Review of the Spirobolida on Madagascar, with descriptions of twelve new genera, including three genera of ' fire millipedes' (Diplopoda) Author Wesener, Thomas urn:lsid:zoobank.org:author: Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, U. S. A. & Zoologisches Forschungsmuseum Alexander Koenig, Museumsmeile Bonn, Adenauerallee 160, D- 53113 Bonn, Germany. & B 54 D 3623309 Author Enghoff, Henrik urn:lsid:zoobank.org:author: Natural History Museum of Denmark, Universitetsparken 15, DK- 2100 Copenhagen, Denmark Author Sierwald, Petra urn:lsid:zoobank.org:author: Field Museum of Natural History, 1400 S. Lake Shore Drive, Chicago, IL 60605, U. S. A. text ZooKeys 2009 2009-09-04 19 19 1 128 journal article 10.3897/zookeys.19.221 e6ea8eea-7156-4785-8313-f4bd88dadad1 1313–2970 576503 C473F9F6-1AE7-4B3F-B17F-CA1C2709010C Granitobolus Wesener , gen. n. urn:lsid:zoobank.org:act: 81CDE7E7-992A-42FD-9FCB-16D88FA8E7B9 Type species : Granitobolus endemicus sp. n. Other species included : G. andohahelensis sp. n. Diagnosis : telopodite tip of posterior gonopods in Granitobolus uniquely shaped, resembling a duck’s head with an acutely projecting beak ( Fig. 37M ). Adults with 37 or 38 body rings, lowest number of any Malagasy genus of Spirobolida . Natural colour usually contrasting, with red appendages, dark mesozonites and greyish or light brown metazonites ( Figs 37 A–D, 38A). Shares numerous characters only with species of the genus Pseudocentrobolus : a stout preanal process which extends beyond anal valves ( Fig. 37F ), well-developed lips on the anal valves, thick moniliform body rings which widely overlap each other reaching the metazonite ( Fig. 37C ), rectangular extensions ( Fig. 37F ) on the posterior margins of the rings and a greatly enlarged collum, which is ventrally longer than body ring 2. Gnathochilarium with a subdivided mentum and a single sclerotized ledge on each stipites ( Fig. 37H ), like in Flagellobolus , Riotintobolus , Pseudocentrobolus , Caprobolus gen. n. , Alluviobolus gen. n. , Ostinobolus gen. n . Both genera possess, however, completely different posterior gonopods. Distribution and ecology : species of Granitobolus have been recorded from numerous rainforest and littoral rainforest sites in Southeast Madagascar . Unfortunately, mature males could be collected only from three sites (Andohahela-Vasiha, Manantantely and the Ambatotsirongorongo Mountain, Fig. 30 ). At least single immature specimens found in the littoral forest of Mandena and the Sainte Luce fragments S8 and S9 might represent another undescribed species because of their uniformly dark, almost black coloration. The specimens from Isaka-Ivondro (Andohahela), Ebosika (Andohahela) and Ivorona (Anosyenne) closely resemble G. andohahelensis sp. n. Except for two or three isolated specimens, which were collected on trails, all other specimens of Granitobolus were taken from large, massive, partly overgrown granitic outcrops. Specimens were collected in large numbers from thin layers of roots covering the stones. On a single rock, specimens of all sizes and both sexes were found; however, mature males remained rare. This might be related to the beginning of the dry season (June), when the collections were conducted. Both holotypes were infested with rather long nematodes (P, Fig. 38B ). The legs and body rings of G. endemicus sp. n. holotype are also covered with fungi. Figure 37. Granitobolus endemicus sp. n. , A–C, G–J male paratype, K female paratype, D–F, L–O male holotype. A living specimen, uncurling B living specimen, curled C dead male, curled D head, lateral view E legs 1–7, ventral view F telson G head, frontal view H gnathochilarium, ventral view; I midbody ring with legs Į, K left midbody legs L anterior gonopod, anterior view M left posterior gonopod, anterior view N anterior gonopod, posterior view O left posterior gonopod, posterior view. av = anal valves; Co = collum; Cx = coxite; fs-p = finger-shaped process; Gn = gnathochilarium; L = ledge; Md = mandible; Me = mentum; ms = marginal setae; Pre = preanal ring; re = rectangular extension; sf = setiferous foveolae; St = sternite; ST = stipites; T = telopodite. Scale bars = 1 mm. Description . Males : length up to 35 mm (broken), diameter up to 3.3 mm . 37 or 38 podous rings. Females: length up to 48 mm , diameter up to 4.85 mm . 37 or 38 podous rings. Length-width ratio in both sexes 1:10. Apodous rings absent in all adult specimens. Colour : mesozona dark olive green or blackish ( Fig. 37A ), metazona much light- er, brown to yellow ( Fig. 37C ). Head, antennae and legs red (faded to light brown in some specimens). Head : each eye large, with circa 35–40 ocelli arranged in 6 or 7 vertical rows ( Figs 37D, G ). Labrum with standard three irregular teeth and one row of 10–12 stout marginal setae ( Fig. 37E ). Clypeus with two setiferous foveolae on each side ( Fig. 38B ). Length of antennae sexually dimorph, protruding back to ring 2 (females) or 4 (males, Figs 37A, D , 38A ). Relative lengths of antennomeres: 1<2=3=4=5<6 ( Fig. 37G ). Terminal antennomere with four large sensory cones located together inside a membranous area. Antennomere 5 latero-apically with three or four rows, antennomere 6 with two rows of sensilla basiconica. Gnathochilarium unusual ( Fig. 37H ). Lamellae linguales each with two standard setae located behind one another. Stipites each with three apical setae. Mentum basally subdivided by a well-visible suture ( Fig. 37H ). Stipites each towards mentum with a wide sclerotized ledge ( Fig. 37H ). Hypopharyngeal crest with a field of spine-like structures. Central pads of endochilarium separated by a step into two levels. Medial palps with a large group of sensilla each, located towards hypopharyngeal crest. Mandible : external tooth simple, rounded; mesal tooth with three cusps. Four pectinate lamellae. Molar plate with numerous (10 or 11) small and shallow transverse furrows. Collum: smooth, laterally longer than body ring 2. Lobe enlarged, covering antenna and part of head ( Figs 37D , 38A ). Body rings: moniliform, overlapping up to anterior half of mesozona. Metazona of swollen appearance, width larger than pro- or mesozona ( Figs 37 A–C). Posterior margin of rings slightly excavated into rectangular extensions ( Fig. 37F ). Ozopores starting at ring 6, touching suture between mesozona and metazona ( Fig. 37D ). Telson : anal valves with well-developed lips and micropunctation, but with neither grooves, nor setae. Preanal process well-rounded or sharp, extending beyond anal valves ( Figs 37F , 38C ). Subanal scale inconspicuous. Legs : coxae 1 and 2 elongated, fused with sternum, podomeres from prefemur to tarsus in both sexes each with 4–10 ventral/mesal setae. Length of midbody legs circa 1.0 times body diameter in males ( Fig. 37J ), 0.6 times body diameter in females ( Fig. 37K ). Each podomere with an apical, ventral seta ( Fig. 37K ). Coxae 3 and beyond of rectangular shape ( Fig. 37I ). Tarsus with a stout dorso-apical seta and a pair of stout ventro-apical ones in males, three pairs of setae in females. In females apical setae pair stout, others successively much smaller. Male sexual characters : tarsi starting at leg 3 up to midbody legs wide and with a large tarsal pad not protruding past base of claw ( Fig. 37J ). Tibia shorter than in females ( Figs 37J, K ). Coxae 3–7 unmodified ( Figs 37E , 38B ). Anterior gonopods : median sternal projection triangular, with a wide, rectangular tip ( Figs 37L , 38D ). Sternite never protruding as high as coxite. Coxite wide, smoothly rounded, laterally curving around telopodite, but only extending slightly at posterior side ( Figs 37N , 38F ). Coxite anteriorly almost completely covering telopodite. Coxite processes absent, but mesal margin towards sternite sometimes projecting ( Fig. 37L ). Telopodite with a short, well-rounded, laterally curved process, which only slightly projects above telopodite margin and never protrudes above coxite margin ( Figs 37N , 38F ). Figure 38. Granitobolus andohahelensis sp. n. , male holotype. A head, lateral view B legs 1–7, ventral view C telson D anterior gonopod, anterior view; E left posterior gonopod, anterior view F anterior gonopod, posterior view G left posterior gonopod, posterior view. av = anal valves; Co = collum; Cx = coxite; fs-p = finger-shaped process; Gn = gnathochilarium; Md = mandible; ms = marginal setae; P = parasitic nematode?; Pre = preanal ring; re = rectangular extension; sf = setiferous foveolae; St = sternite; T = telopodite. Scale bars = 1 mm. Posterior gonopods in situ almost completely covered by anterior gonopods. Suture between coxite and telopodite well-visible ( Fig. 37O ), sternite triangular and sclerotized but often hidden by a large, finger-shaped coxite process ( Fig. 37O ). Finger-shaped process not curved, running parallel to basal margin ( Figs 37O , 38G ). Telopodites arranged face-to-face with one another ( Fig. 37M ). Shape of apical part resembling ‘beak’ ( Figs 37M , 38E ). Beak acutely projecting, partly sclerotized and carrying sperm canal ( Fig. 38G ). Telopodite between apex and beak with a small membranous area ( Fig. 37M ), in basal half mesally of sperm canal with a lower elevated area ( Fig. 37M ). Female sexual characters: vulva simple, with a small, poorly sclerotized operculum at base, bivalve-like. Posterior valve apically slightly overlapping anterior one. Both valves smooth, sensory cones absent. Each valve basally towards opening with two rows of setae. Etymology : Granitobolus , masculine, after Granit (German for granite) for the special habitat, granitic outcrops, on which species of this genus were found, and -bolus .