Revision of the Philippine genus Cyrano Needham & Gyger (Odonata, Chlorocyphidae)*

BAnnales Entomologici Fennici. 1989 Vol. 55, pp. 121-127. Revision of the Philippine genus Cyrano Needham & Gyger (Odonata, Chlorocyphidae)* Matti Hamalainen Hamaiainen, M. 1989: Revision of the Philippine genus Cyrano Needham & Gyger (Odonata, Chlorocyphidae). Ann. Entomol. Fennici 55:121-127. A review is presented of the two taxa placed in the genus Cyrano, Rhinocypha unicolor Hagen in Selys, 1869 and Libellago asiatica Selys, 1879. Libellago asiatica is shown to be a composite species, the male being conspecific with C. unicolor, but the female not A male specimen is selected as lectotype in order to stabilize the already established view of the synonymy of these two species. Cyrano angustior sp.n. (holotype male from Mindanao) is described on new material of both sexes and is compared with C. unicolor. A key to the two known Cyrano spp. is presented. Matti Hamalainen, Department of Agricultural and Forest Zoology, University of Helsinki, 00710 Helsinki, Finland Index words: Odonata, Chlorocyphidae, Cyrano, Cyrano angustior sp.n., taxonomy, lectotype, Philippines Introduction The genus Cyrano was erected by Needham & Gyger (1939) to contain a single species, Rhinocypha unicolor Hagen in Selys, 1869. A brief description of the species, based on a single female collected from Manila and preserved in Coll. Hagen, was given by Hagen in a letter and quoted by Selys Longchamps (1869), who had not seen the specimen. Ten years later, Selys Longchamps (1879) described in detail Libellago asiatica, on material from Luzon and Mindanao. Unfortunately, the length of wings given in his description was 10 mm too short. Probably due to this error, Needham & Gyger (1939), who described unicolor properly in both sexes for the first time, were unable to link asiatica with Cyrano. This was done by Eraser (1949, 1950) and Laidlaw (1950), who correctly transf erred Libellago asiatica to Cyrano, and who further considered asiatica and unicolor to be conspecific. However, Eraser and Laidlaw *Results of the Roland Miiller Zoological Expeditions to the Philippines, No. 3. disagreed about the valid name of their sole species of Cyrano. Eraser took asiatica as the type species of the genus, since he considered unicolor to be a nomen nudum, due to the inadequate description. This was incorrect, however, and Laidlaw was right in considering unicolor a valid name and the type species of Cyrano. Laidlaw's (1950) conclusion that asiatica was a synonym of unicolor was accepted by subsequent workers (Davies & Tobin, 1984 and Tsuda, 1986). While working on Roland A. Miiller's Cyrano material collected on different Philippine islands, I noticed that it contained two distinct species. Further, the collection of Rijksmuseum van Natuurlijke Historic in Leiden includes material of the same two Cyrano species. After comparing the material with the description of Libellago asiatica by Selys Longchamps (1879), I began to suspect that asiatica was a composite species. Study of the type series proved that this suspicion was correct. The male sex is conspecific with C. unicolor, but the female is different. The late Dr. M. A. Lieftinck has also studied the type series and come to the same conclusion, as his determination labels, dated 1969, indicate. Lieftinck placed the lectotype label on the male specimen, which would make asiatica a

122 Hdmdlainen: Revision of the Philippine genus Cyrano... Figs. 1-2: Shape of male abdomen in Cyrano unicolor (Hagen) and C. angustior sp.n., dorsal view. 1: C. unicolor (male from Negros Occ., Mt. Canla-on). 2: C. angustior (paratype male from Leyte). Figs. 3-4: Colour pattern of female pterothorax in Cyrano unicolor (Hagen) and C. angustior sp.n., lateral view. 3: C. unicolor (female from Luzon, Bulacan province, Angat-Dam). 4: C. angustior (paratype female from Leyte). synonym of unicolor and he furnished one of the female specimens with the label "Cyrano spec. nov.". However, Lieftinck never published his designation or a description of the new species. To stabilize the current nomenclatural usage, I am accepting Lieftinck's opinion and designating the male as the lectotype of asiatica (thus confirming the synonymy of asiatica with unicolor), and describing a new species (to which the syntype females of asiatica belong). Genus Cyrano Needham & Gyger Cyrano Needham & Gyger, 1939:250 = Neocypha Cowley in Tillyard & Fraser, 1939: 207, nomen nudum. Type species: Rhinocypha unicolor Hagen in Selys, 1869. Diagnosis Rather large species with hind wing longer than abdomen. Wings in both sexes uncoloured, with only a small brownish area on the extreme base in males. Sectors of arculus distinctly separated at origin; arculus situated distal to the distal primary (= third) antenodal. Fore wing with 13-22 antenodals and 17-23 postnodals. IA and CuP short, reaching to level of nodus or 1-2 crossveins distal to it. IA leaving posterior border of wing distal to level of first antenodal. MA and CuP diverging strongly distally, intervening intercalary vein arising well before level of nodus. In his discussion on the early history of the family Chlorocyphidae and on the relationship of the present genera, Laidlaw (1951) considered Cyrano and Indocypha Fraser, 1949, to represent the earliest and most unspecialized grade of the Rhinocypha genus group. Key to the species Males: 1. Abdomen broadened, segment 3 considerably broader than segment 1. (Fig. 1) C. unicolor Abdomen narrow, segments 1 and 3 of about the same width (Fig. 2) C. angustior sp.n. Females: 1. Mesepistemum with a long narrow yellow anterior stripe and two yellow posterior markings (Fig. 3) C. unicolor Mesepistemum with four yellow dots, anterior one largest and rather quadrangle in shape, middle dots one below the other (Fig. 4) C. angustior sp.n. Cyrano unicolor (Hagen) The following list of references includes those for Libellago asiatica ssp. vittata Selys, 1891, from Burma up to Fraser (1934), who confirmed its status as a distinct species. Fraser (1949) took L. vittata as the type species of his new genus Indocypha. Rhinocypha unicolor Hagen in Selys Longchamps, 1869:666 (reprint p. 21) brief original description of a female from Manila. Libellago asiatica Selys Longchamps, 1879:384-385 (reprintpp. 38-39) original description of male and female from Luzon and Mindanao. Rhinocypha unicolor: Selys Longchamps, 1882:20 reference to Selys Longchamps (1869). Libellago asiatica: Selys Longchamps 1882:20 reference to Selys Longchamps (1879). Libellago asiatica: Kirby 1890:112 "Philippines". Rhinocypha unicolor: Kirby, 1890:114 "Manilla". Libellago asiatica: Selys Longchamps 1891a:490-491 descr. of a new ssp. vittata from Burma, comparison with the Philippine male specimens. Rhinocypha unicolor: Selys Longchamps 1891b:214 a note on the measurements of the type specimen.

Ann. Entomol. Fennici 55:3. 1989 123 Libellago asiatica: Selys Longchamps 1891b:216 a note on a male from Dolores (Luzon); comments on ssp. vittata. Libellago asiatica: Martin 1904:220 "Philippines, Tonkin" (the Tonkin material belongs to ssp. vittata}. (Libellago vittata: William son 1905:173 vittata appears for the first time as a species name, without further comments on its status). Libellago asiatica: Laidlaw 1917:39-40 reference to comments in Selys Longchamps (1891a). Libellago asiatica asiatica: Eraser 1928:458-459 comments on the distribution; redescription of L. asiatica vittata (male and first female) from Burma. Chlorocypha asiatica: Eraser 1934:56-57 C. vittata from Burma considered to be distinct from C. asiatica and the differences discussed. Chlorocypha asiatica: Cowley 1937:1,6,16 penis described and figured; generic status discussed. Rhinocypha unicolor: Lieftinck 1938:51-52 Philippines Cyrano unicolor: Needham & Gyger 1939:250-252, pi. 12: figs. 149-150 (male anal appendages) and pi. 20: fig. 290 (fore wing). both sexes described, find localities (all in Luzon) listed. Libellago asiatica: Needham & Gyger 1939:255 brief comments on the original description of the species, not seen by the authors. Neocypha asiatica: Tillyard & Fraser, 1939:207 asiatica stated to be the type species of the genus Neocypha Cowley, which is, however, a nomen nudum. Cyrano asiatica: Fraser 1949:9, 13, 15 genus diagnosed; asiatica listed as the type species; wing base figured. Cyrano asiatica: Fraser 1950:16-17 note on the selection of the type species for the genus Cyrano (cf. Fraser 1949). Cyrano unicolor: Fraser 1950:16-17 the name unicolor considered a nomen nudum; the synonymy of asiatica and unicolor discussed; the number of specimens in Needham & Gyger's (1939) material is wrongly given as 1700 (merely ca. 50!). Cyrano unicolor :Lsa&av/1950:268,269,270,278 listed as the type (and only) species of the genus. Cyrano asiatica: Laidlaw 1950:270 synonymized with unicolor. Cyrano asiatica: Fraser 1957:74 note on the type species of Cyrano; Neocypha Cowley synonymized with Cyrano. Cyrano unicolor: Lieftinck 1961:121 Philippines. Cyrano unicolor: Asahina 1968:351,355,366 recorded from Mt. Maquiling and Los Banos in Luzon. Cyrano unicolor: Davies & Tobin 1984:16 "Philippines". Cyrano asiatica: Davies & Tobin 1984:16 "syn. of unicolor". Cyrano unicolor: Tsuda 1986:66 "Philippines". C. unicolor was adequately described by Needham & Gyger (1939). In the following some additions and remarks are presented. The dark brownish colouration of the thorax in immature males, seems first to become dark metallic greenish, and then velvety black in fully mature males. The metallic violet touches on the sides of the thorax appear during maturation. In immature males abdominal segments 1-2 are brown with yellow lateral patches. During maturation, these segments turn black with metallic greenish and violet touches. There are two types of mature male specimens as far as the colour of the abdomen is concerned. In the one type, the dorsum of segments 3-8 is brick red, (1-2 and 9-10 black) and in the other type the abdomen is wholly black. Needham & Gyger (1939) link the black colouration of the abdomen with senescence of the individuals. However, it may equally well be a question of two different colour forms. One of the males with a black abdomen, from the Ifugao mountains in Luzon, is not quite mature, judged from the pale markings on the thorax and head. Moreover, none of the 111 males from Mt. Canla-on in Negros Occidental have a black abdomen (in 3 males the abdomen is somewhat darkened). Further studies on field populations are necessary to determine the nature of the red / black colouration of the male abdomen. If two different colour forms exist, both forms seem to be common in the Luzon populations, but in the populations of Mt. Canla-on in Negros the red form seems to dominate. As seen below, two types of abdomen colouration also exist in C. angustior sp.n. Moreover, two colour forms (red and black) are known to occur in mature males in some populations of the endemic Palawan euphaeid Cyclophaea cyanifrons Ris. The shape of the male abdomen of C. unicolor is presented in Fig. 1, the colouration of the female thorax in Fig. 3, and the male wings in Fig. 6. The colouration of the thorax in immature males is the same as in the female. C. unicolor has formerly been reported only from Luzon, where it seems to be widely distributed. The new material from Marindique and Negros extends its known range. Further material from different islands is required before an analysis of the possible intraspecific variability can be attempted. Specimens from Negros seem to be somewhat larger than those from Luzon. In the Luzon males of the present material, the length of the hind wing is 26-30.5 mm (mostly 28-29 mm) and the length of the abdomen (incl. the appendages) is 20-22 mm. In the Negros males, the equivalent measurements are 29-33 mm and 21-23 mm. In the Luzon females, the hind wing measures 27-31 mm (mostly 29-31 mm) and the abdomen 19-21 mm, whereas in the Negros females the measurements are 30-33 mm and 20-22 mm. Type materials Rhinocypha unicolor Hagen in Selys Longchamps, 1869: Dr. Charles Vogt of the Museum of Comparative Zoology of Harvard University has kindly studied the holotype female in Coll. Hagen and informed me that its thoracic colour pattern definitely fits to a figure draft drawn on a female from Luzon.

124 Hamalainen: Revision of the Philippine genus Cyrano... Libellago asiatica Selys Longchamps, 1879: Lectotype designated hereby: male specimen labelled "Luzon" / "509" / "Lib. asiatica Br. <? "/ "Cyrano unicolor Sel. del M. A. Lieftinck" / "Cyrano unicolor Hag.-Sel. (lectotype Rhinocypha (Sic!) asiatica Selys) del M.A. Lieftinck 1969" / "Collection Ede Selys- Longchamps". The specimen is complete and the red colour of abdominal segments 3-8 is fairly well preserved. The lecto type is conspecific with Cyrano unicolor (Hagcn in Selys 1869) and L. asiatica is thus a synonym of C. unicolor. Paralectotypes: One female labelled "Libellago asiatica Brauer. Luzon" / "Cyrano spec. nov. (inedite), (9 Rhin. asiatica Sel.), del M.A. Lieftinck 1969" / "Collections Ede Selys-Longchamps"; one female labelled "Mind" / "29" / "Libellago asiatica Selys" / "Collection Ede Selys-Longchamps"; both paralectotypes are conspecific with the new species described below. The type series is preserved in Coll. de Selys Longchamps at Institut Royal des Sciences Naturelles in Brussels. Note. I am sure that the locality of the first paralectotype is erroneously given as Luzon, instead of Mindanao. The second paralectotype is a broken specimen, with only four abdominal segments, so heavily glued to the crushed thorax that it is impossible to determine the sex on the basis of the genitalia. However, the specimen is mature and has the thorax colouration of the female, and the base of its wings is clear. Selys Longchamps's original description indicates that he was in possession of at least two males, and only one female. Comparison of the nodal index in the description with the type specimens reveals that he has regarded this specimen as male in his measurements, although the description of the colouration of the male sex is based only on the lectotype male. Additional material studied Coll. RMNH Leiden: 10 d1,13 9 from Luzon (Ifugao province, Mayayao, ML Proninee / Camarincs Sur province, ML Iriga / Laguna province, Los Banos). Coll. Roland A. Muller: 7 tf, 2 9 from Luzon (Nueva Vicaya, SL Fe, Dalton Pass / Benguet, ML Sto-Tomas / Bulacan, Angat-Dam), 1 c? from Boac area in Marindique and 111 <3, 18 9 from ML Canla-on in Negros Occidental. The male of Libellago asiatica from Dolores (Luzon) reported by Selys Longchamps (1891b) and preserved in his collection also belongs to C. unicolor. Cyrano angustior sp.n. Libellago asiatica Selys Longchamps, 1879:384-385 (reprint pp. 38-39). The description of the female concerns this species. Description of the holotype male Measurements: hind wing 32.5 mm, abdomen (incl. appendages) 23 mm. Head. B ase of labium yellowish. Rest of lower part of head shining black with brownish portions. Frontal part of anteclypeus, postclypeus, frons and occiput mat black, with two small light brown spots on sides of the lateral ocelli and small triangle-shaped spots on occiput. An obscure dull brownish area between eye margin and second antennal segment. Thorax. Prothorax black. Pterothorax all black with metallic greenish and violet touches. Second lateral suture very narrowly yellow in middle. Legs with spines blackish brown; hind femora reaching to base of segment 2. Wings. Uncoloured, only the base diffusely spotted with brownish to the level of the first antenodal. Venation blackish brown. Pterostigma black, covering 4.5-5 underlying cells, only very slightly fusiform. Quadrangle with 3-4 crossveins. Arculus situated between third (second primary) and fourth antenodal, in most wings just proximal to fourth antenodal. Fore wing with 17-21 antenodals and 21-22 postnodals; hind wing with 17-18 antenodals and 17 postnodals. Abdomen. Narrow, not broadened in apical segments; shaped as in Fig. 2. The greatest width of segment 3 about the same as that of segment 1. Segments 1, 9 and 10 black, the intermediate segments brick red; only the sternites (incl. genitalia) black, and lower side of segment 2 and that of the apical part of segment 8 obscurely black. Joints of segments very finely black. Strong lateral carina on segments 3-8 and a trace of it on segment 2. Anal appendages black and of the typical chlorocyphid shape, similar to those of C. unicolor. Superior appendages twice as long as segment 10, cylindrical and pincer-shaped and slightly dilated at tip. Inferior appendages very swollen at base, extending to about half length of superiors when viewed from the side. Tip of inferiors strongly toothed. Variation among the males The other males are somewhat smaller; Ley te paratype specimens: hind wing 28-31 mm, abdomen 20-22 mm; Dinagat specimens: hind wing 27-28 mm, abdomen 19-20 mm. The basal brown patch on the wings is slightly larger extending to the level of the second antenodal in most of the males. In the Leyte males segment 2 is also black, with an obscure reddish area on the dorsum. The lower side of the abdominal segments, below the lateral carina, is somewhat darkened on segments 3-6 and almost black on segments 7-8. In the apical 1/3 of segment 8, the black colour extends upwards over the lateral carina. In the Leyte males the colouration of the head, legs and thorax does not differ much from that in the holotype. The one incompletely mature male in the series has a pale area in the anterior half of the metepisternum, above the spiracle, and on the posterior half of the metepimeron. Immature males probably have the same thorax colouration as the females.

Ann. Entomol. Fennici 55:3.1989 125 The colouration of the Dinagat males differs more markedly from that of the holotype. The thorax has a stronger metallic greenish tinge with violet reflections, and the abdomen is wholly black; only the sides below the lateral carinae are brownish on segments 3-7. Segments 1-2 have a strong metallic tinge and the base of segment 3 has strong violet reflections. In the Leyte and Dinagat males, the antenodals number 15-18 in the fore wing and 13-17 in the hind wing; the postnodals number 17-22 and 14-21, respectively. The quadrangle has 2-4 crossveins. Female The female was well characterized by Selys Longchamps (1879), as Libellago asiatica. Measurements. Specimens in Coll. Selys: hind wing 28 mm, abdomen, 19 mm. Paratype from Agusan province (Mindanao): hind wing 31 mm, abdomen 20 mm. Paratypes from Leyte: hind wing 30-31 mm, abdomen 19-20 mm. Head black (shining black below, becoming mat black upwards on dorsal part of anteclypeus), extensively marked with chrome yellow. Base of labium, base of mandibles, lateral corners of frontal part of anteclypeus, genae, whole eye margin broadly from genae up to top of second antennal segment yellow; this yellow area extending medially over lateral third of postclypeus. Two small yellow dots on top of postclypeus just below the frontoclypeal suture. In the one somewhat immature specimen of the present series, these spots are connected to the main yellow area in the postclypeus, forming a?- shaped mark. A round yellow patch of variable size in the centre of the labrum becomes smaller in mature specimens. Surface of whole second antennal segment yellow. Dots on sides of lateral ocelli and a pair of triangular spots on occiput yellow. Prothorax. Anterior lobe broadly bordered with chrome yellow, the yellow band continuing to sides of middle lobe. Ventrolateral corners of posterior lobe broadly yellow. Thorax. Black with distinct chrome yellow markings as follows (Fig. 4). A large somewhat quadrangular mark in anterior part of mesepisternum bordering mesinfraepisternum. Two parallel smaller marks in middle of mesepisternum, the upper linear and the lower rounded. A small mark on upper part of mesepisternum below antealar sinus. Triangular oblong stripes just below humeral and first lateral sutures. Large yellow patch in anterior part of metepisternum and similar patch covering most of posterior half of mete- Fig. 5: Apex of female abdomen in Cyrano angustior sp.n. (paratype female from Leyte), lateral view. pimeron. Poststernum yellow, encircled with dark brown band, which crosses the apical part of poststernum. Legs. Coxae black, with large yellow patches on sides. Trochanters yellowish brown. Anterior femora black, middle and hind femora brownish. All tibiae black. Hind femora reaching to middle of segment 2. Wings. No basal brown markings. Otherwise as in male. Fore wing with 15-19 antenodals and 17-22 postnodals, and hind wing with 13-16 and 15-20, respectively. Abdomen. More robust than in male. Distinct lateral carina extending from segment 3 to segment 7. Dorsum of abdomen black, with narrow yellowish brown stripe on median carina. Large yellow lateral patch on segment 1. Sides of segments 2-6 yellowish brown below lateral carina. The pale colour narrowly exceeding the carina in the apical 3/4 of each segment, but extending upwards to form a triangle in the apical 1/4. In segment 7 the black colour extending somewhat below the lateral carina, but the apical yellow triangle distinct. Segment 8 mostly black, but paler in posterolateral area, and with a distinct round yellow mark. Segments 9-10 black, obscurely brownish on lower side. Genital valves and anal appendages as in Fig. 5. Material Holotype male: Philippine Islands, Mindanao, Misamis Or., Minalwang, 1.IV.1961, H. Torrevillas leg. (Rijksmuseum van Natuurlijke Historic, Leiden). Paratypes: 5 c?, 4 9, Philippine Islands, Leyte, Mahaplag, Hilusig, Mt. Balocane, 600 m, 29.Vm.-14.IX. 1986, Th. Borromeo leg.; 1 cf, same locality and collector as above, but May 1986; 1 o, Philippine Islands, Mindanao, Agusan Prov., Esperanza, Bayugan, Yashimoto leg. The last paratype in Rijksmuseum van Natuurlijke Historie, Leiden, the others in Roland A. Mailer's (St. Gallon, Switzerland) and the author's collections. Other material: Coll. Selys: 2 o, Mindanao (see above on p. 122). Coll. Miiller: 1 <? Philippine Islands, Dinagat, Loreto, Mt. Redondo, Midas River, 2-5.6.1988, A. Buonafe leg.; 1 <?, Philippine Islands, Dinagat, Loreto, Esperanda, Bali-Bali River, 25.5.1988, A. Buonafe leg. Etymology. Angustior (= narrower) chosen because the male abdomen of the new species is narrower than in unicolor.

126 Hdmdlainen: Revision of the Philippine genus Cyrano... Figs.6-7: Wings of Cyrano unicolor (Hagen) and C. angustior sp.n. 6: C. unicolor(male from NegrosOcc., ML Canla-on). 7: C. angustior (paratype male from Leyte). Comparative notes Besides the pronounced differences pointed out in the key, the two Cyrano species also differ in minor details, some of which are presented below. The arculus is more distal, usually nearer the fourth than third antenodal, in angustior, whereas in unicolor it is usually only somewhat distal to the third antenodal (Figs. 6-7). The pterostigma is slightly less fusiform in angustior than in unicolor. In the female unicolor, the yellow poststernum is basally divided by a broad black transverse band in its apical part; in angustior the base of the apical part of the poststernum is yellow. In the female angustior, the middle and hind femora are brown, but in unicolor they are black. In angustior the hind femur is shorter than in unicolor, in the male angustior, it reaches only the base of segment 2, whereas in unicolor it reaches the middle of segment 2. In the female unicolor, the hind femur extends to the base of segment 3, but in angustior only to the middle of segment 2. Acknowledgements. I am very grateful to Mr. Roland A. Miiller (St. Gallen, Switzerland) for the unique chance of studying his Philippine material. I wish to express my sincere gratitute to Dr. Jan van Tol for permission to study the Cyrano material in Rijksmuseum van Natuurlijke Historic, Leiden, to Dr. P. Grootaert (Institut Royal des Sciences Naturelles de Belgique, Brussels) for the loan of the type series oflibellago asiatica and to Dr. Charles Vogt (Museum of Comparative Zoology of Harvard University) for informal)on on the type of C. unicolor. References Asahina, S. 1968: Records and notes on Philippine Odonata. Japan. J. Zool. 15:349-376, pis. 1-2. Cowley, J. 1937: The penis of the Chlorocyphidae (Odonata) as a group-character. Trans. R. Entomol. Soc. London 86:1-17. Davies, D. A. L. & Tobin, P. 1984: The Dragonflies of the world: A systematic list of the extant species of Odonata. Vol. 1. Zygoptera, Anisozygoptera. Soc. Int. Odonatol. Rapid Communications (Suppl.) 3:ix + 127 pp. Fraser, F. C. 1928: Indian dragonflies. Part XXIX. J. Bombay Nat. Hist Soc. 32:450-459, pis. 1-3. 1934: The faunaof British India, including Ceylon and Burma. Odonata. Vol. JJ. Taylor & Francis, London, xxiv+398 pp., pis. 1-4. 1949: A revision of the Chlorocyphidae with notes on the differentiation of the Selysian species rubida, glauca, cyanifrons and curia. Bull. InsL R. Sci. Nat Belg. 25(6): 1-50. 1950: A revision of the Chlorocyphidae: addenda, with key to the Platycypha and Chlorocypha and descriptions of new species. Bull. InsL R. Sci. Nat. Belg. 26(18): 1-22. 1957: A rectification of the order Odonata. Zool. Soc. N.S.W., Sydney, 134 pp., 1 pi. Kirby, W. F. 1890: A synonymic catalogue of Neuroptera Odonata or dragonflies. With an appendix of fossil species. Gurney & Jackson, London, x + 202 pp. Laidlaw, F. F. 1917: A list of the dragonflies recorded from ihe Indian Empire with special reference to the collection of Ihe Indian Museum. Rec. Ind. Mus. 13:23-40, pi. 2. 1950: A survey of the Chlorocyphidae (Odonata: Zygoptera), with diagnosis of proposed new genera, and description of a new geographical subspecies. Trans. R. Entomol. Soc. London 101(8):257-280. Lieftinck, M. A. 1938: The dragonflies (Odonata) of New Guinea and neighbouring islands V. Descriptions of new and little known species of the families Libellaginidae, Megapodagrionidae, Agrionidae (sens, lat.), and Libellulidae (Genera Rhinocypha, Argiolestes, Drepanosticta, Notoneura, Palaiargia, Papuagrion, Teinobasis, Nannophlebia, Synthemis, and Anacordulia). Nova Guinea (N.S.) 2:47-128. 1961: New and interesting Odonata from the Philippines. Fieldiana Zool. 42:119-149. Martin, R. 1904: Liste des Nevropteres de 1'Indo-Chine. Mission Pavie (Zool.), pp. 204-221. Needham, J. G. & Gyger, M. K. 1939: The Odonata of the Philippines, II Suborder Zygoptera. Philipp. J. Sci. 70:239-307, pis. 11-22. Selys Longchamps, E. de 1869: Secondes additions au Synopsis des Calopterygines. Bull. Acad. Belg. (2) 27:645-680. 1879: Quairiemes additions au Synopsis des Calopterygines. Bull. Acad. Belg. (2) 47:349-409.

Ann. Entomol. Fennici 55:3. 1989 127 1882: Odonates des Philippines. Ann. Soc. Hist Nat Esp. 11:1-32,1 pi. 1891a: Viaggio di Leonardo Fea in Birmaniaeregioni vicine. XXXII. Odonates. Ann. Mus. Civ. Stor. Nat Geneva(2) 10 (30):433-518. 1891b: Additions aux Odonates des Philippines. Ann. Soc. Hist Nat Esp. 20:209-218. Tillyard, R. J. & Eraser, F. C. 1939: A rectification of the order Odonata based on some new interpretations of the venation of the dragonfly wing. II. The suborders Zygoptera (cont) and Protanisoptera. Australian Zool. 9:195-221. Tsuda, S. 1986: A distributional list of world Odonata. Preliminary edition. Osaka, 6 + 246 pp. Williamson, E. B. 1905: Thedragonflies (Odonata) of Burma and Lower Siam. I. Subfamily Calopteryginae. Proc. U.S. Natn. Mus. 28:165-187. Received 21.X.1988 Printed 15.VIII.1989