrolateral. Scale bars, 1, 8, 9, 10, 11 = 0.5 mm, 2, 3, 4 = 0.25 mm, 5, 6, 7 = 0.1 mm.

FIGURES 12–14. Spinnerets of female Phyxelididae. 12, 13. Vytfutia labalaba new species, Mt. Kinabalu, Borneo, CASENT9023842. 14. Ambohima ranohira Griswold, Wood and Carmichael, 2012, Ranohira, Madagascar, CASENT9019986. 12, 14. Ventral. 13. Lateral. Arrow and inset in 14 to stout, curved apicodorsal seta, which occurs in Phyxelidini and Vidoleini but not in Vytfutiini. Scale bar = 0.2 mm.

FIGURES 56–61. Vytfutia spp., male leg I with clasping modification of metatarsus. 56, 58, 61. Dorsal. 57, 59, 60. Retrolateral. 56, 57. Vytfutia bedel Deeleman-Reinhold, 1986, Holotype male from Gunung Leuser, border of National Park at Bohorok in primary rainforest, N. Sumatra, Indonesia, left. 58, 59. Vytfutia labalaba new species, Holotype male from the wall of a building at 1550m elevation on Mt. Kinabalu, Sabah-Borneo, Malaysia, CASENT9023842, right. 60, 61. Vytfutia pallens Deeleman-Reinhold, 1989, male paratype from Niah Cave, Sarawak, Malaysia, right. Scale bar = 1mm.

FIGURES 31–35. Vytfutia labalaba new species, Mt. Kinabalu, Borneo, CASENT9023842, female, habitus photos. 31. Dorsal. 32. Ventral. 33. Cephalothorax, ventral. 34. Cephalothorax, dorsal. 35. Cephalothorax, lateral. Scale bars = 0.5 mm.

FIGURES 26–30. Vytfutia labalaba new species, Holotype male, Mt. Kinabalu, CASENT9023842, habitus photos. 26. Carapace, dorsal. 27. Dorsal. 28. Ventral. 29. Cephalothorax, ventral. 30. Lateral. Scale bar = 0.5 mm.

FIGURES 50–55. Vytfutia halandrefana new species, Berenty, CASENT9012420, female legs and palps, SEM. 50. Metatarsus IV, retrolateral, showing calamistrum. 51. Metatarsus IV, retrolateral, showing calamistral setae. 52. Tarsus IV, apical, showing STC and ITC. 53. Palpal femur, anterolateral, showing thorns (arrow). 54. Palpal femur, anterolateral thorn, close up. 55. Tarsus IV, retrolateral showing STC and ITC. Scale bars: 50, 300 µm; 51, 20 µm, 52, 55, 50 µm, 53, 200 µm, 54, 30 µm.

FIGURE 62. Vytfutia spp. Distributions on map of Eurasia and Australasia. Filled circle. Vytfutia bedel Deeleman-Reinhold, 1986, Holotype male from Gunung Leuser, border of National Park at Bohorok in primary rainforest, N. Sumatra, Indonesia. Triangle. Vytfutia labalaba new species, Holotype male from the wall of a building at 1550m elevation on Mt. Kinabalu, Sabah-Borneo, Malaysia. Diamond. Vytfutia halandrefana new species, Madagascar. Star. Vytfutia pallens Deeleman-Reinhold, 1989, male paratype from Niah Cave, Sarawak, Malaysia.

FIGURES 20–23. Vytfutia halandrefana new species, female, Berenty, Madagascar, CASENT9012513, habitus photos. 20. Dorsal. 21. Ventral. 22. Left metatarsus-tarsus IV and apex of abdomen with spinnerets, ventral. 23. Epigynal region, dorsal. Scale bars 20–22 = 0.5 mm, 23 = 0.25 mm.