M. A. SCHOUTEN & J. P. DUFFELS

Institute for Biodiversity and Ecosystem Dynamics (Zoological Museum), University of Amsterdam, The Netherlands

A REVISION OF THE OF THE CARMENTE GROUP (HOMOPTERA, ) FROM THE ORIENTAL REGION

Schouten, M. A. & J. P. Duffels, 2002. A revision of the cicadas of the Purana carmente group (Homoptera, Cicadidae) from the Oriental region. – Tijdschrift voor Entomologie 145: 29-46, figs. 1-20, table 1. [ISSN 0040-7496]. Published 1 June 2002. The Purana carmente group is proposed for a supposedly monophyletic group of seven species from the Oriental region. Two species of this group are redescribed: P. carmente from Java and Sumatra, and P. barbosae from Jolo (); the latter species is taken out of syn- onymy with P. carmente. Four species are described here for the first time: P. hermes sp. n. from Sabah and Sarawak, P. infuscata sp. n. from , P. obducta sp. n. from the Malayan Penin- sula, Sabah, and Sarawak, and P. sagittata sp. n. from the Malayan Peninsula. P. dimidia, which was recently described from China and Vietnam, also belongs to this group. A key to identify the males and distribution maps of the species are provided. Correspondence: M. A. Schouten, Institute for Biodiversity and Ecosystem Dynamics (Zoo- logical Museum), University of Amsterdam, The Netherlands, Plantage Middenlaan 64, NL- 1018 DH Amsterdam, The Netherlands. Key words. – Cicadidae; Purana; carmente group; phylogeny; ; new species; ; Oriental region.

Distant (1905a) erected the Purana when he Purana is paraphyletic. Kos & Gogala (2000) sup- divided Leptopsaltria Stål, 1866 in three genera: Lep- posed that Purana ubina Moulton, 1923 and its rela- topsaltria, Purana, and . In Distant’s catalogue tives and Maua quadrituberculata (Signoret, 1847) (1906) these genera were placed in the division Dun- and its relatives form one monophyletic group. dubiaria. In 1923, Moulton transferred Purana and The present paper proposes another supposedly the genera Leptopsaltria, Maua, Nabalua Moulton, monophyletic group, the Purana carmente group, for 1923, and Distant, 1905 to the new section seven species: P. carmente (Walker, 1850), P. barbosae Leptopsaltriaria, characterised by the presence of tu- (Distant, 1889), and P. dimidia Chou & Lei, 1997, bercles on the ventral side of the male abdomen. Met- and 4 new species: P. obducta, P. hermes, P. infuscata, calf (1963) classified the Leptopsaltriaria as a subtribe and P. sagittata. The species of the group are described of the tribe Dundubiini, and added several genera and relationships within the group are investigated. that lack the characteristic tubercles on the male ab- domen to that group. In this paper, we follow Moul- MATERIAL AND METHODS ton’s restricted concept of the Leptopsaltriaria. Moulton (1923) distinguished the genera Leptopsal- The following abbreviations for the institutions, tria, Purana, and Maua, largely upon morphometric which are the depositories of the material studied, characters as relative length and width of head, thorax, have been used in the lists of material and throughout and abdomen. In our view it is highly questionable the text: whether these characters can be used to unravel the phylogenetic relationships within the Leptopsaltriaria. BMNH The Natural History Museum (formerly: The recent recognition of the monophyletic Purana British Museum (Natural History)), Lon- nebulilinea group by Kos and Gogala (2000) was a don first step to a classification of Purana and related gen- BPBM Bernice P. Bishop Museum, Honolulu era based on phylogenetic reconstruction. Their mor- CAS California Academy of Sciences, Depart- phological study of the male genitalia of a number of ment of Entomology, San Francisco species of Purana and Maua suggested that the genus NMWC National Museum of Wales, Cardiff

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Table 1. Character state matrix for the species of the Purana MNM Muzium Negara Malaysia, Kuala Lumpur carmente group and the outgroup used in the cladistic analy- PMSL Prirodoslovni Muzej Slovenije, Ljubljana sis. Numbers refer to the characters as discussed in the text. UKM Pusat Sistematik Serangga, Universiti Ke- bangsaan Malaysia, Bangi, Selangor 123456 RMNH Nationaal Natuurhistorisch Museum (for- merly Rijksmuseum van Natuurlijke His- P. carmente 110001 torie), Leiden P. barbosae 001100 P. obducta 011100 UMS Universiti Malaysia Sabah, Kota Kinabalu P. hermes 110011 ZMAN Zoölogisch Museum, Amsterdam P. infuscata 000010 P. sagittata 000011 The following geographical sources were used for P. dimidia 10100? tracing the co-ordinates of localities: The Times Atlas P. tigrina 000000 of the World (Anonymous 1994), GEOnet Names Server of the U.S. Defence Mapping Agency (http:// gnpswww.nima.mil/geonames/GNS/index.jsp), An- analysis. The characters used are discussed below and drees allgemeiner Handatlas (Anonymous 1906), the matrix is given in table 1. Nelles Road Atlas Southeast Asia excluding Indonesia 1. – Medial margin of male operculum: (0) shaded (Anonymous 1992), Aardrijkskundig woordenboek with black (figs. 6, 9, 13, 14); (1) with broad black van Nederlandsch Oost-Indië (Dumont 1917), Atlas band (figs. 3, 12, 19). van Tropisch Nederland (Anonymous 1938). The lo- 2. – Male operculum: (0) short, reaching no fur- calities and other label data of the specimens studied ther than half or two thirds of 3rd abdominal seg- for this revision were filed in a FileMaker Pro 4.0 data- ment; (1) long, reaching beyond two thirds or even base. The maps of the species distributions were printed beyond posterior margin of 3rd abdominal segment. from this database using the programme MapInfo for 3. – Basal pygofer lobes: (0) long, only basally con- Power Mac, version 4.0.3, on maps of ADC-Worldmap nected to lateroventral part of pygofer (figs. 4, 15- version 2.0 vol. 4 Southern Asia & Australia. 17); (1) short, for the greater part connected to py- Random selections of specimens (or all available gofer (figs. 7, 10, 20). specimens) were used for calculating the ranges, aver- 4. – Uncus: (0) uncus with median lobe, but with- ages and standard deviations of the size of the body out lateral lobes (figs. 4, 15, 16, 17, 20); (1) uncus and body parts mentioned in the descriptions. Mea- with a median and two lateral lobes (figs. 7, 10). surements were taken using a sliding calliper. 5. – Claspers: (0) absent (figs. 4, 7, 10, 20); (1) pre- sent (figs. 15-17). 6. – Dark patches on pronotal collar just in front of PHYLOGENY the anterior ends of the lateral mesonotal fasciae: (0) The Purana carmente group is erected to accom- small, not connected with lateral fasciae of mesono- modate P. carmente and its related species. The tum; (1) broad, usually connected with lateral fasciae monophyly of this group is based on three presumed of mesonotum. apomorphic characters: (1) male timbal cover with triangular, black marking (2) basal veins of the 2nd The analysis (exhaustive search) resulted in one sin- and 3rd apical areas of the tegmina not or very weak- gle most parsimonious tree with a length of nine ly infuscated, and (3) male opercula relatively long steps, which is given in fig. 1. and slender, reaching or passing the posterior margin of the 3rd abdominal segment. BIOGEOGRAPHY The species centred around Purana tigrina form the most likely sister group of the P. carmente group. According to Moulton (1923), Metcalf (1963), Purana tigrina and its relatives are characterised by Duffels & Van der Laan (1985), Chou et al. (1997), the prominently infuscated 2nd and 3rd apical areas and Kos & Gogala (2000), the genus Purana compris- of tegmina and the elongated, first pair of tubercles. es at least 35 species, which are distributed mainly in The sister group relationship of the P. carmente group Southeast Asia: Myanmar (Burma), , Cam- and the P. tigrina group is supported by the following bodia, Vietnam, the Malayan Peninsula, Philippines, characters: the predominantly black lower part of and the Greater Sunda Islands, but also occur in , postclypeus and mandibular plate, and the small first Sri Lanka, China, Taiwan, and Japan. So far, only two apical cell of the tegmina. species of Purana have been recorded from east of A phylogenetic analysis was carried out in order to Wallace’s Line: P. celebensis (Breddin, 1901) from Su- investigate the relationships within the P. carmente lawesi and P. carolettae (Esaki, 1936) from the Caro- group. Purana tigrina was used as outgroup for this line Islands in the Pacific.

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