Research Article |
Corresponding author: Sergei I. Golovatch ( sgolovatch@yandex.ru ) Academic editor: Robert Mesibov
© 2017 Sergei I. Golovatch, Jean-Jacques Geoffroy, Nesrine Akkari.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Golovatch SI, Geoffroy J-J, Akkari N (2017) Revision of the Vietnamese millipede genus Annamina Attems, 1937, with descriptions of three new species (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 669: 1-18. https://doi.org/10.3897/zookeys.669.12561
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The hitherto monotypic diplopod genus Annamina contains now four species, including the revised type-species A. xanthoptera Attems, 1937, as well as A. attemsi sp. n., A. irinae sp. n. and A. mikhaljovae sp. n., all from central or southern Vietnam. The genus is rediagnosed and a key to its constituent species given.
Diplopoda , Paradoxosomatidae , Annamina , taxonomy, new species, Vietnam
The monotypic millipede genus Annamina Attems, 1937, and its type species A. xanthoptera Attems, 1937, were described twice nearly simultaneously from Vietnam, first in a global review of the family Paradoxosomatidae (= Strongylosomatidae) (
The genus Annamina belongs to the tribe Sulciferini Attems, 1898 and differs from the other, mostly Asian genera by a remarkably short flagelliform solenomere and an even shorter, chitinous, tooth-shaped solenophore devoid of membranous elements (
Recently, three new Annamina species were discovered in southern and central Vietnam. One of these, A. attemsi sp. n., was incidentally found by NA when documenting the type material of A. xanthoptera in the Naturhistorisches Museum Wien (
New material that contained two new species of Annamina was taken in 2015 and 2016 during field trips of I. I. Semenyuk (Moscow, Russia) to southern Vietnam in the framework of the research activities of the Joint Russia-Vietnam Tropical Centre. The types of both are deposited in the Zoological Museum of the Moscow State University, Russia (
The
Several paralectotypes of A. xanthoptera, preserved in 70% alcohol, are housed in the Muséum national d’Histoire naturelle, Paris (
Medium-sized (ca 2–3 cm long) Sulciferini with 20 body segments, distinct, thin and mostly subhorizontal paraterga, evident transverse sulci on metaterga 5–17(18), and a very high, tongue-shaped, setose, subtruncate lobe between ♂ coxae 4.
Gonopod mostly stout, prefemoral (= densely setose) region small, much shorter than femorite, the latter usually with evident, sometimes hyline, mesal and/or ventral lobes, clearly set off by sulci from both pre- and postfemoral parts; seminal groove mostly dorsal, not mesal, running onto a short flagelliform solenomere on mesal face near distal (= postfemoral) sulcus; acropodite consisting of a prominent central spine sometimes flanked by a mesal and/or a lateral process/outgrowth and carrying parabasally or near midway an inconspicuous, short, dentiform, ventral solenophore devoid of membranous elements and subtending the distal part of solenomere.
Annamina xanthoptera Attems, 1937
A. attemsi sp. n., A. irinae sp. n. and A. mikhaljovae sp. n.
Lectotype designation was necessary so that the species is based on a complete male that fully matches the original description of A. xanthoptera by
Differs from other members of the genus primarily by showing both the median lobe and the lateral process of the gonopod telopodite strongly microdenticulate-serrate. See also Key below.
Measurements (mm): Males (both
General coloration after many years of preservation in alcohol apparently somewhat faded, rather uniformly light to castaneous brown, without a distinct pattern, sides lighter; telson, legs and ventral parts light brown to yellowish (Fig.
Antennae long, slender and moderately clavate, slightly extending back behind segment 3 (♂) (Fig.
Sterna flat, sparsely setose, cross-impressions faint, without modifications other than a prominent, very high, narrow, triangular, truncate lobe between ♂ coxae 4 (as in Fig.
Gonopods (Figs
Annamina xanthoptera Attems, 1937, ♂ paralectotype (
Annamina xanthoptera Attems, 1937, ♂ paralectotype (
Annamina xanthoptera Attems, 1937, ♂ paralectotype (
A complete catalogue of references to A. xanthoptera is available in
Holotype ♂,
Differs from other species of the genus primarily by an unusually slender telopodite of the gonopod which is only supplied with an apical process. See also Key below.
Honours the famous Austrian myriapodologist Carl Attems (1868–1952), one of the most prominent taxonomists of Diplopoda of his time.
Measurements (mm): Body length ca 18.7 (holotype) or 21.6 (♂ paratype), width of midbody prozonae 1.2 (holotype) or 1.3 (♂ paratype), width of midbody metazonae 1.8 (holotype) or 2.0 (♂ paratype).
General coloration after many years of preservation in alcohol light, almost whitish to yellowish brown, sides, telson, legs and ventral parts pale whitish (Fig.
All other characters (see Figs
Caudal corner of paraterga dentiform and acute-angled starting with segment 4, drawn behind rear tergal margin starting with segment 11 (♂); a second lateral denticle on paraterga completely absent from segments 18 and 19. Transverse sulci fully developed on metaterga 5–17, deep, beaded at bottom and almost reaching the bases of paraterga, weaker on segment 18, absent from 19th.
Gonopods (Figs
Annamina attemsi sp. n., ♂ paratype (
Holotype ♂,
Honours Irina Semenyuk, the collector.
Differs from other species of the genus primarily by the presence of a small ventral lobe and a large mesal lobe on the gonopod femorite, coupled with a small, simple, mesal process, a long, spiniform, lateral process and a prominent, simple, unciform, acuminate, apical process in the postfemoral portion of the gonopod. See also Key below.
Measurements (mm): Body length of all types ca 24, width of midbody pro- and metazona 1.5 and 1.9, respectively.
General coloration in alcohol light brownish to brown, but with a characteristic pattern of a vague, lighter, subtriangular, central spot on each postcollum metatergum flanked on each side by marbled brown patches fused into a complete transverse band in anterior 1/3; sides mostly brown to light brown, lighter closer to coxae; strictures between pro- and metazonae, paraterga both dorsally and ventrally, telson, legs and venter pale yellowish to whitish (Fig.
All characters (see Figs
In width, segments 5–16 > head > 2 > collum = 3 = 4 (♂); body gradually tapering towards telson on segments 17–19 (Fig.
Gonopods (Figs
Annamina irinae sp. n., ♂ paratype (
Holotype ♂,
Honours Elena Mikhaljova, a prominent specialist in the systematics of Asian Diplopoda.
Differs from other species of the genus primarily by the presence of a small ventral lobule and a large mesal lobe on the gonopod femorite, coupled with, much like in A. irinae sp. n., a small, simple, mesal process, a similarly short, but spiniform, clearly serrate lateral process and a prominent, lobe-shaped, apical process in the postfemoral portion of the gonopod. See also Key below.
Measurements (mm): Length ca 22 mm, width of midbody pro- and metazonae 1.8 and 2.7 mm, respectively. Coloration uniformly light brownish to yellow-brown, only antennomeres 6 and 7 contrasting dark brown; tegument largely thin and translucent (Fig.
All characters (see Figs
In width, collum = 3 = 4 < segment 2 < head < 5–16 (♂); thereafter body gradually tapering towards telson. Caudolateral corner of paraterga subrectangular until segment 8, thereafter increasingly well drawn caudad, but always remaining narrowly rounded, clearly projecting behind rear tergal margin only on segments 17–19 (Fig.
Gonopods (Figs
Annamina mikhaljovae sp. n., ♂ holotype (
Even with such a considerable increase in species diversity as described above, Annamina remains a well-defined and quite homogeneous group within the paradoxosomatid tribe Sulciferini. The genus is remarkably uniform in most of the characters of its constituent species, both somatic and gonopodal, as well as in distribution which is confined to south-central and central Vietnam. Annamina is unique amongst the contribal genera in the high, well-developed, laterally mostly monodentate and caudally largely triangular (but never pointed) paraterga; the strongly ribbed strictures between pro- and metazonae; the microgranulate sides of metazonae; the inconspicuous, nearly missing pleurosternal carinae; the uniformly roundly subtriangular hypoproct; the unusually high, subtriangular, setose and apically truncate sternal lobe between ♂ coxae 4; the absence both of adenostyles and ventral brushes on remarkably long and slender legs; and, above all, the special conformation of the gonopodal telopodite.
It is the latter that provides most of the characters useful for a confident separation of species in Annamina (see Key below). The gonotelopodite always shows an enlarged basal part of the femorite which is clearly flattened dorsoventrally. The femorite is set off from the acropodite by a distinct subtransverse sulcus or cingulum (su) that marks a postfemoral region more distally, a trait that, together with the presence of a number of femoral and postfemoral outgrowths or processes, allows the placement of the genus in Sulciferini (cf.
Based on gonopodal structure alone, this latter species seems to be the most disjunct among congeners (see its diagnosis above and Key below). Since it seems to co-occur with A. xanthoptera near Danang, this observation agrees well with the general wisdom that two sympatric or even syntopic congeners tend to differ more strongly than others.
The above outline of the diversity of and variations in gonopodal characters in Annamina spp. helps us not only to better redefine the genus against the other Sulciferini, but also to key all of its four presently known species. Because the millipede fauna of Vietnam is the richest in Indochina, but still quite poorly known, it seems very likely that further Annamina species will be found in the future.
1 | Postfemoral region of gonopod with a distinct, slender, slightly curved, mesal process (mp); apical process (a) broad and lobe-shaped (Fig. |
A. mikhaljovae sp. n. |
– | Mesal process either very short or absent | 2 |
2 | Mesal process (mp) very short, mesal lobe (ml) of femorite ear-shaped (Fig. |
A. irinae sp. n. |
– | Mesal process absent | 3 |
3 | Femorite slender, ventral lobe (vl) hypertrophied, postfemoral lateral process (lp) rudimentary (Figs |
A. attemsi sp. n. |
– | Femorite stout, ventral lobe (vl) small and inconspicuous, postfemoral lateral process (lp) very strong and serrate (Figs |
A. xanthoptera |
The authors are most grateful to Irina Semenyuk (Moscow) for giving us her material for study, to Kirill Makarov (Moscow) who skillfully took all pictures of the