Glyphiulus granulatus, Gervais, 1847

Golovatch, Sergei I., Geoffroy, Jean-Jacques, Mauriès, Jean-Paul & Spiegel, Didier Van Den, 2007, Review of the millipede genus Glyphiulus Gervais, 1847, with descriptions of new species from Southeast Asia (Diplopoda, Spirostreptida, Cambalopsidae). Part 1: the granulatus-group, Zoosystema 29 (1), pp. 7-49 : 11-20

publication ID	https://doi.org/ 10.5281/zenodo.5392079
persistent identifier	https://treatment.plazi.org/id/03F4C378-A22B-3046-FCA9-0270FEDAFB79
treatment provided by	Marcus
scientific name	Glyphiulus granulatus
status

Treatment

The granulatus View in CoL -group

REMARKS Species of the granulatus- group are distinguished by the following characters:

1. Male legs 1 are usually very strongly reduced, represented solely or mainly by a sternum lacking any median structures but bearing strongly separated, evidently curved prongs with a leg vestige on each side at base (usually represented by just a few setae, more seldom also one or two rudimentary segments). Very seldom, a nearly complete telopodite persists, but the sternum still supports a pair of widely separated and curved prongs. [Male legs 1 with central structures on the sternum in other groups].

2. Other male legs normal or nearly so, not enlarged. [Male legs 2 sometimes incrassate in other groups].

3. A typical carinotaxy pattern of the collum is I- VI+7a+pc+ma+pc+7a+VI-I. Quite often the pattern is different, either due to crest reduction (especially anteriorly) or hypertrophy, but a median crest is always traceable, at least near the caudal margin. [Usually typical carinotaxy formulae of the collum are different in other groups].

4. A typical carinotaxy pattern of the metaterga is 3(2)+I+3+I+3(2). The crests usually being divided transversely into two halves, while the median crest is often doubled anteriorly, the formula becomes 3(2)+I+4+I+3(2) and 3(2)+i+3+i+3(2). If the pattern is different, it is usually readily derived from the typical one (or vice versa), i.e. the lateral crests are reduced to two or the crests are undivided, or some of the crests are divided into three, rather than two, transverse rows of tubercles (see also below), etc. A median crest, even when strongly reduced, is always present as well. [Usually typical carinotaxy formulae of the metaterga are different in other groups].

5. The anterior gonopods are reduced to a plate-like coxosternum with moveable, lateral, 1-segmented telopodites. [Shared with some other species groups].

6. The posterior gonopods are highly compressed, showing a subflagelliform (rarely subspatuliform), often plumose, distal process. [Shared with some other species groups].

7. The pleural flaps behind the gonopod opening on male segment 7 usually do not form an apparent transverse ventral ridge. [Shared with some other species groups].

The group thus roughly corresponds to the concept of Glyphiulus in the sense of Verhoeff (1936) and

Loksa (1960). At present it contains the following species: G. adeloglyphus , G. anophthalmus ,

G. balazsi , G. capucinus , G. granulatus , G. lipsorum ,

G. melanoporus , G. quadrohamatus , G. rayrouchi ,

G. septentrionalis , G. superbus , as well as all nine new species described below.

The remaining known cambalopsid species are considered as belonging to other groups and will be treated elsewhere.

Below a description is given of the available material, followed by a key to all of the constituent species of the granulatus -group, as well as a brief analysis of their variation and distribution.

Glyphiulus granulatus ( Gervais, 1847) View in CoL ( Figs 1-6 View FIG View FIG View FIG View FIG View FIG View FIG )

Iulus granulatus Gervais, 1847: 170 , pl. 44, fig. 10.

Glyphiulus granulatus View in CoL – Karsch 1881: 14.

Cambala nodulosa Butler, 1876: 444 , synonymized by Jeekel (2004: 53).

Formosoglyphius tuberculatus Verhoeff, 1936: 57 View in CoL , figs 7-9, synonymized by Korsós (2004: 18, 19).

Glyphiulus vulgatus Zhang & Li, 1982: 85-87 View in CoL , figs 1-6, n. syn.

TYPE MATERIAL. — Réunion (= Île Bourbon). La Bonite, 1838, leg. Eydoux, lectotype ♀ (incomplete, lacking caudal body end), 5 ♀♀ paralectotypes (all fragmented), all here designated ( MNHN GA 013 View Materials ) ; material in rather poor condition (most legs missing), apparently dried before being placed in alcohol.

Mauriès (1983) mentioned that only part of the type material of this species is still preserved in the MNHN, with the series from Île de France (= Mauritius) presumably being lost. The lectotype selection made here is necessary to fix the exact type locality ( Réunion Island).

TYPE LOCALITY. — Réunion.

OTHER MATERIAL EXAMINED. — Réunion. 10.II.1972, leg. J. Travé, strict topotype juv. ♀ ( MNHN GA 022).

Mauritius. Near Mahébourg, sugar cane debris, 6.II.1972, leg. J. Travé, 1 ♀ (det. J.-P. Mauriès, MNHN GA 022).

Comoros. Mayotte Island, Mt. Combani, 12°48’S, 045°08’E, 470 m, Winkler extraction of forest litter, 22-24.II.1999, leg. Jocqué & De Smet, 3 ♂♂, 1 ♀, 1 juv. ( ZMUM, ex coll. MRAC No. 017.940), several ♂♂ and ♀♀ ( SEM).

China. Hong Kong, leg. M. Crosland, det. J.-P. Mauriès, 2 ♀♀ ( MNHN GA 013).

Fiji. Viti Levu Island, Suva, park, in litter, 5.VI.1980, leg., det. & don. S. Golovatch, 1 ♂ ( MNHN GA 013).

Seychelles. Mahé Island, 1892, leg. C. Alluaud, det. H. W. Brölemann, 1 complete ♀, 1 incomplete ♀ ( MNHN GA 013).

New Caledonia. Dumbea, leg. Mme Pruvot, det. H. W. Brolemann, 4 ♂♂, 2 ♀♀ ( MNHN GA 013).

Loyalty Islands. Lifou, leg. Mme Pruvot, det. H. Brolemann, 1 incomplete ♀ ( MNHN GA 013).

DIAGNOSIS. — Differs from congeners primarily by the peculiar anterior gonopods, each showing an unusually high median outgrowth of the coxosternum, combined with the typical patterns of carinotaxy.

Review of Glyphiulus : the granulatus -group ( Diplopoda, Cambalopsidae )

DESCRIPTION

Length of adults of both sexes 10-18 mm, midbody segments round in cross-section ( Fig. 3B View FIG ), their width (horizontal diameter) and height (vertical diameter) similar, 0.6-0.9 mm. Coloration yellowbrown; non-faded specimens variegated, with a dark brown vertex, blackish ocellaria, mainly brown crests on collum and brownish lateral longitudinal stripes beginning from dark brown ozoporiferous tubercles. Sometimes a thin axial line traceable due to darker median crests.

Adult body with 30-53 podous segments + 4-2 apodous ones + telson (formula 30-38p+4a+T in smaller specimens, up to 44-53p+2a+T in larger ones; formulae generally following Enghoff et al. 1993). Clypeus with 3-6 teeth anteromedially. Eye patches transversely ribbon-shaped (smaller specimens) to ovoid (larger individuals), each composed of 11-18 rather flat ocelli in 5 or 6 irregular longitudinal rows ( Fig. 4A View FIG ). Antennae short and clavate ( Figs 2B View FIG ; 3A View FIG ; 4A View FIG ), antennomeres 6 and 7 with a small distodorsal field or corolla of bacilliform sensilla ( Fig. 4B, C View FIG ). Gnathochilarium usually, but not always (n = 10:1), with a separate promentum ( Fig. 3A View FIG ).

Head width = segment 2 <collum = midbody segments (close to 18th to 20th)> segment 3 = 6> 4 = 5 <7 <8 = 10; body abruptly tapering toward telson on a few posteriormost segments. Postcollar constriction very evident ( Fig. 2C View FIG ).

Collum with 7+7 longitudinal crests starting from fore edge, but both median crests interrupted in about caudal 1/3, being replaced there by similar 1+1+1 crests ( Fig. 2 View FIG A-C) (formula I- VI+7a+pc+ma+pc+7a+VI-I).

Subsequent metaterga similarly strongly crested ( Fig. 2 View FIG B-E), especially so from segment 5 onwards, whence enlarged pore-bearing tubercles commence, less evident on legless segments in front of telson due to loss of ozopores ( Fig. 2E View FIG ). Ozoporiferous tubercles round, wider than high; midbody metatergal crests divided into two about midway, each half neither especially high nor sharp ( Fig. 2 View FIG B-F). Carinotaxy formulae 3+I+4+I+3 and 3+i+3+i+3, the former standing for front row of crests, the latter for caudal one, both fairly independent ( Figs 2 View FIG D-F; 3B).

Tegument delicately alveolate-areolate, dull throughout. Fine longitudinal striations in front of stricture between pro- and metazona, remaining surface of prozona very delicately shagreened. Metatergal setae absent. Segment 2 with long pleural flaps ( Fig. 2B View FIG ). Limbus extremely finely and more or less regularly denticulate. Epiproct simple, devoid of tuberculation, like a transverse rounded ridge in caudal part ( Figs 2E, F View FIG ; 3C, D View FIG ). Paraprocts rather regularly convex, each with a row of several setae at medial margin and 2+2 setae more laterally ( Fig. 3C, D View FIG ). Hypoproct rather evidently but broadly emarginate caudally to receive ventral edges of paraprocts, with 1+1 strongly separated setae near caudal margin ( Fig. 3C View FIG ).

Ventral flaps behind gonopod opening on male segment 7 barely distinguishable as low swellings, not forming a marked transverse ridge ( Fig. 3E, F View FIG ).

Legs short, on midbody segments about 3/4 length of segment height ( Figs 2B, D, E View FIG ; 3B View FIG ). Claw at base with a strong accessory spine almost half as long as claw itself ( Fig. 5D View FIG ). Tarsi characteristically fringed terminally, some of terminal setae with scattered denticles ( Fig. 5D View FIG ).

Male legs 1 highly characteristic ( Figs 4D View FIG ; 6A View FIG ) in being very strongly reduced, represented only by a sternum devoid of any median or paramedian structures but carrying 1+1 strongly separated prongs, both evidently curved posteriad and bearing several strong setae and a minute tubercle (vestige of legs) at base on caudal face. Male legs 2 very slightly hypertrophied, only claw and, anteriorly, coxa somewhat reduced; penes broad, oblong-subtrapeziform, each with 2 or 3 strong setae distolaterally ( Figs 4E View FIG ; 6B View FIG ). Male legs 3 modified in having coxa especially slender and elongate ( Fig. 4F View FIG ).

Anterior gonopods ( Figs 5A, B View FIG ; 6C View FIG ) with a typical shield-like coxosternum which is modestly setose on caudal face and provided with a pair of con- spicuous, high, terminally coiled, mesal processes (visible also in situ, Fig. 3E View FIG ). Telopodite small but movable, 1-segmented, lateral in position, with 2 or 3 strong apical setae and a field of small setae at base, slightly longer than adjacent lateral corner of coxosternum. Posterior gonopods ( Figs 5C View FIG ; 6D View FIG ) very compact, coxite medio-apically with a plumose distal process and a hyaline lobe, telopodite setose both laterally and medially, lower than both coxal process and lobe.

Vulvae very simple, bare, modestly emarginate medially ( Fig. 6G, H View FIG ). Female legs 1 and 2 as in Figure 6E, F View FIG , female coxa 3 as slender as in male ( Fig. 6G View FIG ).

Early juvenile stadia recognised by normal, large ozoporiferous tubercles on segments 5 and 6, these tubercles being considerably reduced in size on subsequent segments.

REMARKS

The above is the first formal record of G. granulatus in Fiji.

Based on the original description and illustrations of G. vulgatus alone ( Zhang & Li 1982), it is clear that this taxon does not differ in any way from G. granulatus , thus justifying the above new synonymy. The far-inland record of G. granulatus (= G. vulgatus ) in Guangxi Province, southern China suggests a continental Chinese origin of this species, which has since become established on numerous tropical islands all over the globe. The port of Hong Kong seems to have been a plausible outlet for the recent expansion of G. granulatus through human agency. The fact that, of all the numerous congeners, only this species that has attained such a vast distribution seems to be rooted in its biology and ecology, the most relevant traits being its fairly small size (on average the smallest) and, apparently, a relatively rapid development, suggested indirectly by the observations of Enghoff (1993) and Mauriès & Nguyen Duy-Jacquemin (1997). The ontogeny of certain cambalopsids is unique among Diplopoda in containing a haplopodous early juvenile stage (third?). At this stage, all body segments after the sixth are only equipped with a single pair of legs!In G. granulatus , the total number of such segments is only up to 13, as opposed to 18 in G. subgranulatus n. sp. and G. semigranulatus n. sp. (see below), 19 in Trachyjulus tjampeanus (Attems, 1903) , from Java, Indonesia (NB: in the abundant material of cambalopsids taken from numerous caves from all over Java and accumu- lated at MNHN, we have only encountered this species!), and 28 in Glyphiulus zorzini , a southern Chinese cavernicole. In addition, in the Hawaiis, G. granulatus has been recorded as a myrmecophile ( Crosland 1994).

Similarly, based solely on the original descriptions of G. septentrionalis and G. multicarinus (cf. Murakami 1975; Zhang & Li 1982), it is clear that these species cannot be separated, agreeing perfectly with each other in every character. Hence the new synonymy advanced: G. septentrionalis Murakami, 1975 = G. multicarinus Zhang & Li, 1982 . The occurrence of this species on Okinawa, Ryukyus, both in caves and epigeically ( Murakami 1975), seems only to represent a recent introduction through human agency, with southern China definitely having served as the source area. Much like G. granulatus , G. septentrionalis is a relatively small species (adults 17-30 mm long), but the number of segments is usually considerably higher (43-66p+4-1a+T), implying a longer development, which apparently reduces its vagility.