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Downloaded from Brill.Com09/30/2021 07:33:50PM Via Free Access T  E,  145, 2002 1 2 DIRK GASSMANN and MATTI HÄMÄLÄINEN 1Institute of Evolutionary and Ecological Sciences, Leiden University / National Museum of Natural History Leiden 2Department of Applied Biology, University of Helsinki A REVISION OF THE PHILIPPINE SUBGENUS RISIOCNEMIS (IGNEOCNEMIS) HÄMÄLÄINEN (ODONATA: PLATYCNEMIDIDAE)* Gassmann, D. & M. Hämäläinen, 2002. A revision of the Philippine subgenus Risiocnemis (Ig- neocnemis) Hämäläinen (Odonata: Platycnemididae). – Tijdschrift voor Entomologie 145: 213-266, figs. 1-140, tables 1-3. [ISSN 0040-7496]. Published 1 December 2002. The subgenus Igneocnemis Hämäläinen, 1991 of the Philippine damselfly genus Risiocnemis Cowley, 1934, is revised. Descriptions and diagnoses of both sex of all 15 previously recognized species are provided, and five new taxa are described: R. antoniae sp. n. and R. rubricercus sp. n. from northeastern Mindanao, R. pistor sp. n. from southeastern Mindanao, and R. kaiseri sp. n. and R. nigra sp. n. from Samar. The females of 11 species are described for the first time. Keys to males and females are provided. Based on extensive new collections from across the Philip- pine archipelago, the distribution of all species is mapped. Characters of the male ligula and ap- pendages and the female prothorax were studied by scanning electron microscopy. Correspondence: D. Gassmann, Institute of Evolutionary and Ecological Sciences, Section Theoretical Biology & Phylogenetics, Leiden University c/o National Museum of Natural His- tory (Naturalis), P.O. Box 9517, NL-2300 RA Leiden, The Netherlands. E-mail: [email protected] Key-words. – Platycnemididae; Calicnemiinae; Igneocnemis Hämäläinen; new species; Philip- pines. The genus Risiocnemis Cowley, 1934 is the most terial from the Roland Müller Expeditions to the speciose genus of the damselfly subfamily Calicnemi- Philippines was pouring in until 1997. Whereas some inae in the Indo-Pacific region. It is endemic to the 1000 specimens of the genus Risiocnemis were present Philippine archipelago, but does not occur in the Sulu in the Müller collection in 1991, by 1997 the amount archipelago and the Palawan island chain. Generally, had increased to over 8300 of which some 2800 spec- the species of the genus are confined to small clear imens belong to Igneocnemis. Thus, a complete taxo- streams in shady rain forest environment, occurring nomic treatment of the subgenus Igneocnemis has be- from lowland up to lower montane forest. come timely and it is presented here. The present The genus is defined mainly by wing venational paper may be considered as the second part of the se- characters like the very distal position of the R4, IR3 ries ‘The Philippine genus Risiocnemis Cowley (Zy- and IR2 veins (cf. Hämäläinen 1991a; see below for a goptera: Platycnemididae)’ (Hämäläinen 1991a). It more thorough diagnosis). also forms part of a comprehensive taxonomic and Two subgenera within the genus Risiocnemis were phylogenetic revision of the Indo-Pacific Calicnemi- recognized by Hämäläinen (1991a) based on the inae carried out by the first author (Gassmann 1999, species-groups defined by Lieftinck (1961): Risiocne- 2000). The genus Risiocnemis now includes 36 de- mis Cowley, 1934 s. str., and Igneocnemis Hämäläinen, scribed species, of which 20 are assigned to the sub- 1991. A revision of the subgenus Risiocnemis Cowley, genus Igneocnemis. It is worth mentioning that over 1934, was presented by Hämäläinen (1991a), but the 4500 specimens of the subgenus Risiocnemis have be- revision of the subgenus Igneocnemis Hämäläinen, come available for study and one new species has been 1991, was postponed since a constant flow of new ma- described after the revision in 1991 (Hämäläinen 2000). The new material also includes two other new * Results of the Roland Müller Zoological Expeditions to species and one first male and one first female, not yet the Philippines, No. 17. described. 213 Downloaded from Brill.com09/30/2021 07:33:50PM via free access T E, 145, 2002 Monophyly of Igneocnemis and Risiocnemis s. l. History (Leiden) in 1998. For an itinerary of Roland Müller’s Philippine Expeditions and a general history Several putative synapomorphies of Igneocnemis of dragonfly research in the Philippines see Hämäläi- species point to the monophyly of the subgenus: the nen & Müller (1997). only smoothly crenulate wing margin (fig. 1), which is The present study is based on specimens in the fol- unique within the Malesian Calicnemiinae, the posi- lowing collections: tion of the arculus at the second antenodal vein, the CUIC Cornell University Insect Collection, Ithaca angulate postclypeus and the subequal length of the ISNB Institut Royal des Sciences Naturelles, Brussels first and second antennal segments. However, the MCZ Museum of Comparative Zoology, Harvard monophyly of the whole genus, i. e., Risiocnemis sensu University lato, is uncertain; several features of ligula morpholo- RMNH National Museum of Natural History (Natu- gy, colour pattern and female prothorax relate some ralis; formerly: Rijksmuseum van Natuurlijke species of the subgenus Risiocnemis to the Oriental Historie), Leiden (including the former genus Coeliccia Kirby, 1890, rather than to Igneocne- Roland Müller collection) mis. This question will soon be treated in detail with- SMFD Senckenberg-Museum, Frankfurt in the scope of a phylogenetic study of the Malesian Calicnemiinae (Gassmann in prep.). The following collection is referred to in the text (no material examined): FMNH Field Museum of Natural History, Chicago Relationships within Igneocnemis Although the males of Igneocnemis species often dif- Specimens examined fer remarkably in their coloration, in particular of the All specimens listed in the ‘material’ section of the legs, structural differences are usually less conspicuous descriptions below were examined by the senior au- within the group. Consequently, establishing species- thor, except for the holotype of R. atripes (Needham groups based on the shape of the male ligula (penis), as & Gyger, 1941), holo- and allotype of R. tendipes done for the subgenus Risiocnemis (Hämäläinen (Needham & Gyger, 1941), both in MCZ, and the al- 1991a), proved to be difficult. In most species the lotype of R. rubripes (Needham & Gyger, 1939) in ligula structure is quite uniform; however, some CUIC; these specimens were studied only by the junior species show distinct peculiarities in the shape of the author. A database of additional specimens, seen only apical margin of the terminal lobes or the median in- by the second author, who has preliminarily identified cision between them; R. ignea (Brauer, 1868) differs the entire Roland Müller odonate collection, is avail- from all other species by having the terminal lobes al- able from the authors upon request most entirely reduced. Except for R. flammea (Selys, 1882) which was figured by Hämäläinen (1991a), the Scanning electron microscopy ligula structures of Igneocnemis species are examined The male ligula and the appendages of all species and illustrated here for the first time. were studied by scanning electron microscopy (SEM). Contrary to the males, the females of the group are Since the ligula of several Igneocnemis species proved quite uniform in coloration. However, they can be to be quite uniform, only a selection of species is illus- distinguished by the structure of the posterior prono- trated here to show the most important structural dif- tal lobe and the length of valvae. ferences. The male appendages of all species are illus- Preliminary groupings within the subgenus based trated in lateral, posterolateral and dorsal view. The on, for instance, the male appendages, appeared to be female prothorax of all species was studied by SEM, ex- weakly supported by other characters, e. g. coloration. cept for R. melanops and R. polilloensis of which only Therefore, the definition of species-groups is post- one female specimen each was available. The anterior poned until a thorough phylogenetic study of the view of the posterior pronotal lobe of R. fuligifrons, R. group will be available (Gassmann in prep.). haematopus and R. ignea could not be illustrated due to the oblique angle of the lobe. All scanning electron micrographs were taken by the first author using a MATERIALS AND METHODS JEOL SEM 6400 microscope. The objects were cleaned Collections in 70% ethanol, air-dried and subsequently sputtered Most of the material examined was collected by with gold for 1.5-2 min at 20 mA before examination. Roland A. Müller (St. Gallen, Switzerland) and his as- sistants and co-workers during his Philippine Expedi- Terminology tions (1985-1997), which since 1987 were focused The terminology of odonate wing venation applied mainly on odonates. The Roland Müller collection here follows the modified Tillyard-Fraser system was acquired by the National Museum of Natural (Watson & O’Farrell 1991). 214 Downloaded from Brill.com09/30/2021 07:33:50PM via free access G H: Subgenus Igneocnemis Table 1. Distinguishing features of the two subgenera of Risiocnemis Cowley. Character Igneocnemis Hämäläinen Risiocnemis Cowley Wing: position of arculus at Ax2 distal to Ax2 Wing: shape of apex smoothly crenulate (fig. 1) distinctly crenulate Antenna: length of pedicellus subequal to scapus about 2 times length of scapus Male postclypeus: shape distinctly angulate in lateral view smoothly angulate or rounded in lateral view Female posterior pronotal lobe: shape not (entirely) divided trilobate Measurements Diagnosis The measurements of wing and abdomen were tak-
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