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The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae (Odonata, Platycnemididae)1

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The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae (Odonata, Platycnemididae)1 Bonner zoologische Beiträge Band 53 (2004) Heft 1/2 Seiten 37–80 Bonn, Juni 2005 The Phylogeny of Southeast Asian and Indo-Pacific Calicnemiinae (Odonata, 1 Platycnemididae) Dirk GASSMANN Institute of Biology, Leiden University / National Museum of Natural History, Leiden Abstract. Phylogenetic relationships of Southeast Asian and Indo-Pacific damselflies of the subfamily Calicnemiinae (Odonata: Platycnemididae) are examined by cladistic analyses using morphological characters. The strict consensus cladogram of the resulting equally most parsimonious trees supports the monophyly of the Papuan genus Idiocnemis Se- lys, the Philippine genus Risiocnemis Cowley and its subgenera, but leaves the basal relationships of the African genera and the Palawan genus Asthenocnemis Lieftinck partly unresolved. A preferred phylogenetic hypothesis is presented showing a well supported 'Indo-Pacific clade' consisting of Philippine, New Guinean and Solomon island taxa, and as sister group Asthenocnemis. Risiocnemis turns out to be a sister group of Lieftinckia/Salomocnemis (Solomon Islands), the sister taxon of those being the central New Guinean Arrhenocnemis Lieftinck. Together, these form a monophyletic group with the remaining Papuan taxa. Idiocnemis leonorae Lieftinck is transferred to Rhyacocnemis Lieftinck comb. nov. The possible effects of taxon sampling are discussed. Key words. Taxonomy, biogeography, morphology, Southeast Asia, phylogeny, cladistic analysis, Rhyacocnemis leon- orae comb. nov. 1. INTRODUCTION apically zigzagged), and size (small to medium-sized). Another character of Platycnemididae, long ciliae on the The odonate subfamily Calicnemiinae Fraser, 1957 repre- legs, was already mentioned by Selys (1886) for his le- sents small to medium-sized damselflies (Zygoptera) in- gion Platycnemis; however, this trait is shared by other habiting small running waters throughout the old world. zygopteran families (SCHMIDT 1951b). The group is almost exclusively tropical. Due to the con- finement to stable habitats and lacking dispersal capaci- The Platycnemididae now consist of two subfamilies, ties in the rainforest environment, the majority of species Platycnemidinae Yakobson & Bianki, 1905, and Calic- shows a high degree of endemism, both at species and nemiinae Fraser, 1957. genus level. Therefore, the group is well suited for his- torical biogeographical analyses (GASSMANN in prep.). The subfamily Platycnemidinae SELYS (1863) distinguished seven subgenera in his ‘le- FRASER (1957) was the first to define the subfamily in gion Platycnemis’. Together, those subgenera, which its present form, containing the two genera Platycnemis were later given generic rank by the same author (SELYS Burmeister, 1839, and Copera Kirby, 1890. The dilation 1886) represented a large part of the Afrotropical, of the male tibiae, which is the most distinct feature Southeast Asian and Indo-Pacific Calicnemiinae known Fraser used to characterize the subfamily, has turned out today. TILLYARD (1917) raised SELYS’ legions to sub- to be not diagnostic for the group. Several species cur- family level resulting among others in a subfamily rently assigned to Platycnemidinae, for example, some Platycnemidinae (he used the incorrect name ‘Platy- Madagascan ones (SCHMIDT 1951a) have undilated tib- cneminae’). However, the family-group name had al- iae. Contrarily, one species of Calicnemiinae, Risiocne- ready been introduced earlier (YAKOBSON & BIANKI mis atropurpurea (Brauer), does have dilated tibiae. 1905). TILLYARD & FRASER (1938/40) finally erected the family Platycnemididae. In the most recent account Within the genus Platycnemis, 31 species are currently on the group, MARTENS (1996) provided five diagnostic recognized (derived from MARTENS 1996 and characters (at least with regard to Coenagrionidae) for HÄMÄLÄINEN 2003). The group exhibits a disjunct pat- the family: the comparatively long and obtuse discoidal tern of distribution, involving the western Palaearctic cell (quadrilateral), the length of the pterostigma region, eastern Asia, northwestern/ western Africa as (matching no more than one wing cell), the variable po- well as the Madagascan region (SCHMIDT 1951a; MAR- sition of arculus (at or distal to Ax2), the course of the TENS 1996). No representatives were known from longitudinal veins MA and IR3 (mainly straight, only Southeast Asia until recently. However, the recent dis- covery of a new species from Laos (P. phasmovolans 1 In commemoration of Clas Michael Naumann zu Königsbrück Hämäläinen) extends the distribution of the group far (26.06.1939 – 15.02.2004) into the Indo-Chinese region (HÄMÄLÄINEN 2003). 38 Bonner zoologische Beiträge 53 (2004) Within the genus Copera (Copera spec., Plate 1b) cur- yan region (LIEFTINCK 1984); however, its distributional rently nine species are recognized (MARTENS 1996). range further extends southeast into the Indo-Chinese The group is largely replacing Platycnemis in Southeast mainland as well as to Taiwan and Hainan. Interest- Asia, but there is an area of overlap in eastern Asia ingly, no representatives of Calicnemia are known from (MARTENS 1996). The generic borders between Copera the Greater Sunda Islands. Indocnemis Laidlaw, 1917, is and Platycnemis are weakly defined, and their identity a monotypic genus; I. orang (Förster in Laidlaw, 1907) recently has been put in question by HÄMÄLÄINEN occurs in India and mainland Southeast Asia. Coeliccia (2003). Kirby, 1890 is by far the largest genus of Calicnemii- nae, currently comprising 59 species according to The subfamily Calicnemiinae SCHORR et al. (2004). Its distribution area ranges from the Himalayan region via the Southeast Asian mainland The damselfly subfamily Calicnemiinae was erected by into the Greater Sunda Islands and even into the western FRASER (1957) to accomodate species of platycnemidid and southwestern Philippines; south to the Philippines, damselflies which did not fit into the subfamily Platy- along the Greater Sunda Islands, the eastern border of cnemidinae. Fraser himself stated: “This subfamily is Coeliccia matches the Wallace line, except for the is- formed mainly for convenience to separate the true land of Bali from where no representative of the genus Platycnemiines, and one has only to compare the genera is known. A taxonomic and phylogenetic revision of a composing it, to note how artificial it appears to be...”. presumably monophyletic subgroup of the genus, con- Unfortunately, Fraser did not provide diagnostic charac- sisting of Sundaland and Philippine taxa, is in work ters for the subfamily, except for the absence of dilated (DIJKSTRA & GASSMANN, in prep.). The fourth Oriental tibiae, which on the other hand is a striking feature in genus, Sinocnemis, comprises two species and was es- the second subfamily Platycnemidinae. BECHLY (1996) tablished quite recently (WILSON & WEN-BAO 2000); found no autapomorphies for the group. Later authors however, it could not be studied in detail for the present (MARTENS 1996) have provided additional distinguish- analysis. ing characters for Calicnemiinae, i.e. the comparatively acute discoidal cell (quadrilateral) and the length of the anal vein. The Indo-Pacific Calicnemiinae The Calicnemiinae reach their highest diversity in Recently, a large part of the New Guinean and Philip- Southeast Asia and the Indo-Pacific region. Remarka- pine taxa have been revised taxonomically (GASSMANN bly, the group is absent from Wallacea (Sulawesi, 1999, 2000; GASSMANN & HÄMÄLÄINEN 2002; Smaller Sunda Islands, Moluccas), which has been con- HÄMÄLÄINEN 1991a/b, 2000). Based on recently col- firmed, at least for Sulawesi, by several odonatological lected specimens from different sources, several new surveys during the last decades (VAN TOL 2000). species were recognized. The New Guinean genus Idiocnemis Selys now amounts to 19 species and thus The African genera of Calicnemiinae represents the largest genus in the Papuan region, fol- lowed by Lieftinckia Kimmins (Solomon Islands; 6 spe- The African genera of the subfamily are a rather hetero- cies), Rhyacocnemis Lieftinck (New Guinea; 3 species, genous assemblage comprising only about a dozen spe- see below), Paramecocnemis Lieftinck (New Guinea, 2 cies in total (SCHORR et al. 2004). The family status of species). The remaining Papuan genera Cyanocnemis some of them, in particular that of Mesocnemis Karsch, Lieftinck, Lochmaeocnemis Lieftinck, Thaumatagrion 1891 (Western Africa, 4 species) and Metacnemis Selys, Lieftinck, Torrenticnemis Lieftinck (New Guinea) and 1863 (Southern Africa, Madagascar; 3 species), has Salomocnemis Lieftinck (Solomon Islands) are all been doubted (MARTENS 1996; WATERSTON 1984). Ex- monotypic (for Arrhenocnemis see below). The Philip- cept for Allocnemis Selys, 1863 (Southern Africa, 2 pine genus Risiocnemis Cowley now comprises 36 de- species), all remaining African genera are monotypic: scribed species, with subgenus Igneocnemis Hämäläinen Leptocnemis Selys, 1886 and Paracnemis Martin, 1903 (20 species) being somewhat larger than the nominal (Madagascar), Oreocnemis Pinhey, 1971 (eastern Af- subgenus Risiocnemis Cowley (GASSMANN & rica) and Arabicnemis Waterston, 1984 (Arabian Penin- HÄMÄLÄINEN 2002). Within the Philippine Calicnemii- sula). Allocnemis and Arabicnemis also extend into sub- nae, the Palawan genus Asthenocnemis Lieftinck is an tropical areas. outstanding representative. Originally, Lieftinck thought that the peculiar female specimen from the Martin col- The Oriental genera of Calicnemiinae lection of which he
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