Zatavua, Huber, 2003

Huber, Bernhard A., 2003, Cladistic analysis of Malagasy pholcid spiders reveals generic level endemism: Revision of Zatavua n. gen. and Paramicromerys Millot (Pholcidae, Araneae), Zoological Journal of the Linnean Society 137 (2), pp. 261-318 : 263-266

ZATAVUA , NEW GENUS

Type species. Zatavua griswoldi , new species.

Etymology. Named for Zatavu, a great magician in Malagasy mythology who created himself and was therefore allowed to marry a goddess. Gender feminine.

Note. Millot (1946) described several species belonging to this genus (under Spermophora ). Unfortunately, most of the material studied by him has not been available to me (it could not be found at the MNHN in Paris; C. Rollard, pers. comm.). Therefore, the diagnosis and description given below may not perfectly cover all the species included. Nevertheless, judging by Millot’s excellent illustrations, some of his species seem very closely related to some of the species newly described herein (see Specific Relationships below), so the diagnosis and description may in fact be tolerably accurate.

Diagnosis. Long-legged, six-eyed pholcids with globular or elongate opisthosoma and variable size (total length ~ 2–8 mm). Distinguished from Paramicromerys and other six-eyed genera by the lateral cheliceral apophyses pointing backwards ( Figs 62 , 84, 86 and 98 ), the presence of a retrolateral notch on the cymbium ( Figs 34 , 48 and 57 ), and by the shift of the tibia-tarsus joints, resulting in a dorsal position of the retrolateral joint and in a ventral position of the prolateral joint ( Figs 29, 30 , 33 and 34 , etc.; two latter characters missing in Z. kely , mahafaly ). Distinguished from Paramicromerys also by the closeness of the triads (e.g. Fig. 67 ; PME-PME <180% of PME diameter, vs.>230% in Paramicromerys ), and by the presence of several spigots on the ALS in addition to the basic set of two ( Figs 39, 40, 46 , 74, 75 and 77 ).

Description. Total length in males usually ~ 3–5 mm; only Z. zanahary and punctata (Millot) up to 8 mm, Z. kely only ~ 1.5 mm. Carapace oval or round, with or without thoracic groove, often with dark pattern that may be distinctive ( Figs 1–18 ). Six eyes in two triads, on moderately elevated ocular area; no trace of AME. Distance PME-ALE small (~20–40% of PME diameter), distance PME-PME also small (~90–180% of PME diameter). Clypeus in some species with paired unsclerotized projections at rim. Male chelicerae with pair of lateral apophyses pointing backwards ( Figs 62 , 84, 86 and 98 ), and frontal apophyses variable in position and shape; never with modified hairs, never with stridulatory ridges. Male palps moderately large in relation to overall size; coxa and trochanter without retrolateral apophyses, femur usually cylindrical, sometimes with small ventral apophysis proximally; tibia moderately to highly expanded, with two trichobothria; tibia-cymbium joints distinctively shifted (resulting in dorsal position of retrolateral joint and ventral position of prolateral joint); cymbium with distinctive notch on retrolateral side ( Figs 34 , 48 and 57 ); procursus usually complex but never with hinged process; with capsulate tarsal organ ( Fig. 73 ), often in very distal position ( Figs 30 , 33 , 47 , 52 , 56 , 61 and 78 ); bulb consisting of proximal globular part and embolus that is partly sclerotized and often provided with sclerotized spine. Legs usually long (leg 1 about 8– 11 ¥ body length; in Z. kely only 6 ¥), medium-thin (tibia 1 L/d ~40–90), leg 1 always longest, legs 2 and 4 about same length, leg 3 shortest. Legs usually without spines, with few vertical hairs, without curved hairs; retrolateral trichobothrium of tibia 1 usually at 5–15%, in Z. kely at 27%. Prolateral trichobothrium missing on tibiae 1, present in all others. Tarsus 1 with over 20 pseudosegments, but only ~10–20 distal pseudosegments easily visible in dissecting microscope. Opisthosoma either globular or elongate, often with posterior elongation over spinnerets ( Figs 4, 14 and 16 ). Male gonopore with four epiandrous spigots in all species examined ( Z. zanahary , griswoldi , voahangyae ; Figs 41 and 66 ); ALS with several spigots in addition to basic set of two (examined: Z. griswoldi , vohiparara , voahangyae , isalo , kely ; Figs 39, 40, 46 , 74, 75 and 77 ); other spinnerets typical for family (e.g. Fig. 38 ; cf. Huber, 2000).

Sexual dimorphism slight. In some species females with median conical elevation posteriorly on carapace; corresponding side of opisthosoma not visibly modified. Epigynum shape very variable, often with pair of pockets, in Z. vohiparara with short scape ( Fig. 45 ); internally with pair of relatively large pore plates and complex system of sclerites and membranes of unknown function.

Monophyly. The monophyly of this genus is supported by the following three synapomorphies: the backward facing lateral cheliceral apophyses (shared by all species included), the shift of the tibia-cymbium joints, and the notch on the cymbium (the latter two characters are not shared by Z. kely and Z. mahafaly ).

Generic relationships. As discussed above, Zatavua is here considered the sister taxon of all other pholcines. The presence of lateral apophyses on the male chelicerae places the genus within the subfamily, but the genus lacks the retrolateral trochanter apophysis characteristic of other pholcines.

Specific relationships. Several synapomorphies support species groups within Zatavua . A core group of species is characterized by the presence of a conical elevation on the female carapace [ Z. griswoldi , tamatave , vohiparara , maybe including andrei (Millot) and ankaranae (Millot)]. This group shares with Z. analalava the projections on the male clypeus. The resulting group shares with several further species [ Z. zanahary , punctata (Millot) , voahangyae , talatakely , probably impudica (Millot)] the distal position of the male palpal tarsal organ. Within this latter group, Z. zanahary and punctata share their unusual size and the unique pattern on the carapace ( Fig. 9 ). All species listed so far share with Z. isalo and madagascariensis (Fage) the shift of the tibia-cymbium joints and the notch on the cymbium. The latter two species have almost identical palps (not coded) and are therefore seen as closely related. Zatavua fagei (Millot) is only known from the female, but was collected near the type locality of Z. madagascariensis (Fage) and is either a junior synonym or a closely related species. Finally, Z. mahafaly and kely are very unusual in several aspects, but share with all other species the backward facing lateral cheliceral apophyses on the male chelicerae. Of these, Z. kely might be close to imerinensis (Millot) , judging by the similar shapes of the palps.

Natural history. No species has ever been studied in any detail, and little can be inferred from notes on the collection labels. Several species have been collected in caves [ Z. andrei , ankaranae, impudica, madagascariensis (Fage) , mahafaly ], two more were classified as ‘semi-cavernicole’ by Millot (1946) ( Z. fagei , punctata ), and one as ‘semi-domestique’ ( Z. imerinensis ). All new species described below (except Z. mahafaly ) were apparently collected in forests. Zatavua vohiparara was collected on Pandanus , an undescribed species from the AMNH was collected by beating vegetation.

Distribution. Known from Madagascar only. The genus is widely distributed in Madagascar (Map 1), and probably covered most of the island before human destruction of a large part of the primary vegetation. As in Paramicromerys , no species has been collected at two localities more than a few kilometres apart. A possible exception is Z. punctata (Millot) , which has been recorded from the Antonibe Peninsula and on Nosy- Komba Island (about 200 km apart), but the latter record is based on a female and a juvenile specimen and should be checked.

Composition. The genus includes a total of 17 described species. Of these, six were described by Millot (1946), and are not treated below ( Z. andrei , ankaranae, fagei , imerinensis , impudica, punctata ). The remaining 11 species [10 new species and Z. madagascariensis (Fage) ], are (re)described below. The collections studied contain four additional new species that are not described due to their poor state of preservation. Considering the facts that (1) most or all species seem to have small distributional ranges, and that (2) all areas that were subjected to intense collecting yielded between one and three species, it seems reasonable to expect several dozen additional species not yet discovered.

ZATAVUA GRISWOLDI , NEW SPECIES

( FIGS 1, 2 , 19 , 29–32 , 38–44 )

Type. Male holotype from Marojejy Res. , 8.4 km NNW Manantenina (14∞26¢S, 49∞45¢E), 700 m a.s.l., Antsiranana, Madagascar; November 10–16, 1993 (C. E. Griswold, J. Coddington, N. Scharff, S. Larcher, R. Andriamasimanana); in CAS .

Etymology. Named for the first collector, Charles Griswold from the California Academy of Sciences, who kindly made his superb collection of Malagasy pholcids available to me.

Diagnosis. Medium size species with elongated opisthosoma, distinguished from known congeners by the shape of procursus and embolus ( Figs 29, 30 : overall shape and several unnamed details), and by the shape of the epigynum ( Figs 19 and 31 ). The AMNH has a male from a nearby locality (10.5 km NW Manantenina) that differs slightly by having a shorter procursus and more profoundly by the shape of the embolus. The MRAC has a further close relative from

Figures

Figures 60–65. Zatavua voahangyae. Left male palp in prolateral (60) and retrolateral (61) views, male chelicerae in lateral (62) frontal (63) views, and cleared epigynum in ventral (64) and dorsal (65) views. ‘e’: embolus; ‘p’: pocket; ‘s’: spine on embolus; ‘to’: tarsal organ. Scale lines: 0.5 mm (60–61), 0.2 mm (62–63), 0.3 mm (64–65).

Figures 81–86. Zatavua isalo (81–84) and Z. madagascariensis (85, 86). Left male palp in prolateral (81) and retrolateral (82) views, and male chelicerae in frontal (83, 85) and lateral (84, 86) views. ‘e’: embolus; ‘f’: distinctive flap. Scale lines: 0.5 mm (81, 82), 0.3 mm (83–86).

Figures 96–101. Zatavua kely. Left male palp in prolateral (96) and retrolateral (97) views, male chelicerae in lateral (98) and frontal (99) views, and cleared epigynum in ventral (100) and dorsal (101) views. ‘a’: apophysis on embolus; ‘e’: embolus; ‘p’: pocket; ‘pr’: procursus; ‘r’: putative receptacle; ‘to’: tarsal organ. Scale lines: 0.2 mm.

Figures 33–37. Zatavua vohiparara. Left male palp in prolateral (33) and retrolateral (34) views, male chelicerae in frontal view (35), and cleared epigynum in ventral (36) and dorsal (37) views. ‘a’, ‘b’: distal structures on procursus that differ in male from Mahira (see text); ‘e’: embolus; ‘l’: lamellae on procursus; ‘n’: notch on cymbium; ‘s’: scape; ‘sp’: spine on embolus; ‘to’: tarsal organ. Scale lines: 0.5 mm (33–34, 36–37), 0.3 mm (35).

Figures 47–50. Zatavua tamatave. Left male palp in prolateral (47) and retrolateral (48) views, and cleared epigynum in ventral (49) and dorsal (50) views. ‘b’: branch of embolus; ‘e’: embolus; ‘n’: cymbium notch; ‘p’: pocket; ‘to’: tarsal organ; arrow points to distinctive spine on procursus. Scale lines: 0.5 mm.

Figures 56–59. Zatavua zanahary. Left male palp in prolateral (56) and retrolateral (57) views, and cleared epigynum in ventral (58) and dorsal (59) views. ‘e’: embolus; ‘n’: cymbium notch; ‘p’: pocket; ‘pa’: papilla on bulb; ‘to’: tarsal organ. Scale lines: 1 mm.

Figures 29–32. Zatavua griswoldi. Left male palp in prolateral (29) and retrolateral (30) views, and cleared epigynum in ventral (31) and dorsal (32) views. ‘e’: embolus; ‘l’: lamellae on procursus; ‘p’: pocket; ‘s’: spine on embolus; ‘to’: tarsal organ. Scale lines: 0.5 mm.

Figures 66–77. Zatavua voahangyae (66–75) and Z. analalava (76–77). 66, Male gonopore with epiandrous spigots. 67, Female prosoma, frontal view. 68, Epigynum with pockets. 69, Male pedipalpal coxa serrula. 70, Tip of female palp with tarsal organ. 71, Procursus, retrolatero-dorsal view. 72, Embolus with sclerotized spine (‘s’) and sperm duct opening (‘o’). 73, Male palpal tarsal organ. 74, Female ALS. 75, Male ALS and PMS. 76, Epigynum with pockets (‘p’). 77, Female ALS. Scale lines: 40 Mm (66), 400 Mm (67), 100 Mm (68, 71), 30 Mm (69–70, 75), 20 Mm (73–74, 77), 200 Mm (76).

Figures 38–46. Zatavua griswoldi (38–44) and Z. vohiparara (45–46). 38, Female spinnerets. 39, Male ALS and PMS. 40, Female ALS. 41, Male gonopore with epiandrous spigots. 42, Pockets (‘p’) on epigynum. 43, Claws of third male leg. 44, Procursus, showing position of tarsal organ (‘to’). 45, Epigynum with scape (‘s’). 46, Female ALS. Scale lines: 100 Mm (38), 30 Mm (39–40), 50 Mm (41), 80 Mm (42), 40 Mm (43), 300 Mm (44–45), 20 Mm (46).

Figures 1–18. Zatavua habitus, males in dorsal and lateral views. 1–2, Z. griswoldi. 3–4, Z. analalava. 5–6, Z. mahafaly. 7–8, Z. vohiparara. 9–10, Z. zanahary. 11–12, Z. isalo. 13–14, Z. tamatave. 15–16, Z. voahangyae. 17–18, Z. talatakely.

Figures 51–55. Zatavua analalava. Left male palp in prolateral (51) and retrolateral (52) views, male chelicerae in frontal view (53), and cleared epigynum in ventral (54) and dorsal (55) views. ‘e’: embolus; ‘p’: pocket; ‘to’: tarsal organ. Scale lines: 0.5 mm (51–52), 0.2 mm (53), 0.3 mm (54–55).

Figures 78–80. Zatavua talatakely. Right male palp in retrolateral (78) and prolateral (79) views, and male chelicerae in frontal view (80). ‘e’: embolus; ‘to’: tarsal organ; arrow points to distinctive spine. Scale line: 1 mm (78, 79).

Figures 19–28. Zatavua, female opisthosomata, ventral views with epigynum. 19, Z. griswoldi. 20, Z. vohiparara. 21, Z. tamatave. 22, Z. analalava. 23, Z. zanahary. 24, Z. voahangyae. 25, Z. madagascariensis. 26, Z. isalo. 27, Z. mahafaly. 28, Z. kely.