The alluaudi-group of Gíomeris, another Macaronesian species swarm in miliipedes (Diplopoda: Glomeridae)

SERGEI 1. GOLOVATCH

Goiovatch, S.I.: The alluaudi-group of , another Macaronesian species swann in mii- lipedes (Diplopoda: Glomeridae). Ent. yand Ent. scand. 17: 503-509. Copenhagen. Denmark 3 April 1987. ISSN 0013-8711. Glomeris alluaudi Brolemann, 1900 (type-specics of Trichoglomeris Verhoeff, 1M)and G. gomerana Attems, 1911. both redescribed upon newiy coliected topotypes, as weil as G. COM- riensis sp.n.. al1 endemic in the Canary Islands, are shown to form a distinct spccies-group within Glomeris Latreilie, 1802-03 (Trichoglomerk being another of its numerous junior synonyms). Like several other genera, the olluaudi-group seems to form a species swarm in Macaronesia. S.I. Goiovatch, Institute of Evolutionary Morphology and Ecoiogy of , USSR Academy of Sciences, Moscow V-71,Leninsky prospekt 33, USSR.

The first glomerid millipede described from In short, the finding of male topotypes of Glo- Macaronesia was Glomeris alluaudi Brolemann , meris alluaudi was crucial for solving the whole 1900 from Tenenfe, Canary Islands. Despite the riddle, while the existence of G. gomerana, exter- fact that Brolemann (1900) had erected his taxon nally a good Trichoglomeris, but anatomically an for a single female, Verhoeff (1906) regarded the indisputable Glomeris, was only an indirect and peculiar tergal pilosity in G.alfuaudi sufficient for often ignored indication. creating an independent , Trichoglomeris. Soon after that Attems (191 1) described a second Material and acknowledgements. Through the kind mis- Canarian Trichoglomeris, G. gomerana, from tance of Dr. H. Enghoff. Universitetcts Zooio@&e Gomera, though cautiously referred to Verhoeffs Museum, Copenhagen (ZMUC), Dr. J. Gruber, Natur- historisches Muscum Wien (NHMW),Dr. LP. Mau- category as a subgenus of Glomeris. Indeed, ries, MusCum Nationai d’Histoirc Naturcile, Paris Attems (1911) had good reasons to do so, because (MNHP), and Dr. K. Thaier, Institut für Zoologie der he had at hand a series comprising males. He noti- Universitat Innsbruck, 1 have been pnviiglled not oniy to restudy the type specimens of both G. alluaudiand G. ced that the telopod structure of his G.gumerann, gomerana, but aiso to examine new, topotypicai materi- the crucial character for generic allocation in the ai. The fresh sampie from Tenerife, made by Dr. K. , was practically identical to that of the Thaier, has turned out to contain a maie of G. alluaudi, numerous European Glomeris species. thiis making it possibie to judge on the true status of Trichoglomeris. No less interesting. two sampies from However, later on both Attems (1926) and Ver- Gomera, both beionging to the Copcnhagen Museum, hoeff (192fj-32) resurrected Trichoglomeris to full comprised, besides G. gomerana, a new spccia. generic status. though the latter author still re- Before going further, 1 should iike to extend my sin- tained it for the type-species only. In his latest re- cere thanks to the abovementioned penons. classification of the Diplopoda. Hoffman (1979) practically agreed with Verhoeffs decision, al- though Mauries (1971) had expressed serious doubts as regards the generic rank of Trichoglo- ’ Pcrhap on geographical rather ihan morpholoyical considcnrionr. MauriC (1971) regarded Trichoglomrru (non TrichomtN- this must nieris and had even discarded this name from his be a typographical error) a nomen dubium sub Loboglomeris, a gcnur own systern of the Glomerida‘. rrrtricied 10 ihe Pyrenees and Canrabrie Mountainr.

Eniomologca scandinavrca (Grp. 8) ENT. SCAND. VOL. 17 (1986)

TAXONOMY caudo-femorai process with a weli-deveioped dis- tal membranous sac. basally more Hyleoglomeris- Glomeris alluaudi Brolemann, 1900 like. Figs. 1, 2, &6. Glomeris gomerana Attems, 1911 Glomeris Alluaudi Brolemann. 1900: 439; type-locality: Figs. 3,7-9. Tenerife, forest "Las Mercedes" near La Laguna. Glomeris (Trichoglomeris) gomeranu Attems, 191 1: Murerial srudied: Holotype P (MNHP). Tenerife, forest 11 1; type-locality: Gomera. Cumbre del Carbonero. "Las Mercedes". 750 m, 30.V.1890. Alluaud. - 1 d,2 9 (RHMW, Thaler ded.), Tenenfe. Orotava, Aguaman- Materiulstudied: 1 d, herewith designated as lectotype3. sa, 1100-1400 m. 13, 16 & 17.11.1982,Thaler. Gomera, Cumbre del Carbonero, 14.1.1908, May (NHMW). - Paralectotypes: 1 d (dissected). 1 P (in- tact), 1 head, 2slides. same data as lectotype (NHMW)4. Diagnosis. Differs from other Canarian species by the - 1 8,1 O (ZMUC), Gomera, pine forest, 1200 m. X- particularly small body size (width less than 3.0 mm), J. RaMl. relatively low outer coxa1 lobes of leg-pair 17 in ¿, nar- XI.1978, rowly ogive syncoxite notch of leg-pair 18 in d, smaller Diugnosk. Differs from Canarian congeners by the suboval median lingual lamina of the telopods, etc. medium-sized body (width 3.4-4.1 mm), higher outer coxai lobes of leg-pair 17 in d, roundly triangular median lingual lamina of the telopod syncoxite, papillate telo- Notes pod femur, etc. The original description of G. alluaudi is rather accurate (see Broiemann 1900), though incom- Notes plete. The holotype is now considerabiy faded, Attems (1911) must have been misied by Broie- yet the colour pattern is stili eiucidabie (cp. Figs. mann's (1900) description of the coiour pattern in 1 & 2). the fresh topotypes, the coiour is bright In G. alluaudi to state that his G. gomerana was (especially in d), background dark brown, spots quite distinct by this character. On the contrary, a or markings whitish-yellow, ventrum and legs (re)study of pertinent material shows that by colo- pale yeliowish-brown. Body particuiariy small ration G. gomerana is barely distinguishable from (up to 4.9 mm long in topotypes, 2.5 mm wide in G. alluaudi Figs. 2 & 3). However. in G. gome- holotype. 2.2 and 2.9 mm wide in O topotypes, (cp. rana the thoracic shield is generaiiy much more 2.9 mm wide in d topotype). Ocelii 6 1 (holo- + broadly pale along lateral and antero-lateral mar- type') or 7 1 (topotypes). Antennae slender and + gins, ventrum and legs are brown. background long, antennomere 6 2.5-2.6 times as long as coiour of terga is blackish-brown, spots and mark- wide. Head beset with rather long anddense setae ings duii yeliowish. This pattern concerns the below levei of antennai sockets. Coiium with two fresh specimens oniy, while the types are aii much usual transverse striae. Thoracic shieid with three faded, though, e.g., the wide paie patches the (holotype and topotypes) or four (topotype on P) thoracic shield are stili traceable. pody medíum- striae. of which oniy the iast one starts a bit dorsad sized, 3.4-4.1 mm wide, up to5.61ñm Ocelii of the schism and the two or three posteriormost long. 7. 1 (lectotype) or 8 1 (both paralecto- and ones cross the dorsum; hyposchism reaching to + + topotypes). Antennae iong and slender, antenno- hind tergal contour, but not exceeding it. Terga 5- mere 6 c. 3.2-3.3 times longer than wide. Coilum 11 with a very siight median sinuosityof hind mar- gin. Entire tergal surface finely pilose. O : Pygidium without niodifications, roundly convex like in 9. Leg-pair 17 (Fig. 4) with rela- . Even the faded occlli in the holotypc distinctly show [he numberof 6 tiveiy high outer coxa1 iobes, telopodite 4-jointed, + I íperhaps even 7 + l. bccause the anterionn

Figs. 1-9. GIomeris spp. - 1-3. Habitus. dorsal; (1) G. alluaudi Brolernann, holotype 9.2.5 rnrn wide; (2) same. topotypic ? ,2.9 rnrn wide; (3) G. gomerana Atterns. topotypic P. 4.0 rnrn wide. -4-6. G. affuaudi,topotypic 8; (4)leg -pair 17: (5) leg-pair 18; (6) telopods. - 7-9. G.gomerana, topotypic 8;(7) leg-pair 17; (8) leg-pair 18; (9) telopods. - Scalr (4-9): 0.2 nirn. 506 Golovatch, S. I. ENT.SCAND. VOL. 17 (1986) with two usual transverse striae. Thoracic shield ogive syncoxite notch, telopodite 4-jointed. Telo- with three or four striae, of which the posterior pods (Fig. 12) with a particularly large, rounded, one starts a bit above the schism and the one or finely setose lingual lamina of syncoxite, lateral two middle ones cross the dorsum; hyposchism homs thick, setose, apically pointed, subapically reaching to hind tergal contour, though not pro- with a tiny lobule (alike G. gomerano); caudo- jecting beyond it. Terga 5-11 with a very slight femoral process huge, without membranous sac, medial sinuosity of hind margin. Surface of terga almost concealing fronto-femoral flagelliferous finely pilose. finger; tarsus not pointed, rounded. d : Pygidium without modifications, roundly 9: Length c. 10.2 mm, width 6.0 mm. Other convex like in 9. Leg-pair 17 (Fig. 7) with higher non-sexual characters as in d. outer coxal lobes. Leg-pair 18 (Fig. 8) with a wide ogive syncoxite notch. Telopods (Fig. 9) with a The alluaudi-group high and more or less roundly triangular linguai lamina of syncoxite, caudo-femoral process mas- The Canarian Giomeris alluoudi, G. gomerana, sive, typically Gfomeris-like, but with a distal and G. canariensis certainly fonn a weil-defined membranous sac. species-group within this large Euro-Meditema- nean genus. This group may be called the alha- udi-group, with the following characters: (1) mi- Glomeris cananensis sp.n. cropilosity of the terga; (2) unusually long and Figs. 10-12. slender antennae (like in some cavemicoles); (3) outer coxal lobes of leg-pair 17 in O unusually Muter¿af studied: 1 8 (holotype), 1 P (paratype) high (rather reminding of Hyleoglomeris); (4) (ZMUC),Gomera, launsilva, under stoncs and leaves, X-X1.1978,J. Rabsl. Tomosvary’s organs almost transverse. However, this set of characters seems hardly Diagnosir. Differs from Canarian congeners by larger sufficient to warrant a separate (sub)genenc rank, body size (width 5.26.0 mm), uniform blackish colora- tion devoid of spots or markings, higher outer coxal because the groundplan telopod structure is indis- lobes of leg-pair 17 in d. larger syncoxite median Iingual putably Glomeris-like. Thus the name Trichoglo- lamina of the telopods. etc. meris Verhoeff, 1906 must join the long list of junior synonyms of Glomeris Latreille, 1802-03 (see Hoffman 1979). Description What seems particularly interesting is the fact 8 : Length c. 10 mm, width 5.2 mm. Colour black- that within the alluaudi-group we find aparallel to ish-brown, without any spots or markings even on the general trend of complication of the primi- ’ collum; only lateral and hind tergal margins pale tive Glomeris-like telopod type into that met with whitish; ventrum, legs and labium pale brownish. in the more advanced genus Hyleogiomeris. The 9 + 1 black convex ocelli on each side. Head latter is known to compnse a nch variety of usu- setose below leve1 of antennal sockets. Antennae ally medium- to smaller-sized species in the long and slender, with usual four apical cones; Oriental realm, the Himalayas, Mjddle Asia, the antennomere 6 c. 3.2 times as long as wide. Caucasus, Asia Minor and evei the Balkans. Tomosvary’s organs almost transverse, as in both Such forms possess usually an increased number G. alluaudi and G. gomerana. Collum with two of striae on the thoracic shield, a more pronoun- usual transverse striae. Thoracic shield with three ced reduction of ieg-pair 17 in d, and a strongly striae, al1 beginning beneath schism, only the 2nd differentiated caudo-femoral process of the telo- cross the dorsum; hyposchism rounded regularly, pods. reaching to hind tergal contour, but not project- In G. alluaudi, this process is practically Hyle- ing caudad beyond it. Terga 5-10 medially very oglomeris-like, in both G. alluaudi and G. gome- slightly sinuate at hind margin. Tergal surface rana it is provided with a well-developed distal very finely and densely pilose. Pygidium regularly membranous sac (as in Hyleoglomeris), and in al1 rounded. without modifications. the three Canarian species ieg-pair 17 in 8 is re- Leg-pair 17 (Fig. 10) with particularly high duced in a more Hyleoglomeris- than Glomeris- outer coxal lobes, telopodite somewhat reduced, like way. 4-jointed. Leg-pair 18 (Fig. 11) with a broadly Biogeographically, two reasonabie alternatives ENT. SCAND. VOL. 17 (1986) The Gomeris alliiaudi group 507

, 0.2 ,

Figs. 10-12. GIomeris canariemir sp.n., 8 holotype; (10) leg-pair 17; (11) leg-pair 18; (12) telopods. - Scale: 0.2 mm.

can be proposed to explain the distnbution of seems reasonable to surmise that an early Glom- Glomeris and Hyleoglomeris, both implying an ris-libe typc first become establihcd throughout ancient age of the Macaronesian swarm. One Eurasia and later spiit into Hykogldin the alternative seems particularly consistent with the east and present-day Glomeris in the wgt. If so, traditional view of the origin of the Mediterra- Hyleogiomeris may be seen as a relatively young nean nemoral biota from the Oriental one (see (monophyletic?) taxon which, besides Rhopolo- Heptner 1936; Wulff 1944; etc.), and thus is pos- meris, is the only genus of the advanced glomerid tulating the predominance of ancient migrations tribe Trachysphaerini semu Maurits (1971, westwards. Such a view has quite a solid paleon- 1984) that populates the Oriental Region (along tological background as well; the hardwood with the Mediterranean, this is the major centre genera Quercus, Fagus, Pyrus and others are of generic radiation in the entire order Glomer- good examples of Oriental elements migrating ida), and is the only glomerid genus occumng along the northern coast of the declining Tethys between Indochina and the Aegean Sea (the lar- Sea in Early-Middle Tertiary (see Menitsky 1982; gest known generic range in Glomerida). In other Zhilin 1984; etc.). words, the old idea of its Oriental origin and Taking into account the recent discovenes of expansion to the west (probably in Late Pale- two H-vleoglomeris species in the Balkans (see ogene-Early Neogene) may be regarded as plaus- Mauries 1984) linking the modern ranges of Glo- ible (see Golovatch 1975). meris and the more advanced Hyleoglomeris, it On the other hand, though the followingsuppo- 508 Golovarch. s. 1. ENT. SCAND. VOL. 17 (1986) sition is less consistent with biogeographical and Key to the species of the alliraitdi-group of paleontological evidence, it seems not too im- Glomeris probable that an early Glomeris-type, perhaps during Early Tertiary in Europe, could have given 1. Terga with a distinct colour pattern (Figs. 2.3); body much less than 5 mm wide: outer coxal rise to Hyleoglomeris and allied genera (see Mau- lobes of leg-pair 17 in d not high (Figs. 4, 7); ries 1984). An eastward dispersa1 could then have telopod femur with a iarge caudal processprovid- followed the northern coast of the regressing ed with a membranous sac, syncoxite median Tethys Sea. Several westem relicts, including both lingual lamina not broad, at most semi-circular (Figs. 6,9) ...... 2 the Balkan species, obviously of the most ancient - Terna without distinct colour uattern. uniformlv stock, may have survived only as cavernicoles. blackish-brown; body more than 5 mm wide (ih However, in the context of the origin of the adults); outer coxal lobes of leg-pair 17 in d ailuaudi-group of it matters little whe- extrcmeiy high (Fig. 10); telopod caudal femorai Glorneris, proces devoid of membranous sac, syncoxite ther Hyleoglomeris, nowadays almost exclusively median lamina very broad and regularly rounded Asian, originated in situ or is migrant from the (Fig. 12) ...... canariensis west. What seems important is the fact that the 2. Body lcss than 3 mm wide; antennomere 6 c. 2.5 “hyleoglomerization” within the Macaronesian times ionger than wide; outer coxal lobes of leg- pair 17 in 8 low (Fig. 4); telopod median lingual species swarm must have taken place independ- lamina of syncoxite semi-circular, femur not ently and long ago, perhaps in Miocene-Early ~auillatc. caudo-femoral urocess weil differ- Pliocene, and from a true Glomeris-type. At least kiiated (Fig. 6) ...... alluoudi among the 13 currently known Glorneris species - Body width 3.4-4.1 mm;antennomere 6 more than 3 times as long as wide; outer coxal lobes and subspecies of continental Spain (see Vicente of leg-pair 17 in d high (Fig. 7): telopod median 1981), there is no parallel to the development in lingual lamina of syncoxite roundly triangular, the alluaudi-group. femur at least proximally papillate, caudo-fe- moral process poorly differentiated (Fig. 9) ... It is no news that the Macaronesian millipede ...... gomerana fauna is rich (about a hundred species and subspe- cies), comprising not only a lot of endemic forms, but also a number of (sub)endemic species-groups References and (sub)genera. Such are the huge tnudeirae- group of Cylindroiulus chiefly restricted to Attems, C. 1911. Myriopoden von Gomera. Gesammelt von Prof. W.May. -Arch. Naturgesch. 77. Suppl. Madeira (Enghoff 1982b, 1983b), the genus Aci- 2, 1: 107-118. pes containing six species from Madeira and one - 1926. . - In: Kükenthal & Krumbach, each from the Canary Islands and mainland Spain Handb. Zool., Berl. 4: 1402. (Enghoff 1983a, 1986), as well as Dofichoiulus BrBlemann, H.W. 1900. Voyage de M. Ch. Alluaud aux Iles Cananes. (Novembre 1889 Juin 1890). Myri- encompassing a good of species from Sici- - number apodes. -MCm. Soc. 2001. Fr. 13: 131453. ly ,France, Northwest Africa, mainland Spain and Enghoff, H. 1982a. (Dipiopoda) from the about a dozen species from the Canaries, two Cape Verde Isiands. - Cour. ForehInst. Sencken- from Madeira and one from the Cape Verde bera 52: 137-138. Islands (cf. Mauries 1970, 1982; Enghoff 1982a, 1982b. The millipede genus Cylindroirtliccon Madeira - an insular species swarm (Diplopoda. Julida: b). And now the alluaudi-group of Glomeris must Julidae). - Ent. scand.. Suppl. 18: 142 pp. join such examples of Macaronesian species 1983a. Acipes- a Macaronesian genus of millipedes swarms. (Diplopoda, Julida, Bianiulidae). - Steenstrupia 9: 137-179. Interestingly, within the Canarian alluaudi- 1983b. Adaptive radiation of the millipede genus group, body size diversity is great like in the other Cylindroiulus on Madeira: habitat. body size. and known instances of insular species swarms in mil- morphology (Diplopoda, Julida: Julidae). - Revue lipedes (see Enghoff 1983b). Most probably this is Ecoi. Biol. Sol 20: 403115. 1986. A continental species of thc .-iciprs (Dipiopo- also due to niche segregation. But another da: Julida: Blaniulidae). - Scnclcnberg. hiol. 67: interesting point is the question whether the 207-209. “hyleoglomerization” within the alíuaudi-group Golovatch, S.I. 1975. Two new to the USSR fauna went along with a decrease in body size. The genera of Oniscomorpha (Dipiopoda) found in larger G. canariensis (and its habitat?) might then Transcaucasia and their zoogeopraphical connec- tions. - Zool. Zh. 54: 1566-1571 (in Russian). be regarded as the most ancestral species of the Heptner, W.G.1936. General zoogeography. -518 pp. alluaudi-group. Moscow & Leningrad (in Russian). EKT SCAXD. L'OL. 17 (1986) The Comeris nlluaiidi group 509

Hoffman. R.L. 1979. Classification of the Diplopoda. - Verhoeff, K.W. 1906. Über Diplopoden. 4. (24.) 237 pp. Geneve. Aufsatz: Zur Kenntnis der Glomeriden (zugleich Slauriks, J.-P. 1970. Diplopodes récoltés a Maderc par Vorlaufer einer Glomeris-Monographie). (Beitrage C. Allunud en 1938. Description d'une espece nou- zur Systematik, Geographie, Entwicklung, vergleich- wiie du genre Nesopochyiirlus Attems. - Boca- enden Morphologie und Biologie). - Arch. Natur- gianri7J: 14. gesch. 72. l(2): 107-226. - 1971. Diplopodes épigés et cavernicoles des Pyré- - 192632. Diplopoda. - In: Bronn, Klassen und nCes espagnoles et des Monts Cantabriques. VII. Ordnungen des Tierreichs. wissenschaftiich darge- GlomCridrs. Essai de classification des Glomeridea. stellt in Wort und Biid. 5?, 2: 2084 pp. - Bull. Soc. Hist. nat. Toulouse 107: 423436. Vicente, M.C. 1981. Diplópodos epigeosde Cataiuña, 1 - 1%2. Doliclioiiclw rongiorgii (Strasser). Diplopode (Gloméridos, Craspedosómidos y Polidésmidos). - halophile nouveau pour la faune de France. Remar- Eos, Madr. 57: 279-315. qurs sur la classification des Pachyiulini (Mynapo- Weidner, H. 1960. Die Entomologischen Sammlungen da. Diplopoda. Iulida). - Bull. Mus. natn. Hist. des Zooiogisches Staatsinstituts und Zoologischen nat. Paris (4)4A: 43M. Museums Hamburg. 111. Teil. Chilopoda und Progo- - 1954. Deux especes nouvelles de Diplopodes cavemi- neata. - Mitt. hamb. zool. Mus. Inst. 58 57-104. coles des Cyclades: Hyleoglomeris beroni (Glomeri- Wulff, E.W. 1944. Historical geography of plants. - da) et SFrioiiilus undreevi (Iulida). -Biologia gailo- 546 pp. Moscow & Leningrad (in Russian). hellen. 11: 3739. Zhiiin, S.G. 1984. The main stages of the formation of Xlenitsky. Y.L. 1982. Review of the species of the genus the temperate forest flora in the Oligocene-Early Qucrci(s L. of Eurasia. - Komarovskie chteniya 32: Miocene of Kazakhstan. - Komarovskie chteniya 5s pp. (in Russian). 33: 112 pp. (in Russian).

.~lnnicscriptnccepted January 1986.

Jt1urn;il .inJ riibwriprion 1,) reprznh o! pmicul;irgri>ups rii he orikred trom: ,-\POLLO I300K.S. Lundhyvcj 36. DK-57íYI Svendhor~.Venrnnrl.