Research Article |
Corresponding author: Sergei I. Golovatch ( sgolovatch@yandex.ru ) Corresponding author: Somsak Panha ( somsak.pan@chula.ac.th ) Academic editor: Didier Vanden Spiegel
© 2016 Natdanai Likhitrakarn, Sergei I. Golovatch, Somsak Panha.
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:
Likhitrakarn N, Golovatch SI, Panha S (2016) Review of the Southeast Asian millipede genus Antheromorpha Jeekel, 1968 (Diplopoda, Polydesmida, Paradoxosomatidae). ZooKeys 571: 21-57. https://doi.org/10.3897/zookeys.571.7566
|
The genus Antheromorpha is redefined and shown to comprise 11 valid species: A. miranda (Pocock, 1895), A. bistriata (Pocock, 1895), A. comotti (Pocock, 1895), A. festiva (Brölemann, 1896), A. harpaga (Attems, 1937), A. mediovirgata (Carl, 1941), A. minlana (Pocock, 1895), A. pardalis (Pocock, 1895), A. paviei (Brölemann, 1896), comb. n., A. rosea Golovatch, 2013 and A. uncinata (Attems, 1931). Three new synonymies are proposed: A. bivittata (Pocock, 1895) and A. melanopleuris (Pocock, 1895) are synonymized under A. miranda (Pocock, 1895), and A. orophila (Carl, 1941) under A. comotti (Pocock, 1895). Detailed descriptions and illustrations of fresh material from Thailand and Malaysia are given, especially regarding colour patterns which appear to be crucial for accurate species identifications. Two Antheromorpha species proposed by Attems are redescribed, based on type material. The genus is rediagnosed and a key and a distribution map are also provided. At least in Thailand, adult A. rosea have been found to occur every year only for one or two weeks in September or October, disappearing thereafter.
Review, Paradoxosomatidae , taxonomy, new synonymy, key, distribution, map
The Southeast Asian millipede genus Antheromorpha Jeekel, 1968 is currently known to comprise 13 medium-sized to very large species showing moderately developed to very prominent paraterga and, above all, unlike the other, numerous genera of the basically Oriental tribe Orthomorphini Brölemann, 1916 to which it belongs, a very deeply bifid gonopod tip (
Because of the remarkably bright colour patterns and large bodies, unidentified Antheromorpha species have repeatedly been reported throughout Thailand (http://siamensis.org/webboard/topic/35582#comment-34142; http://thailandwildlife.photoshelter.com/gallery-image/Other-Arthropods/G0000OdCpTbz8ENY/I0000PMJm_Wnsl8E; http://www.projectnoah.org/spottings/10657453; https://www.flickr.com/photos/lennyworthington/sets/72157628909951579/). Moreover, one of the species shows swarming behaviour (http://www.manager.co.th/Local/ViewNews.aspx?NewsID=9490000084506).
The only attempt at reviewing Antheromorpha and outlining its diagnosis (
A. bistriata (Pocock, 1895), A. bivittata (Pocock, 1895), A. festiva (Brölemann, 1896), A. mediovirgata (Carl, 1941), A. melanopleuris (Pocock, 1895), A. miranda (Pocock, 1895).
A. comotti (Pocock, 1895), A. orophila (Carl, 1941).
A. uncinata (Attems, 1931).
A. harpaga (Attems, 1937).
A. minlana (Pocock, 1895).
A. pardalis (Pocock, 1895).
The only subsequent addition to the list seems to be A. rosea Golovatch, 2013, the first species of the genus to be reported from southern China, yet not placed into any of the species groups (
The present paper provides an updated review of Antheromorpha, based on abundant new samples from Thailand and Malaysia. In addition, type material of two species of
New material was taken throughout Thailand and from Malaysia between 2006 and 2015 by SP and members of the Animal Systematics Research Unit, Chulalongkorn University. Animals, both live and alcohol material, were photographed in the laboratory. Specimens were preserved in 75% ethanol and morphological investigations were carried out in the laboratory using an Olympus stereomicroscope. Scanning electron micrographs (SEM) of gonopods coated with gold were taken using a JEOL, JSM–5410 LV microscope, and the gonopods were returned to alcohol after examination. Digital images of the specimens were taken in the laboratory and assembled using the “CellD” automontage software of the Olympus Soft Imaging Solution GmbH package. In addition, line drawings of gonopods were prepared. Type material of two Attemsian species of Antheromorpha from Thailand and Vietnam, housed in the
Collecting sites were located by GPS using the WGS84 datum.
In the synonymy sections, D stands for the original description or subsequent descriptive notes or appearance in a key, R for subsequent record or records, whereas M for a mere mention.
Brachytropis Silvestri, 1896: 198 (D) (preoccupied).
Brachytropis –
Antheromorpha Jeekel, 1968: 57 (M).
Antheromorpha –
Body medium-sized to very large (ca 19–44.5 mm long, ca 2.3–6.1 mm wide), composed of 18 podous and one apodous ring, plus telson. Paraterga from moderately to very strongly developed. Sterna without modifications. Sternal lobe or cone(s) between male coxae 4 present. Pleurosternal carinae usually well-developed. First pair of male legs without femoral adenostyles. Legs without particular modifications except for at least some ♂ legs bearing ventral brushes on tarsi, sometimes also on tibiae.
Gonopods long and rather slender; coxa slightly curved and long, with several setae distoventrally; prefemoral (= setose) part of telopodite short to very short, 1/3–1/4 as long as acropodite (= remaining part of telopodite); femorite slender to rather stout, straight to evidently curved, sometimes enlarged distally, with a strong distolateral sulcus (s) demarcating a “postfemoral” part; seminal groove running entirely mesally along femorite, the latter devoid of processes. Solenophore (sph) (= tibiotarsus) and solenomere relatively short to rather long; sph curved rather strongly caudad, consisting of a well-developed lamina medialis (lm) and a rather small lamina lateralis (ll); lm about halfway bearing a well-developed process d; sph usually bilobate to bifid, with a mesal process (m, or the end part of lm) and a ventral process (v, or the end part of ll), both supporting a long and flagelliform solenomere (sl).
Orthomorpha miranda Pocock, 1895, by direct substitution.
A. bivittata (Pocock, 1895), A. comotti (Pocock, 1895), A. festiva (Brölemann, 1896), A. harpaga (Attems, 1937), A. mediovirgata (Carl, 1941), A. minlana (Pocock, 1895), A. orophila (Carl, 1941), A. pardalis (Pocock, 1895), A. paviei (Brölemann, 1896), comb. n., A. rosea Golovatch, 2013, A. uncinata (Attems, 1931).
Brachytropis Silvestri, 1896, was originally established to distinguish several species of Orthomorpha Bollman, 1893 which occurred in Myanmar and Indochina (
Orthomorpha miranda Pocock, 1895: 812 (D).
Orthomorpha miranda –
Orthomorpha (Orthomorpha) miranda –
“Orthomorpha” miranda –
Brachytropis miranda –
Antheromorpha miranda –
Orthomorpha bivittata Pocock, 1895: 814 (D), syn. n.
Orthomorpha bivittata –
“Orthomorpha” bivittata –
Antheromorpha bivittata –
Orthomorpha melanopleuris Pocock, 1895: 813 (D), syn. n.
Orthomorpha melanopleuris –
“Orthomorpha” melanopleuris Jeekel, 1963: 269 (M).
Antheromorpha melanopleuris –
This species was described from Yangon (Rangoon) (the type locality); Tharrwaddy, Bago Division; Palon in Pegu (state/region); Thigian, upper Irrawaddy and Minhla, Myanmar (
In addition, according to H. Enghoff (in litt.), the
Based solely on
Antheromorpha miranda (Pocock, 1895), ♂ (A–D), ♀ (F) non-type material from Palon in Pegu, ♂ non-type material from Minhla (E, G), ♀ lectotype of Orthomorpha bivittata Pocock, 1895 (H, I), ♀ lectotype of Orthomorpha melanopleuris Pocock, 1895 (J, K). A anterior part of body, lateral view B, C, G, H, I , J, K segments 10 and 11, dorsal, lateral, dorsal, dorsal, lateral, dorsal and lateral views, respectively D, E right gonopod, mesal view F segment 10, dorsal view (after
Orthomorpha bistriata Pocock, 1895: 814 (D).
Orthomorpha bistriata –
“Orthomorpha” bistriata –
Antheromorpha bistriata –
Antheromorpha bistriata –
This species was described from Bhamo, Myanmar (
Antheromorpha bistriata (Pocock, 1895), ♂ holotype (A–C); Antheromorpha comotti (Pocock, 1895), ♀ holotype of Orthomorpha comotti Pocock, 1895 (D–F), ♂ holotype of Orthomorpha (Orthomorpha) orophila Carl, 1941 (G–I). A, H, I right gonopod, mesal, lateral and submesal views, respectively B, C, E, F segments 10 and 11, dorsal, lateral, dorsal and lateral views, respectively D anterior part of body, lateral view G sternal cones between coxae 4, subcaudal view view (after
Orthomorpha comotti Pocock, 1895: 814 (D).
Orthomorpha Comotti –
Orthomorpha comotti –
“Orthomorpha” comotti –
Antheromorpha comotti –
Orthomorpha (Orthomorpha) orophila Carl, 1941: 361 (D), syn. n.
“Orthomorpha” orophila –
Antheromorpha orophila –
This species was described and still remains known only from Minhla, Myanmar (
Orthomorpha (Orthomorpha) mediovirgata Carl, 1941: 364 (D).
“Orthomorpha” mediovirgata –
Antheromorpha mediovirgata –
This species was described and still remains known only from the northern Chin Hills, Myanmar (
Antheromorpha mediovirgata (Carl, 1941), ♂ holotype (A, B); Antheromorpha minlana (Pocock, 1895), ♂ holotype (C); Antheromorpha pardalis (Pocock, 1895), ♀ holotype (D, E). A, B right gonopod, mesal and lateral views, respectively C left gonopod, mesal view D, E segments 10 and 11, dorsal and lateral views, respectively (after
Orthomorpha minlana Pocock, 1895: 816 (D).
Orthomorpha minlana –
Orthomorpha (Orthomorpha) minlana –
“Orthomorpha” minlana –
Orthomorpha miuhlana (sic!) –
Antheromorpha minlana –
This species was described and still remains known only from Minhla, Tharrawaddy District, Myanmar (
An indefinite number of ♂ and ♀ syntypes of A. minlana must be deposited in the London Museum, UK (
Orthomorpha pardalis Pocock, 1895: 815 (D).
Orthomorpha pardalis –
“Orthomorpha” pardalis –
Antheromorpha pardalis –
This species was described and still remains known only from a single ♀, the holotype which comes from Palon in Pegu, Myanmar (
Orthomorpha Paviei Brölemann, 1896: 1 (D).
Orthomorpha Paviei –
?Prionopeltis Paviei –
Pratinus Paviei –
Orthomorpha paviei –
Redescribed based on new material from Khone Phapen Waterfall, Laos (
Antheromorpha paviei (Brölemann, 1896), ♂ from Laos. A habitus, live coloration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F–H posterior part of body, dorsal, ventral and lateral views, respectively I, J sternal cones between coxae 4, subcaudal and sublateral views, respectively (After
Orthomorpha (Orthomorpha) uncinata Attems, 1931: 117 (D).
Orthomorpha uncinata –
Orthomorpha (Orthomorpha) uncinata –
“Orthomorpha” uncinata –
Antheromorpha uncinata –
♂ of Orthomorpha uncinata (
1 ♀ (
Lectotype designation proposed herewith is necessary to ensure the species is based on a male.
4 ♂, 9 ♀ (
Length 30.0–42.5 (♂) or 34.0–44.5 mm (♀), width of midbody pro- and metazonae 2.6–4.0 and 2.9–4.4 mm (♂), 3.9–5.3 and 4.4–6.1 mm (♀), respectively.
Coloration of live animals red, orange to yellow (Fig.
Antheromorpha uncinata (Attems, 1931), ♂ lectotype. A, B anterior part of body, dorsal and lateral views, respectively C, D segments 10 and 11, dorsal and lateral views, respectively E–G posterior part of body, lateral, dorsal and ventral views, respectively H, I sternal cones between coxae 4, subcaudal and sublateral views, respectively.
Antheromorpha uncinata (Attems, 1931), ♂ from Wat Tham Phromlok Khaoyai (A–I), ♂ from Tham Prakaiphet (J), ♂ from Tham Phet Phothong (K), male from Thap Sakae (L). A, B anterior part of body, dorsal and lateral views, respectively C, D, J–L segments 10 and 11, dorsal, lateral, dorsal, dorsal and dorsal views, respectively E–G posterior part of body, lateral, dorsal and ventral views, respectively H, I sternal cones between coxae 4, subcaudal and sublateral views, respectively.
Clypeolabral region densely, vertex sparsely setose; epicranial suture distinct. Antennae long (Figs
Tegument smooth and shining, prozonae finely shagreened, metaterga leathery and faintly rugulose, surface below paraterga finely microgranulate. Postcollum metaterga with two transverse rows of setae traceable at least as insertion points when setae broken off: 2+2 in anterior (pre-sulcus) and 3+3 in posterior (post-sulcus) row. Tergal setae long, strong, slender, about 1/3 of metatergal length. Axial line visible only on metaterga. Paraterga very strongly developed (Figs
Sterna sparsely setose, without modifications; a large, central, setose cone between ♂ coxae 4 (Figs
Gonopods (Figs
This species was described from Muok Lek, Thailand (
Antheromorpha rosea Golovatch, 2013a: 23 (D).
Antheromorpha rosea –
5 ♂, 9 ♀ (
Length 33.5–38.0 (♂) or 34.0–44.5 mm (♀), width of midbody pro- and metazonae 2.6–3.5 and 4.4–5.0 mm (♂), 3.2–4.2 and 4.9–5.8 mm (♀), respectively.
Coloration of live animals pinkish (Fig.
Antheromorpha rosea Golovatch, 2013, ♂ from Queen Sirikit Botanic Garden. A habitus, live coloration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F–H posterior part of body, lateral, dorsal and ventral views, respectively I, J sternal cones between coxae 4, subcaudal and sublateral views, respectively.
Antennae long (Fig.
The available descriptions (
At least in Thailand, adult A. rosea have been found to occur every year only for one or two weeks in September or October, disappearing thereafter.
Orthomorpha festiva Brölemann, 1896: 1 (D).
Orthomorpha festiva –
Orthomorpha (Orthomorpha) festiva –
“Orthomorpha” festiva –
Antheromorpha festiva –
3 ♂ (
Length 23.0–29.5 (♂) or 26.0–34.5 mm (♀), width of midbody pro- and metazonae 1.8–2.5 and 2.9–3.7 mm (♂), 2.7–3.1 and 3.6–4.4 mm (♀), respectively (vs length 28–30 mm, as given in the available descriptions (
Coloration of live animals dark red to red-brownish, with contrasting light red to pale pinkish paraterga and epiproct; a complete inverted V-shaped line on collum, a pair of parallel oblique bands on metazonae and a pair of parallel bands on prozonae of following segments; legs and venter brownish to pale brown; coloration of alcohol material after one year of preservation faded to castaneous or pale brown; paraterga, epiproct and parallel bands faded to pale pinkish or pale yellow, legs and venter paler brown to yellowish (Fig.
Antheromorpha festiva (Brölemann, 1896), ♂ from Sungai Bekok (A), ♂ from Kampung S. Ramasamy (B–J). A habitus, live coloration B, C anterior part of body, dorsal and lateral views, respectively D, E segments 10 and 11, dorsal and lateral views, respectively F–H posterior part of body, lateral, dorsal and ventral views, respectively I, J sternal cones between coxae 4, subcaudal and sublateral views, respectively.
Clypeolabral region sparsely setose, epicranial suture distinct. Antennae short (Fig.
Tegument smooth and shining, prozonae very finely shagreened, metazonae smooth and delicately rugulose; surface below paraterga finely microgranulate. Postcollum metaterga with two transverse rows of setae, these being always abraded and traceable as insertion points: 2+2 in anterior (pre-sulcus) row, 3+3 in posterior (post-sulcus) one. Tergal setae simple, slender, about 1/3 of metatergal length. Axial line visible both on pro- and metazonae, starting with collum. Paraterga very strongly developed (Fig.
Sterna sparsely setose, without modifications; a high paramedian pair of evident, high, nearly pointed, fully separated, setose cones between ♂ coxae 4 (Fig.
Gonopods (Figs
The new specimens fully agree with the most detailed and beautifully illustrated redescription of the species as given by
The above samples thus derive from the first specified localities in Thailand. Moreover, A. festiva appears to be not only new to the fauna of Malaysia, but it also seems to be quite widespread across the southern half of Malay Peninsula both within lowland southern Thailand and Western Malaysia, being confined there to elevations not exceeding 60 m a.s.l. (Fig.
Orthomorpha harpaga Attems, 1937: 77 (D).
Orthomorpha harpaga – Attems 1938: 211 (D).
“Orthomorpha” harpaga –
Antheromorpha harpaga –
Antheromorpha harpaga –
♂ of Orthomorpha harpaga (
1 ♂ (
The lectotype is designated here to ensure that the name-bearing specimem is a complete ♂.
Length 19–21 mm (♂), width of midbody pro- and metazonae 1.8–1.9 and 2.3–2.6 mm, respectively (vs 1.8 and 2–2.5 mm in width of pro- and metazonae, respectively, as given in the available descriptions (
Antheromorpha harpaga (Attems, 1937), ♂ lectotype. A, B anterior part of body, dorsal and lateral views, respectively C, D segments 10 and 11, dorsal and lateral views, respectively E–G posterior part of body, lateral, dorsal and ventral views, respectively H, I sternal cones between coxae 4, subcaudal and sublateral views, respectively.
Clypeolabral region sparsely setose, epicranial suture distinct. Antennae short (Fig.
Tegument smooth and finely shargreened, metaterga smooth and delicately rugulose; surface below paraterga finely microgranulate. Postcollum metaterga with two transverse rows of setae traceable at least as insertion points when setae broken off: 2+2 in anterior (pre-sulcus), 3+3 in posterior (post-sulcus) row. Tergal setae simple, slender, about 1/3 of metatergal length. Axial line barely visible, starting with collum. Paraterga very strongly developed (Fig.
Sterna sparsely setose, without modifications; a high, subcordiform, sternal lobe between ♂ coxae 4 (Fig.
Gonopods (Fig.
Distribution of Antheromorpha species (11 species). Open triangle A. rosea Golovatch, 2013 Crossed circle A. bistriata (Pocock, 1895) Open square A. comotti (Pocock, 1895) and A. mediovirgata (Carl, 1941) Filled diamond A. miranda (Pocock, 1895) Crossed square A. comotti (Pocock, 1895), A. miranda (Pocock, 1895) and A. minlana (Pocock, 1895) Inverted open triangle A. pardalis (Pocock, 1895) Filled Circle A. uncinata (Attems, 1931) Open Circle A. paviei (Brölemann, 1896) Filled square A. harpaga (Attems, 1937) Open diamond A. festiva (Brölemann, 1896).
This is the only species in this genus that has been reported from Vietnam (
1 | Colour pattern of metaterga: yellowish paramedian spots in front of transverse sulcus, the latter visible starting with segment 2 | A. pardalis |
– | Colour pattern of metaterga otherwise. Transverse sulcus present starting with segment 4 or 5 | 2 |
2 | Colour pattern of metaterga: yellowish paramedian stripes | 3 |
– | Colour pattern of metaterga otherwise | 8 |
3 | Gonopod femorite relatively short (Figs |
4 |
– | Gonopod femorite longer (Figs |
6 |
4 | Metazonae ca 2.0 mm wide. Pleurosternal carinae poorly developed, in ♂ slightly projecting caudad behind rear tergal margin only until segment 5 | A. mediovirgata |
– | Metazonae ≥ 2.9 mm wide. Pleurosternal carinae well-developed, in ♂ slightly projecting caudad behind rear tergal margin at least until segment 10 | 5 |
5 | Sternal lamina between ♂ coxae 4 with a paramedian pair of evident, high, nearly pointed, fully separated, setose cones (Fig. |
A. festiva |
– | Sternal lamina between ♂ coxae 4 with only single small cone. Gonopod process d shorter (Fig. |
A. bistriata |
6 | Sternal lamina between ♂ coxae 4 a large, cordiform, ventrally evidently concave lobe (Fig. |
A. comotti |
– | Sternal lamina between ♂ coxae 4 a simple, rounded, conical knob. Gonopod solenophore evidently curved (Figs |
7 |
7 | Metazonae ca 2.6 mm (♂) or ca 3.0 mm wide (♀). Gonopod femorite rather straight, process d longer than solenophore (sph) (Fig. |
A. minlana |
– | Metazonae 3.2–3.7 mm (♂) or 3.6–4.6 mm wide (♀). Gonopod femorite strongly curved caudad, process d shorter than solenophore (sph) (Fig. |
A. miranda |
8 | Sternum between ♂ coxae 4 with a single lamina or cone (Figs |
9 |
– | Sternum between ♂ coxae 4 with a pair of separated cones (Figs |
10 |
9 | Colour pattern: a light axial stripe flanked on each side by a dark stripe on collum to epiproct (Figs |
A. uncinata |
– | Colour pattern indistinct, with a pale yellowish median stripe against a uniformly brown background (Fig. |
A. harpaga |
10 | Colour pattern: paraterga and epiproct contrasting dark yellow on a blackish body (Fig. |
A. paviei |
– | Colour pattern: a dark band present only on posterior halves of proterga and posterior halves of metaterga (Fig. |
A. rosea |
As a result of our review, the genus Antheromorpha now comprises 11 species ranging from southern China, through Myanmar, Thailand, Laos and Vietnam, to Western Malaysia (Fig.
The pair A. uncinata and A. festiva shows a remarkable geographical gap in southern Thailand, more specifically, in the northern half of the Malay Peninsula (Fig.
There can be no doubt that further collecting efforts, especially in still very poorly explored regions such as Laos, China, Malaysia and Vietnam, will reveal more Antheromorpha species, as well as further records of the known congeners. Cambodia remains an especially poorly prospected country in Indochina whence no Antheromorpha has been documented yet (
This project was partly funded through grants received from the Office of the Royal Development Projects Board, while most of the financial support was obtained from The Thailand Research Fund, The TRF Senior Research Scholar RTA 5880002 (2015–2018) to SP. We thank the members of the Animal Systematics Research Unit for their invaluable assistance in the field. Both Henrik Enghoff (