A reclassification of the millipede superfamily Trichopolydesmoidea, with descriptions of two new species from the Aegean region (Diplopoda, Polydesmida)

Abstract Two new species are described from caves in several Greek islands in the Aegean Sea: Galliocookia gracilis sp. n., a presumed troglobite from Rhodes, Dodecanese Islands, and Sphaeroparia simplex sp. n., likely a troglophile from Kithnos, Cyclades, and Chios, Eastern Sporades. These genera are assigned to the family Trichopolydesmidae Verhoeff, 1910. Because Sphaeroparia Attems, 1909, an Afrotropical genus, nicely bridges the gap, both morphologically and geographically, between the Euro-Mediterranean Trichopolydesmidae and the much more diverse, pantropical Fuhrmannodesmidae Brölemann, 1916, the latter family is considered as a new junior subjective synonym of the former, syn. n. Thus expanded, the family Trichopolydesmidae is rediagnosed and its position within the superfamily Trichopolydesmoidea refined. Because the families Mastigonodesmidae Attems, 1914, Macrosternodesmidae Brölemann, 1916 and Nearctodesmidae Chamberlin & Hoffman, 1950 are also formally synonymized with Trichopolydesmidae, syn. n., the Trichopolydesmoidea currently contains only two families, Trichopolydesmidae and Opisotretidae Hoffman, 1980.

Prompted by the discovery of two new species of trichopolydesmoids in caves on three Aegean islands of Greece, this superfamily is reclassified. One of the genera appears to so nicely bridge the gap between the Euro-Mediterranean Trichopolydesmidae and the much more diverse, pantropical Fuhrmannodesmidae, that the latter family is formally synonymized here with the former.

Material and methods
The material treated below had been taken in 1987 by Petar Beron, of the National Museum of Natural History, Sofia, Bulgaria (NMNHS), in caves on a few Aegean islands of Greece and was offered to me for treatment in August 2013, when I visited the NMNHS in the framework of the Russian-Bulgarian interacademician exchange programme. Most of the type material will be returned to the NMNHS, with only two paratypes retained for the collection of the Zoological Museum, State University of Moscow, Russia (ZMUM).
Gonopod aperture transversely oblong-oval, taking up most of ventral part of metazonite 7. Gonopods (Figs 5, 6) with small, subglobose, evidently exposed, medially fused coxae, each carrying only 1 long seta distolaterally and a long, curved cannula distomesally; a scaly texture on lateral face absent. Telopodite suberect, slender and long, up to about its half being taken up by an elongate, setose prefemoral part; acropodite slightly helicoid, its apical lobe subacuminate, with a pre-apical, lobe-shaped, subtriangular solenomere (sl) and a short, papillate, rounded stump a little proximally. Both prefemoral part and acropodite strictly coaxial, in situ directed forward and parallel to each other. No traces of accessory seminal chamber or hairy pulvillus.
Remarks. Among the Euro-Mediterranean genera of Trichopolydesmoidea, only some show a deeply bipartite and strongly curved gonopod telopodite, the prefemoral part of which is quite elongate, but lies more or less transversely to strongly angular, largely (sub)parallel telopodites and extends across the nearly entire ventral width of segment 7. Such are Trichopolydesmus Verhoeff, 1898 (together with Banatodesmus Tabacaru, 1980), Bacillidesmus Attems, 1898, Napocodesmus Ceuca, 1974 andCaucasodesmus Golovatch, 1985. In contrast, the gonotelopodites in Verhoeffodesmus Strasser, 1959, Cottodesmus Verhoeff, 1936and Occitanocookia Mauriès, 1980 have increasingly shortened prefemoral parts, being enlarged and laterally flattened distad, unipartite and mostly less strongly curved, in Cottodesmus and Occitanocookia also devoid of a solenomere, but sometimes supplied instead with what can be seen as a primordial accessory seminal chamber. In Trichopolydesmus, Heterocookia Silvestri, 1898, Ingurtidorgius Strasser, 1974 and, especially, Mastigonodesmus Silvestri, 1898, the solenomere is flagelliform, branching off near the base of the femorite. In all these genera, the gonotelopodites are strongly exposed, not sunken inside an obvious central coxal cavity (= gonocoel). A modest gonocoel seems to only be observed in Ingurtidorgius and Haplocookia Brölemann, 1915. This latter genus does resemble Galliocookia, but its gonopod telopodite is clearly curved, there is a small gonocoel and both sexes have 20 body segments (see also review by Mauriès 1983).
The new species definitely belongs to Galliocookia, sharing with the other three congeners (see Diagnosis above) a small coxa and a slender, simple and suberect telopodite, the latter showing an extended prefemoral part strictly coaxial with the acropodite. Moreover, the solenomere is likewise modest, lobe-shaped and located distally. Even the presence of 19 or 20 body segments in the male and female, respectively, as well as of a normal ozopore formula, setose metaterga and laterally serrate/indentate paraterga coincide.
Biogeographically, the discovery of a Galliocookia in the Aegean region, so far away from France, is remarkable, emphasizing a pan-Mediterranean distribution pattern of this currently purely cavernicolous, likely troglobitic genus.   Mauriès and Heymer (1996), by 20 body segments in both sexes and the presence of an axial sternal process on the especially simple gonopods, including a nearly fully suppressed solenomere.
Gonopod aperture evident, transversely oblong-oval, taking up most of ventral part of metazonite 7. Gonopods (Figs 12-14, 17) with large, transversely subglobose, evidently exposed, medially fused, medially clearly excavate coxites, each microgranular/scaly and micropilose laterally, carrying only 1 long seta distofrontally and a long, curved cannula distomesally; sternum plate-like, with a slender, central, not too strongly chitinized process (k) between coxites. Telopodites short, stout, subglobose, sac-shaped, very simple, only moderately exposed below coxites, deeply sunken inside a prominent gonocoel; prefemoral part as usual, setose, quite elongate, about as long as, but clearly set at an angle to, acropodite, the latter with a distinct seminal groove running entirely on mesal side and terminating on a vestigial lobuliform solenomere (sl) at base of a rather strong, only faintly curved, apical spine (d). Neither an accessory seminal chamber nor a hairy pulvillus.
The discovery of a Sphaeroparia in the Aegean region is even more remarkable than that of a Galliocookia. Indeed, Sphaeroparia is a rather large genus hitherto believed to be strictly Afrotropical, ranging from Liberia and Benin in western Africa, through Gabon and Zaire, to Kenya and Tanzania in eastern Africa (see review by Mauriès and Heymer 1996). Furthermore, it has been assigned to the large pantropical family Fuhrmannodesmidae (Hoffman 1980, Mauriès andHeymer 1996), as opposed to the Holarctic Macrosternodesmidae, the Nearctic Nearctodesmidae (often referred to as only a subfamily of Macrosternodesmidae), and the Euro-Mediterranean Trichopolydesmidae. The Fuhrmannodesmidae has long been considered as an artificial, composite group (e.g. Hoffman 1980, Mauriès and Heymer 1996, Golovatch et al. 2013, being distinguished from the above allies solely by its being tropical. Golovatch (1994) provided an evolutionary scenario for the genera of Fuhrmannodesmidae known from South America, accepting as the basalmost those genera showing rather small, subglobose gonopod coxae that form no significant gonocoel in which to hinge the largely exposed, usually rather simple and elongate telopodites. Moreover, as in some true Trichopolydesmidae (see above), the prefemoral (= setose) part of the gonopod is mostly orientated transversely to the body axis, extending mesally across the entire width of the coxae. Following a series of transitional states, such forms ultimately culminate in having the gonopod coxae strongly enlarged, forming a large gonocoel in which to conceal the clearly shortened, usually highly complex and deeply sunken telopodites. Their prefemoral parts already tend to be positioned increasingly parallel to the body's main axis, thus providing a transition between the usually small-sized Trichopolydesmoidea (= so-called "micropolydesmoids") to the normally medium-to large-sized Polydesmoidea (= so-called "macropolydesmoids").
Naturally, similar general trends can be surmised to have occurred in the fuhrmannodesmids of Central America and the Afrotropical and Oriental realms, which also support fairly diverse faunas of this family. Remarkably, none of the genera of Trichopolydesmidae (see above), all confined to Europe and the Mediterranean, though demonstrating a certain degree of variation in the length and orientation of the gonopod prefemoral part, has a deep gonocoel. Furthermore, Galliocookia as one of the few genera where the gonocoxae are particularly small while the prefemoral part is still elongate, but already strictly coaxial with a very strongly exposed acropodite represents an evolutionary extreme, apparently the basalmost situation. Figures 11-14. Sphaeroparia simplex sp. n., holotype (12, 13) and ♂ paratype (11, 14) from Kithnos 11 antenna, lateral view 12 leg 9 and both gonopods, caudal view 13, 14 both gonopods, oral and ventral views, respectively. Scale bars: 11 0.2 mm; 12-14 0.1 mm.
The discovery of a Sphaeroparia in the eastern Mediterranean, of an Afrotropical genus demonstrating a very large and deep gonocoel for the small and only poorly exposed telopodites to be hinged into, clearly represents the opposite, evolution-arily obviously the most advanced extreme. Therefore, the Euro-Mediterranean Trichopolydesmoidea appear to show basically the same full range of evolutionary trends in the development of gonopod structures as do at least the properly assessed South American representatives now assigned to Fuhrmannodesmidae! Among the Afrotropical Trichopolydesmoidea, Sphaeroparia is certainly the most speciose genus and Mauriès and Heymer (1996) divide it into as many as six subgenera, most of which they themselves admit to be ill-grounded. A Sphaeroparia species is likely to be present also in the Seychelles (Golovatch and Gerlach 2010). In contrast to the Euro-Mediterranean region, the Afrotropical realm is thus dominated by trichopolydesmoids showing a deep gonocoel between strongly hypertrophied gonopod coxae. Only Peronorchus parvicollis Attems, 1907, the type and still sole species of Peronorchus Attems, 1907, shows a far more basal gonopod structure quite comparable to that of Trichopolydesmus. That species was originally described from near Buitenzorg (= Bogor), Java, Indonesia, and it has since been redescribed from material from Mauritius, Indian Ocean and formally assigned to the family Trichopolydesmidae (Mauriès and Geoffroy 1999). However, Peronorchus has recently been transferred to Fuhrmannodesmidae (Golovatch et al. 2013). Considering the above new evidence, I no longer hesitate to formally synonymize the family Fuhrmannodesmidae Brölemann, 1916under Trichopolydesmidae Verhoeff, 1910 n.

A reclassification
The following refined classification and diagnosis of the superfamily Trichopolydesmoidea, as well as of its two constituent families can be proposed (see also Golovatch 2011, Golovatch et al. 2013.

Superfamily Trichopolydesmoidea Verhoeff, 1910
Diagnosis. Body largely polydesmoid, only 2-16 mm long, exceptionally up to 30 mm, usually ≤ 10 mm long, with 18-20 segments. Antennomeres 2-6 usually subequal in length, but 6 th normally highest; 5 th often, 6 th only exceptionally, devoid of an apicodorsal field of sensilla. ♂ head sometimes with vertigial modifications (humps, fossae etc.). Paraterga at most with only thin calluses, often devoid of these, lateral margin entire, only exceptionally lobulate (Trilobodesmus Golovatch & Mauriès, 2007), typically at least faintly incised and setigerous; metaterga devoid of a cerategument, more often with 3-4, more or less regular, transverse rows of low, polygonal or rounded, setigerous bosses, but rather usually these latter either obliterated or pronounced and more numerous. Ozopores flush, opening on dorsal surface of paraterga to be located at their lateral margin, pore formula normal. Sphaerotrichomes rather often present, usually affecting both tibiae and tarsi.
Gonopod aperture large, transversely ovoid, not extending onto prozonite, but sometimes spreading caudad between coxae 9; gonocoxae subglobose, sometimes with a distolateral projection; prefemoral parts usually lying transversely to main body axis and, together with coxae, taking up most of ventral extent of ♂ ring 7, quite often with an obvious gonocoel formed by enlarged coxae; cannula and seminal groove nearly always present, both absent only exceptionally; entire telopodites or, more usually, acropodites distal to prefemoral parts usually complex, parallel to main body axis and directed forward, but often either crossing each other or, more rarely, directed more or less laterad, dorsolaterad or, exceptionally, even caudad, also exceptionally perforating lateral walls of coxites. A solenomere, an exomere and/or an endomere often present, sometimes these being prominent. An accessory seminal chamber and/or a hairy pulvillus at its orifice only seldom developed, mostly tend to be absent.
Distribution. All continents except Australia, mostly a Northern Hemisphere group only marginally represented in the Southern Hemisphere (Indonesia + New Guinea and Melanesia, Afrotropical realm north of Malawi and Madagascar, Neotropical realm north of Bolivia, Paraguay and southern Brazil), apparently fully allopatric with the superfamily Dalodesmoidea.
The diversity of gonopod structural plans in nearctodesmids+macrosternodes mids (Shelley 1994, Shear andShelley 2007) appears to be quite modest and uniform, since all of their constituent genera such as Nearctodesmus Silvestri, 1910, Kepolydesmus Chamberlin, 1910, Bistolodesmus Shelley, 1994, Tidesmus Chamberlin, 1943 show clearly transverse and elongated prefemoral portions which are set subrectangular to the subparallel, usually bi-or triramous, elaborate, normally clearly curved and well exposed acropodites. The gonocoel if any is moderate at most. According to Shear and Shelley (2007), the differences between these "families" lie only in the number and location of branches on the acropodite, a distinction that by far fails to exceed the diversity of gonopod plans observed among the South American fuhrmannodesmids alone (Golovatch 1994).
Contents and distribution. Thus refined, the Trichopolydesmidae currently contains ca 20 genera and about 60 species in the Holarctic, as well as ca 55 genera and about 80 species in the tropics.

Family Opisotretidae Hoffman, 1980
http://species-id.net/wiki/Opisotretidae Diagnosis. Sphaerotrichomes absent. Gonopod aperture large, transversely oval, with small and largely exposed coxites. Telopodites elongate, directed strongly dorsolaterad; seminal groove running along most of telopodite extent on caudal face to terminate distally either on a special branch or tooth (= solenomere), flush on caudal surface, or debauch inside an accessory seminal chamber which normally is supplied with a hairy pulvillus.
Type genus. Opisotretus Attems, 1907 Contents and distribution. Only seven genera and 29 described species ranging from the Ryukyu Islands, Japan, Taiwan, southern mainland China and the Himalayas in the North and Northwest, through Indochina and across Indonesia, to Papua New Guinea in the Southeast (Golovatch et al. 2013