Corresponding author: Anh D. Nguyen (
Academic editor: M. Short
The East Asian millipede genus
The millipede genus
Following several discussions on the genus in the past (
Fresh material of
Line drawings were made by using an Olympus SZX10 (ADN), and a Leica M125 (ZK) stereo microscope with drawing tube attached. SEM images were made by using a Hitachi S4800 scanning electron microscope. Colour photographs were taken by ZK using a Nikon D90 digital camera with macro lens and Leica microscope photo tube attached. The distribution map was generated using the software DIVA-GIS version 7.0.
Gonofemorite strongly curved, distal part somewhat swollen and membraneous. Postfemoral region demarcated from femorite by obvious cingulum, and bent continuously forming almost a complete circle with femorite. Postfemoral regions consisting of a thick, strong and free solenomere, and an extremely short solenophore (= tibiotarsus).
This genus is relatively close to the genus
Holotype male, in fragments – only 11 segments in 5 pieces – with segments around gonopods missing, Reg.-Nr. ZSMA20052252, and two slides with gonopods, Reg.-Nr. ZSMA20035204, and legpairs 1–7, Reg.-Nr. ZSMA20035205 (all BSCZ).
New records: 2 males, 2 females, Japan, Ryukyu Archipelago, Miyako-jima Island, Rinko-abu (cave), 21 August 1979; 8 males, 2 females, Japan, Ryukyu Archipelago, Miyako-jima Island, Fukumine-no-kara (cave), 25 August 1979; 1 male, 1 female, 1 juv., 1 fragment, Japan, Ryukyu Archipelago, Miyako-jima Island, Nishibe zuzaga (cave), 26 August 1979, all leg. M. Shimojana (in the collection of M. Shimojana, Okinawa).
Japan, Ryukyu Archipelago, Miyako-jima island.
Although after the description of
Holotype male, NSMT-My 358, “Japan, Okinawa Pref., Is. Ishigaki-jima, Ibaruma, Sabichi-go Cave, 31 July 1973, coll. by S. Uéno” (NMNS).
New records: 2 males, 1 female, Japan, Ryukyu Archipelago, Yaeyama Island Group, Ishigaki-jima Island, Banna-dake, secondary forest,
After studying the type and freshly collected specimens, it became clear that the species
This genus,
A feminine noun to emphasize the simple gonopod conformation.
In fact, the solenophore of
Comparison of genera of
No. | Characters |
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1 | Paraterga | Well-developed | Well-developed | Modestly-developed | Poorly-developed | Poorly-developed | Poorly-developed |
2 | Sternal process between coxae 4 | Missing | Missing | Two separate cones | A single or bifid process | A single process | Linguiform |
Gonopod conformation | |||||||
3 | Coxa | Long, cylindrical but not thick | Long, thick and subcylindrical | Thick and subcylindrical | Long and subcylindrical | Long and subcylindrical | Long and subcylindrical |
4 | Femorite | Long and slender, with a rugose membranous lamina at distal part of lateral side | Especially long, slender, simple without any outgrowths, however, sometimes, with a lamina at mesal side | Especially long, but simple and slender, devoid of any lobes or lamina. | Long, but broadened parabasally at dorsal side, without any lamina or processes | Long, slender and erected, a little bit broadened distally, devoid of any modifications | Long, slightly curved, not broadened at base |
5 | Postfemoral part | Demarcated by cingulum | Demarcated by cingulum | Demarcated by cingulum | Demarcated by obvious sulcus at lateral side | Set off by oblique sulcus at lateral side, apical part with a big tooth. | absent |
6 | Solenomere | Long, thick and simple, not sheathed by solenophore | Long, thick and simple, not sheathed by solenophore | Thick, but slender, and strongly curved downwards | Flagiliform, almost completely sheathed by solenophore, sometimes, only tip exposed | Long, slender and coiled, subfiliform | Long, flagiliform, almost sheathed by solenophore |
7 | Solenophore | Very short, simple, without any additional processes | As long as solenomere, with a basal dentiform outgrowth | Completely missing | Base with a small, obvious lobe. Tip with bifid lobe | Laminate, coiled, and as long as solenomere | Long and coiled, with a process at base |
3 males, 2 females, (IEBR-166), Vietnam, Lam Dong prov., Bi Doup-Nui Ba National Park, corn field, 1400m a.s.l., pitfall traps, 2–9 April 2008, leg. Anh. D. Nguyen; 1 male, 1 female, 1 juvenile, (IEBR-167), same locality, grasslands, 1400m a.s.l., pitfall traps, 2–9 April 2008, leg. Anh. D. Nguyen; 1 female, (IEBR-168), same locality, bushes near stream, 1400m a.s.l., pitfall traps, 25 April 2008, leg. Anh. D. Nguyen; 4 males, 2 females, (IEBR-169), same locality, evergreen forest, 1800m a.s.l., 25 March–23 April 2008, leg. Anh. D. Nguyen;1 male (IEBR-125), Vietnam, Khanh Hoa Prov., Hon Ba Mts., 1300–1500m a.s.l., primary forest, 15–24 April 2006, leg. Anh. D. Nguyen; 1 male, 1 female, (HNHM), same data as sample IEBR-125.
Surface of metaterga general fine in posterior part, but with small oblique or longitudinal rugulose in anteriormost part. Stricture dividing pro- and metazona deep, obvious and beaded. Prozona surface shagreened with fine microgranulation
Transverse sulcus on metaterga starting from somite 5 and more evident on subsequent somites. Metaterga with a row of 2+2 setae in pre-sulcus part. Axial line vague.
Paraterga not well-developed, small, look like small keels in poreless segments, but more developed in pore-bearing somites. Ozopore located on lateral side, near tip of angular paraterga of segments 5, 7, 9–10, 12–13 and 15–19.
Pleura shagreened with fine microgranulation. Pleurosternal carinae rather well-developed in anteriormost segments, gradually decreasing posteriorly.
Epiproct truncated and curved down ventrad, with 4 strong setae on tip. Anal valves sub-semicircular with 1+1 long setae, and a deep emargination inbetween. Hypoproct trapeziform, with 1+1 setae.
Prostatic groove runs mesally along femorite, distolaterad, and turns to lateral side, then running mesally, and ending at tip of solenomere.
Male gonopods
Midbody (11th-13th) segments, dorsal view:
Habitus of millipedes:
Distribution of three species
Vietnam, Lam Dong province, Bi Doup-Nui Ba National Park; Khanh Hoa province, Hon Ba Mountain; Laos PDR, Xieng Khoang
New material does not much differ from Attems’s description. Only minor difference is the presence of two separate cones between coxae 4 instead of only one small conal process in Attems’s description. Recently,
1 | Paraterga very weak, body looks almost cylindrical, colour uniformly light brown-yellowish. Gonofemorite very long and slender, without any modifications or processes. Postfemoral region consisting only a simple, strong, long solenomere. Solenophore totally absent ( |
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– | Paraterga well-developed with strong, triangular, posterio-lateral processes. Dorsal metazona colouration divided into dark brown anterior and much lighter posterior half, transversely separated by a deep sulcus | 2 |
2 | Gonofemorite large, slender and strongly concave, with a lamina on the mesal side. Postfemoral region with a long, large solenomere and a shorter solenophore, the latter basally with two processes, a laminar mesal ( |
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– | Gonofemorite only slightly curved, with a weak dorsal lamina, distal part swollen and membraneous. Postfemoral region consisting a thick, strong, free solenomere, and a short, somewhat hidden solenophore ( |
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In the key above,
As a result of our character comparisons, the taxonomic status of the following three species:
We would like to express our deepest thanks to the Vietnamese–Russian Tropical Centre (Vietnam), and the Bi Doup-Nui Ba National Park for supporting the first author during his field expeditions in Vietnam. Special thanks also to the AusAID program of the Australian government for providing financial support through its International Seminar Support Scheme for the first author to participate and present the paper at the 15th International Congress of Myriapodology.
The second author would like to express his sincere gratitude to Prof. H. Ota and M. Toda (Tropical Biosphere Research Center, University of the Ryukyus, Okinawa) for inviting him to study millipedes in the Ryukyus. His study is also supported by the Hungarian Research Fund (OTKA No. 69235). H. Ono (National Museum of Nature and Science, Tokyo) and S. Friedrich (Bavarian State Collection of Zoology, Munich) are deeply acknowledged for arranging the loans of appropriate type material. We also thank M. Shimojana (Urasoe, Okinawa) for letting us study his valuable unidentified material collected in Miyako-jima Island, and Y. Nakamura (Okinawa) for his help in the field and in literature search.