Paramicromerys, MILLOT, 1946

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 : 283-286

PARAMICROMERYS MILLOT, 1946

Paramicromerys Millot, 1946: 145 ; type species by original designation: Micromerys madagascariensis Simon, 1893 .

Diagnosis. Long-legged, six-eyed pholcids of medium size (total length ~ 1.5–4 mm), with oval or slightly elongate opisthosoma. Distinguished from Zatavua by lateral cheliceral apophyses not pointing backwards ( Figs 136 , 179 , 193 and 201 ), absence of notch on cymbium, wide distance between eye triads (PME- PME> 230% of PME diameter, vs. <180% in Zatavua ) and by reduction of spigots on ALS to basic set of two ( Figs 148, 149 , 160 and 188 ). Distinguished from other related genera by shift of tibia-tarsus joints, resulting in dorsal position of prolateral joint and ventral position of retrolateral joint ( Figs 135 , 155 , 169 , 178 , 192 , 199 and 205 ).

Description. Total length in males about 1.5–3.5 mm. Carapace oval or roundish, thoracic groove present but often very shallow and restricted to frontal half, often with dark pattern that may be distinctive for species. Six eyes in two triads, on moderately elevated ocular area. Distance PME-ALE small (~30–50% of PME diameter), distance PME-PME large (~230– 500% of PME diameter); no trace of AME. Clypeus never modified. Male chelicerae with pair of simple lateral apophyses and frontal apophyses in variable position and shape; the latter in some species with modified hairs imbedded in tips ( P. nampoinai , manantenina ; Figs 189, 190 and 195 ). Male palps small to very large in relation to overall size (compare Figs 109 and 113 ); coxa without apophysis, in some species with ventral hump; trochanter complex, with several sclerotized apophyses; femur very short, in some species with dorso-distal apophysis (arrows in Figs 178 , 192 and 199 ); patella triangular in lateral view; tibia cylindrical, not highly expanded, with two trichobothria; tibia-cymbium joints distinctively shifted, resulting in dorsal position of prolateral joint and ventral position of retrolateral joint ( Figs 135 , 155 , 169 , 178 , 192 , 199 and 205 ); procursus usually complex, with cup-shaped tarsal organ (e.g. Figs 145 , 150 and 165 ), in proximal position (e.g. Figs 135 and 174 ), with hinged process prolatero-ventrally (‘hp’ in Figs 154 , 168 and 209 ); bulb consisting of proximal globular part and membranous embolus that is sometimes provided with distal transparent spine. Legs usually long (leg 1 about 10–13 ¥ body length), medium-thin (tibia 1 L/d ~50–90), leg 1 always longest, legs 2 and 4 about same length, leg 3 shortest. Legs sometimes with macrosetae ventrally on femora (only femora 1, rarely also 2), with few vertical hairs, without curved hairs; retrolateral trichobothrium of tibia 1 at 7–12%. 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 oval or slightly elongate, rarely with slight posterior elongation over spinnerets ( Figs 105 and 107 ). Male gonopore with four epiandrous spigots in all species examined ( P. betsileo , coddingtoni , manantenina ; Figs 146 , 162 and 186 ); ALS with only basic set of two spigots (examined: P. betsileo , coddingtoni , ralamboi , manantenina , nampoinai , scharffi , rothorum , marojejy , rabeariveloi ; Figs 148, 149 , 160 and 188 ); other spinnerets typical for family ( Fig. 147 ; cf. Huber, 2000).

Sexual dimorphism slight. Colour pattern often more variable in females than in males. Female legs never with spines. Epigynum shape very variable ( Figs 122–130 ), often with pair of pockets.

Monophyly. The synapomorphy supporting this genus is the shift of the tibia-cymbium joints (see Description above).

Generic relationships. As discussed above, the sister group of Paramicromerys is not resolved in any of the 12 most parsimonious cladograms obtained. Instead, Paramicromerys is part of a polytomy that always includes East African ‘ Spermophora ’, along with either Spermophorides or Metagonia and Micromerys (Appendix 5). All these genera share a hinged process on the procursus that is here interpreted as homologous, and the reduction of the spigots on the ALS to the basic set of two.

Specific relationships. A close relationship between P. nampoinai and manantenina is supported by several synapomorphies: modified hairs on the tips of the cheliceral apophyses, short side branch on embolus, sclerotized cone-shaped structure proximally on procursus ( Figs 178 and 192 ) that carries the tarsal organ ( Fig. 187 ). Both species share with P. scharffi a proximo-dorsal apophysis on the male palpal femur (arrows in Figs 178 , 192 and 199 ). A second speciesgroup is characterized by a transparent spine distally on the embolus [ P. coddingtoni and the putatively close relatives combesi (Millot) , ralamboi , quinteri , mahira ], but this might be a plesiomorphy. Both species groups share the spines ventrally on the frontal male femora [unknown in P. quinteri , mahira , and combesi (Millot) ]. The type species P. madagascariensis (Simon) and P. betsileo have almost identical palps (compare Figs 131 and 135 ) and are therefore considered closely related. The other species included ( P. rothorum , marojejy , rabeariveloi ) all share the shift of the tibia-cymbium joints, but their exact position is unclear. Finally, P. megaceros (Millot) is tentatively assigned to Paramicromerys because it shares the wide distance between the eye triads and the simple lateral cheliceral apophyses, but it may be misplaced.

Natural history. No species has ever been studied in any detail, and little can be inferred from notes on the collection labels. In contrast to Zatavua , only one known species is cavernicole [the tentatively assigned P. megaceros (Millot) ]. Paramicromerys betsileo has been collected abundantly on low foliage and saplings, by fogging of dead leaves on fallen trees, on roadside vegetation, and by beating low vegetation (often together with ‘Spermophora’ ranomafana n. sp.).

Distribution. Known from Madagascar only. Most records are from just three localities (Map 2), but the high diversity at each of these localities suggests that the genus has a much wider distribution. No species has been collected at two localities more than a few kilometres apart. This is especially striking in the north, where two locations less than 200 km apart (Montagne d’Ambre and Marojejy Reserve ) yielded four and five species, respectively (one species from Marojejy is not described herein), with no overlap.

Composition. The genus includes a total of 14 described species. Of these, two were described by Millot (1946), and are not treated below ( P. combesi , megaceros ). The remaining 12 species [11 new species and P. madagascariensis (Simon) ], are described

Map 2. Known distribution of Paramicromerys and of Malagasy ‘ Spermophora ’ species.

below. The collections studied contain three 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 (see above) and that (2) all areas that were subjected to intense collecting yielded between four and five species, it seems reasonable to expect several dozen additional species not yet discovered.

Figures

Figures 131–138. Paramicromerys madagascariensis (131–132) and P. betsileo (133–138). 131, Left male palp, retrolateral view; arrows point to distinctive apophyses on trochanter. 132–133, Male chelicerae, frontal views. 134–135, Left male palp, pro- and retrolateral views. 136, Male chelicerae, lateral view. 137-138, Cleared epigynum, ventral (137) and dorsal (138) views. ‘d’: distinctive element of procursus; ‘e’: embolus; ‘p’: pocket; ‘to’: tarsal organ. Scale lines: 0.4 mm (131, 134–135), 0.2 mm (132, 133, 136), 0.3 mm (137, 138).

Figures 177–182. Paramicromerys manantenina. Left male palp in prolateral (177) and retrolateral (178) views, male chelicerae in lateral (179) and frontal (180) views, and cleared epigynum in ventral (181) and dorsal (182) views. ‘hp’: hinged process; ‘pa’: papilla on bulb; arrow points to proximo-dorsal femur apophysis. Scale lines: 0.5 mm (177, 178), 0.2 mm (179, 180), 0.4 mm (181, 182).

Figures 191–197. Paramicromerys nampoinai. Left male palp in prolateral (191) and retrolateral (192) views, male chelicerae in lateral (193) and frontal (194) views, tip of male cheliceral apophysis with two modified hairs (195), and cleared epigynum in ventral (196) and dorsal (197) views. ‘a’: apophysis; ‘e’: embolus; ‘hp’: hinged process; ‘p’: pocket; arrow points to proximo-dorsal apophysis on femur. Scale lines: 0.5 mm (191, 192), 0.3 mm (193, 194), 50 Mm (195), 0.3 mm (196, 197).

Figures 198–203. Paramicromerys scharffi. Left male palp in prolateral (198) and retrolateral (199) views, male chelicerae in frontal (200) and lateral (201) views, and cleared epigynum in ventral (202) and dorsal (203) views. ‘e’: embolus; ‘hp’: hinged process; ‘p’: pocket; arrow points to proximo-dorsal apophysis on femur. Scale lines: 0.5 mm (198, 199, 202, 203), 0.3 mm (200, 201).

Figures 147–153. Paramicromerys betsileo. 147, Female spinnerets. 148, Male ALS and PMS. 149, Female ALS. 150, Tip of female palp with tarsal organ. 151, Male pedipalpal coxa serrula. 152, Reduced claw at tip of female palp. 153, Epigynum, showing pockets (arrows) and protrusions. Scale lines: 70 Mm (147), 20 Mm (148, 150, 151), 10 Mm (149), 6 Mm (152), 100 Mm (153).

Figures 159–167. Paramicromerys coddingtoni. 159, Male cheliceral apophyses. 160, Female ALS and PMS. 161, Epigynum with pockets (arrows) and posterior plate (‘p’). 162, Male gonopore with (damaged) epiandrous spigots. 163, Male palpal trochanter, ventral view, showing two prolateral (‘p1’, ‘p2’) and two retrolateral (‘r1’, ‘r2’) apophyses. 164, Procursus and embolus, retrolatero-dorsal view (arrow points to distal hinged structure of procursus). 165, Male palpal tarsal organ. 166, Male tarsus 1, showing pseudosegmentation. 167, Tip of female third leg, showing tarsal claws. Scale lines: 30 Mm (159, 166), 20 Mm (160, 165, 167), 200 Mm (161, 164), 40 Mm (162), 100 Mm (163).

Figures 183–190. Paramicromerys manantenina. 183, Male palpal trochanter (‘t’) and tips of procursus (‘p’) and embolus (‘e’). 184, Procursus with hinged process (‘hp’). 185, Structure on tip of procursus. 186, Male gonopore with epiandrous spigots. 187, Conical elevation on male palpal cymbium with tarsal organ. 188, Male ALS and PMS. 189–190, Modified hairs imbedded in tips of male cheliceral apophyses. Scale lines: 200 Mm (183, 184), 50 Mm (185), 20 Mm (186), 40 Mm (187), 30 Mm (188), 5 Mm (189, 190).

Figures 154–158. Paramicromerys coddingtoni. Left male palp in prolateral (154) and retrolateral (155) views, male chelicerae in frontal view (156), and cleared epigynum in ventral (157) and dorsal (158) views. ‘e’: embolus; ‘hp’: hinged process. Scale lines: 0.3 mm (154, 155, 157, 158), 0.2 mm (156).

Figures 168–172. Paramicromerys ralamboi. Left male palp in prolateral (168) and retrolateral (169) views, male chelicerae in frontal view (170), and cleared epigynum in ventral (171) and dorsal (172) views. ‘e’: embolus; ‘hp’: hinged process; ‘p’: pocket. Scale lines: 0.5 mm (168, 169), 0.2 mm (170–172).

Figures 204–208. Paramicromerys rothorum. Left male palp in prolateral (204) and retrolateral (205) views, male chelicerae in frontal (206) view, and cleared epigynum in ventral (207) and dorsal (208) views. ‘e’: embolus; ‘p’: pocket; ‘t’: trochanter apophysis. Scale lines: 0.5 mm (204, 205), 0.2 mm (206), 0.3 mm (207, 208).

Figures 102–121. Paramicromerys habitus, males in dorsal and lateral views. 102–103, P. betsileo. 104–105, P. mahira. 106–107, P. rothorum. 108–109, P. rabeariveloi. 110–111, P. coddingtoni. 112–113, P. nampoinai. 114–115, P. scharffi. 116– 117, P. ralamboi. 118–119, P. manantenina. 120–121, P. marojejy.

Figures 139–146. Paramicromerys betsileo. 139, Female prosoma, frontal view. 140, Tip of male cheliceral apophysis. 141, Male chelicerae. 142, Male palpal trochanter, ventral view, showing the prolateral (‘p’), ventral (‘v’), and retrolateral (‘r’) apophyses. 143, Male prosoma with palp, showing the shift of the tibia-cymbium joints (arrows). 144, Male palpal trichobothrium base. 145, Male palpal tarsal organ. 146, Male gonopore with epiandrous spigots. Scale lines: 300 Mm (139, 143), 10 Mm (140), 200 Mm (141), 80 Mm (142), 6 Mm (144), 20 Mm (145), 50 Mm (146).

Figures 173–176. Paramicromerys quinteri (173–174) and P. mahira (175–176). Left male pedipalps in prolateral (173, 175) and retrolateral (174, 176) views. ‘hp’: hinged process; ‘to’: tarsal organ; arrow points to distinctive bend in embolus. Scale lines: 0.5 mm.

Figures 209–212. Paramicromerys marojejy. Left male palp in prolateral (209) and retrolateral (210) views, and cleared epigynum in ventral (211) and dorsal (212) views. ‘e’: embolus; ‘hp’: hinged process; ‘p’: pocket; arrow points to distinctive femur apophysis. Scale lines: 0.4 mm (209, 210), 0.2 mm (211, 212).

Figures 122–130. Paramicromerys, female opisthosomata, ventral views with epigynum.122, P. betsileo. 123, P. coddingtoni. 124, P. nampoinai. 125, P. manantenina. 126, P. ralamboi. 127, P. scharffi. 128, P. rothorum. 129, P. marojejy. 130, P. rabeariveloi.