Tropical Natural History 10(1): 27-36, April 2010 ©2010 by Chulalongkorn University

The Family Haplodesmidae (Diplopoda, ) Recorded in Taiwan for the First Time, with the Description of a New Species

SERGEI I. GOLOVATCH1*, ELENA V. MIKHALJOVA2, ZOLTÁN KORSÓS3 AND HSUEH-WEN CHANG4

1Institute for Problems of Ecology and Evolution, Russian Academy of Sciences, Leninsky pr. 33, Moscow 119071, RUSSIA 2Institute of Biology and Soil Science, Far Eastern Department, Russian Academy of Sciences, Prospekt Stoletiya 159, Vladivostok 690022, RUSSIA 3Department of Zoology, Hungarian Natural History Museum, Baross u. 13, H-1088 Budapest, HUNGARY 4Deparment of Biological Sciences, National Sun Yat-Sen University, 70 Lien-hai Rd., Kaohsiung 804 Taiwan, ROC * Corresponding author. E-mail: [email protected] Received: 14 January 2010; Accepted: 1 February 2010

ABSTRACT.– The East Asian to Australasian millipede family Haplodesmidae is reported from Taiwan for the first time on the basis of Eutrichodesmus taiwanensis n. sp. This new species joins the newly established peculiaris-group which also includes the Japanese Eutrichodesmus peculiaris (Murakami, 1966), E. nodulosus (Verhoeff, 1939) and E. silvaticus (Haga, 1968), as well as E. pectinatidentis (Zhang, 1995), E. anisodentus (Zhang, 1995) and E. soesilae Makhan, 2010, this latter trio from continental China. The species group is characterized by complete volvation often showing an unusual overlap pattern which becomes typical from segment 3 or 4, the presence of only two transverse rows of metatergal tuberculations, the broad and dorsoventrally flattened epiproct (sometimes unusually spatuliform in the male), and the gonopods slender, fully to nearly fully devoid of a distofemoral process, but at least sometimes supplied with an accessory seminal chamber. All of these seven species are keyed. The genera Kylindogaster Verhoeff, 1939 and Thelodesmus Miyosi, 1951 are formally allocated in Haplodesmidae, this being confirmed in the former case and newly established in the latter one. The following new synonymies and combinations are proposed: Kylindogaster Verhoeff, 1939, Thelodesmus Miyosi, 1951, Eucondylodesmus Miyosi, 1956 and Nanocondylodesmus Zhang, 1995 = Eutrichodesmus Silvestri, 1910, all syn. n.; E. nodulosus (Verhoeff, 1939), comb. n. ex Kylindogaster; E. armatus (Miyosi, 1951), comb. n. ex Thelodesmus; E. elegans (Miyosi, 1956), comb. n. ex Doratodesmus Cook in Cook and Collins, 1895; E. silvaticus (Haga, 1968), comb. n. ex Dimorphodesmus Murakami, 1966; E. pectinatidentis (Zhang, 1995) and E. anisodentus (Zhang, 1995), both comb. n. ex Nanocondylodesmus; Prosopodesmus similis (Haga, 1968), comb. n. ex Rhipidopeltis Miyosi, 1958.

KEY WORDS: Haplodesmidae; taxonomy, Eutrichodesmus; new species; species group; Taiwan

INTRODUCTION synonymization of Agathodesmus Silvestri, 1910 with Atopogonus Carl, 1926 (Mesibov, Both the East Asian to Australasian 2009) and the description of E. soesilae millipede family Haplodesmidae and its Makhan, 2010 from continental China largest constituent genus Eutrichodesmus (Makhan, 2010). Prompted by the discovery Silvestri, 1910 have been reviewed very of the first haplodesmid in Taiwan, another recently (Golovatch et al., 2009a, b). The new Eutrichodesmus, we take the only changes made since have been the opportunity not only to describe it, but also to 28 TROPICAL NATURAL HISTORY. 10(1), APRIL 2010 update and refine the classification of this ♀♀ (IBSS), same locality and date, together family through proposing a new family with holotype. allocation and a number of synonymies and combinations. The first species group in Non-types.– 3 ♀♀, 3 juveniles (with 18 Eutrichodesmus is also delimited here, with segments) (HNHM), Taiwan, Taichung all of its seven constituent species keyed. County, Mts Dashue-san, logging road No. 210, Cryptomeria plantation, 2000 m a.s.l., MATERIALS AND METHODS 14 October 2007, leg. Z. Korsós. 2 ♂♂, 1 ♀ (NMNS), Taichung County, Heping The material serving as the basis for the Township, Sihuyuan Wind Gap, 2050 m, present contribution was preserved in 75% 21.VIII.2002, collector unknown. 1 ♂ alcohol and is currently shared between the (NSYSUB), Kaohsiung County, Liouguei, collections of the National Museum of Shanping Workstation, May 2004, leg. Mei- Natural Science, Taichung, Taiwan (NMNS), Jhu Hung. Department of Biological Sciences, National Sun Yat-Sen University, Kaohsiung, Taiwan Name.– To emphasize the first haplodesmid (NSYSUB), Zoological Museum, State formally recorded in Taiwan. University of Moscow, Russia (ZMUM), Hungarian Natural History Museum, Diagnosis.– Differs from the closest Budapest, Hungary (HNHM), Natural congeners in the peculiaris-group mainly in History Museum of Denmark, University of the bifid gonopod endomere, coupled with Copenhagen, Denmark (ZMUC), Muséum the presence of a rudimentary distofemoral National d’Histoire Naturelle, Paris, France process (see also Key below). (MNHN), and Institute of Biology and Soil Science, Far Eastern Branch, Russian Description.– Length ca 5.0 (♂) to 6.0 mm Academy of Sciences, Vladivostok, Russia (♀), width ca 1.0 mm, body broadest on (IBSS), as indicated thereafter. Specimens segments 5-16 (♂) or 5-17 (♀). Holotype ca were studied and illustrated using standard 5.0 mm long and 1.0 mm wide. Coloration stereomicroscopic, photographic and drawing uniformly pallid to yellow (Fig. 1A). equipment. Adults with 19 (♂) or 20 segments (♀), conglobation complete, pattern of volvation SYSTEMATICS unusual for Haplodesmidae in overlap switching to typical starting already from Eutrichodesmus taiwanensis n. sp. segment 3 (Fig. 1B, C) (cf. Golovatch (Figures 1 and 2) 2003). Head slightly transverse (wider than Holotype.– ♂ (NMNS-6242-001), Taiwan, high), rather densely pilose, microgranular Taipei City, Wenshan Distr., Chih-Nan and microvillose just below antennae and on Temple, March 2002, leg. C.C. Chen et al. vertex; epicranial suture shallow and wide. Antennae rather short and clavate; Paratypes.– 19 ♂♂, 19 ♀♀ (NMNS-6242- antennomere 6 longer than 5th, both with an 002), 2 ♂♂, 3 ♀♀ (NSYSUB), 3 ♂♂, 3 ♀♀ evident dorso-apical pit containing a tight (ZMUM), 2 ♂♂, 3 ♀♀ (HNHM), 2 ♂♂, 3 group of minute bacilliform sensilla; ♀♀ (ZMUC), 1 ♂, 1 ♀ (MNHN), 1 ♂, 2 antennomere 8 with the usual four sensory GOLOVATCH ET AL. — MILLIPEDE FAMILY HAPLODESMIDAE IN TAIWAN 29

FIGURE 1. Habitus of Eutrichodesmus taiwanensis n. sp., paratypes ♀ (A, B) and ♂ (C), dorsal, lateral and lateral views, respectively. Photographed not to scale. cones apically. Collum subtrapeziform, covered with a cerotegumental crust held by rather large, slightly broader than head, abundant microvilli. Metatergum 2 with flattened mid-dorsally, not covering the three, subsequent metaterga with two head from above; entire surface transverse and mixostictic (i.e. irregular in microvillose, with four transverse rows of axial direction) rows of subequal, low, round bosses/tubercles (Fig. 1). Prozona rounded bosses, each boss obviously very finely alveolate, collum and metaterga supporting an abraded seta traced only as an 30 TROPICAL NATURAL HISTORY. 10(1), APRIL 2010 insertion point (Fig. 1A-C). Metaterga 2-4 longer than coxite, rather slender evidently flattened mid-dorsally (Fig. 1). throughout, setose in its basal one-third, Paramedian mid-dorsal bosses in fore row with a small, subtriangular, low, lateral lobe or even in both rows on metatergum 5 often obviously corresponding to a distofemoral slightly higher than those on following process (dp) at about midway of seminal metaterga. Paraterga strongly declivous, groove. No traces of a transverse sulcus near rather broad, rounded and very faintly point of seminal groove’s recurvature bilobate laterally, evidently surpassing level laterad. Seminal groove debouching in an of venter, caudolaterally at base with one evident accessory seminal chamber showing distinct lobulation (Fig. 1B, C); paraterga 2 a pilose pad area and terminating near base strongly enlarged, with a series of four both of a small, short, lobe-shaped exomere bosses near anterolateral edge, schism and (ex) and a longer, slightly twisted, bifid hyposchism both very small; paraterga 3 endomere (en), latter clearly separated by a and 4 slightly shorter than others (Fig. 1B, medial constriction and carrying a short, C), overlap of following paraterga typical. distally tuberculate, flagelliform outgrowth Paraterga 17 and 18 either directed at base on lateral face. ventrocaudad (♂) or, like 19th, small and as usual strongly declined (♀) (Fig. 1B, C). Remarks.– This pallid species seems to Pore formula normal (5, 7, 9, 10, 12, 13, 15- inhabit the entire island and occurs at 19), ozopores evident, located near base of various elevations (up to the midmontane caudolateral lobulation. Pleurosternal ridges subtropical forest belt above 2000 m). We absent. Epiproct strongly broadened, either face a typical “doratodesmid” (capable of especially clearly flattened, subrectangular volvation, see Golovatch et al. 2009a) in lateral view, bare and roundly spatuliform showing especially close affinities with in dorsal view, and carrying a minute three congeners: Eutrichodesmus peculiaris middle knob as base (♂), or with three rows (Murakami, 1966), from Oshima, Niihama of bosses dorsally, dome-shaped, regularly Prefecture, Shikoku Island, Japan (Muraka- sloping in lateral view and subtriangular in mi, 1966); E. nodulosus (Verhoeff, 1939), caudal view (Fig. 1B, C), i.e. just as from a cave in the Ryukyu island of described and depicted for E. peculiaris by “Fukafugusa” (Verhoeff, 1939), a dubious Murakami (1966). Hypoproct and locality because no such island exists in the paraprocts typical of the genus. Ryukyus, later reported also from Ishigaki Sterna usually with a deep, narrow, and Iriomote islands (Omine, 1982), both transverse depression between coxae. Legs lying south of Okinawa (ca 150 km east of rather long and relatively slender, barely Taiwan), as well as on Okinawa Island reaching tips of paraterga; femoral and proper (Omine and Ito, 1998), the Ryukyus, tarsal segments longest, subequal in length; Japan; and E. silvaticus (Haga, 1968), from claw normal, simple, very slightly curved Gooya, Hojoo-machi, Tagawa City, ventrad; some setae with microdenticula- Fukuoka Prefecture, Kyushu Island, Japan tions. ♂ coxa 8 clearly enlarged. (Haga, 1968). These four species seem to Gonopods (Fig. 2A-D) rather simple. share complete volvation (albeit at least Coxae subquadrate, large, microtuberculate sometimes with an unusual overlap pattern on lateral face and with two macrosetae which becomes typical already from latero-apically. Telopodite only slightly segment 3), the presence of only two GOLOVATCH ET AL. — MILLIPEDE FAMILY HAPLODESMIDAE IN TAIWAN 31

FIGURE 2. A-D. Gonopods of Eutrichodesmus taiwanensis n. sp., paratype ♂, lateral, submedial, lateral and submedial views, respectively. Scale bars: 0.1 mm. transverse rows of metatergal tubercula- with Eutrichodesmus Silvestri, 1910 tions, the remarkably broad and dorsoven- (Golovatch et al., 2009a), while Kylindo- trally flattened epiproct (especially so in ♂), gaster was simply overlooked. Its only and the slender gonopods virtually or nearly constituent species, K. nodulosa Verhoeff, devoid of a distofemoral process, but at least 1939, was based on a single female sometimes supplied with an accessory specimen with 19 body segments, seminal chamber. This seems to warrant apparently adult because the body lumen their assignment to a single species group, was said to be filled with eggs (Verhoeff, the peculiaris-group, the first to be 1939). Judged from the general body shape, delimited in Eutrichodesmus. the presence of only two transverse rows of Murakami (1966), when describing his bosses on most of the metaterga, albeit Dimorphodesmus peculiaris, correctly isostictic (i.e. regular in axial direction) and compared his new genus with Eucondylo- somewhat differentiated, the peculiar, desmus Miyosi, 1956, Kylindogaster strongly broadened epiproct, and prove- Verhoeff, 1939 and Doratodesmus Cook in nance (the Ryukyus), there can be no doubt Cook and Collins, 1895. Both Eucondylo- whatever that this species belongs in the desmus and Doratodesmus have since been peculiaris-group. This confirms the family placed in Haplodesmidae, the genus allocation [in Doratodesmidae, a family Eucondylodesmus has been synonymized currently considered as a junior synonym of 32 TROPICAL NATURAL HISTORY. 10(1), APRIL 2010

Haplodesmidae, see Golovatch et al. spines as well. This implies the following (2009a)] which seems to have first been new synonymy and combinations: proposed by Murakami (1993). This also Nanocondylodesmus Zhang, 1995 is still implies the following formal nomenclatural another junior subjective synonym of changes: Kylindogaster Verhoeff, 1939 is Eutrichodesmus Silvestri, 1910, syn. n., another junior subjective synonym of while Nanocondylodesmus pectinatidentis Eutrichodesmus Silvestri, 1910, syn. n., Zhang, 1995 and N. anisodentus Zhang, while Kylindogaster nodulosa Verhoeff, 1995 are formally to be transferred to 1939 is to be referred to as Eutrichodesmus Eutrichodesmus, both comb. n. nodulosus (Verhoeff, 1939), comb. n. Another congener, E. soesilae Makhan, As regards Dimorphodesmus silvaticus 2010, which superficially is especially Haga, 1968, also overlooked by Golovatch similar to E. anisodentus, has just been et al. (2009a), Haga (1968) quite correctly described from Mt. Jinyun, Beibei District, emphasized the especially evident Chongqing Municipality, China (Makhan, similarities of this species to D. peculiaris 2010). Its description, however poor (e.g. and noted certain variation in the shapes of the number of body segments was both exo- and endomere in D. silvestris and miscounted as 19, actually being 20) and D. peculiaris. Now that D. silvaticus, too, based on a single female specimen, still joins the peculiaris-group, the following allows to treat it as a distinct species. Even new combination results: Eutrichodesmus though the gonopod structure of E. soesilae silvaticus (Haga, 1968), comb. n. ex remains unknown, in all probability it also Dimorphodesmus Murakami, 1966. belongs in the peculiaris-group. Among further omissions in Golovatch The following key can serve to separate et al. (2009a, b), the status of all of the seven currently known species of Nanocondylodesmus Zhang, 1995 deserves the peculiaris-group: special attention. This genus is known from two species, N. pectinatidentis Zhang, 1995, 1a Transverse rows of metatergal tubercula- the type species from Mt Tianmu, Lin’an tions isostictic. Adult ♀ with 19 body County, Zhejiang Province, China, and N. segments, < 4.5 mm long. The Ryukyus, anisodentus Zhang, 1995, from Mt Wuyi, Japan. ………………….…. E. nodulosus Fujian Province, China (Zhang, 1995a, b), 1b Transverse rows of metatergal both so strongly resembling the members of tuberculations mixostictic. Adult ♀ with the peculiaris-group that we are also 20 body segments, > 4.5 mm long. ...… 2 inclined to place them therein. Thus, they 2a Metatergal tuberculations usually well- also show complete volvation (albeit with differentiated, mid-dorsal ones evidently the overlap pattern becoming typical starting to very strongly enlarged at least on some from segment 4 or 5), two transverse rows segments. Mainland China. …………... 3 of more (N. anisodentus) or less (N. 2b Metatergal tuberculations uniform, like pectinatidentis) strongly differentiated flat bosses. Japan and Taiwan. ……..… 5 tubercles on most of the metaterga, a 3a 1+1 mid-dorsal tubercles only slightly flattened epiproct and, above all, slender higher than others and located only in 2nd gonopods devoid of a distofemoral process, row on segments 4-6(7). ………………. but supplied with subequal and short exo- ……………………….. E. pectinatidentis and endomere, the latter carrying several GOLOVATCH ET AL. — MILLIPEDE FAMILY HAPLODESMIDAE IN TAIWAN 33

3b Mid-dorsal tubercles much higher than description and fine illustrations of E. others and located in both rows at least elegans as presented by Miyosi (1956), on segments 4-16(17). ………………... 4 especially in the light of the variation in 4a Most of tuberculations on collum gonopod structure, both inter- and obliterated, retained only near lateral intraspecific, which is observed within the edge. Mid-dorsal tubercles on penul- peculiaris-group of Eutrichodesmus, we are timate segment low but evident, like a now inclined to reconsider the above small crest. Chongqing Municipality. ….. synonymy and combination. Indeed, like in ………………………..…….. E. soesilae some species of the peculiaris-group, the 4b Almost entire collum covered with gonopods of E. elegans are especially tuberculations. Mid-dorsal tubercles on slender, each devoid of a distofemoral penultimate segment nearly wanting, flat, process, but supplied with a conspicuous not crest-shaped. Fujian Province. ……... hairy pad (apparently marking the orifice of …………………………. E. anisodentus an accessory seminal chamber), as well as 5a Gonopod distofemoral process present, with a spiniform exomere near the pad and a albeit rudimentary, like a small and low long, subfalcate endomere carrying lobe; endomere simple and bifid, numerous spiniform processes. Although exomere stout and lobiform (Fig. 2. A- formally this species cannot be treated as a D). Taiwan. …..….. E. taiwanensis n. sp. member of the peculiaris-group because 5b Gonopod distofemoral process obviously most of its metaterga show three, not two absent; endomere usually more complex, transverse rows of undifferentiated especially so distad; exomere not tuberculations, while the epiproct is not too lobiform. Japan. ……………………… 5 conspicuously flattened, the following 6a Exomere elongate rounded to unciform. nomenclatural changes seem to be ……………………………. E. peculiaris warranted: Eucondylodesmus Miyosi, 1956 6b Exomere stout and unciform. …………... is one more junior subjective synonym of …………………………..… E. silvaticus Eutrichodesmus Silvestri, 1910, syn. n., while Doratodesmus elegans (Miyosi, 1956) Because Rhipidopeltis Miyosi, 1958 has ought to be transferred to Eutrichodesmus, already been synonymized with Prosopo- comb. n. desmus Silvestri, 1910 (Golovatch, et al. The genus Thelodesmus Miyosi, 1951 2009a), the likewise omitted species was first proposed in the family Rhipidopeltis similis Haga, 1968, from Leptodesmidae (Miyosi, 1951), then moved Kyushu, Japan (Haga, 1968), is to be to Vanhoeffeniidae (Miyosi, 1959), cur- transferred to Prosopodesmus, formally rently referred to Pyrgodesmidae (Hoffman, resulting in Prosopodesmus similis (Haga, 1980; Murakami, 1993; Wang and Mauriès, 1968), comb. n. 1996). Its type and only constituent species, In the latest review of Haplodesmidae T. armatus Miyosi, 1951, was originally (Golovatch et al., 2009a), the Japanese described from “Yosihuzi-Mura, Kaminada- genus Eucondylodesmus Miyosi, 1956 was Mati (Ehimé-Ken)”, i.e. Ehime Prefecture, synonymized with, and its sole constituent Shikoku Island, Japan, and has since been species E. elegans Miyosi, 1956 transferred reported also from Urai, Taiwan (Wang, to, Doratodesmus Cook in Cook and 1958; Korsós, 2004) and from Okinawa, the Collins, 1895. Based on the detailed Ryukyus, Japan (Karasawa et al., 2008). 34 TROPICAL NATURAL HISTORY. 10(1), APRIL 2010

The original description alone (Miyosi, the identity of a number of hitherto dubious 1951) unequivocally implies, however, that East Asian micropolydesmidans. Still much Thelodesmus is actually another member of more work is needed to properly assess the Haplodesmidae, again unfortunately over- real diversity and distribution of Haplodes- looked by Golovatch et al. (2009a). Indeed, midae, as well as of all Polydesmida in the the collum in T. armatus is not flabellate to region as a whole and in Taiwan in cover the head from above, i.e. a condition particular. uncharacteristic of Pyrgodesmidae; the In the large genus Eutrichodesmus metaterga show 4-7 transverse rows of which at present encompasses already 32 setigerous tuberculations, both abundant species, the first species group is delimited, tergal setation and mid-dorsally undifferen- the peculiaris-group. All of its seven species tiated tuberculations likewise being are keyed. Previously, only a few pairs of untypical of Pyrgodesmidae; the paraterga especially similar species were revealed are so small that the body is not capable of therein, whereas the bulk remains yet too volvation; paraterga 2 are only slightly difficult to group into some coherent enlarged compared to the subsequent ones. assemblages (Golovatch et al., 2009b). In Both latter features commonly occur both in general, if with our growing knowledge the Haplodesmidae and Pyrgodesmidae. Yet the peculiaris-group is to be promoted into a most remarkable character is gonopod separate genus or subgenus, several genus- conformation, which appears to be that of a group names are readily available. The typical Eutrichodesmus in showing both an oldest is certainly Kylindogaster Verhoeff, evident distofemoral process and a 1939, but we still do not know the male of transverse sulcus at about its midway, as its only constituent species, E. nodulosus well as an apparently subapically termina- (Verhoeff, 1939). Moreover, even its type ting seminal groove. As a result, in addition locality remains obscure. The second to the new family allocation, the following possible option is Dimorphodesmus Mura- nomenclatural changes seem to be kami, 1966, because its type species D. appropriate: Thelodesmus Miyosi, 1951 is peculiaris Murakami, 1966, unlike that of yet one more junior subjective synonym of Kylindogaster, is nicely documented. Eutrichodesmus Silvestri, 1910, syn. n., One must be very critical before accepting while Thelodesmus armatus Miyosi, 1951 is at least some of the original or subsequent also to be transferred as Eutrichodesmus records of dubious forms, such as those by armatus (Miyosi, 1951), comb. n. Verhoeff (1939), Omine (1982), Omine and Ito (1998) and Karasawa et al. (2008) in the CONCLUSION Ryukyus, as well as by Wang (1958) in Taiwan, until a restudy and a proper The first formal record of the millipede description of pertinent material has been family Haplodesmidae in Taiwan has made. brought about not only the description of a new species, but also a new family ACKNOWLEDGEMENTS allocation and numerous new synonyms and combinations. All this not only considerably We are most grateful to the National refines the family’s classification, which Science Council, Taiwan, Republic of China alone is important enough, but also clarifies and to the Russian Academy of Sciences, GOLOVATCH ET AL. — MILLIPEDE FAMILY HAPLODESMIDAE IN TAIWAN 35

Moscow, Russian Federation for the support Hoffman, R.L. 1980 (for 1979). Classification of the rendered to the Taiwanese and Russian Diplopoda. Muséum d’histoire naturelle, Genève, 237 pp. teams, headed by H. W. Chang and S.I. Karasawa, S., Beaulieu, F., Sasaki, T., Bonato, L., Golovatch, respectively, to actively colla- Hagino, Y., Hayashi, M., Itoh, R., Kishimoto, T., borate in our joint ecofaunistic studies on the Nakamura, O., Nomura, S., Nunomura, N., of Taiwan. Zoltán Korsós’ Sakayori, H., Sawada, Y., Suwa, Y., Tanaka, S., Tanabe, T., Tanikawa, A. and Hijii, N. 2008. collecting trip to Taiwan in 2007 was also Bird’s nest ferns as reservoirs of soil supported in part by the National Science biodiversity in a Japanese subtropical rainforest. Council and hosted by I-Min Tso (Tunghai Edaphologia, 83: 11-30. University, Taichung, Taiwan), as well as by Korsós, Z. 2004. Checklist and bibliography of the Hungarian Academy of Sciences and the (Diplopoda) of Taiwan. Collection and Research (NMNS, Taichung), 17: 11-32. Hungarian Scientific Research Fund (OTKA Makhan, D. 2010. Eutrichodesmus soesilae sp. nov., a No. 69235). We are much indebted to Chao- new millipede from Mt. Jinyun, Beibei, Chun Chen and Mei-Jhu Hung (both Chongqing, China (Diplopoda, Polydesmida, National Sun Yat-Sen University, Kaoh- Haplodesmidae). Calodema, 110: 1-5. Mesibov, R.E. 2009. Revision of Agathodesmus siung, Taiwan) for allowing us to study their Silvestri, 1910 (Diplopoda, Polydesmida, collections. Kwen-Shen Lee (National Haplodesmidae). ZooKeys, 12: 87-110. Museum of Natural Science, Taichung) Miyosi, Y. 1951. Beiträge zur Kenntniss japanischer provided type entry numbers. Kirill Makarov Myriopoden. 1. Aufsatz: Ueber eine Gattung von (Moscow, Russia) skillfully took the Leptodesmidae. Zoological Magazine, 60: 149- 150. (In Japanese with German summary). photographs. We are also obliged to Natdanai Miyosi, Y. 1956. Beiträge zur Kenntnis japanischer Likhitrakarn (Chulalongkorn University, Myriopoden. 17. Aufsatz: Über eine neue Gattung Bangkok, Thailand) for his technical von Oniscodesmidae und eine neue Art von assistance. Richard Hoffman (Virginia Monotarsobius. Zoological Magazine, 65: 311- 314. (In Japanese with German summary). Museum of Natural History, Martinsville, Miyosi, Y. 1959. Über japanische Diplopoden. U.S.A.) very kindly edited the text. Special Number. Arachnological Society of Japan, Osaka, 223 pp., 19 pls. (In Japanese). LITERATURE CITED Murakami, Y. 1966. Postembryonic development of the common Myriapoda of Japan XXI. A new Golovatch, S.I. 2003. A review of the volvatory genus of the family Oniscodesmidae and a new Polydesmida, with special reference to the species of the genus Archandrodesmus patterns of volvation (Diplopoda). African (Cryptodesmidae). Zoological Magazine, 75: 30- Invertebrates, 44: 39-60. 33. (In Japanese with English summary). Golovatch, S.I., Geoffroy, J.-J., Mauriès, J.-P. and Murakami, Y. 1993. Diplopoda, Pauropoda, VandenSpiegel, D. 2009a. Review of the Symphyla. A list of Japanese species. Inverte- millipede family Haplodesmidae Cook, 1895, with brates 1. Shizen-Kankyô-Kenkyû Center. Envi- descriptions of some new or poorly-known ronmental Agency Japan, Tokyo: 95-106. (In species (Diplopoda, Polydesmida). ZooKeys, 7: 1- Japanese). 53. Omine, T. 1982. Preliminary survey of soil fauna in Golovatch, S.I., Geoffroy, J.-J., Mauriès J.-P. and the center of Iriomote-Island (riverside of the VandenSpiegel, D. 2009b. Review of the Itarashiki and Urauchi rivers). Main collection of millipede genus Eutrichodesmus Silvestri, 1910 Cryptostigmata, Myriapoda and Formicidae. (Diplopoda, Polydesmida, Haplodesmidae), with Okinawa Daigaku Kiyo (Journal of Okinawa descriptions of new species. ZooKeys, 12: 1-46. University), 2: 83-139. (In Japanese). Haga, A. 1968. Japanese Millipedes. Private Omine, T. and Ito, Y. 1998. Abundance and diversity publication, 11 pp., 6 pls. (In Japanese). of soil macrofauna of forests of Yanbaru, northern montane part of Okinawa Island, with special 36 TROPICAL NATURAL HISTORY. 10(1), APRIL 2010

reference to removal of undergrowth. Okidai Zhang, C.-Z. 1995a. Small Myriapoda in soil from Ronso (Journal of Okinawa University), 15: 131- China II. A new genus and species of the family 159. Doratodesmidae from Zhejiang Province Verhoeff, K.W. 1939. Zur Kenntnis ostasiatischer (Diplopoda: Polydesmida). Acta Zootaxonomica Diplopoden IV. Zoologischer Anzeiger, 127: 273- Sinica, 20: 411-415. (In Chinese with English 285. summary). Wang, Y.M. 1958. Serica 1i: On Diplopoda from Zhang, C.-Z. 1995b. Small Myriapoda in soil from Taiwan with a new strongylosomid. Quarterly China III. A new species of the milliped genus Journal of the Taiwan Museum, 11: 340-344. Nanocondylodesmus Zhang (Diplopoda: Polydes- Wang, D. and J.-P. Mauriès. 1996. Review and mida: Doratodesmidae). Acta Zootaxonomica perspective of study on myriapodology of China. Sinica, 20: 416-419. (In Chinese with English Mémoires du Muséum National d’Histoire summary). Naturelle, 169: 81-99.