Ambohima, GRISWOLD 1990

Griswold, Charles E., Wood, Hannah Marie & Carmichael, Anthea D., 2012, The lace web spiders (Araneae, Phyxelididae) of Madagascar: phylogeny, biogeography and taxonomy, Zoological Journal of the Linnean Society 164 (4), pp. 728-810 : 765-770

AMBOHIMA GRISWOLD 1990

Ambohima Griswold 1990: 126 ; Platnick 2011.

Type species: Ambohima sublima Griswold 1990 , by original designation.

Diagnosis: Male palpal bulb ( Figs 10A–B , 22A–D ) lacking median apophysis, with conductor greatly hypertrophied, apex directed mesad, origin of embolus retrolateral, embolus concealed in conductor groove completely encircling bulb; male palpal tibia with unique, backwards-pointing prolateral hook ( PTA) ( Figs 24D, E , 25E, F ); male metatarsi I ( Fig. 42A–O ) and II ( Fig. 1 ) having prolateral, spinule-lined concavity at midsegment, at base of concavity a short, prolateral process surmounted by stout clasping spine. Epigynum with broad, oval posterior median lobe ( ML), copulatory openings slit-like and converging anteromedially ( Fig. 30D, F ).

Synapomorphies: Synapomorphies for Ambohima include the loss of the male palpal median apophysis ( Figs 10A–B , 22A–D ) and the backwards-pointing prolateral hook ( PTA) on male palpal tibia ( Figs 24D, E , 25E, F ), both unique in Phyxelididae ; and epigynal copulatory openings that are slit-like and converging anteromedially ( Fig. 30D, F ) and spermathecae with an anterior straight or spiral blind-ending duct ( Figs 31B , 33E ).

Description: Total length 3.70–10.90. Typical markings with carapace orange-brown, margin of pars thoracica dusky, with broad, dusky, V-shaped mark anteriad of thoracic fovea and dusky longitudinal band on each side extending anteriorly along pars cephalica, with dusky reticulations laterad of eyes; black pigment surrounding each eye and extending between AM and between lateral eyes ( Fig. 6A–F ); chelicerae, sternum, labium, coxae, legs, and palpi yellow-brown to orange-brown, sternum dusky, labium and palpal coxa lighter near tips ( Fig. 47C ), legs unmarked or with annulations ( Figs 3A, B , 7B ); abdomen grey-brown, dorsum with anterior light longitudinal markings that may comprise a herringbone pattern, chevrons, or a median band that is narrowed posteriorly with lateral extensions ( Fig. 7A–H ), dorsum unmarked in A. maizina sp. nov. ( Fig. 6G ), venter with pair of bold, entire, longitudinal light bands ( Fig. 6H ); differences from these markings are noted in species descriptions. Carapace length 1.15– 1.48 times width, height 0.21–0.45 times width, thoracic fovea narrow oval, length 0.14–0.28 carapace length; PER straight to slightly recurved, AER straight, PER width 2.00–3.22 times OA length; clypeal height 1.23–5.51 times AM diameter; chelicerae smooth or with very weak wrinkles, length 4.06–8.00 times clypeal height, pro- and retromargins of fang furrow with 5–8 heterogeneous teeth ( Fig. 49A, B ); sternum length 1.10–2.53 times width, base narrowed, apex may be prolonged, pointed, a right angle, or blunt; labium broad, notched at base, length slightly greater than width ( Fig. 47C ). Legs long to elongate, femur I of females 1.18–2.72, of males 1.57–3.62 times carapace length, metatarsus I of males 1.38–3.77 times carapace length; leg formula 1423; male metatarsi I and II with prolateral, spinule-lined concavity at midsegment, at base of concavity a short, prolateral process surmounted by stout clasping spine ( Fig. 1 ), legs otherwise unmodified; metatarsi lacking apical combs; calamistrum subapical, origin at 0.40–0.50 distance from metatarsus base, length about one-third metatarsus length ( Fig. 46F ). Spination (based on Ambohima sublima : different species and even individuals within species exhibit small variations in the presence/absence and position of spines). Male: palp: femur d0-0-0-1; leg I: femur p0-1-1–1, r0-1-0-1, tibia p0-1-1-0, v1-0-1-0, r0-1-1-0, metatarsus p0-1(clasper)-0-1, v1-1-1-1, r0-1- 0-1; leg II: femur d1-0-1-0, p0-1-1-1, r0-1-0-1, tibia p0-1-1-0, v0-0-0-1, r0-1-1-0, metatarsus p0-1(clasper)- 0-1, v2-2-0-1, r0-1-0-1; leg III: femur p0-0-0-1, tibia d1-0-0-0, p0-1-1-0, v0-0-0-2, r0-1-1-0, metatarsus p0-1-0-2, v2-2-0-1, r0-1-0-2; leg IV: femur d1-0-0-0, p0-0-0-1, r0-0-0-1, tibia d1-0-0-0, p0-1-1-0, v0-0-0-2, r0-1-1-0, metatarsus v2-1-1-2. Female: palp: femur d0-0-0-1, patella d0-0-1, tibia d0-0-0-1, p0-1-1-0, tarsus p1-0-1-0, v0-0-1-2-1-2-3-2, r1-0-0-0-0; leg I: femur d1-0-0-0, p0-1-1-1, r0-0-1-1, tibia p0-1-1-0, v0-1-1-1, r0-1-1-0, metatarsus p0-1-0-1, v2-0-2-1, r0-1- 0-1; leg II: femur d0-1-1-0, p0-1-1-1, r0-1-1-1-1, tibia p0-1-1-0, v0-1-0-2, r0-1-1-0, metatarsus p0-2-0-1, v2-2-0-1, r0-1-0-1; leg III: femur d1-0-1-0, p0-1-1-1, r0-1-0-1, tibia p0-1-1-0, v0-1-0-2, r0-1-1-0, metatarsus p0-1-0-2, v2-0-2-1, r0-1-0-2; leg IV: femur d1-1-0-0, p0-0-0-1, r0-0-0-1, tibia p0-1-1-0, v0-1-0-2, r0-1-1-0, metatarsus p0-1-0-1, v2-0-2-1, r0-0-0-1. Abdomen dorsal markings as in Figures 6A–G , 7C–H , typically with posterior chevrons dorsally although markings obscure to uniform in some species, venter with longitudinal bands narrow and entire. Tracheae (observed in penultimate male Ambohima sublima, CASENT 9016997) with medians and laterals simple. Spinnerets (based on SEM examination of Ambohima ranohira and A. sublima ): Female ALS ( Fig. 52B ) with two MAP spigots, the anterior largest, with large tartipore adjacent to these; PI spigot field of 30 to more than 80 spigots with rounded bases, interspersed with numerous small tartipores; male ALS ( Fig. 53B ) with posterior MAP spigot reduced to nubbin, but with comparable number of PI spigots as female; female PMS ( Fig. 52C ) with anteriorly encircling row of 15–25 PC spigots, these crowded together so that the bases are laterally flattened, centrally a large mAP spigot with a broad, squat base and conical shaft, mesad and posteriad of this are 5–7 AC spigots, and posteriorly one CY spigot with a long, conical base and conical shaft; males ( Fig. 53C ) retain the AC spigots but lack the CY spigot, PC spigots are replaced by nubbins, and the central mAP spigot is replaced by a large nubbin; female PLS ( Fig. 52D ) with retroapical stout, curved seta (black in life), one subapical MS spigot with 1–2 flanking nubbins, a field of 12–20 AC spigots, and one basal and 1 anteromedian CY spigot; males ( Fig. 53D ) lack the CY spigots, have fewer AC spigots, and the MS spigot region is replaced by a huge nubbin. Palpal femur with anterobasal group of 3–8 setae set in enlarged bases, at least some distal setae shortened and enlarged as thorns, these setae forming a row or triangular group ( Fig. 46C ); male palpal tibia ( Figs 24D, E , 25E, F ) with a sclerotized dorsal bladeshaped process ( DTA), a dorsally prolonged prolateral hook ( PTA) and a prominent retrodorsal lobe ( RL); cymbium narrowed for apical half; palpal bulb ( Fig. 45C, D ) with small, sclerotized petiole at base of subtegulum, this with 4–5 anneli, tegulum central, small relative to hypertrophied conductor, reservoir course a simple curve within tegulum, without switchbacks, median apophysis absent; embolus slender, a narrow tapering blade to a thread-like spine, lamellar pars pendula ( PP) large ( Figs 21B, C , 22A, C ), readily detached from truncus of embolus (E), conductor (C) greatly hypertrophied, with embolic groove completely encircling bulb, apex ( CA) directed mesad ( Fig. 23B ). Epigynum ( Figs 31A , 32D , 34E ) with broad, oval to trapezoidal ML, copulatory openings slit-like or hidden beneath recurved lateral transverse ridges ( LTR); spermathecae with an anterior straight or spiral blind-ending duct, fertilization duct posterior ( Figs 31B , 33E ).

Composition: Ten species.

Distribution: Madagascar ( Figs 57–59 )

AMBOHIMA ANDREFANA SP. NOV. ( FIGS 4A–D , 6A, B , 10A–C , 21A–E , 31E–I , 39G–L, 42A–C, 46C, 57)

Types: Holotype male ( CASENT9005932 ) and paratype female ( CASENT9019919 ) collected in dry forest on sandy soil at an elevation of 770 m at Parc National Zombitse , 19.8 km 84° E Sakaraha, 22°50′36″S, 44°42′36″E, Toliara Province, Madagascar, 5–9 February 2003, by C. Griswold, D. Silva, and J. J. Rafonomezantsoa, deposited in CASC.

Etymology: The specific name is from the Malagasy word for west; a noun in apposition.

Figure 39. Ambohima female genitalia. A–F, A. ranohira . G–L, A. andrefana . A–C, G–I, epigyna, ventral. D–F, J–L,

Figures

Figure 1. Habitus of Ambohima sublima male from Ambohimanga, CASENT9016199; inset shows metatarsus II prolateral clasping spine. Illustration by JS.

Figure 3. Dorsal habitus of Ambohima males. A, A. ranohira from Ranohira, CASENT9005938. B, A. sublima from Anjozorobe, CASENT9004080.

Figure 4. Mating of Ambohima andrefana. A, male (top), female (bottom). B, male (top), female (bottom); note female biting male leg I. C, male (bottom), female (top); note female biting male leg I and male leg II claspers grasping base of female palps (arrow). D, male (bottom), female (top); note male leg II claspers grasping base of female palps (arrow). A, CASENT9005931 (male), 9019919 (paratype female). B, C, CASENT9005931 (male), 9019918 (female). D, CASENT9005932 (holotype male), 9019920 (female).

Figure 6. Habitus of Ambohima. A, A. andrefana, male, dorsal, Analavelona, CASENT9020321. B, A. andrefana, male, dorsal, Zombitse, CASENT9005931. C, A. vato, male, dorsal, Itremo, CASENT9016755. D, A. zoky, holotype male, dorsal, CASENT9017018. E, A. sublima, male, dorsal, Ambohimanga, CASENT9016200. F, A. sublima, male, dorsal, Talatakely, CASENT9024485. G, I, A. maizina, holotype male, CASENT9016206. G, dorsal. I, ventral. H, A. ranohira, female, ventral, Ranohira, CASENT9019986.

Figure 7. Habitus of Ambohima females. A, B, A. antsinanana from Andringitra, CASENT9029887. A, dorsal. B, lateral. C, A. vato from Itremo, CASENT9020002, dorsal. D, A. ranohira from Analalava, CASENT9019989, dorsal. E, A. avaratra from Montagne d’Ambre, CASENT9017012, dorsal. F, A. zandry, Montagne d’Ambre, CASENT9017021, dorsal. G, A. zoky, Montagne d’Ambre, CASENT9020320, dorsal. H, A. pauliani, holotype, dorsal.

Figure 10. Left male palpus of Ambohima andrefana from Zombitse (CASENT9005930), tibia and tarsus. A, prolateral. B, ventral. C, retrolateral. Illustrations by GM.

Figure 21. Right male palpus of Ambohima andrefana from Zombitse (SEM images flipped to appear left) CASENT9005933. A, prolateral. B, ventral. C, retrolateral. D, tibia, dorsal, showing DTA. E, bulb, apicoventral. F, tibia, prolateral, showing DTA and PTA. Scale bars: A = 200 Mm, B–F = 100 Mm. E: embolus, PP: pars pendula of embolus.

Figure 22. Right male palpus of of Ambohima ranohira from Betroka (SEM images flipped to appear left) CASENT9016178. A, prolateral. B, retrolateral. C, proapical. D, ventral. Scale bars: A, B, D = 430 Mm, C = 250 Mm. E: embolus, PP: pars pendula of embolus.

Figure 23. Right male palpus of of Ambohima ranohira from Betroka (CASENT9016178). A, bulb with embolic base, retroapical. B, apex of conductor, with pars pendula and truncus separated. C, tibia, dorsal. D, tibia, proapical. Scale bars: A–D = 200 Mm. C: conductor, CA: conductor apex, E: embolus, PP: pars pendula of embolus.

Figure 24. Right male palpus of Ambohima zandry from Montagne d’Ambre (SEM images flipped to appear left), CASENT9006874. A, prolateral. B, ventral. C, retrolateral. D, tibia, prolateral, showing RL, DTA, and PTA. E, tibia, dorsal. Scale bars: A–D = 100 Mm, E = 30 Mm. DTA: dorsal tibial apophysis, PTA: prolateral tibial apophysis, RL: retrolateral lobe of tibial apex.

Figure 25. Right male palpus of Ambohima zoky holotype from Antsiranana (SEM images flipped to appear left) CASENT9017018. A, prolateral. B, ventral. C, retrolateral. D, tibia, prolateral, showing DTA and PTA. E, tibia, prodorsal, showing RL, DTA, and PTA. F. Tibia, dorsal, apices of DTA and PTA. Scale bars: A–C = 430 Mm, D, F = 158 Mm, E = 200 Mm. DTA: dorsal tibial apophysis, PTA: prolateral tibial apophysis, RL: retrolateral lobe of tibial apex.

Figure 30. Female genitalia of Ambohima sublima. A–C, E, vulvae, dorsal. D, F, epigyna, ventral. A, Perinet, CASENT9016184. B, Ambohitantely, CASENT9016213. C, D, Ambohimanga, CASENT9016204. E, F, Ranomafana, CASENT9016193. Illustrations by JS.

Figure 31. Ambohima female genitalia. A, G, epigyna, ventral. B–F, H, I, vulvae, dorsal. A–D, A. ranohira. E–I, A. andrefana. A–D, Ranohira, CASENT9019985. E, F, Analalava, CASENT9019978. G–I, Zombitse, CASENT9019919. CD: copulatory duct, FD: fertilization duct, LL: lateral lobe of epigynum, ML: median lobe of epigynum, SP: spermathecal poreplate.

Figure 32. Ambohima sublima female genitalia. A–C, Talatakely, CASENT9024053. D–F, Ambohimanga, CASENT9020323. A, D, epigyna, ventral. B, E, vulvae, dorsal. C, F, close-up of spermathecae. LL: lateral lobe of epigynum, LTR: lateral transverse ridge along anterior margin of median lobe of epigynum, ML: median lobe of epigynum, SP: spermathecal poreplate.

Figure 33. Ambohima female genitalia. A, D, G, epigyna, ventral. B, C, E, F, H, I, vulvae, dorsal. A–C, A. vato, Itremo, CASENT9005552. D–F, A. zoky, Montagne d’Ambre, CASENT9017016. G–I, A. antsinanana, Andringitra, CASENT9029887. CD: copulatory duct, FD: fertilization duct, SP: spermathecal poreplate.

Figure 34. Ambohima female genitalia. A, E, epigyna, ventral. F, epigynum, posterior. B–D, G–I, vulvae, dorsal. A–D, A. avaratra, Montagne d’Ambre, CASENT9017012. E–I, A. zandry, Montagne d’Ambre, CASENT9017019. LL: lateral lobe of epigynum, LTR: lateral transverse ridge along anterior margin of median lobe of epigynum, ML: median lobe of epigynum, SP: spermathecal poreplate.

Figure 42. Ambohima male leg I metatarsal claspers, left. A, D, G, J, M, clasper close up, dorsal. B, E, H, K, N, metatarsus, dorsal. C, F, I, L, O, clasper close up, prolateral. A–C, A. andrefana, Analavelona, CASENT9018656. D–F, A. andrefana, Zombitse, CASENT9005930. G–I, A. maizina, holotype, CASENT9016206. J–L, A. ranohira, Ranohira, CASENT9005788. M–O, A. sublima, Ambohimanga, CASENT9016227.

Figure 45. Expanded male palps of Rahavavy fanivelona, A, B (CASENT9016967) and Ambohima ranohira, C, D (CASENT9005790). A, C, retrolateral. B, D, prolateral. AN: anneli of subtegulum, CA: conductor apex, DTA: dorsal tibial apophysis, E: embolus, P: petiole of subtegulum, PP: pars pendula of embolus, ST: subtegulum, T: tegulum.

Figure 46. Morphology of Ambohima. A, A. sublima male from Ambohimanga, CASENT9016225, right coxa-trochanter I, ventral. C, A. andrefana, male from Zombitse, CASENT9005930, palpal femur prolateral showing thorns. B, D–F, A. ranohira female from Ranohira, CASENT9019986. B, trochanter I, right. D, pedicle, dorsal. E, trochanter IV, left. F, metatarsus IV, retrolateral, showing calamistrum.

Figure 47. Morphology of Ambohima ranohira, female from Ranohira, CASENT9019986. A, face. B, clypeus and bipartite chilum. C, cephalothorax, ventral. D, spinnerets.

Figure 49. Morphology of Ambohima ranohira female from Ranohira, CASENT9019986. A, left chelicera, promargin. B, left chelicera, retromargin. C, right endite, dorsal. D, labrum, dorsal. E, serrula on right endite, close up. F, cheliceral gland pores (arrow).

Figure 52. Left spinnerets of female Ambohima sublima from Ambohimanga, CASENT9020323. A, overview. B, ALS. C, PMS. D, PLS. Scale bars: A = 150 Mm, B = 43 Mm, C, D = 30 Mm. CY: cylindrical gland spigots, MAP: major ampullate gland spigot, mAP: minor ampullate gland spigot, MS: PLS modified spigot, n: nubbin, PC: paracribellar spigot.

Figure 53. Right spinnerets of male Ambohima sublima from Ambohimanga, CASENT9020323. A, overview. B, ALS. C, PMS. D, PLS. Scale bars: A = 200 Mm, B = 43 Mm, C, D = 25 Mm. MAP: major ampullate gland spigot, MAPn: major ampullate gland spigot nubbin, mAPn: minor ampullate gland spigot nubbin, n: nubbin of PLS modified spigot, PCn: paracribellar spigot nubbins.

Figure 57. Distribution of Madagascar Phyxelididae.