A revision of the cicadas of the genus Maua Distant (Hemiptera, Cicadidae) from Sundaland

Transcription

1 A revision of the cicadas of the genus Maua Distant (Hemiptera, Cicadidae) from Sundaland J.P. Duffels This revision of the genus Maua presents descriptions, illustrations and distribution maps of 8 species of the genus occurring in Sundaland: the Malayan Peninsula, Java, Sumatra, Borneo and Palawan. Six species are redescribed: M. affinis Distant, 1905, M. albigutta (Walker, 1857), M. latilinea (Walker, 1868), M. linggana Moulton, 1923, M. platygaster Ashton, 1912 and M. quadrituberculata (Signoret, 1847). Two species are described as new to science: M. borneensis sp. n. and M. palawanensis sp. n. Two other species proved to be synonyms: M. ackermanni Schmidt, 1924 is a junior synonym of M. quadrituberculata and M. dohrni Schmidt, 1912 is a junior synonym of M. latilinea. Three species of Maua, two from China and one from the Philippines are not included in this paper. A key for the identification of the Sundaland species of Maua is presented. J.P. Duffels, Zoological Museum (Department of Entomology), University of Amsterdam, Plantage Middenlaan 64, NL-1018 DH Amsterdam, the Netherlands. j.p.duffels@uva.nl Introduction The genus Maua was erected by Distant (1905) for Cicada quadrituberculata Signoret, 1847 and M. affinis Distant, The genus is currently placed in the tribe Dundubiini and the subtribe Leptopsaltriina (Duffels & Van der Laan 1985; Moulds 2005). In 1923, Moulton erected the new section Leptopsaltraria [sic] for the genera Leptopsaltria Stål, 1866, Maua Distant, 1905, Nabalua Moulton, 1923, Purana Stål, 1866 and Tanna Distant, The new section was characterised by the presence of one to three pairs of tubercles on the ventral side of the male abdomen. A revision of Nabalua was recently presented by Duffels (2004), and three, presumed monophyletic, groups of the genus Purana were revised: the P. nebulilinea group (Kos & Gogala 2000), the P. carmente group (Schouten & Duffels 2002) and the P. tigrina group (Duffels et al. 2007). In 1963, Metcalf added several genera, with and without abdominal tubercles, to the subtribe; none of these genera are recorded from Sundaland. This paper presents a revision of the genus Maua from Sundaland: the Malay Peninsula, Java, Sumatra, Borneo, Palawan and several smaller islands inbetween. Two species of Maua, M. borneensis from Borneo and M. palawanensis from Palawan and Balabac are described as new to science, and six species from Sundaland are redescribed: M affinis, M. albigutta (Walker, 1857), M. latilinea (Walker, 1868), M. linggana Moulton, 1923, M. platygaster Ashton, 1912, and M. quadrituberculata. Two species, M. ackermanni Schmidt, 1924 and M. dohrni Schmidt, 1912, both described from Sumatra, proved to be junior synonyms of respectively M. quadrituberculata and M. latilinea. Three species of Mau are not treated in this paper: M. albistigma (Walker, 1850) and M. fukienensis Liu, 1940 from China, and M. philippinensis Schmidt, 1924 from the Philippines. This study aims to contribute to a better knowledge of cicada biodiversity in Southeast Asia. Basic systematic studies providing illustrations and descriptions of species and keys to genera and species are regarded as a prerequisite for biodiversity studies. Biodiversity studies of cicadas in other tropical areas, such as Sulawesi, New Guinea, and the West Pacific, Tijdschrift voor Entomologie 152: , Figs [ISSN ] Nederlandse Entomologische Vereniging. Published 1 August 2009.

2 304 Tijdschrift voor Entomologie, volume 152, 2009 have already shown that cicadas can be instrumental in recognizing hot spots of species richness and areas of endemism with unique biota (Duffels & De Boer 1990, De Boer & Duffels 1997). Material and methods The institutions listed below are the depositories of the material studied. The abbreviations have been used in the lists of material and throughout the text. BMNH Natural History Museum, London (former British Museum (Natural History)), London KBIN Koninklijk Belgisch Instituut voor Natuurwetenschappen, Brussels MNHN Muséum National d Histoire Naturelle, Paris MNKM Muzium Negara Malaysia, Kuala Lumpur MIZ Museum and Institute of Zoology, Polish Academy of Sciences, Warszawa MZB Museum Zoologicum Bogoriense, Cibinong MZHF Zoological Museum, Finnish Museum of Natural History, Helsinki NHRS Swedish Museum of Natural History, Stockholm PSML Prirodoslovni Muzej Slovenije, Ljubljana RMNH Nationaal Natuurhistorisch Museum (formerly Rijksmuseum van Natuurlijke Historie), Leiden ROM Royal Ontario Museum, Toronto UKM Pusat Sistematik Serangga, Universiti Kebangsaan Malaysia, Bangi, Malaysia UMS Universiti Malaysia Sabah, Kota Kinabalu ZMAN Zoölogisch Museum, Universiteit van Amsterdam, Amsterdam Data on the distribution of the species were derived from the author s Biodiversity Database of the Cicadas of South East Asia and the West Pacific, and plotted on maps of ADC-Worldmap version 2.0 vol. 4 Southern Asia & Australia with the program MapInfo for Power Mac, version The localities and other data from the specimen labels in the database are filed in the program FileMaker Pro 4.0. Geographical information has been retrieved from the following sources: Atlas van Tropisch Nederland (Anonymous 1938), Gazetteer Nasional Nama-nama Geografi for Indonesia (Anonymous 1978), Nelles Road Atlas Indonesia (Anonymous 1992), The Times Comprehensive Atlas of the World (Anonymous 1999) and GEOnet Names Server of the U.S. Defense Mapping Agency ( The terminology adopted in this paper for external features of the body and the male genitalia follows that of Duffels (Duffels 1977, 1983; Duffels & Turner 2002) and Moulds (2003, 2005). Taxonomy Genus Maua Distant Maua Distant 1905: 61. Type-species by original designation: Maua quadrituberculata (Signoret, 1847). Maua: Moulton 1923: ; Metcalf 1963: ; Duffels & Van der Laan 1985: 113; Moulds 2005: 391, 432. [For further references before 1980 see: Metcalf 1963 and Duffels & Van der Laan 1985]. The species of Maua studied have one character in common viz., the broad, more or less parallel-sided male abdomen with about equally wide segments 3 to 4 or 2 to 5. The shape of the male abdomen was traditionally used to separate Maua from other genera of the subtribe (Distant 1906; Moulton 1923). The genus Maua in its present concept is probably not monophyletic. Distribution The genus Maua is found in China, Thailand, the Philippines and Sundaland (Malay Peninsula, Java, Sumatra, Borneo and Palawan) (Metcalf, 1963; Duffels & van der Laan 1985; Sanborn et al. 2007). The genus is certainly most divers in Borneo (seven species). Four species are probably Borneo endemics: M. affinis, M. borneensis, M. linggana and M. platygaster. Maua affinis seems to be restricted to Palawan and Balabac Island between Borneo and Palawan. Maua latilinea is known from Borneo and Sumatra. The two remaining species have a wider distribution: M. albigutta is found in the Malay Peninsula and Sumatra, and M. quadrituberculata is known from China, Thailand, the Malay Peninsula, Java (?), Sumatra, Borneo and Nias Island. Relationships Three genera of the subtribe Leptopsaltriina occurring in Sundaland have two pairs of tubercles on the male sternites 3 and 4: Leptopsaltria, Maua and Purana. Maua is distinguished from the other two genera by the broad, more or less parallel-sided male abdomen; in Purana and Leptopsaltria the male abdomen gradually narrows from segment 2 backwards. The genus Maua is divided in two groups that are presumed to be monophyletic: (1) the M. quadrituberculata group with a male body length of mm, comprising: M. affinis, M. borneensis, M. latilinea, M. palawanensis, M. quadrituberculata

3 Duffels: Revision of Maua (Cicadidae) 305 (type species of the genus) and (2) the M. albigutta group with a male body length of mm, comprising: M. albigutta and M. linggana. The Maua quadrituberculata group and the M. albigutta group are readily separated by the following characters: the lateral fasciae on the mesonotum are continuous from anterior margin to at least three fifths of mesonotum length in the M. quadrituberculata group, and consist of three spots or short lines in the M. albigutta group; the male abdomen is very broad, and widest across the segments 3 and 4, in the M. quadrituberculata group, and fairly broad, and about equally wide across the segments 2 5, in the M. albigutta group; the male sternites 3 and 4 have a pair of thick, nipple-shaped tubercles in the M. quadrituberculata group, and a pair of narrow protuberances in the M. albigutta group. The albigutta group is allocated in the genus Maua because of the fairly broad male abdomen, but allocation in the genus Purana can also be considered because of the marking on the mesonotum and the weakly developed protuberances on the male sternites 3 and 4. The Maua albigutta group as well as two species groups of the genus Purana, the P. carmente group (Schouten & Duffels 2002) and the P. nebulilinea group (Kos & Gogala 2000), might represent new genera. However, a cladistic analysis of the species of Purana and Maua and related genera of the subtribe Leptopsaltriina is needed to provide a basis for a new generic division of the subtribe. One species, M. platygaster, is not attributed to one of the species groups and is not described in the same detail as the other Sundaland species. I have seen only one female of M. platygaster, the male of this species is only known to me by the original description of Ashton (1912). Maua platygaster can be separated from the other species of the genus by its characteristic marking of anastomosing spots on the tegmina and small body size (body length male: 21 mm). Key to the species of Maua The principal use of this key is for the identification of males. It is of limited value for the identification of females. The females of three species, M. affinis, M. latilinea and M. palawanensis are unknown. 1. Body length male: mm Body length male: mm Postclypeus with black spot or black marking at clypeal suture Postclypeus without black spot or black marking at clypeal suture Tegmen with distinct brown spots at bases of 2nd, 3rd and 5th apical cells; basis of 7th apical cell without spot. Distal ends of paramedian mesonotal fascia usually connected with median fascia. Palawan; Balabac I M. palawanensis Tegmen with distinct brown spots at bases of 2nd and 3rd apical cells; bases of 5th and 7th apical cells without spots. Distal ends of paramedian mesonotal fascia not connected with median fascia Lateral fascia on mesonotum broad and straight, and reaching from anterior mesonotum margin to almost hind margin of mesonotum (Fig. 17). Abdominal tergite 3 in male and female with triangular median marking. Malay Peninsula (?); Borneo; Karimata I.; Sumatra M. latilinea Lateral fasciae on mesonotum somewhat irregular and reaching from anterior mesonotum margin to three fifths of mesonotum length (Fig. 1). Abdominal tergite 3 in male (female unknown) without triangular median marking. China; Thailand; Malay Peninsula; Java (?); Borneo; Sumatra; Nias I M. quadrituberculata 5. Spot at basis of 2nd apical cell of tegmen continuing along basal vein of 1st apical cell. Male abdomen very broad, segment 3: times as wide as pronotal collar. Body length male: mm. Borneo.. M. affinis Spot at basis of 2nd apical cell of tegmen not continuing along basal vein of 1st apical cell. Male abdomen broad, segment 3: times as wide as pronotal collar. Body length male: mm. Borneo M. borneensis 6. Tegmen with anastomosing spots at bases of 1st and 2nd apical cells and at bases of 3rd and 4th apical cells, with separate spots at bases of 5th and 7th apical cells, and without spots at apices of longitudinal veins of apical cells. Body length male: 21 mm. Borneo M. platygaster Tegmen with separate spots at bases of 2nd and 3rd apical cells, and with small brown spots at apices of longitudinal veins of apical cells 1 4. Body length male: mm Male operculum apically broadly rounded and reaching, or reaching just beyond, posterior margin of abdominal segment 2. Lateral fasciae on mesonotum consisting of three short black lines on a row, posterior two lines sometimes fused (Fig. 23). Body length male: mm. Borneo M. linggana Male operculum tapering to the apex and reaching to half-length abdominal segment 3. Lateral fasciae on mesonotum consisting of

4 306 Tijdschrift voor Entomologie, volume 152, 2009 three distinct black spots on a row (Fig. 22). Body length male: mm. Malay Peninsula; Sumatra; Siberut I M. albigutta Maua quadrituberculata group The Maua quadrituberculata group consists of five medium-sized to fairly large species (body length male: mm): M. affinis, M. borneensis, M. latilinea, M. palawanensis and M. quadrituberculata. The lateral mesonotal fasciae are somewhat irregular or straight but continuous from anterior margin to at least three fifths of mesonotum length or to at most the hind margin of the mesonotum. The apical cell 1 of the tegmen is relatively short. The basal vein of apical cell 1 is shorter than half as long, or at most half as long, as the longitudinal vein of apical cell 1. The male abdomen is very broad, widest across the segments 3 and 4, segment 2 is only slightly narrower; the sternites 3 and 4 have a pair of thick, nipple-shaped tubercles pointing sidewards. Maua latilinea differs from the other species of the group in the body marking and the structure of the male genitalia. The basal pygofer lobes of M. latilinea are large, black-brown and spine-shaped, and reach from two fifths to three fourths of the pygofer length, while the uncus is triangular and strongly sclerotised. In the other four species of this group the basal pygofer are more or less parallel, apically acute, ridges, while the uncus is trapezoid and not strongly sclerotised. Maua quadrituberculata (Signoret) Figs 1 5, 7 Cicada quadrituberculata Signoret, 1847: Leptopsaltria quadrituberculata: Distant 1889: 31; Distant 1891: pl. viii figs 6, 6a-b. Maua quadrituberculata: Distant 1905: 61; Moulton 1923: 124, 126, 168; Metcalf 1963: 494, 495; Duffels & Van der Laan 1985: 113; Zaidi & Ruslan 1998: 153; Boulard 2007: Maua ackermanni Schmidt, 1924: 287 Holotype?: Soekaranda / Januar 1894 / Dohrn [print in black cadre], Maua / Ackermanni / Schmidt /? Edm. Schmidt / determ. 1924, Typus [print in black cadre; red paper], MIZ (MIZ) [examined]. syn. n. [For further references before 1980 see: Metcalf 1963 and Duffels & Van der Laan 1985]. Maua quadrituberculata is a large species (body length?: mm). The males have a very broad abdomen (segment 3: (1.45) times as wide as pronotal collar), though the abdomen of the males of M. affinis is usually even broader (segment 3: times as wide as pronotal collar). Maua quadrituberculata can be distinguished from all other large species of Maua by the unmarked posterior oblique fissures on the mesonotum that are black in the other species. Maua quadrituberculata and M. latilinea from Borneo, Kalimantan are the only species with two black spots, at the bases of the second and third apical cells of the tegmen; the other species of the group have spots at the bases of the second, third and fifth apical cells or at bases of second, third, fifth and seventh apical cells. Maua latilinea is easily distinguished from M. quadrituberculata and all other large species of Maua by the continuous broad lateral fasciae on the mesonotum that reach from anterior to almost posterior mesonotal margin. Synonymy In February 2008, Dr Tomi Trilar of the Prirodoslovni Muzej Slovenije, Ljubljana visited the Museum of the Institute of Zoology, Polish Academy of Science, Warsaw, Poland. He found the Typus and the Co-Typus of M. ackermanni Schmidt, 1924 in the collection of this museum, brought these to my attention and sent me excellent photographs of the type specimens (Figs 3 4). Later, Dr D. Mierzwa (MIZ) kindly gave me on loan the Typus and the Co-Typus of M. ackermanni. Study of this material proved that M. ackermanni is a junior synonym of M. quadrituberculata. Description Ground colour of head, pronotum and mesonotum yellowish to greenish brown, abdomen of male and female ochraceous to brown. Head (Fig. 1). A black mark with three pairs of arms encloses the ocelli: a pair of out curved, narrow, black arms run from median ocellus to the anterior, a pair of somewhat longer, narrow, arms extend from paired ocelli to the anterior, and a pair of straight, tapering, black lines run from paired ocelli to the posterior and sometimes reach black posterior margin of the head. Proximal three fourth of supra-antennal plate black. Pair of black triangular markings extend from just below these plates to level of paired ocelli. Postclypeus with black lines in anterior 5 7 transverse grooves of either side; basal part of postclypeus with broad black transverse mark at clypeal suture, which is medially divided by an upside-down triangle of the ground colour. Anteclypeus black, but triangular basal part, median keel and apical part yellow. Gena with broad black fascia running from antenna to eye. Basal half of lorum black. Rostrum reaching male sternite 3 or female sternite 4; apex brown. Thorax (Fig. 1). Pronotum. Paired central fasciae narrow, strongly widened toward black, anterior pronotum margin and fused at anterior margin of

5 Duffels: Revision of Maua (Cicadidae) Figs 1 2. Maua quadrituberculata, male, Sarawak, Gunung Mulu Nat. Park, Site 7. 1, male body in dorsal view; 2, male abdomen in ventral view. pronotal collar; central fasciae enclose an upsidedown, key-hole shaped median interspace. A pair of low black triangles extends from half-length central fascia along basal half of anterior oblique fissures. Posterior oblique fissures unmarked. Inner side of lateral part of ambient fissure with very broad fasciae that continues in the ambient fissure as a very narrow line reaching to distal ends of posterior oblique fissures; a short, broad, black mark extends from proximal end of broad fascia to black posterior margin of pronotal collar and another (much) smaller black mark is situated just above it. Mesonotum (Fig. 1). Five black fasciae. Median fascia reaching from anterior margin of mesonotum to level of anterior angles of cruciform elevation; the fascia gradually widens from its anterior end to two thirds of its length, where it attains 3 5 times its anterior width, and gradually narrows again to the posterior. Paramedian fasciae extending from anterior mesonotum margin to about half-length of mesonotum. A pair of tiny black spots in front of anterior angles of cruciform elevation. Lateral fasciae somewhat irregular, extending from anterior mesonotum margin to three fifths of mesonotum length and apically narrowed; some specimens have a pair of small dark spots more distally on the mesonotum. Posterior margin of cruciform elevation black to dark brown. Legs. Yellow with greenish tinge and brown to black marking. Fore femora with three spines: a long and narrow, erect, ochraceous, spine with a brown tip at half-length, a somewhat shorter, broad, black spine at four fifths of length from base, and a small, black to brown spine next to the latter. Fore femur with a subapical dark patch on inner and outer side and a dark brown to black fascia on upper side. Fore tibia black but apical one sixth and basal one third of upper side yellow. Middle tibiae with subapical black-brown ring as long as one fourth of tibia. Metatarsus and basal half of mesotarsus of fore and middle legs yellow, apical half of mesotarsus and pretarsus black-brown. Tarsi of hind legs ochraceous. Claws brown.

6 308 Tijdschrift voor Entomologie, volume 152, Figs 3 4. Maua quadrituberculata, male holotype M. ackermanni. 3, body in dorsal view; 4, body in ventral view. Photo s Dr T. Trilar. 4 Tegmina and wings. Hyaline. Tegmina with distinct dark brown spot on transverse veins at bases of second and third apical cells. Small brown spots either at apices of all longitudinal veins, or at apices of apical cells 1 4, or totally missing as in a specimen from Deli, Sumatra. Venation of tegmina light brownish to yellowish variegated with dark brown to blackbrown, venation of wings light to dark brown. Male operculum (Fig. 2). Lateral margin very weakly sinuate and directed slightly inwards. Apical part fairly broadly rounded. Medial margin very weakly convex to medial corner. Operculum reaching to one fourth or one third of abdominal segment 3. Ochraceous. Margin of operculum black, but basal part of lateral margin and basal, incurved, part of medial margin unmarked. Basal part of operculum with black marking next to meracanthus. Male abdomen. Abdomen very broad, widest across segment 3 or 4, segment 3: (1.45) times as wide as pronotal collar, segment 2 only slightly narrower. Dorsal side, including timbal covers, light to dark brown, sometimes with greenish tinge, and with silvery to golden pubescence; basal part of lateral margin of timbal cover dark brown. Tergite 2 with black band of variable width along about two thirds of posterior tergite margin. Posterior margins of tergites 3 6 with a narrow, light brown band that widens to brown to a black-brown triangles at their lateral ends. Tergite 7 laterally dark brown. Tergite 8 brown with irregular dark brown marking. Ventral side light ochraceous, with waxy, pilose, patches. Posterior margin of sternite 2 dark brown and of sternites 3 6 light brown. Sternite 7 brown, turning dark brown to the posterior. Sternite 8 brown but basal parts of lateral margins yellowish. Sternites 3 and 4 with thick, glossy brown, nipple-shaped tubercles; distance between tubercles on sternite times (n=8) as wide

7 Duffels: Revision of Maua (Cicadidae) Figs 5 6. Male pygofer in ventral view. 5, Maua quadrituberculata, Sarawak: Gunung Mulu Nat. Park, Site 7; 6, Maua affinis, holotype. as distance between tubercle and lateral segment margin. Male genitalia (Fig. 5). Pygofer ochraceous, but lateral pygofer lobes and uncus sometimes brown. Basal pygofer lobes consisting of a pair of slightly converging ridges that are broadly connected with lateroventral part of pygofer, reach to three fifths of pygofer length, and are apically acute. Uncus short, trapezoid; apical margin with very weak median incurvation. Basis of uncus inwardly curved and forming two flat and fairly long, about triangular sclerites meeting in midline of pygofer; these sclerites form a short median tube enclosing the aedeagus and a pair of protuberances at their lateral margins. Aedeagus regularly curved, basally very broad, suddenly narrowed at about two thirds of length and ending in a simple acute apex. Female operculum. Greenish ochraceous with mediobasal brown marking and scattered silver pilosity. About as long as wide and just not reaching abdominal segment 3. Lateral margin undulate to rounded rectangular laterodistal angle; posterior margin almost straight; medial margin concave to fairly narrowly rounded mediodistal angle. Female abdomen. Dorsal side light brown with silver pilosity. Tergites 2 6 and epipleurites 2 3 with narrow black bands along their posterior margins; epipleurites 4 6 with somewhat broader and more irregular bands. Tergites 3 6 laterally with small brown triangle. Tergites 7 and 8 dorsally brown, laterally black-brown; posterior margin of tergite 7 dark brown. Segment 9 brown with a pair of irregular, dark brown markings reaching from anterior segment margin to half the segment length. Sternites 3 5 brownish with greenish tinge, and with fairly broad black-brown band along anterior margin and black-brown lateral marking. Sternite 6 shining black-brown with small, median, brownish patch. Sternite 7 shining black-brown. Measurements in mm (6? 1/). Body length?: , /: 31; tegmen length?: , /: 46; head width?: , /: 10.2; pronotum width?: , /: 11.8.

8 310 Tijdschrift voor Entomologie, volume 152, 2009 Fig. 7. Distribution of Maua quadrituberculata (rounds) and M. affinis (triangles). Distribution (Fig. 7) Maua quadrituberculata was originally described from Java, but its occurrence on Java is not confirmed by the present or earlier studies. According to Moulton (1923), the species occurs in Java, Malay Peninsula and Borneo, and is recorded from China and the Philippines. The China and Philippines records of M. quadrituberculata by Moulton (l.c.) and other authors probably all date back to Distant s Monograph of Oriental Cicadidae (1889, 1891). The BMNH has one old specimen of M. quadrituberculata with a label China, but Chou et al. (1991) have not seen a single specimen from China (Chinese text kindly translated by Dr Wei Cong, Yangling, Shaanxi, China). The present study could not confirm the occurrence of M. quadrituberculata in the Philippines and China. The material available for this study confirms the occurrence of M. quadrituberculata in Thailand, Malay Peninsula and Borneo (Sabah, Sarawak and Brunei), and presents new records for Sumatra and Nias Island. Remark My esteemed colleagues Dr Michel Boulard (MNHN) and Mrs Khuankanok Chueata recently collected a new subspecies of M. quadrituberculata in northern Thailand. The subspecies was described as M. quadrituberculata khunpaworensis (Boulard 2007: ). According to its description, the new continental subspecies can be distinguished from the insular nominal subspecies by the more triangular head, the shorter pronotum and the more rounded male opercula. The reader is referred to Boulard s paper for a more detailed taxonomic discussion, the interesting ethological data about the new subspecies and the description of its song. Material examined. 16? 2/. China: Chini (round label with on the reverse side: 63/184, 1? (BMNH). Thailand: Peninsular Siam, Nakon Sri Tamaret, Khao Ram, ft, 23.ii.1922, H.M. Pendlebury, Ex. F.M.S. Museum, B.M , 1? (BMNH), same data but 750 ft, 24.ii.1922, 1? (BMNH). Peninsular Malaysia: Johor: Endau Rompin N.P., Janing Ridge, 20 min N E, primary lowland rainforest, transect primary 4, at light, 27.iii.2001, M.A. Schouten, A. Majanil & M. Renganathan, 1? (ZMAN); Endau Rompin N.P., Janing Ridge, 5 min N E, disturbed forest, transect disturbed 6, at light, 26.iii.2001, M.A. Schouten, 1? (ZMAN). Perak: Doherty, Distant coll , 2? 1/ (BMNH); Batang Padang, Jor Camp, 2000 ft, 2.vi.1923, H.M. Pendlebury,

9 Duffels: Revision of Maua (Cicadidae) 311 Ex. F.M.S. Museum, B.M , 1? (BMNH). Sumatra: Soekaranda, Co-Typus Maua ackermanni Schmidt?, Edm. Schmidt determ. 1924, MIZ , 1? (MIZ); Deli, coll. Dr. D. MacGillavry, Leptopsaltria quadrituberculata, det. McGillavry, 1? (ZMAN). Nias: Noord Nias, Hili, Madjedja, Mitschke, 4 de trim. 95, 1? (BMNH). Borneo: Malaysia: Sabah: Tawau Hill, along Sungai Tawau near trail to Air Panas, river bed in primary riverine forest, 300 m, at light, 28.iii.2001, J.P. & M.J. Duffels, 1?1/ (ZMAN. Sarawak: Gunung Mulu Nat. Park, Site 7, Long Pala (Base), , 50 m, Alluvial/secondary forest, Acl-understorey, J.D. Holloway, RGS Mulu exped., B.M , 1? (BMNH); Gunung Mulu Nat. Park, Site 26, G. Api, Pinnacles, 1200 m, open scrub, MV, , J.D. Holloway, RGS Mulu exped., B.M , 1? (BMNH). Brunei: Waterstradt, 1? (BMNH); Temburong District, ridge NE of Kuala Belalong, approx. 300 m alt, x.1992, J.H. Martin, coll. B.M , 1? (BMNH). Maua affinis Distant Figs 6 9 Maua affinis Distant 1905: 61. Lectotype: Museum Paris / Bornéo / coll. Noualhier 1898, Museum Paris / MNHN (EH) / 4301, Maua / affinis Dist / (type) [examined]. Maua affinis: Distant 1906: 52; Moulton 1912: 133; Moulton 1923: 124, 125, 168 (partim: Borneo); Metcalf 1963: 492 (partim: Borneo). Maua affinis: Zaidi 1996: 101; Zaidi 1997: 113; Zaidi & Ruslan 1998: 352; Zaidi, Ruslan & Azman 1999: 306; Zaidi, Ruslan & Azman 2000: 204; Zaidi, Noramly & Ruslan 2000: 324 (these records of M. affinis by Zaidi and co-authors from various localities in Sarawak and Sabah refer to either M. borneensis or possibly to M. affinis). [Misidentification Maua affinis: Distant 1912: 41, pl. 5 Figs 33a-c (= Maua palawanensis); Chou et al, 1997: 232, Fig. 114 (see remark following the description).] There has been much confusion about the identity of Maua affinis since the specimen depicted by Distant (1912) as Maua affinis belongs to the species that is described in this paper as M. palawanensis. Maua affinis can be separated from M. quadrituberculata by the presence of spots at the basal veins of the fifth and seventh apical cells, the number of 9 10, instead of 5 7, transverse black-brown lines on the postclypeus, the absence of a distinct black mark at the clypeal suture, the larger black spots in front of anterior angles of cruciform elevation, and the presence of black-brown posterior oblique fissures on the pronotum. The types and the identity of Maua affinis The original description of M. affinis by Distant (1906) was followed by Hab. Borneo (Paris and Stockholm) indicating type locality and depositories. The Paris museum has a male specimen with the labels: Museum Paris / Bornéo / coll. Noualhier 1898, Museum Paris / MNHN (EH) / 4301, Maua / affinis Dist / (type) (Distant s handwriting). The Stockholm museum has a male specimen with the labels: Palawan, Staudgr., Maua / affinis / Dist. and NHRS-HEMI (pers. com. Dr G. Lindberg, Stockholm). The London museum has a male specimen with the labels: Balabac / coll. Noualhier 1898, Maua / affinis / Dist, Type [round label with red circle; print], Distant coll / Balabac island, between Borneo and Palawan. All identification labels are in Distant s handwriting The specimen from Paris is designated here as lectotype of M. affinis because it is the only specimen from the type locality Borneo and because it bears the label Maua affinis type in Distant s handwriting. The round Type label of the London specimen was probably not attached by Distant but by a later curator of the museum. The specimens from Stockholm and London are from respectively Palawan and Balabac, an island between Borneo and Palawan, and not from Borneo. Because it is questionable whether these specimens belong to the type series, I have labelled these specimens:?paralectotype Maua affinis Distant, J.P.Duffels Maua affinis is probably endemic to northern Borneo. The identity of M. affinis is established in the description below. The questionable paralectotypes from Palawan and Balabac belong to a new species described in this paper as M. palawanensis which is found on these islands only and not on Borneo. Description of male Ground colour of head, pronotum and mesonotum ochraceous to light brown sometimes with greenish tinge, abdomen of male and female ochraceous to brown. Head (Fig. 8). Black mark enclosing ocelli, marking on supra-antennal plates and triangular markings below these plates as in M. quadrituberculata. Anterior and ventral parts of postclypeus with black-brown transverse lines in the two series of 9 10 grooves; upper 6 pairs of these lines reaching to supra-antennal plates or to lateral clypeal clefts, other streaks shorter. Medial ends of transverse lines of each side connected by a more or less distinct black line; these lines enclose a long oval area at anterior margin of postclypeus and a yellowish wedge-shaped area on ventral side of postclypeus. Lower part of postclypeus at clypeal suture unmarked. Underside of head yellow with black marking. Anteclypeus either black with triangular basal part, median keel and apical part yellow or with a pair of long black

10 312 Tijdschrift voor Entomologie, volume 152, Figs 8 9. Maua affinis, holotype. 8, male body in dorsal view; 9, male abdomen in ventral view. markings, parallel to median keel. Gena with broad black fascia running from antenna to eye. Mediobasal quarter of lorum black. Rostrum reaching almost sternite 3, apex brown. Thorax (Fig. 8). Pronotum. Marking as in M. quadrituberculata, but differing in the following features: A pair of black triangles extends from half-length central fascia along basal half of anterior oblique fissures; each triangle forms a black hook with another black triangle just below the fissure. Posterior oblique fissures black-brown. Mesonotum (Fig. 8). Five black fasciae. Median fascia as in M. quadrituberculata. Paramedian fasciae as in M. quadrituberculata but black spots in front of anterior angles of cruciform elevation larger. Lateral fasciae consisting of an irregular linear part reaching from anterior mesonotum margin to three fifths of mesonotum length, and a pair of distinct black spots. Posterior margin of cruciform elevation black to dark brown. Legs. Yellow with greenish tinge and brown to black marking. Fore femora with three spines: a long and narrow, erect, ochraceous, spine with a brown tip at half-length, a somewhat shorter, broad, mainly black, spine at four fifths of length from base, and a very small, black to brown spine next to the latter. Fore femur with a subapical, irregular, dark marking, a dark brown to black fascia on upper side, and some dark marking at basis. Undersides of middle and hind femora with subapical brown streak. Fore tibia black but apical one sixth and basal two thirds of upper side yellow. Middle tibiae with subapical black-brown ring as long as one fourth of tibia and a light brown ring at short distance from basis. Tarsi and claws as in M. quadrituberculata. Tegmina and wings. Hyaline. Tegmina with an anastomosing spot on transverse veins at bases of first and second apical cell and separate spots at bases of third, fifth and seventh apical cells; some specimens with an anastomosing spot at the bases of the third and fourth apical cells. Tegmina with distinct dark brown marking on transverse veins at bases of the first and second, third, fifth and seventh apical cells; those at bases of first and second apical cells

11 Duffels: Revision of Maua (Cicadidae) 313 connected. Apices of longitudinal veins of apical cells with small brown spots. Venation of tegmina light brownish to yellowish variegated with dark brown to black-brown, venation of wings light to dark brown. Operculum (Fig. 9). Lateral margin sinuate and directed backwards or slightly outwards. Apical part very broadly rounded. Medial margin straight or very weakly convex to medial corner. Operculum reaching to one fourth or half the length of abdominal segment 3. Ochraceous. Lateral margin of operculum, with exception of extreme basal part, with black band that is fairly strongly widened toward its proximal end and continues across the operculum; posterior margin usually black; basal, incurved, part of medial margin unmarked. Abdomen (Fig. 9). Abdomen very broad, widest across abdominal segment 3 or 4, segment 3: times as wide as pronotal collar, segment 2 only slightly narrower. Dorsal side, including timbal covers, light to dark brown and with silvery to golden pubescence; laterobasal corner of timbal cover dark brown. Tergites as in M. quadrituberculata. Ventral side ochraceous. Sternite 3 with a pair of brown triangles at anterior margin. Posterior margins of sternites 3 4 brown. Sternites 5 6 brown but median part ochraceous. Sternite 7 dark brown. Sternite 8 brown but basal parts of lateral margins yellowish. Sternites 3 and 4 with thick, glossy brown, nippleshaped tubercles; distance between tubercles on sternite times (n=4) as wide as distance between tubercle and lateral segment margin. Genitalia (Fig. 6). Pygofer ochraceous to light brown. Basal pygofer lobes consisting of a pair of slightly sinuate ridges that are broadly connected with lateroventral part of pygofer, reach to three fifths of pygofer length, and are apically acute. Uncus rounded, about 1.5 times as long as wide; median shallow furrow reaching to weak median incurvation of distal margin. Basis of uncus inwardly curved and forming two flat, about triangular sclerites meeting in midline of pygofer; these sclerites form a short median tube enclosing the aedeagus and a pair of lateral protuberances at their lateral margins. Aedeagus regularly curved, basally broad, suddenly narrowed at about half its length and ending in a simple acute apex. Measurements in mm (5?). Body length?: ; tegmen length?: ; head width?: ; pronotum width?: Remark on Maua affinis from China The specimen of Maua affinis described and illustrated by Chou et al. (1997: 232, Fig. 114) comes from Mt. Damaosha, Boyang County, Jiangxi province, China ( E, 29 N). Judging from the description and the picture I am convinced that this specimen does not belong to M. affinis but to a new species related to M. palawanensis. The specimen from China differs from M. palawanensis in the shorter and more slender male body (body length: mm; body length of M. palawanensis: 39 mm) and in the lateral fasciae that are broken up in the China specimen and irregular and very variable in width, but not broken up, in M. palawanensis. Dr Wei Cong (Yangling, Shaanxi, China) kindly translated the description of Maua affinis in Chou et al. (1997). Distribution (Fig. 7) This species is probably endemic to Mount Kinabalu. Zaidi and co-authors (see above) recorded M. affinis from various localities in Sarawak and Sabah. These specimens probably belong to either M. borneensis or possibly to M. affinis. Material examined. 5?. Malaysia: Borneo: Sabah: Mt. Kinabalu, Br. North Borneo, Waterstradt, 1? (BMNH) 1? (ZMAN); Mt. Kinabalu, Waterstradt, Leptopsaltria 4-tuberculata Sg., Brit. Mus , 1? (BMNH); Mt. Kinabalu, Mesilau, 8.ii.1964, J. Smart. Royal Soc. Exped. B.M., , 1? (BMNH); Mt. Kinabalu, Mesilau Camp, 31.iii.1964, 5000 ft., Royal Soc. Exped., coll. S. Kueh, B.M , Mau affinis Dist. Michel Boulard det. 1975, 1? (BMNH). Maua borneensis sp. n. Figs 10 12, 14 Zaidi and co-authors recorded M. affinis from various localities in Sarawak and Sabah; these specimens may belong to either M. borneensis or possibly to M. affinis: Maua affinis: Zaidi 1996: 101; Zaidi 1997: 113; Zaidi & Ruslan 1998: 352; Zaidi, Ruslan & Azman 1999: 306; Zaidi, Ruslan & Azman 2000: 204; Zaidi, Noramly & Ruslan 2000: 324. Type material. Holotype?: Borneo: Malaysia: Sarawak: SARAWAK / Gunong Mulu / Nat. Park, Site 27. April / G. Api 1500 m / Pandanus Camp / Scrub / Pandanus Act., J.D. Holloway / RGS Mulu exped. / B.M (BMNH). Paratypes: 8? 2/. Borneo: Malaysia: Sarawak: same data as holotype, 1? (BMNH); Batu Niah, xii.1980, A. Harmann, B.M , 1? (BMNH); Gunong Mulu Nat. Park, Site 1, Camp 4, January, , 1790 m, lower montane (moss) forest, MV- canopy, RGS Mulu exped. B.M , 1? (BMNH); Gunong Mulu Nat. Park, Site 26, April, G. Api, pinnacles, 1200 m, , open scrub, MV, J.D. Holloway, RGS Mulu exped.,

12 314 Tijdschrift voor Entomologie, volume 152, Figs Maua borneensis. 1, male body in dorsal view, holotype; 2, male abdomen in ventral view, paratype, Sabah, 105 km S of Beaufort, airstrip Long Pa Sia. B.M , 1? (UMS); Foot of Mt. Dullit, Junction of rivers Tinjar & Lejok, 3.ix.1932, Oxford Univ. Exp., B.M. Hobby & A.W. Moore, B.M , H185, 1? (BMNH). Sabah: nr Long Pa Sia Long Miau, c m, 27.xi.1987, C. v. Achterberg, 1/ (RMNH); 105 km S of Beaufort, airstrip Long Pa Sia, 4 24 N E, 16.iv.1987, semicultivated area, asl. near disturbed evergreen trop. rainforest, 1000 m, at light, h, Van Tol & Huisman, 1? (RMNH); 15km W Lahad Datu, confl. S. Sabran, S. Danum, S/N, E 4 57 N, 200 m, 23.x.1987, J. Huisman & R. de Jong, 1/ (RMNH); Sandakan dist., Rumidi, R. Labuk, ix.1973, C. Pruett, B.M , 1? (BMNH). Indonesia: Kalimantan: Long Nawang, leg. Mjöberg, 1? (ZMAN). Maua borneensis is alike to M. affinis in the striking similar marking on pronotum and mesonotum and in the marking on the postclypeus, both species lack the black spot at the clypeal suture that is found in the other species of the M. quadrituberculata group. Maua borneensis (body length?: mm) can be separated from M. affinis (body length?: ) by its smaller size and the different marking on the tegmina (see key to the species). Maua borneensis is separated from the other Maua species by the relatively short distance between tubercles on sternite 4, which is times as wide as the distance between tubercle and lateral segment margin; the other species of Maua have ratios ranging from 1.11 to 2.5. Description Ground colour of head, pronotum and mesonotum ochraceous to light brown with greenish tinge, abdomen of male and female ochraceous to brown. Head (Fig. 10). Black mark enclosing ocelli, marking on supra-antennal plates and triangular markings below these plates as in M. affinis, but M. borneensis has an extra pair of drop-shaped brown to black spots proximally of the triangular

13 Duffels: Revision of Maua (Cicadidae) 315 markings. Anterior and ventral parts of postclypeus with black-brown transverse lines in the two series of 9 10 grooves; upper 5 6 pairs of lines reach from a central glabrous area to supra-antennal plates or lateral clypeal clefts; lower 4 5 pairs of lines shorter and with medial ends of lines of each side connected by a black line; the black lines of both sides enclose a yellowish wedge-shaped median area; lowest two pairs of transverse lines sometimes fused to a pair of black markings. Lower part of postclypeus at clypeal suture unmarked. Underside of head yellow with black marking. Anteclypeus black, but triangular basal part, median keel and apical part yellow. Gena with black fascia running from antenna to eye. Basal half of lorum black. Rostrum reaching sternite 3; apex brown. Thorax (Fig. 10). Marking as in M. affinis, but greatest width of median mesonotal fascia 4 7 times its anterior width. Legs. Yellow with greenish tinge and brown to black marking. Fore femora with three spines: a long and narrow, erect, ochraceous, spine with a brown tip at half-length, a somewhat shorter, broad, blackbrown, spine at four fifths of length from base, and a very small, black to brown spine next to the latter. Fore femur with a subapical, irregular, dark marking, a dark brown to black fascia on upper side, and some dark marking at basis. Fore tibia black but apical one sixth and basal one third to basal half of upper side yellow. Middle tibiae with subapical black-brown ring as long as one fourth or one third of tibia. Metatarsus and mesotarsus of fore and middle legs yellow, pretarsus black-brown, tarsi of hind legs ochraceous, claws brown. Tegmina and wings. Hyaline. Tegmina with brown spots on transverse veins at bases of the second, third and fifth apical cells and in most specimens with small brown spot at basis of seventh apical cell. Apices of longitudinal veins of apical cells in most specimens with small to hardly discernible brown spots. Venation of tegmina light brownish to yellowish variegated with dark brown to black-brown, venation of wings light to dark brown. Male operculum (Fig. 11). Triangular. Lateral margin directed inwards, weakly sinuate in basal one fourth, and slightly concave or straight to narrowly rounded apical part of operculum. Medial margin straight or weakly concave to medial corner. Operculum reaching to two fifths or three fifths of abdominal segment 3. Ochraceous, with small brown marking at base, and a black-brown rim except in the sinuate basal part of lateral margin and the basal part of medial margin proximal of medial corner. Male abdomen (Fig. 11). Broad, widest across abdominal segment 2 or 3, segment times as wide as pronotal collar. Dorsal side, including timbal covers, light to dark brown and with greyish pubescence; laterobasal corner of timbal cover dark brown. Tergite 2 with black band of variable width along about two thirds of posterior margin. Posterior margins of tergites 3 5 or 3 6 with narrow, light brown bands. Tergite 7 laterally dark brown. Tergite 8 dark brown to black. Ventral side ochraceous. Posterior margin of sternites 3 4 sometimes brown. Sternite 5 with posterior and lateral parts light brown. Sternite 6 dark brown but median part ochraceous. Sternite 7 black-brown. Sternite 8 brown but laterobasal parts of lateral margins ochraceous. Sternites 3 and 4 with fairly thick, light brown to brown, but basally often ochraceous, nipple-shaped tubercles; distance between tubercles on sternite times (n=8) as wide as distance between tubercle and lateral segment margin. Male genitalia (Fig. 12). Male genitalia as in M. quadrituberculata but lateral sides of uncus less oblique. Female operculum. Ochraceous to brownish with scattered silver pilosity, mediobasally somewhat darker brown. Somewhat wider than long, medially reaching to almost hind margin of abdominal segment 3, laterally reaching to two thirds of segment length. Lateral margin with triangular protrusion in basal half, apical half brown and weakly convex to rounded rectangular laterodistal angle; posterior margin weakly undulate; medial margin concave to narrow mediodistal angle. Female abdomen. Dorsal side (light) brown with silver pilosity. Tergites 2 6 and epipleurites 2 3 with narrow black bands along their posterior margins; epipleurites 4 5 with somewhat broader and more irregular black-brown bands. Tergites 3 5 with brown lateral spot. Lateral parts of tergites 6 8 and epipleurites 6 8 black-brown. Segment 9 brown and with short or fairly long, dark brown marking along posterior margin of segment 8. Sternites 3 5 light brown with fairly broad, black-brown band along posterior margin; sternite 3 with black-brown band along anterior margin, and sternites 4 5 with black-brown laterobasal marking. Sternites 6 7 shining black-brown, sometimes with indistinct median brownish patch. Measurements in mm (8? 2/). Body length?: 31 35, /: ; tegmen length?: 43 47, /: 40; head width?: , /: ; pronotum width?: , /: Distribution (Fig. 14) Maua borneensis seems to be endemic to Borneo (Sabah, Sarawak and Kalimantan). Dr Zaidi Mohd. Isa and co-workers (UKM) published the results of several cicada inventories in Malaysia. The specimens

14 316 Tijdschrift voor Entomologie, volume 152, Figs Male pygofer in ventral view. 12, Maua borneensis, holotype; 13, M. palawanensis, Palawan. from Sabah and Sarawak identified in their papers as M. affinis are not included in the distribution map since their identity could not be confirmed. Maua palawanensis sp. n. Figs Maua affinis Distant 1905: 61 (?partim: the specimens from Balabac (BMNH) and from Palawan (NHRS) listed below as holotype and paralectotype of M. palawanensis may be part of the type series of M. affinis). [Misidentification.] Maua affinis: Distant 1912: 41, pl. 5 figs 33a-c; Moulton 1912: 133; Moulton 1923: 124, 125, 168 (partim: specimens from Palawan and Balabac); Metcalf 1963: 492 (partim: Palawan and Balabac). [Misidentifications.] Type material. Holotype?: Balabac: Balabac / coll. Noualhier 1898, Maua / affinis / Dist [Distant s handwriting], Type [round label with red circle; print], Distant coll/ ,? paralectotype of Maua affinis (BMNH). Paratypes: Palawan: Palawan, Staudgr., Maua affinis Dist. [Distant s handwriting], NHRS-HEMI , 1?? paralectotype of Maua affinis (NHRS); Palawan I., 1? (BMNH). Maua palawanensis can be separated from the other large Maua species by a combination of characters: the presence of a broad, black, transverse mark at the clypeal suture, the irregular, but continuous, lateral mesonotal fasciae reaching from anterior to almost posterior margin of mesonotum, and paramedian mesonotal fasciae that are often apically connected with the median fascia. This species is known from Palawan and from Balabac Island between Palawan and Borneo. Description of male Ground colour of head, pronotum and mesonotum yellowish to greenish brown, abdomen ochraceous to brown. Head (Fig. 15). A black mark with three pairs of arms encloses the ocelli, a pair of out curved, fairly broad,

15 Duffels: Revision of Maua (Cicadidae) 317 Fig. 14. Distribution of Maua borneensis (rounds), M. palawanensis (triangles) and M. latilinea (squares). black arms run from median ocellus to the anterior, a pair of narrow arms extend from paired ocelli to the anterior, and a pair of straight, short, tapering, black lines run from paired ocelli to the posterior. Proximal three fourths of supra-antennal plate black. A pair of black triangular markings extends from just below these plates to level of paired ocelli. Postclypeus ochraceous with a median light coloured fascia almost reaching the clypeal suture and blackbrown lines in all (7 9) transverse grooves. Medial ends of lower three transverse lines of each side connected by a black line. Basal part of postclypeus with a broad, black, transverse mark at clypeal suture, medially divided by an upside-down triangle of the ground colour. Anteclypeus with a pair of paramedian, black markings, but margins and median keel of anteclypeus ochraceous. Gena with fairly broad, black fascia running from antenna to eye. Basal two thirds of lorum black. Rostrum reaching sternite 3, apex brown. Thorax (Fig. 15). Pronotum. Marking as in M. affinis. Mesonotum (Fig. 15). Five black fasciae. Median fascia reaching from anterior margin of mesonotum to level of anterior angles of cruciform elevation; the fascia widens initially gradually, and then suddenly, to three fifths of its length, where it attains 5 times its anterior width, and subsequently narrows from the broadest part to its posterior end. Paramedian fasciae reach from anterior mesonotum margin to about half-length of mesonotum and are apically connected with the suddenly widening part of the median fascia. A pair of medium-sized, round, black spots in front of anterior angles of cruciform elevation. Lateral fasciae reaching from anterior mesonotum margin to level of round black spots in front of cruciform elevation; the fascia is irregular and very variable in width: broad in anterior one seventh, very thin and almost broken up in next one seventh, widest at half-length and narrowing from half-length to posterior end. Posterior margin of cruciform elevation black-brown. Legs as M. quadrituberculata. Tegmina and wings. Hyaline. Tegmina with distinct dark brown spot on transverse veins at bases of second and third apical cells and a small spot at base of fifth apical cell; apices of longitudinal veins of apical cells with small brown spots. Venation of tegmina light brownish to yellowish variegated with dark brown to black-brown, venation of wings light to dark brown.

16 318 Tijdschrift voor Entomologie, volume 152, Figs Maua palawanensis, male paratype, Palawan. 15, body in dorsal view; 16, abdomen in ventral view. Operculum (Fig. 16). Lateral margin weakly sinuate in basal one third, distal two thirds straight and directed slightly inwards to rounded laterodistal corner. Medial margin almost straight to medial corner. Operculum reaching to one third of abdominal segment 3. Ochraceous with greenish tinge, but lowest basal part of operculum dark brown. A broad dark brown band runs from one third of length of lateral margin, along laterodistal corner and posterior margin, and continues in a narrow brown band in the basal, incurved, part of medial margin. Abdomen (Fig. 16). Abdomen very broad, widest across abdominal segment 3 or 4, segment 2 only slightly narrower, segment times as wide as pronotal collar. Dorsal side castaneous, timbal covers with greenish tinge, basal part of lateral margin of timbal cover dark brown. Tergite 2 with black band of variable width along posterior margin, and with narrow median triangle rising from black band to two thirds of segment length. Posterior margins of tergites 3 5 with somewhat narrower, black bands, that widen to brown to black-brown triangles at their lateral ends. Tergite 6 with narrow black band along posterior margin. Lateral sides of tergites 6 7 dark brown. Tergite 8 dark brown with castaneous median marking. Ventral side ochraceous with greenish tinge, with waxy, pilose, patches. Sternite 3 with a pair of brown triangles at anterior margin. Posterior margins of sternite 2 with dark brown band, of sternites 3 5 with light brown bands, that on sternite 5 broader than on sternites 3 4. Sternite 6 brown. Sternite 7 dark brown but laterobasal corners light brown. Sternite 8 brown with basal yellowish triangle covering about half of sternite surface. Sternites 3 and 4 with fairly thick, glossy brown, nipple-shaped tubercles; distance between tubercles on sternite times (n=1) as wide as distance between tubercle and lateral segment margin; tubercles dark brown with exception of the lighter brown proximal sides. Genitalia (Fig. 13). Pygofer ochraceous, but lateral pygofer lobes and uncus light brown. Basal pygofer lobes consisting of a pair of slightly converging ridges

17 Duffels: Revision of Maua (Cicadidae) Figs Maua latilinea, male, Kalimantan, Kubing. 17, body in dorsal view; 18, abdomen in ventral view. that are broadly connected with lateroventral part of pygofer, reach to three fifths of pygofer length, and are apically acute. Uncus short, trapezoid with a median lanceolate elevation close to apical margin. Basis of uncus inwardly curved and forming two flat and short, triangular sclerites meeting in midline of pygofer; these sclerites form a short, thick, median tube enclosing the aedeagus and a pair of round protuberances at their lateral margins. Aedeagus regularly curved, basally very broad, gradually narrowed at about half its length and ending in a simple acute apex. Measurements in mm (1?). Body length: 39; tegmen length: 48; head width: 10.7; pronotum width: Distribution (Fig. 14) This species is known from Palawan and from Balabac Island, between Borneo and Palawan. Maua latilinea (Walker) Figs 14, Dundubia latilinea Walker, 1868: 85, 87. Holotype /: Penang [above horizontal line], Wallace [below horizontal line; handwritten], latilinea [Walker s handwriting], Type [round label with green circle; print] (BMNH) [examined]. Cosmopsaltria latilinea: Distant 1889: 48 (partim, as far as the text refers to the female holotype from Penang). Maua dohrni Schmidt, 1912: 65, 66. Holotype?: Soekaranda / Januar 1894 / Dohrn [print in black box], Maua / dohrni / Schmidt /? Edm. Schmidt / determ. 1912, Type [print in black box; red paper], MIZ (MIZ) [examined]. syn. n. Maua latilinea: Moulton, 1923: 92, 124, 126, 168; Metcalf, 1963: 493, 494. [For further references before 1980 see: Metcalf 1963]. [Misidentification: Cosmopsaltria latilinea (nec Walker); Distant 1889: 48 (partim), Pl. iv, Figs 15, 15a, b. (the specimen described and depicted here belongs to Orientopsaltria padda (Distant, 1897) (see Duffels & Zaidi 2000: 209).]

18 320 Tijdschrift voor Entomologie, volume 152, 2009 Study of this material proved that M. dohrni is a junior synonym of M. latilinea. Fig. 19. Maua latilinea, male Kalimantan Kubing, pygofer in ventral view. This paper presents the first description of the male of M. latilinea. This species is easily separated from the other large Maua species by the shape of the central mark enclosing the ocelli, the characteristic marking on the postclypeus, the absence of a broad, black fascia along inner side of lateral part of ambient fissure, the relatively narrow ochraceous interspace between central pronotal fasciae, the broad and straight lateral fasciae on the mesonotum that reach from the anterior mesonotum margin to almost hind margin of mesonotum, the cup-shaped and triangular markings on respectively tergites 2 and 3 and the very characteristic shape of uncus and basal pygofer lobes. Synonymy In February 2008, Dr Tomi Trilar of the Prirodoslovni Muzej Slovenije, Ljubljana visited the Museum of the Institute of Zoology, Polish Academy of Science, Warsaw, Poland. He found the Type and the Co-Typus of M. dohrni Schmidt, 1912 in the collection of this museum, brought these to my attention and kindly sent me excellent photographs of the type specimens (Figs 20 21). Later, Dr D. Mierzwa (MIZ) was so kind to send me the Type and the Co-Typus of M. dohrni on loan. Description Ground colour of head, pronotum and mesonotum brown; abdomen of male ochraceous to brown, abdomen of female with reddish tergites and blackbrown sternites. Head (Fig. 17). Ocelli enclosed by black mark with a pair of broad, black, diverging fasciae reaching from left paired ocellus and median ocellus to the anterior and from right paired ocellus and median ocellus to the anterior; the fasciae are for the greater part fused but separated at frontoclypeal suture by a median triangle of the ground colour; in some specimens the anterior ends of each fascia enclose a semi-oval spot of ground colour at the frontoclypeal suture but other specimens have a narrow area of the ground colour extending along the frontoclypeal suture. A pair of straight, narrow to fairly broad, black lines run from paired ocelli to posterior margin of head. Proximal two thirds of supra-antennal plate black, and connected with a pair of black triangular markings that extend to level of paired ocelli. Inner margin of eye with irregular black marking. Postclypeus with black lines in upper 8 transverse grooves of either side; upper 5 pairs of lines reaching to supraantennal plates or to lateral clypeal clefts, other lines shorter. Medial ends of transverse lines of each side connected by a black line; the two lines connecting the upper 5 pairs of lines enclose a glabrous oval area of the ground colour at anterior margin of postclypeus; the lower 3 pairs of lines and a pair of large black marks in basal two fifths of postclypeus enclose a narrow median space reaching to clypeal suture. Anteclypeus black, but triangular basal part, median keel and apical part yellow. Gena with very broad black fascia running from antenna to eye. Basal half of lorum black. Rostrum reaching to one sixth of male sternite 3; apex brown. Thorax (Fig. 17). Pronotum. Paired central fasciae fairly broad, strongly widened toward anterior pronotum margin and almost fused at anterior margin of pronotal collar; the central fasciae enclose a narrow median interspace that is equally broad along its whole length. A pair of narrow black triangles extends from half-length central fascia along basal one third of anterior oblique fissures. Anterior oblique fissures unmarked. Posterior oblique fissures black in anterior two third part, but entirely black in specimen from Karimata Island. Inner side of lateral part of ambient fissure without fascia. Pronotal collar with a broad, black mark along medioproximal corner of ambient fissure, medial part of mark reaching to black posterior margin of pronotal collar.

19 Duffels: Revision of Maua (Cicadidae) 321 Mesonotum (Fig. 17). Five black fasciae. Median fascia widens from anterior margin of mesonotum to 2 3 times its anterior width at one third of its length, slightly narrows at about half length, widens again to 3 4 times its anterior width at two thirds of length, and strongly narrows to its posterior end at level of anterior angles of cruciform elevation. Paramedian fasciae fairly broad and extending from anterior mesonotum margin to about half-length of mesonotum. A pair of small black spots in front of anterior angles of cruciform elevation. Lateral fascia on mesonotum broad and straight, and reaching from anterior mesonotum margin to almost hind margin of mesonotum. Posterior margin of cruciform elevation black to dark brown. Legs. Ochraceous with brown to black marking. Fore femora with three spines: a long and narrow, erect, black-brown, spine at half-length, a somewhat shorter, broad black spine at four fifths of length from base, and a small, black to brown spine next to the latter. Fore femur with a subapical dark patch on inner and outer side and a dark brown to black fascia on upper side. Fore tibia dark brown but basal three fourths of upper side yellow. Middle tibiae with subapical black-brown ring as long as one fourth of tibia. Meta- and mesotarsus of fore and middle legs, and basal one third of pretarsus ochraceous, apical part of pretarsus brown; tarsi of hind legs ochraceous. Claws brown. Tegmina and wings. Hyaline. Tegmina with distinct dark brown spot on transverse veins at bases of second and third apical cells and with small brown spots at apices of longitudinal veins of apical cells 1 3. Venation of tegmina and wings light to dark brown. Male operculum (Fig. 18). Triangular. Lateral margin very weakly sinuate in basal one third, apical part straight or slightly concave and directed inwards. Apical part of operculum narrowly rounded. Medial margin weakly convex to medial corner. Operculum reaching to one fifth of abdominal segment 3. Ochraceous, but dark brown to black in medial one third of basal part and along apical and medial margins. Male abdomen (Fig. 18). Abdomen broad, widest across abdominal segment 3 or 4, segment 3: times as wide as pronotal collar, segment 2 only slightly narrower. Dorsal side, including timbal covers, brown, with greyish pubescence. Tergite 2 with cup-shaped or trapezoid median black marking between anterior and posterior tergite margins and with black band of variable width along posterior margin. Tergite 3 with median black triangle, about as wide as cruciform elevation at anterior tergite margin, reaching to three fourths of tergite length, and narrowly (in the paratype: broadly) connected with black posterior tergite margin; tergite 3 also with a pair of sublateral rectangular brown markings. Tergites 4 7 with dark brown to black-brown median, transverse marking at anterior margins reaching to one third or half the segment length, a pair of dark brown to black-brown sublateral markings, and dark brown to black-brown bands, one fifth to one fourth as high as the segment, along posterior margins. Tergite 8 brown with irregular dark brown marking. Ventral side light ochraceous. Posterior margin of sternite 2 dark brown. Sternite 4 with a pair of large brown triangles reaching from anterior sternite margin to four fifths of its length. Sternites 4 5 light brown, sternite 7 8 dark brown. Sternites 3 and 4 with fairly narrow, brown to blackbrown, nipple-shaped tubercles with distinct transverse ridges; distance between tubercles on sternite times (n=4) as wide as distance between tubercle and lateral segment margin. Male genitalia (Fig. 19). Pygofer oval-shaped. Basal pygofer lobes form a pair of large, black-brown, spine-shaped projections reaching from two fifths to three fourths of pygofer length. Uncus triangular with straight lateral margins, and strongly sclerotised. Underside of the uncus forming a pair of fairly short, narrowing lobes with a lateral ridge, that sustain the apex of the aedeagus. Aedeagus regularly curved and gradually narrowing to the simple acute apex. Pygofer ochraceous, but distal half of lateral sides, apical half of basal pygofer lobes and uncus black- brown. Female operculum. Lateral margin straight, laterodistal corner rounded rectangular, distal margin undulate. Medial part of operculum blackish. Female abdomen. Tergites, especially tergites 2 and 3, reddish. Cup-shaped median black marking on tergite 2 and median black triangle on tergite 3 as in male. Segment 4 with a median irregular black marking. Anterior parts of segment 5 7 blackish suffused. Posterior margins of all tergites with black bands, those on segments 5 7 broader than on the other tergites. Sternites black-brown. Valvae blackbrown. Segment 9 with broad, black pattern along lower margins. Measurements in mm (4?). Body length: ; tegmen length: ; head width: ; pronotum width: Distribution (Fig. 14) According to its labels, the female type specimen of Dundubia latilinea was collected in Penang and comes from the Wallace collection. Dr George Beccaloni (BMNH; personal communication 2007) kindly brought to my attention a remark in the introduction of Wallace s Catalogue of the Cetoniidae of the Malayan Archipelago (1868: 522): The specimens collected by Mr. Lamb, and noted as from

20 322 Tijdschrift voor Entomologie, volume 152, 2009 Figs 20. Maua latilinea, male holotype M. dohrni, body in dorsal view. Photo Dr T. Trilar. Penang, were most of them collected, not in the island, but in the main land of the Malay Peninsula opposite, termed the Province of Wellesley. Penang may therefore be held to signify merely a particular sub-district of the Malay Peninsula. Up to now M. latilinea was only known from the Malay Peninsula after the female holotype from Penang (see above) and another female from Pahang (Moulton 1923). The present revision of the genus Maua does not confirm the occurrence of this species in the Malay Peninsula but reveals that Maua latilinea certainly occurs in Borneo and Sumatra. Two specimens of M. latilinea studied come from respectively, the west coast of Borneo viz., Karimata Island and Ketapang, Kubing near the coast of Kalimantan Barat opposite Karimata Island. The type specimens of Maua dohrni, a junior synonym of M. latilinea, from Soekaranda, Sumatra, establish its occurrence in North Sumatra (see next paragraph). Figs 21. Maua latilinea, male holotype M. dohrni, body in ventral view. Photo Dr T. Trilar. The locality Soekaranda of [W. L.] H. Dohrn The type-specimens of Maua dohrni and M. ackermanni were collected by Dohrn in Soekaranda [= Sukaranda], Sumatra. In the Atlas van Tropisch Nederland (Anonymous 1938) Soekaranda is spelled as Soekaradja; in the Gazetteer Nasional NamaNama Geografie (Anonymous, 1978) it is spelled as Sukaraja. Both sources give three localities in Sumatra, all in the province of Bengkulu

21 Duffels: Revision of Maua (Cicadidae) Figs Maua species, male body in dorsal view. 22, M. albigutta, Malaysia, Johor, Endau Rompin N.P.; 23, M. linggana, Sabah, 23 km W Sandakan, Sepilok tree tower. in West Sumatra. A further search for the coordinates of Dohrn s Soekaranda revealed a publication by Schneider (1906) mentioning the visit of Dr G. Schneider in 1897 to Dr [W.L.]H. Dohrn who lived in a village called Sukaranda that is located in Upper Langkat in North Sumatra and not in the province of Bengkulu. These data are in accordance with the coordinates of Sukaranda as given by Van Doesburg (2004: 107): 3 37 N and E. Wolfgang Ludwig Heinrich Dohrn ( ) was the son of the coleopterist Carl August Dohrn ( ) and brother of the well-known hemipterist Anton Dohrn ( ). He collected in Sumatra for the Museum für Naturkunde in Stettin (Horn & Kahle, 1935: 59). The material collected by Wolfgang Dohrn is now in the Museum of the Institute of Zoology, Warszawa, Poland. Material examined. 4?. Indonesia: Sumatra: Soekaranda, i.1894, Co-Typus Maua Dohrni Schmidt?, Edm. Schmidt determ. 1912, MIZ , 1? (MIZ). Borneo: Kalimantan: Kubing, Kec. Ketapang, 80m, 7.xi.1999, S E, Oilpalm plantation, near riverine forest, at light, M. Lammertink 1? (ZMAN); W. Kalimantan, Karimata Island Nat. Ketapang Res, ii.1991, IIS , per Sutrisno Djenal, General Collections primary rainforest, closed canopy, 350 m, LS, 1? (ROM). Mau albigutta group The M. albigutta group consists of two mediumsized species (body length of male: mm): M. albigutta and M. linggana. Lateral mesonotal fasciae consisting of three distinct black spots or short lines on a row. Apical cell 1 of the tegmen is relatively long. The basal vein of apical cell 1 is more than half as long as the longitudinal vein of apical cell 1. The male abdomen is fairly broad and about equally wide across the segments 2 5. Sternites 3 and 4 have a pair of narrow protuberances, which are attached to the sternites along their whole length. The basal pygofer lobes are broadly rounded ridges that widen from the base of the pygofer to three fifths of its length.

22 324 Tijdschrift voor Entomologie, volume 152, Figs M. albigutta, Malaysia, Johor, Endau Rompin N.P. 24, abdomen in lateroventral view; 25, male pygofer in ventral view. Mau albigutta (Walker) Figs 22, Dundubia albigutta Walker, 1857: 83. Holotype?: Sumatra / Sir S. Raffles, / Vigors Coll., albigutta / n s. [handwritten], Type [print in round label with green margin] (BMNH) [examined]. Leptopsaltria albiguttata [sic]: Distant, 1889: 36; Distant, 1891: pl. viii Figs 8, 8a-b; Distant 1892 pl. x Figs 4, 4a-b. Purana albigutta: Metcalf 1963: ; Duffels & Van der Laan 1985: 105; Zaidi & Ruslan 1997: 221; Zaidi, Noramly & Ruslan 2000: ; Zaidi, Azman & Ruslan 2001: 109, 110, 112, 115. Maua albigutta: Moulton 1923: 121, 124, 125, 168; Gogala et al. 2004: 1 14; Schouten et al. 2004: [For further references before 1980 see: Metcalf 1963 and Duffels & Van der Laan 1985]. The males of M. albigutta (body length: mm) are slightly smaller than those of M. linggana ( mm). Maua albigutta and M. linggana can also be distinguished by the shape of the male opercula. The male opercula of M. albigutta (Fig. 24), strongly taper to the apex, possess a distinct medial protrusion and reach to half length abdominal segment 3, while those of M. linggana are broadly rounded apically, have a less distinct median protrusion and reach or reach just beyond posterior margin of abdominal segment 2 (Fig. 27). Maua albigutta can be separated from Purana species of the same size by its broad abdomen, characteristic male operculum, the brownish suffused apical part of the tegmina, the three distinct black spots on a row forming the lateral mesonotal fascia, the dark brown to black posterior margins of the male abdominal segments, and the morphology of the male genitalia. Synonymy Moulton (1923) synonymised Purana jacobsoni Distant, 1912 with Maua albigutta. The type of P. jacobsoni in BMNH comes from Java and has the following labels: Purana / jacobsoni / type Dist, Jacobson / Depok / Java / Nov. 1908, Java / (Jacobson) / , Type [round label with red circle; print], 11/15. I have compared this type with

23 Duffels: Revision of Maua (Cicadidae) 325 M. albigutta, and found that P. jacobsoni and M. albigutta differ e.g., in the male opercula and the basal and lateral pygofer lobes of the male genitalia. Description Ground colour of head, pronotum and mesonotum ochraceous to light brown often with greenish tinge, abdomen of male and female ochraceous to brown. Head (Fig. 22). Each of the ocelli enclosed by a black triangle, apices of the three triangles point to distal end of epicranial suture. Triangle around median ocellus with distinct protrusions pointing laterodistally, triangles around paired ocelli sometimes with weak protrusion pointing to the posterior. Supraantennal plates with distinct black marking. A pair of right angled hook-shaped lines with equally long legs extend from just below supra-antennal plates to level of paired ocelli. Inner margin of eyes with a pair of narrow black spots. A pair of juxtaposed half-round spots in line with paired ocelli at posterior margin of head. Anterior and ventral parts of postclypeus with two series of black-brown transverse lines in all (8 9) transverse grooves; upper 3 4 pairs of lines reach from a central glabrous area to supra-antennal plates or lateral clypeal clefts, the other transverse lines (much) shorter. Medial ends of upper 5 6 pairs of transverse black lines of each side connected by a black line, the connecting black lines of both sides enclose a glabrous oval area at anterior margin of postclypeus; lowest two pairs of transverse lines sometimes fused to a pair of black markings. Lower part of postclypeus at clypeal suture unmarked. Anteclypeus with a pair of paramedian, black-brown, oblong markings parallel to keel. Gena with short black fascia running from postclypeus to half or two thirds of width of gena. Lorum with black mediobasal corner. Rostrum reaching almost posterior margin of abdominal segment 2. Thorax (Fig. 22). Pronotum. Paired central fasciae narrow, strongly widened toward black, anterior pronotum margin and fused at anterior margin of pronotal collar; the central fasciae enclose an upsidedown, key-hole shaped median interspace. A pair of short lines extends from half-length central fascia along basal one third or half of anterior oblique fissures; each line forms a black hook with an oblong spot just below the fissure. Posterior oblique fissures black-brown. Inner side of lateral part of ambient fissure with a relatively broad fascia that continues in the ambient fissure as a very narrow line reaching to distal ends of posterior oblique fissures. Pronotal collar with a pair of light brown markings at lateral side of ambient fissure. Mesonotum (Fig. 22). Five black fasciae. Median fascia, reaching from anterior margin of mesonotum to one third of length of cruciform elevation, very narrow in anterior half, suddenly widened to 3 times its anterior width at two thirds of its length and gradually narrowed again to the posterior. Paramedian fasciae reaching from anterior margin to about half-length of mesonotum, as wide as anterior part of median fascia, but basally wider and also widening to its distal end, which is sometimes recurved. A pair of black spots in front of anterior angles of cruciform elevation. Lateral fasciae on mesonotum consisting of three distinct black spots on a row. Legs. Yellow to light brown with brown to black marking. Fore femora with three dark brown to black-brown spines: a long and narrow, erect spine at one third of length from base, a somewhat shorter and broader spine at four fifths, and a small spine next to the latter; apical part of femur and a fascia on upper side dark brown. Fore tibia with dark brown fascia on underside. Tarsi of fore and middle legs brown sometimes with greenish tinge, hind tarsi ochraceous, claws brown. Tegmina and wings. Hyaline. Tegmina with apical part of apical cells 1 2 brownish suffused, with a brown spot on transverse veins at bases of second and third apical cells, and with small brown spots at apices of longitudinal veins of apical cells 1 4. Venation of tegmina light brownish to yellowish variegated with dark brown to black-brown, venation of wings light to dark brown. Male operculum (Fig. 24). Triangular with a distinct medial protrusion. Lateral margin oblique, sinuate at base and slightly concave at half-length. Operculum reaching to half-length abdominal segment 3. Ochraceous, sometimes with greenish tinge; concave part of lateral margin sometimes brownish. Male abdomen (Fig. 24). Abdomen broad, widest across segment 3 or 4, segment 3: times as wide as pronotal collar. Timbal covers, light brown, sometimes with greenish tinge. Tergites 2 5 light brown, sometimes with greenish tinge, tergite 6 light castaneous brown, tergites 7 8 castaneous, all tergites with silvery pubescence. Posterior margins of tergites 1 7 with narrow, dark brown fasciae. Tergites 3 7 with faint lateral spots. Ventral side light ochraceous. Sternites 6 8 castaneous brown. Sternites 3 and 4 with a pair of narrow, brown protuberances, which are attached to the sternite along their whole length; those on sternite 4 somewhat smaller than on sternite 3. Distance between tubercles on sternite times (n=8) as wide as distance between tubercle and lateral segment margin. Male genitalia (Fig. 25). Pygofer ochraceous, but lateral pygofer lobes dark brown. Basal pygofer lobes rounded and widening from base to three fifths of pygofer length. Uncus short, trapezoid; apical

24 326 Tijdschrift voor Entomologie, volume 152, 2009 Fig. 26. Distribution of Maua albigutta (rounds), M. linggana (triangles) and M. platygaster (squares). margin with very weak median incurvation. Basis of uncus inwardly curved and forming two triangular sclerites meeting in midline of pygofer; these sclerites form a pair of lateral, round, protuberances at their lateral margins and a short median tube enclosing the aedeagus. Aedeagus regularly curved and gradually narrowing to a simple acute apex. Female operculum. Lateral margin convex, laterodistal corner broadly rounded, distal margin very weakly undulate. Female abdomen. Tergites 2 6 with a very narrow brown to black line along posterior margins, but on tergites 3 4 the central part of this line, which is as wide as the cruciform elevation, is about as high as one third of tergite. Lateral parts of tergite 7 with brown to black marking on anterior two thirds. Sternites 4 6 with brownish central marking, sternite 7 black-brown along anterior margin. Valvae blackbrown. Segment 9 dorsally with a pair of paramedian black triangles and ventrally with black-brown marking along lower segment margin. Measurements in mm (8? 5/). Body length?: 24 28, /: ; tegmen length?: 30 34, /: ; head width?: , /: ; pronotum width?: , /: Biology Maua albigutta is a common species in localities like Endau Rompin N.P., Johor, Malaysia (Schouten et. al. 2004) and Langkawi Island (Zaidi, Azman & Ruslan 2001). In Endau Rompin N.P., aggregations of singing males were found low on the tree trunks in March The species was less common from August to November 1999 and males were singing higher up in the canopy. Dr Matija Gogala and Dr Tomi Trilar (PMSL) recorded and described its very characteristic and attractive sound (Gogala et al. 2004). The calling song is complex and has pronounced frequency modulated parts. From video recordings, made in nature in Endau Rompin, the movement of the abdomen could be attributed to particular characteristics of the song. Distribution (Fig. 26) Maua albigutta is widely distributed in the Malay Peninsula and Sumatra, and known from Siberut Island. Zaidi, Noramly & Ruslan (2000) recorded this species from Borneo (Sabah, Tibow, Tawau). In spite of intensive collecting in Sabah and Sarawak in the last decades, this is the only record of M. albigutta from Borneo. I have not seen the Sabah

25 Duffels: Revision of Maua (Cicadidae) 327 specimens, so that the occurrence of M. albigutta on this island could not to be confirmed. Material examined. 79? 17/. Indonesia: Sumatra: Brastagi [= Berastagi], , Corporaal 1/ (ZMAN); Fort de Kock, 920 m, 1925, leg. E. Jacobson, 1? (ZMAN); Labuan Bilik [Labuanbilik], HE 5329, , Palm, 15? 3? (MZHF); Lau Rakit, i-iv.1919, Corporaal, 1? (ZMAN); Medan, Pagger Alam, coll. Dr MacGillavry, vi. 1885, Debussy, 1? (ZMAN); Padang, Ex F.M.S. Museum, B.M , Raffles Museum Singapore, xi.1924, C.B.K. & N.S., 9? (MNKM); Lampung Krui: Lampung Krui: Pahmungan E 5 11 S, Damar garden singing on trunk Shorea javanica, 5.xi.2001, K. Smets, 1? (ZMAN); Penengahan E 5 08 S, Damar garden, 12.xi.2001, K. Smets, 1? (ZMAN) 1? (MZB); Sumatra occ., M.R. Belg., 7866, van Lansberg, 2? (KBIN); Sumatra, 746, R. Mus. Hist. Nat. Belg. I.G , 1? (KBIN). Siberut Island: Ex F.M.S. Museum, B.M , Raffles Museum Singapore, , C.B.K. & N.S., 1? (MNKM). Peninsular Malaysia: Kedah: Kota Walamuir, 2.i.1982, Marzuri, 1? (PMSL). Kelantan: Panjang Kelantan, 21.i.1983, Samsuddin Matzam, 1? (MNKM); Salor, 20.i.1982, Said Tarmizi, 1? (PMSL). Pahang: Cameron Highlands, 16.v.1991, Ismail Yusof, 1? (UKM); Fraser s Hill, ft, , Ex F.M.S. Museum, B.M , 1? (MNKM); Krau Wildlife Reserve, Kuala Lompat sector 0.25 km NW Rangers post, 8 km W Kuala Krau, primary forest disturbed by excessive flooding and treefall, at light, 3.ii.1997, M. Kos, 1/ (ZMAN); Krau Wildlife Reserve, Kuala Lompat sector, 1.25 km NNW rangers post 8 km W Kuala Krau, secondary forest riverbank, Old Orang Asli settlement, at light, 15.xii.1996, M. Kos & S. Azman, 2/ (ZMAN); Kuala Lompat, viii.1990, Zaidi & Ismail, 1? (UKM). Johor: Endau Rompin N.P., m, vi.1996, T. Trilar, M. & N. Gogala, 1? (PMSL); Endau Rompin N.P., road staging point to Kuala Jasin, bridge 9, disturbed forest margin, at light, iii.1999, J.P. & M.J. Duffels, M. Zaidi & M.Y. Ruslan, 1? (ZMAN); Endau Rompin N.P., Junction Sg. Endau / Sg. Senawak 2 32 N E, base camp, disturbed forest, at light, 210 m, 19.iii.1999, J.P. & M.J. Duffels, M.Y. Ruslan, 1? (ZMAN), same data but 18.iii.1999, 1? (ZMAN), 20.iii.1999, 2? (ZMAN), 21.iii.1999, 1? (ZMAN); Endau Rompin N.P., NERC, N E, secondary forest, Transect secondary 5, at light, 21.iii.2001, M.A. Schouten, 1? (ZMAN); Endau Rompin N.P., NERC, N E, secondary forest, Transect secondary 11, at light, 16.v.2001, M.A. Schouten, A.J. de Boer & G. Withaar, 1? (ZMAN). Langkawi Island, 14.iv.1928, H.M. Pendlebury, 1/ (UKM), same data but 15.iv.1928, 10? 3/ (UKM), 16.iv.1928, 2? (UKM), 17.iv.1928, 1? (UKM), , 1? (UKM), 24.iv.1928, 1? 1/ (UKM), 25.iv.1928, 2/ (UKM), 26.iv.1928, 2/ (UKM), 29.iv.1928, 1? (UKM). Singapore: Mandai, 3.viii.1960, 2? 1/ (MZHF). Maua linggana Moulton Figs 23, Maua linggana Moulton, 1923: 69, 125, 168. Holotype?: Lingga / Nov. / 1897, / Brit. Mus., Mau linggana Moulton / Type? 25/3/23 [Moulton s handwriting], Type [print in round label with red margin] (BMNH) [examined]. Maua linggana: Metcalf 1963: 494; Gogala et al. 2004: 4 5. Maua linggana can be separated from M. albigutta by the male opercula that are broadly rounded apically and reach to or just beyond the posterior margin of abdominal segment 2. Maua linggana (body length males: mm) is only slightly larger than M. albigutta (body length males: mm). Both species have a pair of weakly developed protuberances on sternites 3 and 4 instead of the large nipple-shaped tubercles found in the species of the M. quadrituberculata group. Description Ground colour of head, pronotum and mesonotum ochraceous to light brown sometimes with greenish tinge, abdomen of male light castaneous brown, that of female ochraceous to brown. Head (Fig. 23). A black round encloses each of the ocelli, the three rounds are fused; round enclosing median ocellus with small protrusions pointing laterodistally, rounds enclosing paired ocelli sometimes with weak protrusion also pointing laterodistally. Supra-antennal plates with distinct black marking. A pair of right-angled hook-shaped markings, occasionally divided in two spots, extends from just below these plates to level of paired ocelli. Inner margin of eyes with a pair of narrow black spots. A pair of juxtaposed half-round spots in line with the paired ocelli at posterior margin of head. Anterior and ventral parts of postclypeus with two series of black-brown transverse lines in all (8 9) transverse grooves; upper 3 pairs of lines reach from a central glabrous area to supra-antennal plates or lateral clypeal clefts, next two pairs somewhat shorter and the other transverse lines much shorter. Medial ends of some transverse black lines of each side connected by a black line. A pair of paramedian triangles below transverse lines, but lowest part of postclypeus at clypeal suture unmarked. Anteclypeus black-brown with exception of yellowish keel. Gena with short black fascia narrowing from postclypeus to two thirds of width of gena. Basal two thirds of lorum black. Rostrum reaching almost posterior margin of abdominal segment 2. Thorax (Fig. 23). Pronotum. Marking as in M. albigutta.

26 328 Tijdschrift voor Entomologie, volume 152, Figs M. linggana 24, abdomen in lateroventral view, Sabah, 23 km W Sandakan, Sepilok tree tower; 25, male pygofer in ventral view, Sabah, W of Sandakan, 11 km N Sepilok. Mesonotum (Fig. 23). Five black fasciae. Median fascia, reaching from anterior margin of mesonotum to one third of length of cruciform elevation, very narrow in anterior half, suddenly widened to 3 4 times its anterior width at two thirds of its length and gradually narrowed again to the posterior. Paramedian fasciae anteriorly about 4 times as wide as median fascia at anterior margin of mesonotum and equally wide in its apical half, but distinctly narrowed at one fourth of its length; the fasciae reach to just beyond half-length of mesonotum. A pair of black spots in front of anterior angles of cruciform elevation. Lateral fasciae on mesonotum consisting of three short black lines on a row, posterior two lines sometimes fused. Legs. Yellow to light brown with brown to black marking. Fore femora with a black-brown fascia connecting three light brown to black-brown spines: a long and narrow, erect spine at two fifths of length from base, a somewhat shorter and broader spine at four fifths, and a very small spine next to the latter; fore femora with dark brown spot at distal underside and sometimes with longitudinal fasciae on inner, outer- and upper sides. Fore tibia with dark brown fascia on underside. Tarsi of fore and middle legs brown sometimes with greenish tinge, tarsi of hind legs ochraceous. Claws brown. Tegmina and wings. As in M. albigutta, but apical parts of apical cells 1 and 2 of tegmina less distinctly brownish suffused. Male operculum (Fig. 27). Lateral margin oblique, sinuate in basal one fourth and weakly convex to broad, and more strongly convex, margin of apical part of operculum; apical margin becomes more or less straight toward medial corner. Medial corner narrowly rounded, medial margin straight or weakly concave from medial corner to operculum base. Operculum ochraceous with brown to black-brown triangular marking at the basal margin of operculum, and reaching, or reaching just beyond, posterior margin of abdominal segment 2. Male abdomen (Fig. 27). Abdomen broad and parallel-sided, equally wide across segments 2 5, segment 3: times as wide as pronotal collar. Timbal

27 Duffels: Revision of Maua (Cicadidae) 329 covers ochraceous to light brown. Tergites 2 7 light castaneous brown, tergite 8 somewhat darker brown. Posterior margins of tergite 2 with narrow, blackbrown fascia; tergites 3 and 4 with similar, somewhat broader, fasciae. Ventral side light brownish; posterior margins of sternites 3 and 4 dark brown. Sternites 3 and 4 with a pair of narrow, light to dark brown protuberances, which are along their whole length attached to the sternite. Distance between tubercles on sternite times (n=7) as wide as distance between tubercle and lateral segment margin. Male genitalia (Fig. 28). Pygofer ochraceous, lateral pygofer lobes dark brown. Basal pygofer lobes are broadly rounded ridges that slightly widen from base to three fifths of pygofer length. Uncus short, trapezoid with broadly rounded corners; apical margin with very weak median incurvation. Basis of uncus inwardly curved and forming two triangular sclerites meeting in midline of pygofer; these sclerites form a pair of small, round protuberances at their lateral margins and a short median tube enclosing the aedeagus. Aedeagus regularly curved and more or less suddenly narrowing, at half-length, to a simple acute apex. Female operculum. Lateral margin weakly convex, laterodistal corner broadly rounded, distal margin very weakly undulate. Operculum ochraceous with dark brown to black basal triangle. Female abdomen. Tergite 2 with black line along posterior margin, tergites 3 6 with a very narrow black line along posterior margins, but on tergites 3 4 the central part of this line, which is slightly wider than the cruciform elevation, is about as high as one fourth of tergite. Anterior part of tergite 7 with black marking or brownish suffused, lateral parts of tergite with brown to black marking on anterior two thirds. Sternites 3 6 black-brown along anterior and lateral margins, sternite 7 black-brown along anterior margin. Valvae black-brown. Segment 9 dorsally with a pair of paramedian black triangles and ventrally with basal two thirds to three fourths black. Measurements in mm (6? 2/). Body length?: , /: ; tegmen length?: 34 37, /: 33.5; head width?: , /: ; pronotum width?: , /: Distribution (Fig. 26) Maua linggana is known from various places in Borneo (Sarawak, Sabah, Kalimantan and Brunei) and is probably a Borneo endemic. Material examined. 8? 2/. Malaysia: Borneo: Sabah: Siboga Reserve, Kebun Cina, W of Sandakan, 11 km N Sepilok, 25.xi.1989, sample Sab. 42, primary Diptocarp rainforest, at light, J.P. & M.J. Duffels, 1? (ZMAN); 23 km W Sandakan, Sepilok tree tower, E 5 49 N, m, 1.xi.1987, J. Huisman & R. de Jong, 1? (RMNH). Sarawak: Semongok, 11.xi.1966, G.H.L. Rothschild, light trap, 9608, Pres. by Com Inst Ent BM , 1? (BMNH). Brunei: Brunei, Waterstradt, Brit. Mus , 2? (BMNH); Temburong District, ridge NE of Kuala Belalong, approx. 300 m, x.1992, J.H. Martin, 125W m.v. light, BM , 2? 2/ (BMNH). Indonesia: Kalimantan Timur: Tiong Buu (= Nahabuan), 18 km NW of Longkay, S. Mahakkam, 15.iv.1996, R. Sözer, 1? (ZMAN). Maua platygaster Ashton Figs 26, 29 Maua platygaster Ashton, 1912: 156. Syntype /: Matang / 3200 ft / , Pres. Raffles Mus. / Singapore / Brit. Mus. / , Maua platygaster Asht, From Sarawak / Museum (BMNH) [examined]. Maua platygaster: Moulton 1912: 133; Moulton 1923: 124, 126, 168; Metcalf 1963: 494. Types Ashton (1912) described M. platygaster in an appendix to Moulton s (1912) Material for a Fauna Borneensis: a list of Bornean cicadas. In this publication, Moulton wrote the following lines about M. platyptera: The Sarawak Museum possesses two specimens from Mt. Matang taken in July and August and in a footnote he added: This species is described by Mr. Howard Ashton in the Appendix at the end of this paper (p. 156). Maua platygaster was described after one male and one female from Sarawak. It seems most likely that the two specimens from Mt. Matang mentioned by Moulton (l.c.) are the type specimens. Dr M. Zaidi (Universiti Kebangsaan Malaysia) tried to trace the type specimens of M. platygaster in the collection of the Sarawak Museum (Zaidi & Ruslan, 1998), but could not find any. My study of the BMNH collection revealed one immature female from Mt. Matang that comes from the Sarawak Museum and is most probably a syntype of M. platygaster. The identity of M. platygaster The only specimen of this species that I have seen is the alleged female syntype from Mt. Matang (Fig. 29). This specimen can be separated from the other species of the genus by its very characteristic light brown marking on the tegmina that consists of anastomosing spots at the bases of the first and second apical cells and at the bases of the third and fourth apical cells, and separate spots at bases of fifth and seventh apical cells. Ashton s original description of M. platygaster mentioned anastomoses to

28 330 Tijdschrift voor Entomologie, volume 152, 2009 Fig. 29. M. platygaster, female syntype, Sarawak, Mt. Matang. Photo M.J. Duffels van Egmond. 2nd, 3rd and 5th apical cells spotted with fuscous. In spite of the discrepancies between the description and the pattern found in the syntype, the mere existence of anastomoses sustains the syntype status of the specimen. Anastomosing spots are found in only one other species of Maua. Maua affinis has an anastomosing spot on the transverse veins at the bases of the first and second apical cells and separate spots at the bases of the third, fifth and seventh apical cells, but some specimens have one anastomosing spot at the bases of the third and fourth apical cells. Other characters of M. platyptera are the rectangular pronotum, irregular lateral mesonotal fasciae that strongly widen to their distal ends, the relatively long apical cell 1 of the tegmen, and the basal vein of apical cell 1 which is much shorter than the longitudinal vein of apical cell 1. According to the original description the male abdomen is very broad and the male opercula are margined with black. Tubercles on the underside of the male body are not mentioned in the original description. Further material, especially males, is needed to establish the identity and the taxonomic position of this species. Remark Moulton s key to the species of Maua (Moulton 1923) divided the genus in two groups: those with tegmina expansions under 80 mm and those with tegmina expansions over 90 mm. The key is misleading because M. platygaster is arranged among the larger species of the genus with tegmina expansions over 90 mm. According to the original description, the expansion of the tegmina in the male M. platygaster is only 70 mm so that it is one of three species with an expansion of the tegmina under 80 mm (cf. Moulton 1923). The body length of the male type specimen of M. platyptera (21 mm) is even smaller than that of M. albigutta (24 28 mm) and M. linggana ( mm). Distribution (fig. 26) Maua platygaster was described from Mt Matang, Borneo, Sarawak. According to Zaidi & Ruslan (1998) the Sarawak Museum has a male specimen of M. platygaster from Mt. Penrissen, Sarawak, collected by E. Mjöberg. I have not seen this specimen. Mt Matang is situated west of Kuching. Mt Penrissen is part of the mountain range southwest of Kuching close to the border between Sarawak and Kalimantan Material examined. Borneo, Malaysia, Sarawak: The female syntype (see above). Acknowledgments I am very much indebted to the following curators for the loan and gift of material: Mrs Pudji Aswari (MZB), Dr J. Constant & Dr P. Grootaert (KBIN), Dr M. Gogala and Dr T. Trilar (PSML), Dr B. Hubley & Dr D.C. Darling (ROM), Dr G. Lindberg (NHRS), Dr Maryati Mohamed (UMS), Dr D. Mierzwa and Dr J. Szwedo (MIZ), Dr A. Soulier-Perkins (MNHN), Dr J. van Tol (RMNH), Mr M. D. Webb (BMNH), Dr Zaidi Mohd. Isa and Mr Ruslan Mohd. Yusop (UKM) and Dr H. J. Zainal Abidin (MNKM). I thank Dr Masami Hayshi (Saitama University, Urawa, Japan)