Amphicnemis triplex sp. nov. from Central Kalimantan, Indonesia (Odonata: Coenagrionidae)

Amphicnemis triplex sp. nov. from Indonesia 1 st June 2014 67 Amphicnemis triplex sp. nov. from Central Kalimantan, Indonesia (Odonata: Coenagrionidae) Naturalis Biodiversity Centre, P.O. Box 9517, 2300 RA Leiden, The Netherlands; <rory.dow230@yahoo.co.uk> Received 27 th September 2013; revised and accepted 10 th February 2014 Abstract. Amphicnemis triplex sp. nov. is described from locations in Central Kalimantan in Indonesian Borneo. Holotype Indonesia, Kalimantan, Kalimantan Tengah, between Buntok and Ampah, black water stream in shallow peat over sand, 29-vi-2012; to be deposited in RMNH. The related species A. erminea is discussed. Key words. Dragonfly, damselfly, Zygoptera, new species, Borneo Introduction Following the transfer of all species from the Philippines to other genera (Villanueva 2012), the genus Amphicnemis Selys, 1863 has 19 named representatives in southern Thailand, the Malay Peninsula, Sumatra and satellite islands, and Borneo; see the summary in Dow et al. (2010). The genus is particularly well represented in Borneo; Dow et al. (2010: 46) list 12 named species of Amphicnemis from the island. A number of new species of Amphicnemis from Borneo await description, but some sections of the genus present considerable taxonomic difficulties and need to be dealt with in a thorough revision. However, during a trip to Central Kalimantan (Kalimantan Tengah) in Indonesian Borneo in 2012, I found two very distinctive new species at a number of peat swamp forest sites. For one of the new species, every individual collected was teneral, so a description awaits the collection of at least a mature male. The other new species is described here as A. triplex sp. nov. Although the male of A. triplex is very distinctive, the supposed female is similar to that of A. erminea Lieftinck, 1953, which has the same known distribution. In fact, a number of old female specimens of A. triplex were found labelled as A. erminea amongst

68 material of the latter species in the collections of the Naturalis Biodiversity Centre. Amphicnemis erminea is discussed here and an illustration of the female prothorax is provided to facilitate separation of the two species; additionally a lateral view of the male anal appendages of A. erminea, lacking in the original description (Lieftinck 1953a) is provided. Terminology for odonate anatomy used here mostly follows Westfall & May (1996). The standard code RMNH is used for the collection of the Naturalis Biodiversity Centre, Leiden, The Netherlands. If available at the time of publication, collection registration numbers are stated for type material, and reference numbers for material at least initially in coll. Dow are also stated for type material. Amphicnemis erminea Lieftinck, 1953 (Figs 1, 4) Amphicnemis erminea Lieftinck (1953a): 247 251, figs 6c e; Lieftinck (1953b): 382, 392; Lieftinck (1954): 66; Lieftinck (1971): 86; van Tol (1992): 93; Dow et al. (2010): 46; Dow & Silvius (2014): 19, fig. 20. Material studied Type material. 2, 6, Ampah, 0 20 m, iv v-1948, leg. L.S. Liong, includes holotype (JvT number 2674), rest paratypes, in RMNH. Other material (all Kalimantan Tengah, Indonesia)., Sampit, iii-1948, leg. Kostermans, in RMNH; 2,, S. Sampit, 12 13-i-1950, leg. W. Buyn, in RMNH; 2, 2, Sampit, vii viii-1953, leg. M.A. Lieftinck, in RMNH; 9, 4, Pemantan-Sampit, 100 m, vii-1953, leg. M.A. Lieftinck, in RMNH; (KAL12_COE137), 5 (KAL12_COE138 141, 175), swamp by stream accessed from main road Palangkaraya Buntok, Palangkaraya area, 19- -vi-2012, leg. R.A. Dow, 1 in RMNH, rest in coll. R.A. Dow; 3 (KAL12_ COE131 133), (KAL12_COE130), strip of swampy forest surrounding stream accessed from Jalan Tjilik Riwut near Palangkaraya, 20-vi-2012, leg. R.A. Dow, 1, 1 in RMNH, rest in coll. R.A. Dow; (KAL12_COE21), peat swamp forest, Tuanan, 24-vi-2012, leg. R.A. Dow, in coll. R.A. Dow; 11 (KAL12_COE6 13, 34, 152, 182), peat swamp forest, accessed from main road Palangkaraya Buntok, Buntok area, 30-vi-2012, leg. R.A. Dow, 3 in RMNH, rest in coll. R.A. Dow; 2 (KAL12_COE22 23), 3 (KAL12_

Amphicnemis triplex sp. nov. from Indonesia 69 COE24 26), same data, leg. M. Silvius, in coll. R.A. Dow; 70, 59, same data as types, in RMNH. Remarks The envelope marked as containing the holotype actually contains two males, so that it is not clear which specimen is actually the holotype. The lateral view of the male anal appendages (Fig. 4), lacking in the original description, provided here was made from one of the topotypical males listed above. The female of this species was described by Lieftinck (1953a) based on supposition, despite the great dissimilarity between the posterior pronotal lobe of the male and the supposed female; the male posterior pronotal lobe is barely raised in its central rear part, in contrast to the large horn present in the same position in the female (Fig. 1). It would not be unreasonable to wonder if in fact the female associated here with A. triplex sp. nov. is actually that of A. erminea and vice versa, or if Lieftinck s female is actually that of some entirely different species for which the male has not yet been found. Figures 1 3. Posterior pronotal lobe of Amphicnemis spp., lateral views: (1) A. erminea female, adapted from Lieftinck (1953a: fig. 6e); (2) A. triplex sp. nov., holotype; (3) A. triplex sp. nov., paratype female.

70 However, I found females matching Lieftinck s description together with male A. erminea at only three sites; at each of these sites no specimens of A. triplex sp. nov. were found. Although the female of A. triplex is also only associated with the male by reasonable supposition, the posterior pronotal lobe of both sexes is similar not only in the horn but also in the lateral parts (Figs 2 3), reducing further the likelihood that the females of the two species have been transposed by Lieftinck and myself. Amphicnemis triplex sp. nov. (Figs 2, 3, 5 7) Amphicnemis new species 1 Dow & Silvius (2014): 21, 29, Table 2. Material studied Holotype (KAL12_COE134). Indonesia, Kalimantan, Kalimantan Tengah, between Buntok and Ampah, black water stream in shallow peat over sand, 29-vi-2012, leg. R.A. Dow. To be deposited in RMNH. Paratypes (all Indonesia, Kalimantan, Kalimantan Tengah, all leg. R.A. Dow and in coll. R.A. Dow unless otherwise noted).,»at house«, Sampit, 25-i-1950, leg. W. Buyn, in RMNH; 2, Sampit, 20-vii-1953, leg. M.A. Lieftinck, in RMNH; (KAL12_COE126), 2 (KAL12_COE127, 239), peat swamp forest in ex-mega Rice Project block E, 18-vi-2013; 5 (KAL12_ COE142 145, 153), 3 (KAL12_COE146, 160, 221), Sebangau, peat swamp forest, 21-vi-2012, KAL12_COE153, 160 in RMNH; 2 (KAL12_COE1, 40), 5 (KAL12_COE107 111), Begantung, peat swamp forest, 23-vi-2012, KAL12_COE1 in RMNH; (KAL12_COE112), Tuanan, peat swamp forest, 25-vi-2009; 4, Ampah, iv v-1948, leg. L.S. Liong, in RMNH. Other material. (KAL12_COE135), same data as holotype. Etymology The specific name is triplex, an adjective meaning triple, in reference to the three pointed appearance of the lower branch of the male superior anal appendage.

Amphicnemis triplex sp. nov. from Indonesia 71 Diagnosis The male of this species is easily separated from all others except A. annae Lieftinck, 1940 by the combination of having a central horn on the posterior pronotal lobe that is as long as or longer than the middle pronotal lobe and the pterostigma not bicoloured and the same colour in both pairs of wings. It is easily separated from A. annae, and all other species, by form of its anal appendages, in particular the three pointed appearance of the terminal part of the lower branch of the superior anal appendage. Male (Holotype) Head Labium pale. Labrum black with a broad yellowish cream transverse marking at free margin. Genae mostly yellow except immediately adjacent to mandible bases, where dark, mandible bases very dark brown with a pale border except adjacent to genae. Postclypeus black, anteclypeus dark brown and black with a central transverse pale mark and a pair of lateral pale marks. Post frons black, lower part ante frons black with a broad yellow transverse stripe narrowly interrupted centrally. Antennae bases with a broad yellow anterior streak, scape black, pedicel and flagella missing. Vertex and occiput very dark metallic green. Thorax Prothorax with pronotum very dark metallic green except for a pale lateral patch on either side of the anterior lobe. Propleuron cream, with the dark colouring of the middle pronotal lobe extending irregularly below the notopleural suture. Posterior pronotal lobe produced centrally at rear into a long erect horn, slightly longer than the middle pronotal lobe (Fig. 2). Synthorax with mesepisternum entirely dark metallic green, almost all of the mesepimeron and most of the mesinfraepisternum the same colour, which extends onto the upper part of the metepisternum and to the metepleural suture near the wing bases, the rest cream coloured. A patch of medium length setae is present on the rear part of the venter. Legs Almost entirely yellow, except for dark spines and a small dark area around the articulation of femur and tibia, and indistinct darker patches on the tarsi. Wings With arculus just after Ax 2, 1A arising at ca level of arculus, distal to ac. IR 3 just distal to Sn, R 4 ca half a cell distal to Sn. 13 Px in Fw, 11 Px

72 in Hw. Pt dark grey-brown with a faint and irregular pale border, costal side ca ⅔ anal side, covering one or slightly less underlying cell. Abdomen S1 cream coloured laterally, this extending dorsally in the basal half, dark metallic above, extending laterally to rear. A patch of setae ventrally on basal half of S1. S2 dark metallic green above, pale laterally along margin of tergite, this extending narrowly as a basal annulus, interrupted dorsally. S3 S8 brownish, becoming darker on successive segments, with a faint, dorsally interrupted, basal annulus on S3 S6. S9 almost entirely dark metallic, S10 brown above, paler to sides. Anal appendages As illustrated in Figures 5 7, pale except tip of interior tooth of lower branch of superior anal appendage. The lower branch of the superior anal appendage bears a robust subapical interior tooth, black at end, and a ventral tooth at the same position, giving the end of the branch a three pointed appearance (Fig. 5). Three long and relatively robust setae Figures 4 7. Male anal appendages of Amphicnemis spp.: (4) A. erminea, lateral view; (5) A. triplex sp. nov. holotype, lateral view; (6) A. triplex sp. nov. holotype, dorsal view; (7) A. triplex sp. nov. holotype, ventral view.

Amphicnemis triplex sp. nov. from Indonesia 73 present on lower branch of superior appendage clearly visible in lateral view (Fig. 5; in Fig. 6 the most basal of these is obscured by the interior tooth of the upper branch). Measurements [mm] Hind wing 20.5, abdomen without anal appendages 34.5, upper branch of superior anal appendage just over 1. Female (based on KAL12_COE239; as male except as noted) Thorax Prothorax with pronotum and propleuron almost entirely greyish blue. Horn on posterior pronotal lobe as in Figure 3 in lateral view, broad and triangular at base, then of even width until near apex. Most of syn thorax same greyish blue colour, mespisternum a yellow-bronze colour, venter pale. Legs With poorly defined greyish markings on middle and posterior coxae, and a black stripe on the extensor surface of the femora. Wings 12 Px in Fw, 11 Px in Hw. Abdomen S9 dark with a small apical dorso-lateral spot. S10 and anal appendages cream, venter S10 dark brown. Superior anal appendages just shorter than S10, inferior anal appendages very short and formless. Ovipositor mostly cream coloured, tip just beyond end of S10. Measurements [mm] Hind wing 20.5, abdomen without anal appendages or ovipositor 34.5. Variation in paratypes Males In the male paratypes, aside from small variations in markings, the only significant variation is that the transverse stripe on the ante frons is more widely divided in the middle in some specimens, that the horn on the posterior lobe of the pronotum is more curved forward in two individuals and that the pterostigma typically has a better defined pale border and in most individuals covers slightly less than one under-lying cell in each wing. There are two or, more commonly, three long prominent setae on the lower branch of the superior anal appendage. Measurements [mm] Hind wing 19 21, abdomen without appendages 32.5 34.5, 11 13 Px in Fw, 11 Px in Hw.

74 Females Also show variation in the pale markings on the ante frons; in some individuals these marks are fused to form a single band. The pale spot on S9 varies in size, and in some individuals the costal side of the pterostigma is shorter, only just over half the anal side. Some individuals are immature and have the thorax and legs almost entirely red. Measurements [mm] Hind wing 19 22.5, abdomen without appendages or ovipositor 31 36, 12 14 Px in Fw, 10 12 Px in Hw. Remarks The female is associated with the male through reasonable supposition based on having been found together with the male at a number of sites; see remarks under A. erminea above. The female (KAL12_COE135) from the same location and date as the holotype is excluded from the type series because it has an abnormal posterior lobe of the prothorax, with the top ca one third of the horn unevenly expanded, and, on either side of the horn, a short, rearward directed and poorly formed projection. The appearance of these atypical structures and the fact that it agrees in all other characters with the females from the type series, suggests that this is merely a malformed individual, but the possibility that it is from a separate species cannot be excluded. Discussion Both Amphicnemis erminea and A. triplex sp. nov. fall within a group of species best referred to as the A. wallacii-group, after A. wallacii Selys, 1863, the genotype of Amphicnemis. The wallacii-group can most simply be characterised by having males with the pterostigma the same colour in both pairs of wings and not bicoloured, except for a narrow pale border. The other species included in the wallacii-group are A. annae Lieftinck, 1940; A. bebar Dow, Choong & Ng, 2010; A. billitonis Lieftinck, 1940; A. gracilis Krüger, 1898; A. kuiperi Lieftinck, 1937; A. mariae Lieftinck, 1940; A. remiger Laidlaw, 1912; and A. smedleyi Laidlaw, 1926. Amphicnemis triplex falls into the wallacii-group because of the colouration of its pterostigma. However its superior anal appendages are very distinctive within the wallacii-group, especially the strongly three pointed appearance of the lower branch (Figs 5+7). In all other species at most a single,

Amphicnemis triplex sp. nov. from Indonesia 75 small, interior tooth or a single spur is present in the terminal part of the lower branch, so that it appears either un-bifurcated or with a double point. The long setae on the lower branch of the superior anal appendages are distinctive, but setae like this are present in the same position in all species of the wallacii-group; in fact they are present on the A. erminea specimen illustrated in Figure 4, but are directed inwards in that individual so that they are not readily visible in lateral view. Although these setae provide another distinguishing character for the group they are of no diagnostic value within the group. It should be noted that shorter, finer setae occur at other positions on the anal appendages; these are not considered to have any diagnostic value whatsoever. Until now our knowledge of the Odonata of the huge province of Central Kalimantan was based almost entirely on material collected during two rather short periods: a collection from the vicinity of Ampah in the southeast of the province made by Liem Swie Liong (later known as Pariwono) in 1948, and a collection made around and inland from Sampit, further to the west, by M.A. Lieftinck in 1953 (Lieftinck 1953b). Lieftinck described new Amphicnemis species from each of these collections: A. platystyla Lieftinck, 1953 and A. erminea in Lieftinck (1953a); A. dactylostyla Lieftinck, 1953 and A. pandanicola Lieftinck, 1953 in Lieftinck (1953b). Of these species, A. erminea, A. platystyla and A. pandanicola have never yet been found outside of Kalimantan Tengah, and the status of dactylostyla-like specimens from Sarawak (see e.g. Dow & Reels 2013) is not clear at this time. Records of A. erminea from Brunei (Thompson & van Tol 1993; Orr 2001) have proven to be based on misidentifications (see Dow et al. 2010). Lieftinck s claim (1953b: 392, 1954) that A. erminea is associated with large»pandan«plants (Pandanus spp.) is merely an assumption. Firstly the species is frequently found where few or no large plants from this genus are present. Secondly the supposition that larvae found in the leaf axils of large Pandanus plants»which greatly resembled that of Pericnemis stictica«were those of A. erminea rather than some Pericnemis species is highly questionable. This supposition appears to have been founded on the idea that Peri cnemis do not occur in swamp forest; this idea is in fact not correct (see Dow et al. 2013: 14). The larvae of Amphicnemis remain unknown but adults are almost always associated with standing water in swamp forest or

76 swampy areas within generally drier forest formations. Teneral individuals are frequently found at locations where no freely flowing water can be found. Until larvae definitely of the species discussed here are found, their habitat cannot be known with certainty, but it seems very likely that they occur in standing water. Amphicnemis triplex sp. nov. was found with A. erminea, A. pandanicola, A. platystyla and the other new species mentioned in the introduction, in various combinations depending on the location. All of the sites at which it was found were peat swamp forest; the diversity of Odonata and probably other animal groups in this most threatened of south-east Asian forest habitat types appears to have been severely underestimated in the past. Acknowledgements The trip to Kalimantan Tengah during which A. triplex was discovered was made possible by Marcel Silvius from Wetlands International, who accompanied me and was responsible for most of the organisation and planning. I am also grateful for all the assistance provided to Marcel Silvius and myself by Dr Suwido Limin of CIMTROP, University Palangkaraya, staff of the Borneo Orangutan Foundation, staff of Camp Release, Ibu Mayang Meilantina for her advice and help with organising transport, and Alue Dohong for his help with logistical arrangements. Hao-miao Zhang and an anonymous reviewer made useful comments which have greatly improved the manuscript. Dow R.A., Choong C.Y. & Ng Y.F. 2010. A review of the genus Amphicnemis in Peninsular Malaysia and Singapore, with descriptions of two new species (Odonata: Zygoptera: Coenagrionidae). Zootaxa 2605: 45-55 Dow R.A. & Reels G.T. 2013. Previously unpublished Odonata records from Sarawak, Borneo. Part I. Kuching Division excluding Kubah National Park, and Samarahan Division. Faunistic Studies in South-East Asian and Pacific Island Odonata 3: 1-25 References Dow R.A. & Silvius M. 2014. Results of an Odonata survey carried out in the peatlands of Central Kalimantan, Indonesia, in 2012. Faunistic Studies in South-East Asian and Pacific Island Odonata 7: 1-37 Dow R.A., Reels G.T. & Butler S.G. 2013. Odonata of the Dulit Range in Sarawak, Malaysian Borneo. Notulae odonatologicae 8: 1-16

Amphicnemis triplex sp. nov. from Indonesia 77 Lieftinck M.A. 1953a. Additions to the odonate fauna of the Indo-Australian archipelago. Treubia 22: 233-269 Lieftinck M.A. 1953b. New dragonflies (Odonata) from Borneo, with notes on their habits and larvae. Treubia 22: 381-406 Lieftinck M.A. 1954. Handlist of Malaysian Odonata. A catalogue of the dragonflies of the Malay Peninsula, Sumatra, Java and Borneo, including the adjacent small islands. Treubia 22, Supplement: i-xiii, 1-202 Lieftinck M.A. 1971. A catalogue of typespecimens of Odonata preserved in The Netherlands, with a supplementary list of the Odonata types described by Dutch scientists deposited in foreign institutional collections. Tijdschrift voor Entomologie 114: 65-139 Orr A.G. 2001. An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology 4: 167-220 Thompson D.J. & van Tol J. 1993. Damselflies and dragonflies from four forest types in Brunei. Brunei Museum Journal 8: 57-72 van Tol J. 1992. An annotated index to names of Odonata used in publications by M.A. Lieftinck. Zoologische Verhandelingen 279: 1-263 Villanueva R.J.T. 2012. Review of the Philippine taxa formerly assigned to the genus Amphicnemis Selys. Part I: Overview and descriptions of three new genera (Odonata: Coenagrionidae). Zoologische Mededelingen 86: 579-604 Westfall M.J. & May M.L. 1996. Damselflies of North America. First edition. Scientific Publishers, Gainesville

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