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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. [email protected]

Introduction Borneo, Palawan and several smaller islands in- The genus Maua was erected by Distant (1905) between. Two species of Maua, M. borneensis from for Cicada quadrituberculata Signoret, 1847 and Borneo and M. palawanensis from Palawan and Bala- M. affinis Distant, 1905. The genus is currently bac are described as new to science, and six species placed in the tribe Dundubiini and the subtribe Lep- from Sundaland are redescribed: M affinis, M. albi- topsaltriina (Duffels & Van der Laan 1985; Moulds gutta (Walker, 1857), M. latilinea (Walker, 1868), 2005). In 1923, Moulton erected the new section M. linggana Moulton, 1923, M. platygaster Ash- Leptopsaltraria [sic] for the genera Leptopsaltria ton, 1912, and M. quadrituberculata. Two species, Stål, 1866, Maua Distant, 1905, Nabalua Moulton, M. ackermanni Schmidt, 1924 and M. dohrni 1923, Purana Stål, 1866 and Tanna Distant, 1905. Schmidt, 1912, both described from Sumatra, The new section was characterised by the presence proved to be junior synonyms of respectively of one to three pairs of tubercles on the ventral side M. quadrituberculata and M. latilinea. Three species of the male abdomen. A revision of Nabalua was of Mau are not treated in this paper: M. albistigma recently presented by Duffels (2004), and three, pre- (Walker, 1850) and M. fukienensis Liu, 1940 from sumed monophyletic, groups of the genus Purana China, and M. philippinensis Schmidt, 1924 from were revised: the P. nebulilinea group (Kos & Gogala the Philippines. 2000), the P. carmente group (Schouten & Duffels This study aims to contribute to a better knowledge 2002) and the P. tigrina group (Duffels et al. 2007). of cicada biodiversity in Southeast Asia. Basic sys- In 1963, Metcalf added several genera, with and tematic studies providing illustrations and descrip- without abdominal tubercles, to the subtribe; none tions of species and keys to genera and species are of these genera are recorded from Sundaland. regarded as a prerequisite for biodiversity studies. This paper presents a revision of the genus Maua Biodiversity studies of cicadas in other tropical areas, from Sundaland: the Malay Peninsula, Java, Sumatra, such as Sulawesi, New Guinea, and the West Pacific,

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have already shown that cicadas can be instrumental The terminology adopted in this paper for external in recognizing hot spots of species richness and areas features of the body and the male genitalia follows of endemism with unique biota (Duffels & De Boer that of Duffels (Duffels 1977, 1983; Duffels & 1990, De Boer & Duffels 1997). Turner 2002) and Moulds (2003, 2005).

Material and methods Taxonomy The institutions listed below are the depositories of Genus Maua Distant the material studied. The abbreviations have been used in the lists of material and throughout the text. Maua Distant 1905: 61. Type-species by original designa- tion: Maua quadrituberculata (Signoret, 1847). BMNH Natural History Museum, London (former Maua: Moulton 1923: 124–126; Metcalf 1963: 491–495; Duffels & Van der Laan 1985: 113; Moulds 2005: British Museum (Natural History)), 391, 432. London [For further references before 1980 see: Metcalf 1963 and KBIN Koninklijk Belgisch Instituut voor Natuur- Duffels & Van der Laan 1985]. wetenschappen, Brussels MNHN Muséum National d’Histoire Naturelle, The species of Maua studied have one character in Paris common viz., the broad, more or less parallel-sided MNKM Muzium Negara Malaysia, Kuala Lumpur male abdomen with about equally wide segments MIZ Museum and Institute of Zoology, Polish 3 to 4 or 2 to 5. The shape of the male abdomen was Academy of Sciences, Warszawa traditionally used to separate Maua from other gen- MZB Museum Zoologicum Bogoriense, Cib- era of the subtribe (Distant 1906; Moulton 1923). inong The genus Maua in its present concept is probably MZHF Zoological Museum, Finnish Museum of not monophyletic. Natural History, Helsinki NHRS Swedish Museum of Natural History, Distribution Stockholm The genus Maua is found in China, Thailand, the PSML Prirodoslovni Muzej Slovenije, Ljubljana Philippines and Sundaland (Malay Peninsula, Java, RMNH Nationaal Natuurhistorisch Museum Sumatra, Borneo and Palawan) (Metcalf, 1963; Duf- (formerly Rijksmuseum van Natuurlijke fels & van der Laan 1985; Sanborn et al. 2007). The Historie), Leiden genus is certainly most divers in Borneo (seven spe- ROM Royal Ontario Museum, Toronto cies). Four species are probably Borneo endemics: UKM Pusat Sistematik Serangga, Universiti M. affinis, M. borneensis, M. linggana and M. platyg- Kebangsaan Malaysia, Bangi, Malaysia aster. Maua affinis seems to be restricted to Palawan UMS Universiti Malaysia Sabah, Kota Kinabalu and Balabac Island between Borneo and Palawan. ZMAN Zoölogisch Museum, Universiteit van Maua latilinea is known from Borneo and Sumatra. Amsterdam, Amsterdam The two remaining species have a wider distribution: M. albigutta is found in the Malay Peninsula Data on the distribution of the species were derived and Sumatra, and M. quadrituberculata is known from the author’s ‘Biodiversity Database of the Cica- from China, Thailand, the Malay Peninsula, Java (?), das of South East Asia and the West Pacific’, and Sumatra, Borneo and Nias Island. plotted on maps of ADC-Worldmap version 2.0 vol. 4 Southern Asia & Australia with the program Relationships MapInfo for Power Mac, version 4.03. The locali- Three genera of the subtribe Leptopsaltriina occur- ties and other data from the specimen labels in the ring in Sundaland have two pairs of tubercles on database are filed in the program FileMaker Pro the male sternites 3 and 4: Leptopsaltria, Maua and 4.0. Geographical information has been retrieved Purana. Maua is distinguished from the other two from the following sources: ‘Atlas van Tropisch Ned- genera by the broad, more or less parallel-sided male erland’ (Anonymous 1938), ‘Gazetteer Nasional abdomen; in Purana and Leptopsaltria the male abdo- Nama-nama Geografi’ for Indonesia (Anonymous men gradually narrows from segment 2 backwards. 1978), Nelles Road Atlas Indonesia (Anonymous The genus Maua is divided in two groups that are 1992), The Times Comprehensive Atlas of the presumed to be monophyletic: (1) the M. quad- World’ (Anonymous 1999) and GEOnet Names rituberculata group with a male body length of 31– Server of the U.S. Defense Mapping Agency 45.5 mm, comprising: M. affinis, M. borneensis, (http://www.nima.mil/gns/html/index.html). M. latilinea, M. palawanensis, M. quadrituberculata

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(type species of the genus) and (2) the M. albigutta apical cell without spot. Distal ends of para- group with a male body length of 24–29.5 mm, median mesonotal fascia usually connected comprising: M. albigutta and M. linggana. The with median fascia. Palawan; Balabac I .. . . . Maua quadrituberculata group and the M. albigutta ...... M. palawanensis group are readily separated by the following charac- – Tegmen with distinct brown spots at bases ters: the lateral fasciae on the mesonotum are con- of 2nd and 3rd apical cells; bases of 5th and tinuous from anterior margin to at least three fifths 7th apical cells without spots. Distal ends of of mesonotum length in the M. quadrituberculata paramedian mesonotal fascia not connected group, and consist of three spots or short lines in with median fascia ...... 4 the M. albigutta group; the male abdomen is very 4. Lateral fascia on mesonotum broad and broad, and widest across the segments 3 and 4, in the straight, and reaching from anterior mes- M. quadrituberculata group, and fairly broad, and onotum margin to almost hind margin of about equally wide across the segments 2–5, in mesonotum (Fig. 17). Abdominal tergite the M. albigutta group; the male sternites 3 and 4 3 in male and female with triangular median have a pair of thick, nipple-shaped tubercles in the marking. Malay Peninsula (?); Borneo; Kar- M. quadrituberculata group, and a pair of narrow imata I.; Sumatra ...... M. latilinea protuberances in the M. albigutta group. – Lateral fasciae on mesonotum somewhat ir- The albigutta group is allocated in the genus Maua regular and reaching from anterior mesono- because of the fairly broad male abdomen, but allo- tum margin to three fifths of mesonotum cation in the genus Purana can also be considered length (Fig. 1). Abdominal tergite 3 in male because of the marking on the mesonotum and (female unknown) without triangular medi- the weakly developed protuberances on the male an marking. China; Thailand; Malay Penin- sternites 3 and 4. The Maua albigutta group as sula; Java (?); Borneo; Sumatra; Nias I . . . . . well as two species groups of the genus Purana, the ...... M. quadrituberculata P. carmente group (Schouten & Duffels 2002) and 5. Spot at basis of 2nd apical cell of tegmen the P. nebulilinea group (Kos & Gogala 2000), might continuing along basal vein of 1st apical cell. represent new genera. However, a cladistic analysis of Male abdomen very broad, segment 3: 1.37– the species of Purana and Maua and related genera 1.44 times as wide as pronotal collar. Body of the subtribe Leptopsaltriina is needed to provide a length male: 39–45.5 mm. Borneo . . M. affinis basis for a new generic division of the subtribe. – Spot at basis of 2nd apical cell of tegmen not One species, M. platygaster, is not attributed to one continuing along basal vein of 1st apical cell. of the species groups and is not described in the Male abdomen broad, segment 3: 1.09–1.17 same detail as the other Sundaland species. I have times as wide as pronotal collar. Body length seen only one female of M. platygaster, the male male: 31–35 mm. Borneo ...... M. borneensis of this species is only known to me by the original 6. Tegmen with anastomosing spots at bases of description of Ashton (1912). Maua platygaster can 1st and 2nd apical cells and at bases of 3rd be separated from the other species of the genus by and 4th apical cells, with separate spots at its characteristic marking of anastomosing spots on bases of 5th and 7th apical cells, and without the tegmina and small body size (body length male: spots at apices of longitudinal veins of apical 21 mm). cells. Body length male: 21 mm. Borneo ...... M. platygaster Key to the species of Maua – Tegmen with separate spots at bases of 2nd The principal use of this key is for the identification and 3rd apical cells, and with small brown of males. It is of limited value for the identification spots at apices of longitudinal veins of apical of females. The females of three species, M. affinis, cells 1–4. Body length male: 24–29.5 mm . . . 7 M. latilinea and M. palawanensis are unknown. 7. Male operculum apically broadly rounded and reaching, or reaching just beyond, poste- 1. Body length male: 31–45.5 mm ...... 2 rior margin of abdominal segment 2. Lateral – Body length male: 21–29.5 mm ...... 6 fasciae on mesonotum consisting of three 2. Postclypeus with black spot or black marking short black lines on a row, posterior two at clypeal suture ...... 3 lines sometimes fused (Fig. 23). Body length – Postclypeus without black spot or black male: 27–29.5 mm. Borneo...... M. linggana marking at clypeal suture ...... 5 – Male operculum tapering to the apex and 3. Tegmen with distinct brown spots at bases reaching to half-length abdominal segment of 2nd, 3rd and 5th apical cells; basis of 7th 3. Lateral fasciae on mesonotum consisting of

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three distinct black spots on a row (Fig. 22). all other large species of Maua by the unmarked pos- Body length male: 24–28 mm. Malay Penin- terior oblique fissures on the mesonotum that are sula; Sumatra; Siberut I ...... M. albigutta black in the other species. Maua quadrituberculata and M. latilinea from Borneo, Kalimantan are the Maua quadrituberculata group only species with two black spots, at the bases of the second and third apical cells of the tegmen; the other The Maua quadrituberculata group consists of five species of the group have spots at the bases of the sec- medium-sized to fairly large species (body length ond, third and fifth apical cells or at bases of second, male: 31–45.5 mm): M. affinis, M. borneensis, third, fifth and seventh apical cells. Maua latilinea is M. latilinea, M. palawanensis and M. quadrituber- easily distinguished from M. quadrituberculata and culata. The lateral mesonotal fasciae are somewhat all other large species of Maua by the continuous irregular or straight but continuous from anterior broad lateral fasciae on the mesonotum that reach margin to at least three fifths of mesonotum length from anterior to almost posterior mesonotal margin. or to at most the hind margin of the mesonotum. The apical cell 1 of the tegmen is relatively short. Synonymy The basal vein of apical cell 1 is shorter than half as In February 2008, Dr Tomi Trilar of the Pri- long, or at most half as long, as the longitudinal vein rodoslovni Muzej Slovenije, Ljubljana visited of apical cell 1. The male abdomen is very broad, the Museum of the Institute of Zoology, Polish widest across the segments 3 and 4, segment 2 is only Academy of Science, Warsaw, Poland. He found slightly narrower; the sternites 3 and 4 have a pair the ‘Typus’ and the ‘Co-Typus’ of M. ackermanni of thick, nipple-shaped tubercles pointing sidewards. Schmidt, 1924 in the collection of this museum, Maua latilinea differs from the other species of the brought these to my attention and sent me excellent group in the body marking and the structure of the photographs of the type specimens (Figs 3–4). Later, male genitalia. The basal pygofer lobes of M. lati- Dr D. Mierzwa (MIZ) kindly gave me on loan the linea are large, black-brown and spine-shaped, and ‘Typus’ and the ‘Co-Typus’ of M. ackermanni. Study reach from two fifths to three fourths of the pygofer of this material proved that M. ackermanni is a junior length, while the uncus is triangular and strongly synonym of M. quadrituberculata. sclerotised. In the other four species of this group the basal pygofer are more or less parallel, apically acute, Description ridges, while the uncus is trapezoid and not strongly Ground colour of head, pronotum and mesonotum sclerotised. yellowish to greenish brown, abdomen of male and female ochraceous to brown. Head Maua quadrituberculata (Signoret) (Fig. 1). A black mark with three pairs of arms encloses the ocelli: a pair of out curved, narrow, Figs 1–5, 7 black arms run from median ocellus to the anterior, Cicada quadrituberculata Signoret, 1847: 297–298. a pair of somewhat longer, narrow, arms extend from Leptopsaltria quadrituberculata: Distant 1889: 31; Distant paired ocelli to the anterior, and a pair of straight, 1891: pl. viii figs 6, 6a-b. tapering, black lines run from paired ocelli to the pos- Maua quadrituberculata: Distant 1905: 61; Moulton 1923: 124, 126, 168; Metcalf 1963: 494, 495; Duffels & terior and sometimes reach black posterior margin of Van der Laan 1985: 113; Zaidi & Ruslan 1998: 153; the head. Proximal three fourth of supra-antennal Boulard 2007: 101–104. plate black. Pair of black triangular markings extend Maua ackermanni Schmidt, 1924: 287 Holotype ?: from just below these plates to level of paired ocelli. ‘Soekaranda / Januar 1894 / Dohrn’ [print in black Postclypeus with black lines in anterior 5–7 trans- cadre], Maua / Ackermanni / Schmidt / ? Edm. verse grooves of either side; basal part of postclypeus Schmidt / determ. 1924’, ‘Typus’ [print in black cadre; red paper], ‘MIZ 187250’ (MIZ) [examined]. syn. n. with broad black transverse mark at clypeal suture, [For further references before 1980 see: Metcalf 1963 and which is medially divided by an upside-down tri- Duffels & Van der Laan 1985]. angle of the ground colour. Anteclypeus black, but triangular basal part, median keel and apical part Maua quadrituberculata is a large species (body yellow. Gena with broad black fascia running from length ?: 38–41.5 mm). The males have a very antenna to eye. Basal half of lorum black. Rostrum broad abdomen (segment 3: 1.23–1.32 (1.45) times reaching male sternite 3 or female sternite 4; apex as wide as pronotal collar), though the abdomen of brown. the males of M. affinis is usually even broader (seg- Thorax (Fig. 1). Pronotum. Paired central fasciae ment 3: 1.37–1.44 times as wide as pronotal collar). narrow, strongly widened toward black, anterior Maua quadrituberculata can be distinguished from pronotum margin and fused at anterior margin of

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1 2

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 upside- of cruciform elevation. Lateral fasciae somewhat down, key-hole shaped median interspace. A pair of irregular, extending from anterior mesonotum margin low black triangles extends from half-length central to three fifths of mesonotum length and apically nar- fascia along basal half of anterior oblique fissures. rowed; some specimens have a pair of small dark spots Posterior oblique fissures unmarked. Inner side of more distally on the mesonotum. Posterior margin of lateral part of ambient fissure with very broad fasciae cruciform elevation black to dark brown. that continues in the ambient fissure as a very narrow Legs. Yellow with greenish tinge and brown to black line reaching to distal ends of posterior oblique fis- marking. Fore femora with three spines: a long and sures; a short, broad, black mark extends from proxi- narrow, erect, ochraceous, spine with a brown tip at mal end of broad fascia to black posterior margin half-length, a somewhat shorter, broad, black spine of pronotal collar and another (much) smaller black at four fifths of length from base, and a small, black mark is situated just above it. to brown spine next to the latter. Fore femur with Mesonotum (Fig. 1). Five black fasciae. Median fascia a subapical dark patch on inner and outer side and reaching from anterior margin of mesonotum to level a dark brown to black fascia on upper side. Fore of anterior angles of cruciform elevation; the fascia tibia black but apical one sixth and basal one third gradually widens from its anterior end to two thirds of upper side yellow. Middle tibiae with subapi- of its length, where it attains 3–5 times its anterior cal black-brown ring as long as one fourth of tibia. width, and gradually narrows again to the posterior. Metatarsus and basal half of mesotarsus of fore and Paramedian fasciae extending from anterior mes- middle legs yellow, apical half of mesotarsus and pre- onotum margin to about half-length of mesonotum. tarsus black-brown. Tarsi of hind legs ochraceous. A pair of tiny black spots in front of anterior angles Claws brown.

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3 Figs 3–4. Maua quadrituberculata, male holotype M. ackermanni. – 3, body in dorsal view; 4, body in ventral view. Photo’s Dr T. Trilar.

Tegmina and wings. Hyaline. Tegmina with distinct as wide as pronotal collar, segment 2 only slightly dark brown spot on transverse veins at bases of sec- narrower. Dorsal side, including timbal covers, light ond and third apical cells. Small brown spots either to dark brown, sometimes with greenish tinge, and at apices of all longitudinal veins, or at apices of api- with silvery to golden pubescence; basal part of lat- cal cells 1–4, or totally missing as in a specimen from eral margin of timbal cover dark brown. Tergite 2 Deli, Sumatra. Venation of tegmina light brownish with black band of variable width along about two to yellowish variegated with dark brown to black- thirds of posterior tergite margin. Posterior mar- brown, venation of wings light to dark brown. gins of tergites 3–6 with a narrow, light brown Male operculum (Fig. 2). Lateral margin very weakly band that widens to brown to a black-brown tri- sinuate and directed slightly inwards. Apical part angles at their lateral ends. Tergite 7 laterally dark fairly broadly rounded. Medial margin very weakly brown. Tergite 8 brown with irregular dark brown convex to medial corner. Operculum reaching to marking. Ventral side light ochraceous, with waxy, one fourth or one third of abdominal segment 3. pilose, patches. Posterior margin of sternite 2 dark Ochraceous. Margin of operculum black, but basal brown and of sternites 3–6 light brown. Sternite 7 part of lateral margin and basal, incurved, part of brown, turning dark brown to the posterior. Ster- medial margin unmarked. Basal part of operculum nite 8 brown but basal parts of lateral margins yel- with black marking next to meracanthus. lowish. Sternites 3 and 4 with thick, glossy brown, Male abdomen. Abdomen very broad, widest across nipple-shaped tubercles; distance between tuber- segment 3 or 4, segment 3: 1.23–1.32 (1.45) times cles on sternite 4 1.40–1.92 times (n=8) as wide

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5 6

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 almost straight; medial margin concave to fairly nar- margin. rowly rounded mediodistal angle. Male genitalia (Fig. 5). Pygofer ochraceous, but lat- Female abdomen. Dorsal side light brown with sil- eral pygofer lobes and uncus sometimes brown. Basal ver pilosity. Tergites 2–6 and epipleurites 2–3 with pygofer lobes consisting of a pair of slightly converg- narrow black bands along their posterior margins; ing ridges that are broadly connected with lateroven- epipleurites 4–6 with somewhat broader and more tral part of pygofer, reach to three fifths of pygofer irregular bands. Tergites 3–6 laterally with small length, and are apically acute. Uncus short, trap- brown triangle. Tergites 7 and 8 dorsally brown, ezoid; apical margin with very weak median incur- laterally black-brown; posterior margin of tergite 7 vation. Basis of uncus inwardly curved and forming dark brown. Segment 9 brown with a pair of irregu- two flat and fairly long, about triangular sclerites lar, dark brown markings reaching from anterior seg- meeting in midline of pygofer; these sclerites form a ment margin to half the segment length. Sternites short median tube enclosing the aedeagus and a pair 3–5 brownish with greenish tinge, and with fairly of protuberances at their lateral margins. Aedeagus broad black-brown band along anterior margin and regularly curved, basally very broad, suddenly nar- black-brown lateral marking. Sternite 6 shining rowed at about two thirds of length and ending in a black-brown with small, median, brownish patch. simple acute apex. Sternite 7 shining black-brown. Female operculum. Greenish ochraceous with medio- Measurements in mm (6? 1/). Body length ?: basal brown marking and scattered silver pilosity. 38–41.5, /: 31; tegmen length ?: 49.5–53, /: 46; About as long as wide and just not reaching abdomi- head width ?: 10.9–11.8, /: 10.2; pronotum width nal segment 3. Lateral margin undulate to rounded ?: 11.1–13.1, /: 11.8. rectangular laterodistal angle; posterior margin

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Fig. 7. Distribution of Maua quadrituberculata (rounds) and M. affinis (triangles).

Distribution (Fig. 7) collected a new subspecies of M. quadrituberculata in Maua quadrituberculata was originally described northern Thailand. The subspecies was described as from Java, but its occurrence on Java is not con- M. quadrituberculata khunpaworensis (Boulard 2007: firmed by the present or earlier studies. Accord- 101–105). According to its description, the new ing to Moulton (1923), the species occurs in Java, continental subspecies can be distinguished from the Malay Peninsula and Borneo, and is recorded from insular nominal subspecies by the more triangular China and the Philippines. The China and Philip- head, the shorter pronotum and the more rounded pines records of M. quadrituberculata by Moulton male opercula. The reader is referred to Boulard’s (l.c.) and other authors probably all date back to paper for a more detailed taxonomic discussion, the Distant’s Monograph of Oriental Cicadidae (1889, interesting ethological data about the new subspecies 1891). The BMNH has one old specimen of and the description of its song. M. quadrituberculata with a label ‘China’, but Chou et al. (1991) have not seen a single specimen from Material examined. 16? 2/. China: Chini (round label China (Chinese text kindly translated by Dr Wei with on the reverse side: 63/184, 1? (BMNH). Thailand: Cong, Yangling, Shaanxi, China). The present study Peninsular Siam, Nakon Sri Tamaret, Khao Ram, 750– could not confirm the occurrence of M. quadritu- 1260 ft, 23.ii.1922, H.M. Pendlebury, Ex. F.M.S. Mu- seum, B.M. 1955–354, 1? (BMNH), same data but 750 berculata in the Philippines and China. The material ft, 24.ii.1922, 1? (BMNH). Peninsular Malaysia: Johor: available for this study confirms the occurrence of Endau Rompin N.P., Janing Ridge, 20 min. 02°31’19’’N M. quadrituberculata in Thailand, Malay Penin- 103°23’59’’E, primary lowland rainforest, transect prima- sula and Borneo (Sabah, Sarawak and Brunei), and ry 4, at light, 27.iii.2001, M.A. Schouten, A. Majanil & presents new records for Sumatra and Nias Island. M. Renganathan, 1? (ZMAN); Endau Rompin N.P., Janing Ridge, 5 min. 02°31’33’’N 103°24’03’’E, dis- Remark turbed forest, transect disturbed 6, at light, 26.iii.2001, M.A. Schouten, 1? (ZMAN). Perak: Doherty, Distant My esteemed colleagues Dr Michel Boulard coll. 1911–383, 2? 1/ (BMNH); Batang Padang, (MNHN) and Mrs Khuankanok Chueata recently Jor Camp, 2000 ft, 2.vi.1923, H.M. Pendlebury, Downloaded from Brill.com10/10/2021 08:41:27PM via free access Duffels: Revision of Maua (Cicadidae) 311

Ex. F.M.S. Museum, B.M. 1955–354, 1? (BMNH). depositories. The Paris museum has a male speci- Sumatra: Soekaranda, Co-Typus Maua ackermanni men with the labels: ‘Museum Paris / Bornéo / coll. Schmidt ?, Edm. Schmidt determ. 1924, MIZ 187251, Noualhier 1898’, ‘Museum Paris / MNHN (EH) / ? 1 (MIZ); Deli, coll. Dr. D. MacGillavry, Leptopsaltria 4301’, ‘Maua / affinis Dist / (type)’ (Distant’s hand- quadrituberculata, det. McGillavry, 1? (ZMAN). Nias: Noord Nias, Hili, Madjedja, Mitschke, 4de trim. ’95, 1? writing). The Stockholm museum has a male speci- (BMNH). Borneo: Malaysia: Sabah: Tawau Hill, along men with the labels: ‘Palawan’, ‘Staudgr.’, ‘Maua / Sungai Tawau near trail to Air Panas, river bed in pri- affinis / Dist.’ and NHRS-HEMI 000000012 (pers. mary riverine forest, 300 m, at light, 28.iii.2001, J.P. & com. Dr G. Lindberg, Stockholm). The London M.J. Duffels, 1?1/ (ZMAN. Sarawak: Gunung Mulu museum has a male specimen with the labels: ‘Bala- Nat. Park, Site 7, Long Pala (Base), 324450, 50 m, Al- bac / coll. Noualhier 1898’, ‘Maua / affinis / Dist’ , luvial/secondary forest, Acl-understorey, J.D. Holloway, RGS Mulu exped., B.M. 1978–206, 1? (BMNH); Gu- ‘Type’ [round label with red circle; print], ‘Distant nung Mulu Nat. Park, Site 26, G. Api, Pinnacles, 1200 m, coll / 1911–383’. Balabac island, between Borneo open scrub, MV, 428545, J.D. Holloway, RGS Mulu ex- and Palawan. All identification labels are in Distant’s ped., B.M. 1978–206, 1? (BMNH). Brunei: Waterstradt, handwriting 1? (BMNH); Temburong District, ridge NE of Kuala The specimen from Paris is designated here as lec- Belalong, approx. 300 m alt, x.1992, J.H. Martin, coll. totype of M. affinis because it is the only specimen ? B.M. 1992–172, 1 (BMNH). from the type locality Borneo and because it bears the label ‘Maua affinis type’ in Distant’s handwrit- Maua affinis Distant ing. The round Type label of the London specimen was probably not attached by Distant but by a later Figs 6–9 curator of the museum. The specimens from Stock- Maua affinis Distant 1905: 61. Lectotype: ‘Museum Paris holm and London are from respectively Palawan and / Bornéo / coll. Noualhier 1898’, ‘Museum Paris / Balabac, an island between Borneo and Palawan, and MNHN (EH) / 4301’, ‘Maua / affinis Dist / (type)’ not from Borneo. Because it is questionable whether [examined]. Maua affinis: Distant 1906: 52; Moulton 1912: 133; these specimens belong to the type series, I have Moulton 1923: 124, 125, 168 (partim: Borneo); Met- labelled these specimens: ?paralectotype Maua affinis calf 1963: 492 (partim: Borneo). Distant, J.P.Duffels 2009. Maua affinis: Zaidi 1996: 101; Zaidi 1997: 113; Zaidi & Maua affinis is probably endemic to northern Bor- Ruslan 1998: 352; Zaidi, Ruslan & Azman 1999: 306; neo. The identity of M. affinis is established in the Zaidi, Ruslan & Azman 2000: 204; Zaidi, Noramly & description below. The questionable paralectotypes Ruslan 2000: 324 (these records of M. affinis by Zaidi and co-authors from various localities in Sarawak from Palawan and Balabac belong to a new species and Sabah refer to either M. borneensis or possibly to described in this paper as M. palawanensis which is M. affinis). found on these islands only and not on Borneo. [Misidentification Maua affinis: Distant 1912: 41, pl. 5 Figs 33a-c (= Maua palawanensis); Chou et al, 1997: Description of male 232, Fig. 114 (see remark following the description).] Ground colour of head, pronotum and mesonotum ochraceous to light brown sometimes with greenish There has been much confusion about the identity tinge, abdomen of male and female ochraceous to of Maua affinis since the specimen depicted by Dis- brown. tant (1912) as Maua affinis belongs to the species Head (Fig. 8). Black mark enclosing ocelli, mark- that is described in this paper as M. palawanensis. ing on supra-antennal plates and triangular mark- Maua affinis can be separated from M. quadritu- ings below these plates as in M. quadrituberculata. berculata by the presence of spots at the basal veins Anterior and ventral parts of postclypeus with of the fifth and seventh apical cells, the number of black-brown transverse lines in the two series of 9–10, instead of 5–7, transverse black-brown lines 9–10 grooves; upper 6 pairs of these lines reaching on the postclypeus, the absence of a distinct black to supra-antennal plates or to lateral clypeal clefts, mark at the clypeal suture, the larger black spots in other streaks shorter. Medial ends of transverse lines front of anterior angles of cruciform elevation, and of each side connected by a more or less distinct black the presence of black-brown posterior oblique fis- line; these lines enclose a long oval area at anterior sures on the pronotum. margin of postclypeus and a yellowish wedge-shaped area on ventral side of postclypeus. Lower part of The types and the identity of Maua affinis postclypeus at clypeal suture unmarked. Underside The original description of M. affinis by Dis- of head yellow with black marking. Anteclypeus tant (1906) was followed by ‘Hab. Borneo (Paris either black with triangular basal part, median keel and Stockholm)’ indicating type locality and and apical part yellow or with a pair of long black

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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 narrow, erect, ochraceous, spine with a brown tip black fascia running from antenna to eye. Medioba- at half-length, a somewhat shorter, broad, mainly sal quarter of lorum black. Rostrum reaching almost black, spine at four fifths of length from base, and sternite 3, apex brown. a very small, black to brown spine next to the latter. Thorax (Fig. 8). Pronotum. Marking as in M. quad- Fore femur with a subapical, irregular, dark mark- rituberculata, but differing in the following features: ing, a dark brown to black fascia on upper side, and A pair of black triangles extends from half-length some dark marking at basis. Undersides of middle central fascia along basal half of anterior oblique fis- and hind femora with subapical brown streak. Fore sures; each triangle forms a black hook with another tibia black but apical one sixth and basal two thirds black triangle just below the fissure. Posterior oblique of upper side yellow. Middle tibiae with subapical fissures black-brown. black-brown ring as long as one fourth of tibia and Mesonotum (Fig. 8). Five black fasciae. Median fas- a light brown ring at short distance from basis. Tarsi cia as in M. quadrituberculata. Paramedian fasciae as and claws as in M. quadrituberculata. in M. quadrituberculata but black spots in front of Tegmina and wings. Hyaline. Tegmina with an anas- anterior angles of cruciform elevation larger. Lateral tomosing spot on transverse veins at bases of first fasciae consisting of an irregular linear part reach- and second apical cell and separate spots at bases ing from anterior mesonotum margin to three fifths of third, fifth and seventh apical cells; some speci- of mesonotum length, and a pair of distinct black mens with an anastomosing spot at the bases of the spots. Posterior margin of cruciform elevation black third and fourth apical cells. Tegmina with distinct to dark brown. dark brown marking on transverse veins at bases of Legs. Yellow with greenish tinge and brown to black the first and second, third, fifth and seventh apical marking. Fore femora with three spines: a long and cells; those at bases of first and second apical cells

Downloaded from Brill.com10/10/2021 08:41:27PM via free access Duffels: Revision of Maua (Cicadidae) 313 connected. Apices of longitudinal veins of apical cells this specimen does not belong to M. affinis but to with small brown spots. Venation of tegmina light a new species related to M. palawanensis. The speci- brownish to yellowish variegated with dark brown to men from China differs from M. palawanensis in the black-brown, venation of wings light to dark brown. shorter and more slender male body (body length: Operculum (Fig. 9). Lateral margin sinuate and 33.4–36.3 mm; body length of M. palawanensis: 39 directed backwards or slightly outwards. Apical part mm) and in the lateral fasciae that are broken up in very broadly rounded. Medial margin straight or the China specimen and irregular and very variable very weakly convex to medial corner. Operculum in width, but not broken up, in M. palawanensis. Dr reaching to one fourth or half the length of abdomi- Wei Cong (Yangling, Shaanxi, China) kindly trans- nal segment 3. Ochraceous. Lateral margin of oper- lated the description of Maua affinis in Chou et al. culum, with exception of extreme basal part, with (1997). black band that is fairly strongly widened toward its proximal end and continues across the operculum; Distribution (Fig. 7) posterior margin usually black; basal, incurved, part This species is probably endemic to Mount Kina- of medial margin unmarked. balu. Zaidi and co-authors (see above) recorded M. Abdomen (Fig. 9). Abdomen very broad, widest affinis from various localities in Sarawak and Sabah. across abdominal segment 3 or 4, segment 3: 1.37– These specimens probably belong to either M. born- 1.44 times as wide as pronotal collar, segment 2 only eensis or possibly to M. affinis. slightly narrower. Dorsal side, including timbal covers, light to dark brown and with silvery to golden Material examined. 5?. Malaysia: Borneo: Sabah: pubescence; laterobasal corner of timbal cover dark Mt. Kinabalu, Br. North Borneo, Waterstradt, 1? brown. Tergites as in M. quadrituberculata. Ventral (BMNH) 1? (ZMAN); Mt. Kinabalu, Waterstradt, Lep- ? side ochraceous. Sternite 3 with a pair of brown tri- topsaltria 4-tuberculata Sg., Brit. Mus. 1962–168, 1 (BMNH); Mt. Kinabalu, Mesilau, 8.ii.1964, J. Smart. angles at anterior margin. Posterior margins of ster- Royal Soc. Exped. B.M., 1964–250, 1? (BMNH); nites 3–4 brown. Sternites 5–6 brown but median Mt. Kinabalu, Mesilau Camp, 31.iii.1964, 5000 ft., Royal part ochraceous. Sternite 7 dark brown. Sternite 8 Soc. Exped., coll. S. Kueh, B.M. 1964–250, Mau affinis brown but basal parts of lateral margins yellowish. Dist. Michel Boulard det. 1975, 1? (BMNH). Sternites 3 and 4 with thick, glossy brown, nipple- shaped tubercles; distance between tubercles on Maua borneensis sp. n. sternite 4 1.11–1.2 times (n=4) as wide as distance Figs 10–12, 14 between tubercle and lateral segment margin. Genitalia (Fig. 6). Pygofer ochraceous to light brown. Basal pygofer lobes consisting of a pair of slightly sin- Zaidi and co-authors recorded M. affinis from vari- uate ridges that are broadly connected with laterov- ous localities in Sarawak and Sabah; these specimens entral part of pygofer, reach to three fifths of pygofer may belong to either M. borneensis or possibly to length, and are apically acute. Uncus rounded, about M. affinis: 1.5 times as long as wide; median shallow furrow Maua affinis: Zaidi 1996: 101; Zaidi 1997: 113; reaching to weak median incurvation of distal mar- Zaidi & Ruslan 1998: 352; Zaidi, Ruslan & Azman gin. Basis of uncus inwardly curved and forming two 1999: 306; Zaidi, Ruslan & Azman 2000: 204; flat, about triangular sclerites meeting in midline Zaidi, Noramly & Ruslan 2000: 324. of pygofer; these sclerites form a short median tube enclosing the aedeagus and a pair of lateral protu- Type material. Holotype ?: Borneo: Malaysia: berances at their lateral margins. Aedeagus regularly Sarawak: ‘SARAWAK / Gunong Mulu / Nat. Park’, curved, basally broad, suddenly narrowed at about ‘Site 27. April / G. Api 1500 m / Pandanus Camp / half its length and ending in a simple acute apex. 429541 Scrub / Pandanus Act.’, ‘ J.D. Holloway / Measurements in mm (5?). Body length ?: 39–45.5; RGS Mulu exped. / B.M. 1978–206’ (BMNH). tegmen length ?: 53–55.5; head width ?: 10.8– Paratypes: 8? 2/. Borneo: Malaysia: Sarawak: 11.2; pronotum width ?: 12.0–13.2. same data as holotype, 1? (BMNH); Batu Niah, xii.1980, A. Harmann, B.M. 1984–301, 1? Remark on Maua affinis from China (BMNH); Gunong Mulu Nat. Park, Site 1, Camp The specimen of Maua affinis described and illus- 4, January, 452463, 1790 m, lower montane (moss) trated by Chou et al. (1997: 232, Fig. 114) comes forest, MV- canopy, RGS Mulu exped. B.M. 1978– from Mt. Damaosha, Boyang County, Jiangxi prov- 206, 1? (BMNH); Gunong Mulu Nat. Park, Site ince, China (116°66’E, 29°N). Judging from the 26, April, G. Api, pinnacles, 1200 m, 428545, description and the picture I am convinced that open scrub, MV, J.D. Holloway, RGS Mulu exped.,

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Figs 10–11. 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. 1978–206, 1? (UMS); Foot of Mt. Dullit, the other species of the M. quadrituberculata group. Junction of rivers Tinjar & Lejok, 3.ix.1932, Maua borneensis (body length ?: 31–35 mm) can be Oxford Univ. Exp., B.M. Hobby & A.W. Moore, separated from M. affinis (body length ?: 39–45.5) B.M. 1933–254, H185, 1? (BMNH). Sabah: by its smaller size and the different marking on the nr Long Pa Sia – Long Miau, c. 1000 m, 27.xi.1987, tegmina (see key to the species). Maua borneensis is C. v. Achterberg, 1/ (RMNH); 105 km S of separated from the other Maua species by the rela- Beaufort, airstrip Long Pa Sia, 4˚24’N 115˚43’E, tively short distance between tubercles on sternite 16.iv.1987, semicultivated area, asl. near disturbed 4, which is 0.86–1.00 times as wide as the distance evergreen trop. rainforest, 1000 m, at light, 18.20– between tubercle and lateral segment margin; the 20.30 h, Van Tol & Huisman, 1? (RMNH); 15km other species of Maua have ratios ranging from 1.11 W Lahad Datu, confl. S. Sabran, S. Danum, S/N, to 2.5. 117˚41’E 4˚57’N, 200 m, 23.x.1987, J. Huisman & R. de Jong, 1/ (RMNH); Sandakan dist., Rumidi, Description R. Labuk, 16–39.ix.1973, C. Pruett, B.M. 1975– Ground colour of head, pronotum and mesonotum 590, 1? (BMNH). Indonesia: Kalimantan: Long ochraceous to light brown with greenish tinge, abdo- Nawang, leg. Mjöberg, 1? (ZMAN). men of male and female ochraceous to brown. Head (Fig. 10). Black mark enclosing ocelli, mark- Maua borneensis is alike to M. affinis in the striking ing on supra-antennal plates and triangular similar marking on pronotum and mesonotum and markings below these plates as in M. affinis, but in the marking on the postclypeus, both species lack M. borneensis has an extra pair of drop-shaped the black spot at the clypeal suture that is found in brown to black spots proximally of the triangular

Downloaded from Brill.com10/10/2021 08:41:27PM via free access Duffels: Revision of Maua (Cicadidae) 315 markings. Anterior and ventral parts of postclypeus wide as pronotal collar. Dorsal side, including timbal with black-brown transverse lines in the two series covers, light to dark brown and with greyish pubes- of 9–10 grooves; upper 5–6 pairs of lines reach from cence; laterobasal corner of timbal cover dark brown. a central glabrous area to supra-antennal plates or Tergite 2 with black band of variable width along lateral clypeal clefts; lower 4–5 pairs of lines shorter about two thirds of posterior margin. Posterior mar- and with medial ends of lines of each side connected gins of tergites 3–5 or 3–6 with narrow, light brown by a black line; the black lines of both sides enclose bands. Tergite 7 laterally dark brown. Tergite 8 dark a yellowish wedge-shaped median area; lowest two brown to black. Ventral side ochraceous. Posterior pairs of transverse lines sometimes fused to a pair of margin of sternites 3–4 sometimes brown. Ster- black markings. Lower part of postclypeus at clypeal nite 5 with posterior and lateral parts light brown. suture unmarked. Underside of head yellow with Sternite 6 dark brown but median part ochraceous. black marking. Anteclypeus black, but triangular Sternite 7 black-brown. Sternite 8 brown but laterobasal part, median keel and apical part yellow. Gena basal parts of lateral margins ochraceous. Sternites 3 with black fascia running from antenna to eye. Basal and 4 with fairly thick, light brown to brown, but half of lorum black. Rostrum reaching sternite 3; basally often ochraceous, nipple-shaped tubercles; apex brown. distance between tubercles on sternite 4 0.86–1.00 Thorax (Fig. 10). Marking as in M. affinis, but great- times (n=8) as wide as distance between tubercle and est width of median mesonotal fascia 4–7 times its lateral segment margin. anterior width. Male genitalia (Fig. 12). Male genitalia as in M. quad- Legs. Yellow with greenish tinge and brown to black rituberculata but lateral sides of uncus less oblique. marking. Fore femora with three spines: a long and Female operculum. Ochraceous to brownish with narrow, erect, ochraceous, spine with a brown tip scattered silver pilosity, mediobasally somewhat at half-length, a somewhat shorter, broad, black- darker brown. Somewhat wider than long, medially brown, spine at four fifths of length from base, and reaching to almost hind margin of abdominal seg- a very small, black to brown spine next to the latter. ment 3, laterally reaching to two thirds of segment Fore femur with a subapical, irregular, dark marking, length. Lateral margin with triangular protrusion a dark brown to black fascia on upper side, and some in basal half, apical half brown and weakly convex dark marking at basis. Fore tibia black but apical one to rounded rectangular laterodistal angle; posterior sixth and basal one third to basal half of upper side margin weakly undulate; medial margin concave to yellow. Middle tibiae with subapical black-brown narrow mediodistal angle. ring as long as one fourth or one third of tibia. Meta- Female abdomen. Dorsal side (light) brown with sil- tarsus and mesotarsus of fore and middle legs yellow, ver pilosity. Tergites 2–6 and epipleurites 2–3 with pretarsus black-brown, tarsi of hind legs ochraceous, narrow black bands along their posterior margins; claws brown. epipleurites 4–5 with somewhat broader and more Tegmina and wings. Hyaline. Tegmina with brown irregular black-brown bands. Tergites 3–5 with spots on transverse veins at bases of the second, third brown lateral spot. Lateral parts of tergites 6–8 and and fifth apical cells and in most specimens with epipleurites 6–8 black-brown. Segment 9 brown small brown spot at basis of seventh apical cell. Api- and with short or fairly long, dark brown marking ces of longitudinal veins of apical cells in most speci- along posterior margin of segment 8. Sternites 3–5 mens with small to hardly discernible brown spots. light brown with fairly broad, black-brown band Venation of tegmina light brownish to yellowish var- along posterior margin; sternite 3 with black-brown iegated with dark brown to black-brown, venation of band along anterior margin, and sternites 4–5 with wings light to dark brown. black-brown laterobasal marking. Sternites 6–7 shin- Male operculum (Fig. 11). Triangular. Lateral margin ing black-brown, sometimes with indistinct median directed inwards, weakly sinuate in basal one fourth, brownish patch. and slightly concave or straight to narrowly rounded Measurements in mm (8? 2/). Body length ?: apical part of operculum. Medial margin straight or 31–35, /: 26.5–29.5; tegmen length ?: 43–47, /: weakly concave to medial corner. Operculum reach- 40; head width ?: 10.0–10.9, /: 9.5–10.1; prono- ing to two fifths or three fifths of abdominal segment tum width ?: 10.9–12.3, /: 10.0–10.9. 3. Ochraceous, with small brown marking at base, and a black-brown rim except in the sinuate basal Distribution (Fig. 14) part of lateral margin and the basal part of medial Maua borneensis seems to be endemic to Borneo margin proximal of medial corner. (Sabah, Sarawak and Kalimantan). Dr Zaidi Mohd. Male abdomen (Fig. 11). Broad, widest across abdom- Isa and co-workers (UKM) published the results of inal segment 2 or 3, segment 3 1.09–1.17 times as several cicada inventories in Malaysia. The specimens

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Figs 12–13. Male pygofer in ventral view. – 12, Maua borneensis, holotype; 13, M. palawanensis, Palawan.

from Sabah and Sarawak identified in their papers as Paratypes: Palawan: Palawan, Staudgr., Maua affinis M. affinis are not included in the distribution map Dist. [Distant’s handwriting], NHRS-HEMI since their identity could not be confirmed. 000000012, 1? ? paralectotype of Maua affinis (NHRS); Palawan I., 1? (BMNH). Maua palawanensis sp. n. Maua palawanensis can be separated from the other Figs 13–16 large Maua species by a combination of characters: Maua affinis Distant 1905: 61 (?partim: the specimens the presence of a broad, black, transverse mark at the from Balabac (BMNH) and from Palawan (NHRS) clypeal suture, the irregular, but continuous, lateral listed below as holotype and paralectotype of M. pala- mesonotal fasciae reaching from anterior to almost wanensis may be part of the type series of M. affinis). [Misidentification.] posterior margin of mesonotum, and paramedian Maua affinis: Distant 1912: 41, pl. 5 figs 33a-c; Moulton mesonotal fasciae that are often apically connected 1912: 133; Moulton 1923: 124, 125, 168 (partim: with the median fascia. This species is known from specimens from Palawan and Balabac); Metcalf 1963: Palawan and from Balabac Island between Palawan 492 (partim: Palawan and Balabac). [Misidentifica- and Borneo. tions.] Description of male Type material. Holotype ?: Balabac: ‘Balabac / coll. Ground colour of head, pronotum and mesonotum Noualhier 1898’, ‘Maua / affinis / Dist’ [Distant’s yellowish to greenish brown, abdomen ochraceous to handwriting], ‘Type’ [round label with red circle; brown. print], ‘Distant coll/ 1911–383’, ? paralectotype of Head (Fig. 15). A black mark with three pairs of arms Maua affinis (BMNH). encloses the ocelli, a pair of out curved, fairly broad,

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Fig. 14. Distribution of Maua borneensis (rounds), M. palawanensis (triangles) and M. latilinea (squares).

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

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15 16

Figs 15–16. Maua palawanensis, male paratype, Palawan. – 15, body in dorsal view; 16, abdomen in ventral view.

Operculum (Fig. 16). Lateral margin weakly sinu- that widen to brown to black-brown triangles at ate in basal one third, distal two thirds straight and their lateral ends. Tergite 6 with narrow black band directed slightly inwards to rounded laterodistal cor- along posterior margin. Lateral sides of tergites 6–7 ner. Medial margin almost straight to medial corner. dark brown. Tergite 8 dark brown with castaneous Operculum reaching to one third of abdominal seg- median marking. Ventral side ochraceous with green- ment 3. Ochraceous with greenish tinge, but lowest ish tinge, with waxy, pilose, patches. Sternite 3 with a basal part of operculum dark brown. A broad dark pair of brown triangles at anterior margin. Posterior brown band runs from one third of length of lateral margins of sternite 2 with dark brown band, of ster- margin, along laterodistal corner and posterior mar- nites 3–5 with light brown bands, that on sternite gin, and continues in a narrow brown band in the 5 broader than on sternites 3–4. Sternite 6 brown. basal, incurved, part of medial margin. Sternite 7 dark brown but laterobasal corners light Abdomen (Fig. 16). Abdomen very broad, widest brown. Sternite 8 brown with basal yellowish trian- across abdominal segment 3 or 4, segment 2 only gle covering about half of sternite surface. Sternites 3 slightly narrower, segment 3 1.3 times as wide as and 4 with fairly thick, glossy brown, nipple-shaped pronotal collar. Dorsal side castaneous, timbal covers tubercles; distance between tubercles on sternite 4 with greenish tinge, basal part of lateral margin of 1.51 times (n=1) as wide as distance between tuber- timbal cover dark brown. Tergite 2 with black band cle and lateral segment margin; tubercles dark brown of variable width along posterior margin, and with with exception of the lighter brown proximal sides. narrow median triangle rising from black band to Genitalia (Fig. 13). Pygofer ochraceous, but lateral two thirds of segment length. Posterior margins of pygofer lobes and uncus light brown. Basal pygofer tergites 3–5 with somewhat narrower, black bands, lobes consisting of a pair of slightly converging ridges

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17 18

Figs 17–18. Maua latilinea, male, Kalimantan, Kubing. – 17, body in dorsal view; 18, abdomen in ventral view.

Maua latilinea (Walker) that are broadly connected with lateroventral part Figs 14, 17–21 of pygofer, reach to three fifths of pygofer length, and are apically acute. Uncus short, trapezoid with Dundubia latilinea Walker, 1868: 85, 87. Holotype /: a median lanceolate elevation close to apical margin. ‘Penang [above horizontal line], Wallace’ [below hori- Basis of uncus inwardly curved and forming two flat zontal line; handwritten], ‘latilinea’ [Walker’s handwriting], ‘Type’ [round label with green circle; print] and short, triangular sclerites meeting in midline of (BMNH) [examined]. pygofer; these sclerites form a short, thick, median Cosmopsaltria latilinea: Distant 1889: 48 (partim, as far as tube enclosing the aedeagus and a pair of round pro- the text refers to the female holotype from Penang). tuberances at their lateral margins. Aedeagus regu- Maua dohrni Schmidt, 1912: 65, 66. Holotype ?: ‘Soeka- larly curved, basally very broad, gradually narrowed randa / Januar 1894 / Dohrn’ [print in black box], at about half its length and ending in a simple acute Maua / dohrni / Schmidt / ? Edm. Schmidt / determ. 1912’, ‘Type’ [print in black box; red paper], ‘MIZ apex. 187249’ (MIZ) [examined]. syn. n. Measurements in mm (1?). Body length: 39; tegmen Maua latilinea: Moulton, 1923: 92, 124, 126, 168; Met- length: 48; head width: 10.7; pronotum width: 11.7. calf, 1963: 493, 494. [For further references before 1980 see: Metcalf 1963]. [Misidentification: Cosmopsaltria latilinea (nec Walker); Distribution (Fig. 14) Distant 1889: 48 (partim), Pl. iv, Figs 15, 15a, b. This species is known from Palawan and from Bala- (the specimen described and depicted here belongs to Orientopsaltria padda (Distant, 1897) (see Duffels & bac Island, between Borneo and Palawan. Zaidi 2000: 209).]

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Study of this material proved that M. dohrni is a junior synonym of M. latilinea.

Description Ground colour of head, pronotum and mesonotum brown; abdomen of male ochraceous to brown, abdomen of female with reddish tergites and black- brown 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 Fig. 19. Maua latilinea, male Kalimantan Kubing, with black lines in upper 8 transverse grooves of pygofer in ventral view. either side; upper 5 pairs of lines reaching to supra- antennal plates or to lateral clypeal clefts, other lines shorter. Medial ends of transverse lines of each This paper presents the first description of the male side connected by a black line; the two lines con- of M. latilinea. This species is easily separated from necting the upper 5 pairs of lines enclose a glabrous the other large Maua species by the shape of the oval area of the ground colour at anterior margin of central mark enclosing the ocelli, the characteristic postclypeus; the lower 3 pairs of lines and a pair of marking on the postclypeus, the absence of a broad, large black marks in basal two fifths of postclypeus black fascia along inner side of lateral part of ambi- enclose a narrow median space reaching to clypeal ent fissure, the relatively narrow ochraceous inter- suture. Anteclypeus black, but triangular basal part, space between central pronotal fasciae, the broad and median keel and apical part yellow. Gena with very straight lateral fasciae on the mesonotum that reach broad black fascia running from antenna to eye. from the anterior mesonotum margin to almost hind Basal half of lorum black. Rostrum reaching to one margin of mesonotum, the cup-shaped and triangu- sixth of male sternite 3; apex brown. lar markings on respectively tergites 2 and 3 and the Thorax (Fig. 17). Pronotum. Paired central fasciae very characteristic shape of uncus and basal pygofer fairly broad, strongly widened toward anterior pro- lobes. notum margin and almost fused at anterior margin of pronotal collar; the central fasciae enclose a nar- Synonymy row median interspace that is equally broad along In February 2008, Dr Tomi Trilar of the Priro- its whole length. A pair of narrow black triangles doslovni Muzej Slovenije, Ljubljana visited the extends from half-length central fascia along basal Museum of the Institute of Zoology, Polish Academy one third of anterior oblique fissures. Anterior of Science, Warsaw, Poland. He found the ‘Type’ oblique fissures unmarked. Posterior oblique fissures and the ‘Co-Typus’ of M. dohrni Schmidt, 1912 black in anterior two third part, but entirely black in in the collection of this museum, brought these to specimen from Karimata Island. Inner side of lateral my attention and kindly sent me excellent photo- part of ambient fissure without fascia. Pronotal collar graphs of the type specimens (Figs 20–21). Later, with a broad, black mark along medioproximal cor- Dr D. Mierzwa (MIZ) was so kind to send me the ner of ambient fissure, medial part of mark reaching ‘Type’ and the ‘Co-Typus’ of M. dohrni on loan. to black posterior margin of pronotal collar.

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Mesonotum (Fig. 17). Five black fasciae. Median fas- with a pair of sublateral rectangular brown mark- cia widens from anterior margin of mesonotum to ings. Tergites 4–7 with dark brown to black-brown 2–3 times its anterior width at one third of its length, median, transverse marking at anterior margins slightly narrows at about half length, widens again to reaching to one third or half the segment length, a 3–4 times its anterior width at two thirds of length, pair of dark brown to black-brown sublateral mark- and strongly narrows to its posterior end at level of ings, and dark brown to black-brown bands, one anterior angles of cruciform elevation. Paramedian fifth to one fourth as high as the segment, along fasciae fairly broad and extending from anterior mes- posterior margins. Tergite 8 brown with irregular onotum margin to about half-length of mesonotum. dark brown marking. Ventral side light ochraceous. A pair of small black spots in front of anterior angles Posterior margin of sternite 2 dark brown. Sternite of cruciform elevation. Lateral fascia on mesonotum 4 with a pair of large brown triangles reaching from broad and straight, and reaching from anterior mes- anterior sternite margin to four fifths of its length. onotum margin to almost hind margin of mesono- Sternites 4–5 light brown, sternite 7–8 dark brown. tum. Posterior margin of cruciform elevation black Sternites 3 and 4 with fairly narrow, brown to black- to dark brown. brown, nipple-shaped tubercles with distinct trans- Legs. Ochraceous with brown to black marking. verse ridges; distance between tubercles on sternite Fore femora with three spines: a long and narrow, 4 1.26–1.42 times (n=4) as wide as distance between erect, black-brown, spine at half-length, a somewhat tubercle and lateral segment margin. shorter, broad black spine at four fifths of length Male genitalia (Fig. 19). Pygofer oval-shaped. Basal from base, and a small, black to brown spine next pygofer lobes form a pair of large, black-brown, to the latter. Fore femur with a subapical dark patch spine-shaped projections reaching from two fifths on inner and outer side and a dark brown to black to three fourths of pygofer length. Uncus triangular fascia on upper side. Fore tibia dark brown but basal with straight lateral margins, and strongly sclerotised. three fourths of upper side yellow. Middle tibiae with Underside of the uncus forming a pair of fairly short, subapical black-brown ring as long as one fourth of narrowing lobes with a lateral ridge, that sustain the tibia. Meta- and mesotarsus of fore and middle legs, apex of the aedeagus. Aedeagus regularly curved and and basal one third of pretarsus ochraceous, apical gradually narrowing to the simple acute apex. Pygofer part of pretarsus brown; tarsi of hind legs ochra- ochraceous, but distal half of lateral sides, apical half ceous. Claws brown. of basal pygofer lobes and uncus black- brown. Tegmina and wings. Hyaline. Tegmina with distinct Female operculum. Lateral margin straight, laterodis- dark brown spot on transverse veins at bases of sec- tal corner rounded rectangular, distal margin undu- ond and third apical cells and with small brown spots late. Medial part of operculum blackish. at apices of longitudinal veins of apical cells 1–3. Female abdomen. Tergites, especially tergites 2 and Venation of tegmina and wings light to dark brown. 3, reddish. Cup-shaped median black marking on Male operculum (Fig. 18). Triangular. Lateral margin tergite 2 and median black triangle on tergite 3 as very weakly sinuate in basal one third, apical part in male. Segment 4 with a median irregular black straight or slightly concave and directed inwards. marking. Anterior parts of segment 5–7 blackish Apical part of operculum narrowly rounded. Medial suffused. Posterior margins of all tergites with black margin weakly convex to medial corner. Operculum bands, those on segments 5–7 broader than on the reaching to one fifth of abdominal segment 3. Ochr- other tergites. Sternites black-brown. Valvae black- aceous, but dark brown to black in medial one third brown. Segment 9 with broad, black pattern along of basal part and along apical and medial margins. lower margins. Male abdomen (Fig. 18). Abdomen broad, wid- Measurements in mm (4?). Body length: 38.5–40.5; est across abdominal segment 3 or 4, segment 3: tegmen length: 47.5–51; head width: 10.0–11.0; 1.16–1.35 times as wide as pronotal collar, segment pronotum width: 11.6–12.2. 2 only slightly narrower. Dorsal side, including timbal covers, brown, with greyish pubescence. Tergite 2 Distribution (Fig. 14) with cup-shaped or trapezoid median black marking According to its labels, the female type specimen between anterior and posterior tergite margins and of Dundubia latilinea was collected in Penang and with black band of variable width along posterior comes from the Wallace collection. Dr George Bec- margin. Tergite 3 with median black triangle, about caloni (BMNH; personal communication 2007) as wide as cruciform elevation at anterior tergite kindly brought to my attention a remark in the margin, reaching to three fourths of tergite length, introduction of Wallace’s ‘Catalogue of the Cetonii- and narrowly (in the paratype: broadly) connected dae of the Malayan Archipelago’ (1868: 522): ‘The with black posterior tergite margin; tergite 3 also specimens collected by Mr. Lamb, and noted as from

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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.” “Pen- ang” 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 Kali- mantan 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).

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 Tro- pisch Nederland’ (Anonymous 1938) Soekaranda is spelled as Soekaradja; in the ‘Gazetteer Nasional NamaNama Geografie’ (Anonymous, 1978) it is Figs 21. Maua latilinea, male holotype M. dohrni, body spelled as Sukaraja. Both sources give three locali- in ventral view. Photo Dr T. Trilar. ties in Sumatra, all in the province of Bengkulu

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22 23

Figs 22–23. 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 coordi- 7.xi.1999, 01006’00’’S 110028’22’’E, Oilpalm plantation, nates of Dohrn’s Soekaranda revealed a publica- near riverine forest, at light, M. Lammertink 1? (ZMAN); tion by Schneider (1906) mentioning the visit of W. Kalimantan, Karimata Island Nat. Ketapang Res, ii.1991, IIS 910025, per Sutrisno Djenal, General Collec- Dr G. Schneider in 1897 to Dr [W.L.]H. Dohrn 0 tions primary rainforest, closed canopy, 350 m, 108 40’ who lived in a village called Sukaranda that is located – 109010’ 1025’ - 1050’ LS, 1? (ROM). 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 Mau albigutta group Doesburg (2004: 107): 3°37’N and 98°14’E. Wolfgang Ludwig Heinrich Dohrn (1838–1913) The M. albigutta group consists of two medium- was the son of the coleopterist Carl August Dohrn sized species (body length of male: 24–29.5 mm): (1806–1892) and brother of the well-known hemi- M. albigutta and M. linggana. Lateral mesonotal fas- pterist Anton Dohrn (1840–1909). He collected in ciae consisting of three distinct black spots or short Sumatra for the Museum für Naturkunde in Stettin lines on a row. Apical cell 1 of the tegmen is relatively (Horn & Kahle, 1935: 59). The material collected long. The basal vein of apical cell 1 is more than half by Wolfgang Dohrn is now in the Museum of the as long as the longitudinal vein of apical cell 1. The Institute of Zoology, Warszawa, Poland. male abdomen is fairly broad and about equally wide across the segments 2–5. Sternites 3 and 4 have a pair Material examined. 4?. Indonesia: Sumatra: Soeka- of narrow protuberances, which are attached to the randa, i.1894, Co-Typus Maua Dohrni Schmidt ?, sternites along their whole length. The basal pygofer Edm. Schmidt determ. 1912, MIZ 187248, 1? (MIZ). lobes are broadly rounded ridges that widen from the Borneo: Kalimantan: Kubing, Kec. Ketapang, 80m, base of the pygofer to three fifths of its length.

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24 25

Figs 24–25. M. albigutta, Malaysia, Johor, Endau Rompin N.P. – 24, abdomen in lateroventral view; 25, male pygofer in ventral view.

Mau albigutta (Walker) and reach to half length abdominal segment 3, while Figs 22, 24–26 those of M. linggana are broadly rounded apically, Dundubia albigutta Walker, 1857: 83. Holotype ?: ‘Su- have a less distinct median protrusion and reach or matra / Sir S. Raffles’, ‘59–57 / Vigors Coll.’, ‘albigutta reach just beyond posterior margin of abdominal / n s.’ [handwritten], ‘Type’ [print in round label with segment 2 (Fig. 27). green margin] (BMNH) [examined]. Leptopsaltria albiguttata [sic]: Distant, 1889: 36; Distant, Maua albigutta can be separated from Purana species 1891: pl. viii Figs 8, 8a-b; Distant 1892 pl. x Figs 4, of the same size by its broad abdomen, character- 4a-b. istic male operculum, the brownish suffused apical Purana albigutta: Metcalf 1963: 463–464; Duffels & Van part of the tegmina, the three distinct black spots der Laan 1985: 105; Zaidi & Ruslan 1997: 221; Zaidi, on a row forming the lateral mesonotal fascia, the Noramly & Ruslan 2000: 322–323; Zaidi, Azman & dark brown to black posterior margins of the male Ruslan 2001: 109, 110, 112, 115. Maua albigutta: Moulton 1923: 121, 124, 125, 168; Goga- abdominal segments, and the morphology of the la et al. 2004: 1–14; Schouten et al. 2004: 373–374. male genitalia. [For further references before 1980 see: Metcalf 1963 and Duffels & Van der Laan 1985]. Synonymy Moulton (1923) synonymised Purana jacobsoni The males of M. albigutta (body length: 25–26 mm) Distant, 1912 with Maua albigutta. The type of are slightly smaller than those of M. linggana (27– P. jacobsoni in BMNH comes from Java and has the 29.5 mm). Maua albigutta and M. linggana can also following labels: ‘Purana / jacobsoni / type Dist’, be distinguished by the shape of the male opercula. ‘Jacobson / Depok / Java / Nov. 1908’, ‘Java / (Jacob- The male opercula of M. albigutta (Fig. 24), strongly son) / 1912–282’, ‘Type’ [round label with red cir- taper to the apex, possess a distinct medial protrusion cle; print], ‘11/15’. I have compared this type with

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M. albigutta, and found that P. jacobsoni and M. albi- one third of length of cruciform elevation, very nar- gutta differ e.g., in the male opercula and the basal row in anterior half, suddenly widened to 3 times and lateral pygofer lobes of the male genitalia. its anterior width at two thirds of its length and gradually narrowed again to the posterior. Parame- Description dian fasciae reaching from anterior margin to about Ground colour of head, pronotum and mesonotum half-length of mesonotum, as wide as anterior part of ochraceous to light brown often with greenish tinge, median fascia, but basally wider and also widening to abdomen of male and female ochraceous to brown. its distal end, which is sometimes recurved. A pair of Head (Fig. 22). Each of the ocelli enclosed by a black black spots in front of anterior angles of cruciform triangle, apices of the three triangles point to distal elevation. Lateral fasciae on mesonotum consisting end of epicranial suture. Triangle around median of three distinct black spots on a row. ocellus with distinct protrusions pointing laterodis- Legs. Yellow to light brown with brown to black tally, triangles around paired ocelli sometimes with marking. Fore femora with three dark brown to weak protrusion pointing to the posterior. Supra- black-brown spines: a long and narrow, erect spine antennal plates with distinct black marking. A pair at one third of length from base, a somewhat shorter of right angled hook-shaped lines with equally long and broader spine at four fifths, and a small spine legs extend from just below supra-antennal plates next to the latter; apical part of femur and a fas- to level of paired ocelli. Inner margin of eyes with cia on upper side dark brown. Fore tibia with dark a pair of narrow black spots. A pair of juxtaposed brown fascia on underside. Tarsi of fore and middle half-round spots in line with paired ocelli at poste- legs brown sometimes with greenish tinge, hind tarsi rior margin of head. Anterior and ventral parts of ochraceous, claws brown. postclypeus with two series of black-brown trans- Tegmina and wings. Hyaline. Tegmina with apical verse lines in all (8–9) transverse grooves; upper 3–4 part of apical cells 1–2 brownish suffused, with a pairs of lines reach from a central glabrous area to brown spot on transverse veins at bases of second and supra-antennal plates or lateral clypeal clefts, the third apical cells, and with small brown spots at api- other transverse lines (much) shorter. Medial ends ces of longitudinal veins of apical cells 1–4. Venation of upper 5–6 pairs of transverse black lines of each of tegmina light brownish to yellowish variegated side connected by a black line, the connecting black with dark brown to black-brown, venation of wings lines of both sides enclose a glabrous oval area at light to dark brown. anterior margin of postclypeus; lowest two pairs of Male operculum (Fig. 24). Triangular with a distinct transverse lines sometimes fused to a pair of black medial protrusion. Lateral margin oblique, sinuate markings. Lower part of postclypeus at clypeal suture at base and slightly concave at half-length. Opercu- unmarked. Anteclypeus with a pair of paramedian, lum reaching to half-length abdominal segment 3. black-brown, oblong markings parallel to keel. Gena Ochraceous, sometimes with greenish tinge; concave with short black fascia running from postclypeus part of lateral margin sometimes brownish. to half or two thirds of width of gena. Lorum with Male abdomen (Fig. 24). Abdomen broad, wid- black mediobasal corner. Rostrum reaching almost est across segment 3 or 4, segment 3: 1.13–1.24 posterior margin of abdominal segment 2. times as wide as pronotal collar. Timbal covers, light Thorax (Fig. 22). Pronotum. Paired central fasciae brown, sometimes with greenish tinge. Tergites 2–5 narrow, strongly widened toward black, anterior light brown, sometimes with greenish tinge, tergite 6 pronotum margin and fused at anterior margin of light castaneous brown, tergites 7–8 castaneous, all pronotal collar; the central fasciae enclose an upside- tergites with silvery pubescence. Posterior margins of down, key-hole shaped median interspace. A pair of tergites 1–7 with narrow, dark brown fasciae. Tergites short lines extends from half-length central fascia 3–7 with faint lateral spots. Ventral side light ochra- along basal one third or half of anterior oblique fis- ceous. Sternites 6–8 castaneous brown. Sternites 3 sures; each line forms a black hook with an oblong and 4 with a pair of narrow, brown protuberances, spot just below the fissure. Posterior oblique fissures which are attached to the sternite along their whole black-brown. Inner side of lateral part of ambient fis- length; those on sternite 4 somewhat smaller than on sure with a relatively broad fascia that continues in sternite 3. Distance between tubercles on sternite 4 the ambient fissure as a very narrow line reaching to 1.36–1.72 times (n=8) as wide as distance between distal ends of posterior oblique fissures. Pronotal col- tubercle and lateral segment margin. lar with a pair of light brown markings at lateral side Male genitalia (Fig. 25). Pygofer ochraceous, but lat- of ambient fissure. eral pygofer lobes dark brown. Basal pygofer lobes Mesonotum (Fig. 22). Five black fasciae. Median fas- rounded and widening from base to three fifths cia, reaching from anterior margin of mesonotum to of pygofer length. Uncus short, trapezoid; apical

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Fig. 26. Distribution of Maua albigutta (rounds), M. linggana (triangles) and M. platygaster (squares).

margin with very weak median incurvation. Basis of Biology uncus inwardly curved and forming two triangular Maua albigutta is a common species in localities like sclerites meeting in midline of pygofer; these sclerites Endau Rompin N.P., Johor, Malaysia (Schouten et. form a pair of lateral, round, protuberances at their al. 2004) and Langkawi Island (Zaidi, Azman & lateral margins and a short median tube enclosing Ruslan 2001). In Endau Rompin N.P., aggregations the aedeagus. Aedeagus regularly curved and gradu- of singing males were found low on the tree trunks ally narrowing to a simple acute apex. in March 1999. The species was less common from Female operculum. Lateral margin convex, latero- August to November 1999 and males were sing- distal corner broadly rounded, distal margin very ing higher up in the canopy. Dr Matija Gogala and weakly undulate. Dr Tomi Trilar (PMSL) recorded and described its Female abdomen. Tergites 2–6 with a very narrow very characteristic and attractive sound (Gogala et brown to black line along posterior margins, but on al. 2004). The calling song is complex and has pro- tergites 3–4 the central part of this line, which is as nounced frequency modulated parts. From video wide as the cruciform elevation, is about as high as recordings, made in nature in Endau Rompin, the one third of tergite. Lateral parts of tergite 7 with movement of the abdomen could be attributed to brown to black marking on anterior two thirds. Ster- particular characteristics of the song. nites 4–6 with brownish central marking, sternite 7 black-brown along anterior margin. Valvae black- Distribution (Fig. 26) brown. Segment 9 dorsally with a pair of parame- Maua albigutta is widely distributed in the Malay dian black triangles and ventrally with black-brown Peninsula and Sumatra, and known from Siberut marking along lower segment margin. Island. Zaidi, Noramly & Ruslan (2000) recorded Measurements in mm (8? 5/). Body length ?: this species from Borneo (Sabah, Tibow, Tawau). In 24–28, /: 18–20.5; tegmen length ?: 30–34, /: spite of intensive collecting in Sabah and Sarawak 26.5–31; head width ?: 7.3–8.4, /: 6.8–7.4; pro- in the last decades, this is the only record of notum width ?: 7.3–8.5, /: 6.9–7.7. M. albigutta from Borneo. I have not seen the Sabah

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Maua linggana Moulton specimens, so that the occurrence of M. albigutta on Figs 23, 26–28 this island could not to be confirmed. Maua linggana Moulton, 1923: 69, 125, 168. Holotype Material examined. 79? 17/. Indonesia: Sumatra: ?: ‘Lingga / Nov. / 1897’, ‘1923–354 / Brit. Mus.’, Brastagi [= Berastagi], 5.1918, Corporaal 1/ (ZMAN); ’Mau linggana Moulton / Type ? 25/3/23’ [Moulton’s Fort de Kock, 920 m, 1925, leg. E. Jacobson, 1? (ZMAN); handwriting], ‘Type’ [print in round label with red Labuan Bilik [Labuanbilik], HE 5329, 1922–1923, Palm, margin] (BMNH) [examined]. 15? 3? (MZHF); Lau Rakit, i-iv.1919, Corporaal, 1? Maua linggana: Metcalf 1963: 494; Gogala et al. 2004: (ZMAN); Medan, Pagger Alam, coll. Dr MacGillavry, 4–5. vi. 1885, Debussy, 1? (ZMAN); Padang, Ex F.M.S. Muse- um, B.M. 1955–354, Raffles Museum Singapore, xi.1924, Maua linggana can be separated from M. albigutta C.B.K. & N.S., 9? (MNKM); Lampung Krui: Lampung by the male opercula that are broadly rounded api- Krui: Pahmungan 103˚57’E 5˚11’S, Damar garden sing- cally and reach to or just beyond the posterior mar- ing on trunk Shorea javanica, 5.xi.2001, K. Smets, 1? gin of abdominal segment 2. Maua linggana (body (ZMAN); Penengahan 103˚56’E 5˚08’S, Damar garden, 12.xi.2001, K. Smets, 1? (ZMAN) 1? (MZB); Sumatra length males: 27–29.5 mm) is only slightly larger occ., M.R. Belg., 7866, van Lansberg, 2? (KBIN); Sumat- than M. albigutta (body length males: 25–26 mm). ra, 746, R. Mus. Hist. Nat. Belg. I.G. 13.409, 1? (KBIN). Both species have a pair of weakly developed pro- Siberut Island: Ex F.M.S. Museum, B.M. 1955–354, tuberances on sternites 3 and 4 instead of the large Raffles Museum Singapore, 9.1924, C.B.K. & N.S., 1? nipple-shaped tubercles found in the species of the (MNKM). Peninsular Malaysia: Kedah: Kota Walamuir, M. quadrituberculata group. 2.i.1982, Marzuri, 1? (PMSL). Kelantan: Panjang Kelan- tan, 21.i.1983, Samsuddin Matzam, 1? (MNKM); Salor, 20.i.1982, Said Tarmizi, 1? (PMSL). Pahang: Cameron Description Highlands, 16.v.1991, Ismail Yusof, 1? (UKM); Fraser’s Ground colour of head, pronotum and mesonotum Hill, 3600–4600 ft, 6.6.1941, Ex F.M.S. Museum, ochraceous to light brown sometimes with greenish B.M. 1955–354, 1? (MNKM); Krau Wildlife Reserve, tinge, abdomen of male light castaneous brown, that Kuala Lompat sector 0.25 km NW Rangers post, 8 km W of female ochraceous to brown. Kuala Krau, primary forest disturbed by excessive flood- Head (Fig. 23). A black round encloses each of the ing and treefall, at light, 3.ii.1997, M. Kos, 1/ (ZMAN); Krau Wildlife Reserve, Kuala Lompat sector, 1.25 km ocelli, the three rounds are fused; round enclosing NNW rangers post 8 km W Kuala Krau, secondary forest median ocellus with small protrusions pointing lat- riverbank, Old Orang Asli settlement, at light, 15.xii.1996, erodistally, rounds enclosing paired ocelli sometimes M. Kos & S. Azman, 2/ (ZMAN); Kuala Lompat, with weak protrusion also pointing laterodistally. 11–13.viii.1990, Zaidi & Ismail, 1? (UKM). Johor: Supra-antennal plates with distinct black mark- Endau Rompin N.P., 50–70 m, 10–12.vi.1996, T. Trilar, ing. A pair of right-angled hook-shaped markings, ? M. & N. Gogala, 1 (PMSL); Endau Rompin N.P., occasionally divided in two spots, extends from just road staging point to Kuala Jasin, bridge 9, disturbed forest margin, at light, 22–23.iii.1999, J.P. & M.J. Duffels, below these plates to level of paired ocelli. Inner M. Zaidi & M.Y. Ruslan, 1? (ZMAN); Endau Rompin margin of eyes with a pair of narrow black spots. N.P., Junction Sg. Endau / Sg. Senawak 2˚32’N 103˚23’E, A pair of juxtaposed half-round spots in line with the base camp, disturbed forest, at light, 210 m, 19.iii.1999, paired ocelli at posterior margin of head. Anterior J.P. & M.J. Duffels, M.Y. Ruslan, 1? (ZMAN), same and ventral parts of postclypeus with two series of data but 18.iii.1999, 1? (ZMAN), 20.iii.1999, 2? black-brown transverse lines in all (8–9) transverse (ZMAN), 21.iii.1999, 1? (ZMAN); Endau Rompin N.P., NERC, 02˚31’45.4”N 103˚23’52.5”E, second- grooves; upper 3 pairs of lines reach from a cen- ary forest, Transect secondary 5, at light, 21.iii.2001, tral glabrous area to supra-antennal plates or lateral M.A. Schouten, 1? (ZMAN); Endau Rompin N.P., clypeal clefts, next two pairs somewhat shorter and NERC, 02˚31’45.4”N 103˚23’52.5”E, secondary forest, the other transverse lines much shorter. Medial ends Transect secondary 11, at light, 16.v.2001, M.A. Schouten, of some transverse black lines of each side connected A.J. de Boer & G. Withaar, 1? (ZMAN). Langkawi Is- by a black line. A pair of paramedian triangles below / land, 14.iv.1928, H.M. Pendlebury, 1 (UKM), same transverse lines, but lowest part of postclypeus at data but 15.iv.1928, 10? 3/ (UKM), 16.iv.1928, 2? (UKM), 17.iv.1928, 1? (UKM), 18.4.1928, 1? clypeal suture unmarked. Anteclypeus black-brown (UKM), 24.iv.1928, 1? 1/ (UKM), 25.iv.1928, with exception of yellowish keel. Gena with short 2/ (UKM), 26.iv.1928, 2/ (UKM), 29.iv.1928, black fascia narrowing from postclypeus to two 1? (UKM). Singapore: Mandai, 3.viii.1960, 2? 1/ thirds of width of gena. Basal two thirds of lorum (MZHF). black. Rostrum reaching almost posterior margin of abdominal segment 2. Thorax (Fig. 23). Pronotum. Marking as in M. albigutta.

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27 28

Figs 27–28. 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 fas- and sometimes with longitudinal fasciae on inner, cia, reaching from anterior margin of mesonotum outer- and upper sides. Fore tibia with dark brown to one third of length of cruciform elevation, very fascia on underside. Tarsi of fore and middle legs narrow in anterior half, suddenly widened to 3–4 brown sometimes with greenish tinge, tarsi of hind times its anterior width at two thirds of its length legs ochraceous. Claws brown. and gradually narrowed again to the posterior. Para- Tegmina and wings. As in M. albigutta, but apical median fasciae anteriorly about 4 times as wide as parts of apical cells 1 and 2 of tegmina less distinctly median fascia at anterior margin of mesonotum and brownish suffused. equally wide in its apical half, but distinctly nar- Male operculum (Fig. 27). Lateral margin oblique, rowed at one fourth of its length; the fasciae reach sinuate in basal one fourth and weakly convex to to just beyond half-length of mesonotum. A pair of broad, and more strongly convex, margin of apical black spots in front of anterior angles of cruciform part of operculum; apical margin becomes more or elevation. Lateral fasciae on mesonotum consisting less straight toward medial corner. Medial corner of three short black lines on a row, posterior two lines narrowly rounded, medial margin straight or weakly sometimes fused. concave from medial corner to operculum base. Legs. Yellow to light brown with brown to black Operculum ochraceous with brown to black-brown marking. Fore femora with a black-brown fascia con- triangular marking at the basal margin of opercu- necting three light brown to black-brown spines: a lum, and reaching, or reaching just beyond, poste- long and narrow, erect spine at two fifths of length rior margin of abdominal segment 2. from base, a somewhat shorter and broader spine at Male abdomen (Fig. 27). Abdomen broad and paral- four fifths, and a very small spine next to the latter; lel-sided, equally wide across segments 2–5, segment fore femora with dark brown spot at distal underside 3: 1.09–1.13 times as wide as pronotal collar. Timbal

Downloaded from Brill.com10/10/2021 08:41:27PM via free access Duffels: Revision of Maua (Cicadidae) 329 covers ochraceous to light brown. Tergites 2–7 light forest, at light, J.P. & M.J. Duffels, 1? (ZMAN); 23 km castaneous brown, tergite 8 somewhat darker brown. W Sandakan, Sepilok tree tower, 118˚06’E 5˚49’N, 0–100 Posterior margins of tergite 2 with narrow, black- m, 1.xi.1987, J. Huisman & R. de Jong, 1? (RMNH). Sarawak: Semongok, 11.xi.1966, G.H.L. Rothschild, brown fascia; tergites 3 and 4 with similar, some- light trap, 9608, Pres. by Com Inst Ent BM 1974–1, what broader, fasciae. Ventral side light brownish; 1? (BMNH). Brunei: Brunei, Waterstradt, Brit. Mus. posterior margins of sternites 3 and 4 dark brown. 1962–168, 2? (BMNH); Temburong District, ridge NE Sternites 3 and 4 with a pair of narrow, light to dark of Kuala Belalong, approx. 300 m, x.1992, J.H. Martin, brown protuberances, which are along their whole 125W m.v. light, BM 1992–172, 2? 2/ (BMNH). length attached to the sternite. Distance between Indonesia: Kalimantan Timur: Tiong Buu (= Nahabuan), tubercles on sternite 4 1.71–2.5 times (n=7) as wide 18 km NW of Longkay, S. Mahakkam, 15.iv.1996, R. Sözer, 1? (ZMAN). as distance between tubercle and lateral segment margin. Male genitalia (Fig. 28). Pygofer ochraceous, lateral Maua platygaster Ashton pygofer lobes dark brown. Basal pygofer lobes are Figs 26, 29 broadly rounded ridges that slightly widen from base to three fifths of pygofer length. Uncus short, trap- Maua platygaster Ashton, 1912: 156. Syntype /: ‘Matang ezoid with broadly rounded corners; apical margin / 3200 ft / 31.7.1909’, ‘Pres. Raffles Mus. / Singapore with very weak median incurvation. Basis of uncus / Brit. Mus. / 1922–302’, ‘Maua platygaster Asht’, ‘From Sarawak / Museum’ (BMNH) [examined]. inwardly curved and forming two triangular scler- Maua platygaster: Moulton 1912: 133; Moulton 1923: ites meeting in midline of pygofer; these sclerites 124, 126, 168; Metcalf 1963: 494. 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 Types less suddenly narrowing, at half-length, to a simple Ashton (1912) described M. platygaster in an appen- acute apex. dix to Moulton’s (1912) ‘Material for a Fauna Female operculum. Lateral margin weakly convex, Borneensis: a list of Bornean cicadas’. In this pub- laterodistal corner broadly rounded, distal margin lication, Moulton wrote the following lines about very weakly undulate. Operculum ochraceous with M. platyptera: “The Sarawak Museum possesses dark brown to black basal triangle. two specimens from Mt. Matang taken in July and Female abdomen. Tergite 2 with black line along pos- August’ and in a footnote he added: ‘This species is terior margin, tergites 3–6 with a very narrow black described by Mr. Howard Ashton in the Appendix line along posterior margins, but on tergites 3–4 at the end of this paper (p. 156)’. Maua platygaster the central part of this line, which is slightly wider was described after one male and one female from than the cruciform elevation, is about as high as one ‘Sarawak’. It seems most likely that the two speci- fourth of tergite. Anterior part of tergite 7 with black mens from Mt. Matang mentioned by Moulton marking or brownish suffused, lateral parts of tergite (l.c.) are the type specimens. Dr M. Zaidi (Univer- with brown to black marking on anterior two thirds. siti Kebangsaan Malaysia) tried to trace the type Sternites 3–6 black-brown along anterior and lateral specimens of M. platygaster in the collection of the margins, sternite 7 black-brown along anterior mar- Sarawak Museum (Zaidi & Ruslan, 1998), but could gin. Valvae black-brown. Segment 9 dorsally with a not find any. My study of the BMNH collection pair of paramedian black triangles and ventrally with revealed one immature female from Mt. Matang that basal two thirds to three fourths black. comes from the Sarawak Museum and is most prob- Measurements in mm (6? 2/). Body length ?: ably a syntype of M. platygaster. 27–29.5, /: 21.5–22.5; tegmen length ?: 34–37, /: 33.5; head width ?: 8.0–8.8, /: 7.9–8.1; pro- The identity of M. platygaster notum width ?: 8.2–9.4, /: 8.0–8.4. The only specimen of this species that I have seen is the alleged female syntype from Mt. Matang Distribution (Fig. 26) (Fig. 29). This specimen can be separated from the Maua linggana is known from various places in other species of the genus by its very characteristic Borneo (Sarawak, Sabah, Kalimantan and Brunei) light brown marking on the tegmina that consists and is probably a Borneo endemic. of anastomosing spots at the bases of the first and second apical cells and at the bases of the third and Material examined. 8? 2/. Malaysia: Borneo: Sabah: fourth apical cells, and separate spots at bases of fifth Siboga Reserve, Kebun Cina, W of Sandakan, 11 km N Se- and seventh apical cells. Ashton’s original descrip- pilok, 25.xi.1989, sample Sab. 42, primary Diptocarp rain- tion of M. platygaster mentioned ‘anastomoses to

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Fig. 29. M. platygaster, female syntype, Sarawak, Mt. Matang. Photo M.J. Duffels van Egmond.

2nd, 3rd and 5th apical cells spotted with fuscous’. with an expansion of the tegmina under 80 mm In spite of the discrepancies between the description (cf. Moulton 1923). The body length of the male and the pattern found in the syntype, the mere exist- type specimen of M. platyptera (21 mm) is even ence of anastomoses sustains the syntype status of smaller than that of M. albigutta (24–28 mm) and the specimen. Anastomosing spots are found in only M. linggana (27–29.5 mm). one other species of Maua. Maua affinis has an anastomosing spot on the transverse veins at the bases of Distribution (fig. 26) the first and second apical cells and separate spots at Maua platygaster was described from Mt Matang, the bases of the third, fifth and seventh apical cells, Borneo, Sarawak. According to Zaidi & Ruslan but some specimens have one anastomosing spot at (1998) the Sarawak Museum has a male specimen the bases of the third and fourth apical cells. Other of M. platygaster from Mt. Penrissen, Sarawak, col- characters of M. platyptera are the rectangular prono- lected by E. Mjöberg. I have not seen this specimen. tum, irregular lateral mesonotal fasciae that strongly Mt Matang is situated west of Kuching. Mt Penrissen widen to their distal ends, the relatively long apical is part of the mountain range southwest of Kuching cell 1 of the tegmen, and the basal vein of apical cell close to the border between Sarawak and Kalimantan 1 which is much shorter than the longitudinal vein of apical cell 1. According to the original descrip- Material examined. Borneo, Malaysia, Sarawak: The fe- tion the male abdomen is very broad and the male male syntype (see above). 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 Acknowledgments taxonomic position of this species. I am very much indebted to the following cura- tors for the loan and gift of material: Mrs Pudji Remark Aswari (MZB), Dr J. Constant & Dr P. Grootaert Moulton’s key to the species of Maua (Moulton (KBIN), Dr M. Gogala and Dr T. Trilar (PSML), 1923) divided the genus in two groups: those with Dr B. Hubley & Dr D.C. Darling (ROM), tegmina expansions under 80 mm and those with Dr G. Lindberg (NHRS), Dr Maryati Mohamed tegmina expansions over 90 mm. The key is mislead- (UMS), Dr D. Mierzwa and Dr J. Szwedo (MIZ), ing because M. platygaster is arranged among the Dr A. Soulier-Perkins (MNHN), Dr J. van Tol larger species of the genus with tegmina expansions (RMNH), Mr M. D. Webb (BMNH), Dr Zaidi over 90 mm. According to the original description, Mohd. Isa and Mr Ruslan Mohd. Yusop (UKM) the expansion of the tegmina in the male M. platyg- and Dr H. J. Zainal Abidin (MNKM). I thank Dr aster is only 70 mm so that it is one of three species Masami Hayshi (Saitama University, Urawa, Japan)

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for his most valuable comments on an earlier draft pour la sience et une jusqu’ici inédite pour la Thaï- of this paper. I thank my wife Greet Duffels van lande (Rhynchota, Cicadoidea, Cicadidae). – Revue Egmond for her help in the field and for making française d’Entomologie (N.S.) 29: 93–120. photographs of cicadas in the BMNH collection. I Chou I., Z. Lei , L. Li , X. Lu & W. Yao , 1997. The Cica- thank Dr Tomi Trilar (PSML) for the photographs didae of China (Homoptera: Cicadoidae). – Ilustrataj of the type specimens of Maua ackermanni and Insect-Faunoj 2: 1–380, 1–16 pls. M. dohrni. I thank Martjan Lammertink (ZMAN) Distant, W.L., 1889. A monograph of Oriental Cicadidae for the gift of a specimen of M. latilinea. I thank 2: 25–48, pls iii-vi. – West, Newman & Co., London. Dr Wei Cong (Yangling, Shaanxi, China) for trans- Distant, W.L., 1891. A monograph of Oriental Cicadidae lating relevant parts of ‘The Cicadidae of China’. A 4: 73–96, pls vii-ix. – West, Newman & Co., London. visit to Peninsular Malaysia in 1999 was partially Distant, W.L., 1892. A monograph of Oriental Cicadidae financed by the Netherlands Foundation for the 5: 97–120, pls x-xii. – Indian Museum, Calcutta. Distant, W.L., 1905. Rhynchotal notes. - xxix. – Annals Advancement of Tropical Research (WR 84–482). and Magazine of natural History (7) 15: 58–70. I thank Dr Zaidi Mohd. Isa and Ruslan Mohd. Distant, W.L., 1906. A synonymic catalogue of Homop- Yusop (UKM) and Francis Cheong, Chew Keng tera. Part 1. Cicadidae. – Trustees British Museum, Lin, Sing Yun Chin, Lili Bte Tokiman, Hazman Bin London, 207 pp. Md. Zaki, Ivy Abdulla, and Heah Hock Heng of Distant, W.L., 1912. Homoptera, Fam. Cicadidae, Sub- the Endau Rompin team of the Malaysian Nature fam. Cicadinae. – Genera Insectorum 142: 1–64 + Society for logistic support and help in the field in plates 1–7. Endau Rompin N.P. I very much enjoyed the coop- Doesburg, P.H. van, 2004. A taxonomic revision of the eration with Dr M. Gogala and Dr T. Trilar (PSML) family Velocipedidae Bergroth, 1891 (Insecta: Heter- during the fieldwork in Endau Rompin N.P. Dr optera). – Zoologische Verhandelingen, Leiden 347: Maryati Mohamed (UMS) has greatly facilitated my 5–110. fieldwork in Sabah in 2001; the very kind assistance Duffels, J.P., 1977. A revision of the genus Diceropyga Stål, received from the staff of UMS is gratefully acknowl- 1870 (Homoptera, Cicadidae). – Monografieën van edged. I specially thank Dick Langerak (ZMAN) for de Nederlandse entomologische Vereniging 8: 1–227. Duffels, J.P., 1983. Taxonomy, phylogeny and biogeogra- preparing the drawings that form such an essential phy of the genus Cosmopsaltria, with remarks on the part of this publication, Rob Portegies (ZMAN) for historic biogeography of the subtribe Cosmopsaltriaria maintaining the database of cicada records and for (Homoptera: Cicadidae). – Pacific Insects Monograph making the maps of distribution and Gerard Verlaan 39: 1–127. (ZMAN) for technical assistance. Duffels, J.P., 2004. Revision of the cicada genus Nabalua (Hemiptera: Cicadidae) from Southeast Asia. – Orien- tal Insects 38: 463–490. References Duffels, J.P. & A.J. de Boer, 1990. Areas of endemism Anonymous, 1938. Atlas van tropisch Nederland. – Konin- and composite areas in East Malesia. – In: Baas P., klijk Nederlands Aardrijkskundig Genootschap & C. Kalkman & R. Geesink (eds.), The plant diversity Topografische Dienst in Nederlandsch-Indië, Amster- of East Malesia: Proceedings of the Flora Malesiana dam/Batavia, ix, 17 pp, 1–31b maps, 1–31b legends. symposium commemorating Professor Dr. C.G.G.J. Anonymous, 1978. Gazetteer nasional nama-nama ge- van Steenis, Leiden, August 1989. Kluwer Academic ografi. – Badan Koordinasi Survey Dan Pemetaan Na- Publishers, Dordrecht, 249–272. sional, Dokumen 12: 1–217. Duffels, J.P. & P.A. van der Laan, 1985. Catalogue of the Anonymous, 1992. Nelles road atlas Indonesia. – Nelles Cicadoidae (Homoptera, Auchenorhyncha) 1956– Verlag GmbH, München, 127 pp. 1980. – Series Entomologica 34: i-xiv, 1–414. Anonymous, 1999. The Times comprehensive atlas of the Duffels, J.P., M.A. Schouten & M. Lammertink, 2007. world. – Times Books, London. 67, v, 220 pp, 124 pls. A revision of the cicadas of the Purana tigrina group Ashton, H., 1912. 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