A REVISION OF THE CICADAS OF THE PURANA ...The Purana carmentegroup is distributed in Viet-nam, the...

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Distant (1905a) erected the genus Purana when he divided Leptopsaltria Stål, 1866 in three genera: Lep- topsaltria, Purana, and Maua. In Distant’s catalogue (1906) these genera were placed in the division Dun- dubiaria. In 1923, Moulton transferred Purana and the genera Leptopsaltria, Maua, Nabalua Moulton, 1923, and Tanna Distant, 1905 to the new section Leptopsaltriaria, characterised by the presence of tu- bercles on the ventral side of the male abdomen. Met- calf (1963) classified the Leptopsaltriaria as a subtribe of the tribe Dundubiini, and added several genera that lack the characteristic tubercles on the male ab- domen to that group. In this paper, we follow Moul- ton’s restricted concept of the Leptopsaltriaria. Moulton (1923) distinguished the genera Leptopsal- tria, Purana, and Maua, largely upon morphometric characters as relative length and width of head, thorax, and abdomen. In our view it is highly questionable whether these characters can be used to unravel the phylogenetic relationships within the Leptopsaltriaria. The recent recognition of the monophyletic Purana nebulilinea group by Kos and Gogala (2000) was a first step to a classification of Purana and related gen- era based on phylogenetic reconstruction. Their mor- phological study of the male genitalia of a number of species of Purana and Maua suggested that the genus Purana is paraphyletic. Kos & Gogala (2000) sup- posed that Purana ubina Moulton, 1923 and its rela- tives and Maua quadrituberculata (Signoret, 1847) and its relatives form one monophyletic group. The present paper proposes another supposedly monophyletic group, the Purana carmente group, for seven species: P. carmente (Walker, 1850), P. barbosae (Distant, 1889), and P. dimidia Chou & Lei, 1997, and 4 new species: P. obducta, P. hermes, P. infuscata, and P. sagittata. The species of the group are described and relationships within the group are investigated. MATERIAL AND METHODS The following abbreviations for the institutions, which are the depositories of the material studied, have been used in the lists of material and throughout the text: BMNH The Natural History Museum (formerly: British Museum (Natural History)), Lon- don BPBM Bernice P. Bishop Museum, Honolulu CAS California Academy of Sciences, Depart- ment of Entomology, San Francisco NMWC National Museum of Wales, Cardiff 29 M. A. SCHOUTEN & J. P. DUFFELS Institute for Biodiversity and Ecosystem Dynamics (Zoological Museum), University of Amsterdam, The Netherlands A REVISION OF THE CICADAS OF THE PURANA CARMENTE GROUP (HOMOPTERA, CICADIDAE) FROM THE ORIENTAL REGION Schouten, M. A. & J. P. Duffels, 2002. A revision of the cicadas of the Purana carmente group (Homoptera, Cicadidae) from the Oriental region. – Tijdschrift voor Entomologie 145: 29-46, figs. 1-20, table 1. [ISSN 0040-7496]. Published 1 June 2002. The Purana carmente group is proposed for a supposedly monophyletic group of seven cicada species from the Oriental region. Two species of this group are redescribed: P. carmente from Java and Sumatra, and P. barbosae from Jolo (Philippines); the latter species is taken out of syn- onymy with P. carmente. Four species are described here for the first time: P. hermes sp. n. from Sabah and Sarawak, P. infuscata sp. n. from Borneo, P. obducta sp. n. from the Malayan Penin- sula, Sabah, and Sarawak, and P. sagittata sp. n. from the Malayan Peninsula. P. dimidia, which was recently described from China and Vietnam, also belongs to this group. A key to identify the males and distribution maps of the species are provided. Correspondence: M. A. Schouten, Institute for Biodiversity and Ecosystem Dynamics (Zoo- logical Museum), University of Amsterdam, The Netherlands, Plantage Middenlaan 64, NL- 1018 DH Amsterdam, The Netherlands. Key words. – Cicadidae; Purana; carmente group; phylogeny; taxonomy; new species; Southeast Asia; Oriental region.

Transcript of A REVISION OF THE CICADAS OF THE PURANA ...The Purana carmentegroup is distributed in Viet-nam, the...

Page 1: A REVISION OF THE CICADAS OF THE PURANA ...The Purana carmentegroup is distributed in Viet-nam, the southern part of China, the southern Philip-pines, the Malayan Peninsula, Borneo,

Distant (1905a) erected the genus Purana when hedivided Leptopsaltria Stål, 1866 in three genera: Lep-topsaltria, Purana, and Maua. In Distant’s catalogue(1906) these genera were placed in the division Dun-dubiaria. In 1923, Moulton transferred Purana andthe genera Leptopsaltria, Maua, Nabalua Moulton,1923, and Tanna Distant, 1905 to the new sectionLeptopsaltriaria, characterised by the presence of tu-bercles on the ventral side of the male abdomen. Met-calf (1963) classified the Leptopsaltriaria as a subtribeof the tribe Dundubiini, and added several generathat lack the characteristic tubercles on the male ab-domen to that group. In this paper, we follow Moul-ton’s restricted concept of the Leptopsaltriaria.

Moulton (1923) distinguished the genera Leptopsal-tria, Purana, and Maua, largely upon morphometriccharacters as relative length and width of head, thorax,and abdomen. In our view it is highly questionablewhether these characters can be used to unravel thephylogenetic relationships within the Leptopsaltriaria.The recent recognition of the monophyletic Purananebulilinea group by Kos and Gogala (2000) was afirst step to a classification of Purana and related gen-era based on phylogenetic reconstruction. Their mor-phological study of the male genitalia of a number ofspecies of Purana and Maua suggested that the genus

Purana is paraphyletic. Kos & Gogala (2000) sup-posed that Purana ubina Moulton, 1923 and its rela-tives and Maua quadrituberculata (Signoret, 1847)and its relatives form one monophyletic group.

The present paper proposes another supposedlymonophyletic group, the Purana carmente group, forseven species: P. carmente (Walker, 1850), P. barbosae(Distant, 1889), and P. dimidia Chou & Lei, 1997,and 4 new species: P. obducta, P. hermes, P. infuscata,and P. sagittata. The species of the group are describedand relationships within the group are investigated.

MATERIAL AND METHODS

The following abbreviations for the institutions,which are the depositories of the material studied,have been used in the lists of material and throughoutthe text:

BMNH The Natural History Museum (formerly:British Museum (Natural History)), Lon-don

BPBM Bernice P. Bishop Museum, HonoluluCAS California Academy of Sciences, Depart-

ment of Entomology, San FranciscoNMWC National Museum of Wales, Cardiff

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M. A. SCHOUTEN & J. P. DUFFELS

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

A REVISION OF THE CICADAS OF THE PURANA

CARMENTE GROUP (HOMOPTERA, CICADIDAE)

FROM THE ORIENTAL REGION

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

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MNM Muzium Negara Malaysia, Kuala Lumpur PMSL Prirodoslovni Muzej Slovenije, LjubljanaUKM Pusat Sistematik Serangga, Universiti Ke-

bangsaan Malaysia, Bangi, SelangorRMNH Nationaal Natuurhistorisch Museum (for-

merly Rijksmuseum van Natuurlijke His-torie), Leiden

UMS Universiti Malaysia Sabah, Kota KinabaluZMAN Zoölogisch Museum, Amsterdam

The following geographical sources were used fortracing the co-ordinates of localities: The Times Atlasof the World (Anonymous 1994), GEOnet NamesServer of the U.S. Defence Mapping Agency (http://gnpswww.nima.mil/geonames/GNS/index.jsp), An-drees allgemeiner Handatlas (Anonymous 1906),Nelles Road Atlas Southeast Asia excluding Indonesia(Anonymous 1992), Aardrijkskundig woordenboekvan Nederlandsch Oost-Indië (Dumont 1917), Atlasvan Tropisch Nederland (Anonymous 1938). The lo-calities and other label data of the specimens studiedfor this revision were filed in a FileMaker Pro 4.0 data-base. The maps of the species distributions were printedfrom this database using the programme MapInfo forPower Mac, version 4.0.3, on maps of ADC-Worldmapversion 2.0 vol. 4 Southern Asia & Australia.

Random selections of specimens (or all availablespecimens) were used for calculating the ranges, aver-ages and standard deviations of the size of the bodyand body parts mentioned in the descriptions. Mea-surements were taken using a sliding calliper.

PHYLOGENY

The Purana carmente group is erected to accom-modate P. carmente and its related species. Themonophyly of this group is based on three presumedapomorphic characters: (1) male timbal cover withtriangular, black marking (2) basal veins of the 2ndand 3rd apical areas of the tegmina not or very weak-ly infuscated, and (3) male opercula relatively longand slender, reaching or passing the posterior marginof the 3rd abdominal segment.

The species centred around Purana tigrina formthe most likely sister group of the P. carmente group.Purana tigrina and its relatives are characterised bythe prominently infuscated 2nd and 3rd apical areasof tegmina and the elongated, first pair of tubercles.The sister group relationship of the P. carmente groupand the P. tigrina group is supported by the followingcharacters: the predominantly black lower part ofpostclypeus and mandibular plate, and the small firstapical cell of the tegmina.

A phylogenetic analysis was carried out in order toinvestigate the relationships within the P. carmentegroup. Purana tigrina was used as outgroup for this

analysis. The characters used are discussed below andthe matrix is given in table 1.

1. – Medial margin of male operculum: (0) shadedwith black (figs. 6, 9, 13, 14); (1) with broad blackband (figs. 3, 12, 19).

2. – Male operculum: (0) short, reaching no fur-ther than half or two thirds of 3rd abdominal seg-ment; (1) long, reaching beyond two thirds or evenbeyond posterior margin of 3rd abdominal segment.

3. – Basal pygofer lobes: (0) long, only basally con-nected to lateroventral part of pygofer (figs. 4, 15-17); (1) short, for the greater part connected to py-gofer (figs. 7, 10, 20).

4. – Uncus: (0) uncus with median lobe, but with-out lateral lobes (figs. 4, 15, 16, 17, 20); (1) uncuswith a median and two lateral lobes (figs. 7, 10).

5. – Claspers: (0) absent (figs. 4, 7, 10, 20); (1) pre-sent (figs. 15-17).

6. – Dark patches on pronotal collar just in front ofthe anterior ends of the lateral mesonotal fasciae: (0)small, not connected with lateral fasciae of mesono-tum; (1) broad, usually connected with lateral fasciaeof mesonotum.

The analysis (exhaustive search) resulted in one sin-gle most parsimonious tree with a length of ninesteps, which is given in fig. 1.

BIOGEOGRAPHY

According to Moulton (1923), Metcalf (1963),Duffels & Van der Laan (1985), Chou et al. (1997),and Kos & Gogala (2000), the genus Purana compris-es at least 35 species, which are distributed mainly inSoutheast Asia: Myanmar (Burma), Thailand, Cam-bodia, Vietnam, the Malayan Peninsula, Philippines,and the Greater Sunda Islands, but also occur in India,Sri Lanka, China, Taiwan, and Japan. So far, only twospecies of Purana have been recorded from east ofWallace’s Line: P. celebensis (Breddin, 1901) from Su-lawesi and P. carolettae (Esaki, 1936) from the Caro-line Islands in the Pacific.

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Table 1. Character state matrix for the species of the Puranacarmente group and the outgroup used in the cladistic analy-sis. Numbers refer to the characters as discussed in the text.

1 2 3 4 5 6

P. carmente 1 1 0 0 0 1P. barbosae 0 0 1 1 0 0P. obducta 0 1 1 1 0 0P. hermes 1 1 0 0 1 1P. infuscata 0 0 0 0 1 0P. sagittata 0 0 0 0 1 1P. dimidia 1 0 1 0 0 ?P. tigrina 0 0 0 0 0 0

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The Purana carmente group is distributed in Viet-nam, the southern part of China, the southern Philip-pines, the Malayan Peninsula, Borneo, Sumatra, andJava. The cladogram of the group (fig. 1) shows thatthe carmente group can be subdivided in two mono-phyletic groups. The first group is distributed in theGreater Sunda Islands and the Malayan Peninsula. Itconsists of four species: P. carmente from Java (andknown from one locality in southern Sumatra), P. in-fuscata and P. hermes from Borneo and P. sagittatafrom the Malayan Peninsula, Sumatra, and Borneo.The second group has a more northern distribution.This group consists of P. dimidia from Vietnam andChina, P. barbosae from the southern Philippines, andP. obducta from the Malayan Peninsula and Borneo.P. dimidia from the mainland of South East Asia is thesister species of the other two species.

TAXONOMY

Characterisation of the Purana carmente groupThe species of the carmente group show a striking

similarity in external characters but a great diversity instructure of male genitalia and in shape and colorationof the male operculum. The body is relatively small:body length of males 19.1-24.2 mm, of females 17.0-20.8 mm. Body generally ochraceous to dark brown,sometimes with an olivaceous tinge on parts of headand thorax, especially in fresh specimens; pronotal col-lar and vertex lobes somewhat lighter. Markings onhead and thorax distinct. Lower part of postclypeusand mandibular plate predominantly black and cov-ered with long silver setae and white wax. Each of thelateral ocelli enclosed by a black, triangular marking.Pronotum with two narrow, black central fasciae andvariable markings on and along the fissures. Anterior

margin of pronotum medially with narrow blackband. Posterior margin of pronotal collar shaded withblack. Mesonotal markings consisting of five distinct,brown to black fasciae: a median fascia, a pair of para-median fasciae, and a pair of lateral fasciae. A pair ofblack dots in front of anterior angles of cruciform ele-vation. Tegmina hyaline, sometimes slightly bronzedtowards apex, basal cell ochraceous; venation ochra-ceous in basal half, brownish to black in apical part.Dorsal side of abdomen densely covered with fine,short, golden setae. Posterior margins of tergites withnarrow black band. All species have a pair of dark tu-bercles on sternites 3 and 4. Timbal covers with dis-tinct triangular brown to black marking reaching fromposterior margin of timbal cover to the anterior. Maleopercula narrow, elongate, and wide apart, their apicesobtusely angulate, but variable in shape, length andcolour. Female opercula short. Male genitalia with apair of distinct basal pygofer lobes and a median uncuslobe, while some species have a pair of lateral uncuslobes or a pair of claspers.

IdentificationThe species of the Purana carmente group mainly

differ in characters of the male genitalia and the maleoperculum. Identification of the females of the car-mente group is difficult, since the coloration of andmarking on the body are very uniform, while thesmall variation in these characters shows overlap be-tween the species. Females of the carmente group canbe distinguished from females of other Purana groupsby the lack of infuscations on the tegmina, except forP. infuscata. The latter species has weakly developedinfuscations at the basal veins of the 2nd and 3rd api-cal areas, and can easily be confused with otherspecies of Purana or even Maua.

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Fig. 1. Cladogram showing the rela-tionships between the speciesof the Purana carmente group.Numbers refer to charactersdiscussed in the text. Closedbars are synapomorphies (5*is a reversal), open bars arehomoplasies.

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Figs. 2-4. Purana carmente, male. – 2, body in dorsal view, Java, Soekaboemi; 3, abdomen in ventral view, Java, Soekaboemi;4. pygofer in ventral view, holotype, Java (bpl: basal pygofer lobe; ml: median uncus lobe).

2 3

4

bpl

ml

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Key to the males of the Purana carmente group 1. Male genitalia with a pair of claspers (figs. 15-17)

.........................................................................2– Male genitalia without claspers (figs. 4, 7, 10, 20)

.........................................................................42. Basal veins of 2nd and 3rd apical areas of tegmi-

na weakly infuscated..........................P. infuscata– Basal veins of 2nd and 3rd apical areas of tegmi-

na not infuscated .............................................33. Uncus extremely slender, apex not bicuspidate

(fig. 17) ..............................................P. sagittata– Uncus broad, with bicuspidate apex (fig. 15) .....

.............................................................P. hermes4. Medial half of operculum heavily touched with

black (figs. 3, 12, 19). Uncus with a single broad,bicuspidate median lobe (fig. 4) ....................... 5

– Medial margin of operculum with fairly narrowblack marking (figs. 6, 9, 13, 14). Uncus with abroad bicuspidate median lobe and two smallerlateral lobes (figs. 7, 10) ....................................6

5. Opercula long, reaching or passing posterior mar-gin of 3rd abdominal segment (fig. 3)P. carmente

– Opercula short, reaching halfway 3rd abdominalsegment (fig. 19).................................P. dimidia

6. Opercula triangular, tapering towards roundedapex, and long, covering 1st pair of tubercles (fig.9) .........................................................P. obducta

– Opercula broadly rounded at apex and short, notreaching 1st pair of tubercles (fig. 6) ............................................................................ P. barbosae

Purana carmente (Walker)(figs. 2-4, 18)

Dundubia carmente Walker, 1850: 71. – Holotype ?: ‘Dun-dubia carmente’ / ‘W Type’ [handwritten]; ‘Java’ [hand-written]; ‘Type’ [printed on round label with green mar-gin] (BMNH) [examined].

Dundubia carmente; Dohrn 1859: 73; Walker 1868: 91.Leptopsaltria carmente Distant, 1889a: 37, pl. 8 figs. 2, 2a-b;

Distant 1892: ix; Bierman, 1908: 151.Leptopsaltria nigrescens Distant, 1889b: 50. – Holotype ?

Leptopsaltria nigrescens from Java, coll. Van Lansberg [notfound in RMNH]; Distant 1889c: 88; Distant, 1889a: 37(in syn. of Leptopsaltria carmente); Distant 1906: 51 (insyn. of Purana carmente); Moulton 1911: 132 (in syn. ofPurana carmente); Distant 1912: 41 (in syn. of Purana car-mente); Moulton 1923: 122 (in syn. of Purana carmente);Metcalf 1963: 464-465 (in syn. of Purana carmente).

Purana carmente; Distant 1905b: 556; Distant 1906: 51;Moulton 1911: 132; Distant 1912: 41; Distant 1913: 39;Moulton 1923: 120, 122, 168; Schmidt 1928: 108; Kato,1932: 161; Wagner 1960: 115-116, fig. 5, Pl. 4.2; Metcalf1963: 464-465; Wagner 1964: Pl. 12.2; Wagner 1968: Pl.1.2; Duffels & Van der Laan 1985: 106; Chou et al. 1997:230, 368.

Purana carmente carmente; Moulton 1923: 122, Metcalf1963: 465.

The following references probably relate to other species:Purana carmente; Zaidi et al. 1990: 265 (= ?P. obducta).Purana carmente; Zaidi & Ruslan 1998: 349 (= ?P. obducta

or P. hermes).

Males of Purana carmente can be distinguishedfrom other species in the carmente group by a verybroad and triangular uncus and by an operculum thatis black for the greater part, pointed outward andreaches to the 4th abdominal segment. P. carmente isquite similar to P. hermes in shape and coloration ofthe male opercula, though the medial part of the op-erculum of the latter species is less heavily colouredwith black. Specimens collected by J. M. A. vanGroenendael are considerably darker than other spec-imens, probably due to conservation or preparation.

DescriptionHead. – Ochraceous to tawny. Eyes encircled with

black, the black markings tend to narrow or to be in-terrupted at level of the vertex lobes. Genae just belowantennae with a transverse fascia between eyes andpostclypeus. Lower half of mandibular plate black.Ventral part of postclypeus with two paramedian se-ries of 3-4 fairly long and 3-4 short, brown to black,transverse streaks. The transverse streaks are mediallyconnected by an brown to black arcuate line that en-closes a broad midventral part of the ground colourand continues into the black coloration covering basalone fourth of postclypeus at clypeal suture. Lateralsides of anteclypeus black. Lateral parts of anteclypeuswith white pubescence, mandibular plates silvery hir-sute. Black-tipped rostrum reaching posterior marginof 2nd abdominal segment. Each of the lateral ocellienclosed by a distally broadening, and proximally at-tenuating, triangular black-brown marking, whichreaches beyond median ocellus and proximally (al-most) reaches anterior margin of pronotum. These tri-angular markings are more or less broadly connectedat median ocellus. A pair of broad, convex Y-shapedfigures, with outer arms longer than inner arms, runparallel to lateral margins of the triangular spots, butare not very clear in the holotype.

Thorax. Pronotum. – Ochraceous to tawny ordark-brown. Pronotal collar somewhat paler than restof pronotum and mesonotum. Anterolateral corner ofpronotum collar rounded with short and blunt later-al tooth; posterolateral corner bent upward. Two nar-row black central fasciae reach from anterior marginof pronotum to pronotum collar, dilated at anteriormargin and half-moon- to crescent-shaped posterior-ly. A pair of, sometimes very light, brownish to blackspots between anterior and posterior oblique fissures.The brown or black colour of the posterior obliquefissure continues posteriorly in lateral direction into anarrow dark brown or black line on ambient fissure.Pronotal collar with dark patches just in front of

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Figs. 5-7. Purana barbosae, male, holotype, Jolo. – 5, body in dorsal view; 6, abdomen in ventral view; 7, male pygofer in ven-tral view (ll: lateral uncus lobe; ml: median uncus lobe).

7

5 6

ll

ml

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proximal ends of lateral mesonotal fasciae. Pronotalcollar with narrow black line along posterior margin.

Mesonotum. – Ochraceous to tawny or dark brown.Median fascia dark brown to black, narrowing fromanterior margin to two thirds of mesonotum length, af-ter which it broadens to 2-4 times its anterior width,only to narrow again more distally, the fascia inciden-tally continues on the cruciform elevation. The slight-ly oblique paramedian fasciae curve inward to middleof mesonotum and slightly widen posteriorly. A pairof black dots in front of anterior angles of cruciformelevation. Lateral fasciae conspicuous, about 2-4 timesas broad as narrowest part of median fasciae, irregularin shape, and reaching from anterior to nearly posteri-or margin of mesonotum. A narrow black fascia alongposterior rim of cruciform elevation.

Tegmina and wings. – Hyaline, slightly bronzedtoward apex, basal cell hyaline. Venation ochraceousin basal half, brownish to black in apical half. Tegmi-na without infuscations. Fourth apical area of tegmi-na about 3 times as long as broad (n = 21).

Legs. – Coxae ochraceous. Fore femora ochraceouswith brownish lines and spots. Anterior side with threespines, proximal spine at about two fifths from base,long and needle shaped; middle spine at about fourfifths from base, more triangular, shorter and broaderthan proximal spine; distal spine immediately adjacentto middle spine, about one third as long as middlespine, and with blunt apex. Spines and areas betweenand around spines brown to black. Proximal and mid-dle spines ochraceous at tip. Fore tibiae ochraceous totawny, distally darkened. Fore tarsi brown, darkeneddistally, claws brown to black. Mid femora ochra-ceous, mid tibiae ochraceous, distally darkening, withlight dorsal band from base to two thirds of length,mid tarsi and claws brown. Hind legs ochraceous, tib-ia with reddish spines, claws tipped black.

Male operculum (fig. 3). – Fairly long, 1.8-2.5times as long as broad (n = 20), reaching or passingposterior margin of 3rd abdominal segment, triangu-lar, attenuate and pointed outwards. Apex rounded orangularly rounded, situated lateral to midline of oper-culum. Medial margin convex, lateral margin convexbasally, and concave from two thirds of length toapex. Medial half of operculum brown to black, later-al half ochraceous, boundary more or less followingthe curve of lateral operculum margin, basal part ofoperculum black. Distance between opercula atapices 2.9-3.5 times the distance at point of closestapproximation (n = 20).

Male abdomen. – Abdomen 1.0-1.2 times as longas head and thorax together (n = 21). Ground colourochraceous. Timbal cover with triangular black mark-ing reaching from posterior margin of timbal cover tohalf or three fourths of its length. Dorsal side of ab-domen covered with short golden setae, which are

most densely set toward the posterior. Tergite 2 occa-sionally with central triangular black marking, whichis about 3 times as long as wide. Tergites 2-4 with twoparamedian rows of bluntly triangular, black mark-ings at posterior margins, tergites 3-6 with narrowblack fasciae along posterior margins. Ventral sidelight ochraceous, darkening proximally; posteriorthird of sternite 7 and entire sternite 8 black. Ventralside of abdomen with long golden setae. Sternites 3and 4 with a pair of well-developed, glossy, brown toblack tubercles.

Male genitalia (fig. 4). – Pygofer ochraceous andoval shaped, slightly constricted in the middle. Basalpygofer lobes narrow and long, entirely black or dark-ening toward the rounded apices that reach halfwaythe length of the pygofer or further, and touch themedian uncus lobe. Basal half of these lobes broadlyconnected with lateroventral part of pygofer. Medianuncus lobe brown to black, fairly broad, constrictedat base, triangularly attenuated towards bicuspidateapex, rounded in lateral view, and half as long as py-gofer. Median uncus lobe with median suture frombase to bicuspidate apex. Lateral processes of pygofershort with recurved, rounded apex, reaching just be-yond anal valves.

Female operculum. – Very short, reaching to twothirds, or occasionally to posterior margin, of 2nd ab-dominal segment. Lateral and posterior marginsslightly convex; laterodistal corner angularly rounded.Medial half, including lateroproximal corner, brownto black, laterodistal corner ochraceous. Operculumdensely covered with silvery setae and often withpowdery white wax.

Female abdomen. – Abdomen 0.9 times as long ashead and thorax together (n = 6). Ground colourochraceous. Dorsal side of abdomen covered withshort golden to silvery setae. Tergites 2-4 with twoparamedian rows of bluntly triangular, black spots atposterior margins, tergites 3-6(7) with very narrowblack fasciae along posterior margins. Ventral side ofabdomen with brownish marking. Abdominal seg-ment 9 with dark coloration along ventral marginsand a pair of paramedian, triangular spots at anteriormargin of pygofer. Ovipositor sheath dark brown,ventral side with long setae, apex with a bundle oflong setae. Ovipositor light brown.

Measurements in mm (20?, 6/). – Body length?: 20.2-23.7 (21.3 ± 0.9), /: 17.9-20.2 (19.0 ± 0.7);tegmen length ?: 25.3-31.0 (26.9 ± 1.3) /: 26.5-28.9 (27.7 ± 0.8); head width ?: 6.2-7.1 (6.6 ± 0.3),/: 6.5-7.1 (6.8 ± 0.2); pronotum width ?: 7.0-8.0(7.3 ± 0.3), /: 7.0-8.0 (7.4 ± 0.4)

Distribution (fig. 18)This species is widely distributed over Java and

recorded from one locality in South Sumatra. The

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Figs. 8-10. Purana obducta, male, holotype, Malaysia, Taman Negara, Pahang. – 8, body in dorsal view; 9, abdomen in ven-tral view; 10, pygofer in ventral view (ll: lateral uncus lobe; ml: median uncus lobe).

10

8 9

ll

ml

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records of Purana carmente from Peninsular Malaysia(Zaidi et al. 1990) and Borneo, Sarawak (Zaidi &Ruslan 1998) probably should be ascribed to one ofthe new species.

Material examined. – 48? 8/. INDONESIA, JAVA: W. Pri-angan, 27.i.1933, J. M. A. van Groenendael, 1? (UKM),same data but: 1935-1939, 5? (ZMAN), 1?, 1/ (UKM),9.iv.1937, 2? (ZMAN), 28.iv.1937, 1? (UKM) 1/ (ZMAN),vii.1937, 3? (ZMAN), 2.ii.1940, 1? (ZMAN); ZW Priangan,1800’, xii.1941, J. M. A. van Groenendael, 2? (ZMAN);Soekaboemi, 2000’, 14.ix.1936, J. M. A. van Groenendael,1?, same data but: 18.iii.1937, 1/, 28.iii.1937, 1?,21.iv.1937, 5?, 17.v.1937, 1?, 8.iii.1940, 1? (all ZMAN);Soekaboemi, Djampang Kidoel, 1000’-1500’, 15.xii.1939, J.M. A. van Groenendael, 1? (ZMAN); omg. [surroundings]Soekaboemie, iv.1933, F. A. Th. H. Verbeek, 1? (RMNH);Buitenzorg, vi-viii.1881, W. C. v. Heurn, 2? (RMNH);Buitenzorg, 1921, W. C. v. Heurn, 1? (RMNH); Buitenzorg,iii.1923, P. Buitendijk, 1?, 1/ (RMNH); Buitenzorg, dr. J.G. Boerlage, 1? (RMNH); Buitenzorg, 1898, J. B. Ledru, 2?(BMNH); Preanger, 20.vi.1937, J. M. A. van Groenendael,1?, same data but: 10.xi.1937, 1?, 14.xii.1937, 1? (allZMAN); Preanger (?), von Beuker, 1? (ZMAN); Wonosobo,v.1909, E. Jacobson, 1? (RMNH); Gng Gedeh, 3500’,6.ii.1940, J. M. A. van Groenendael, 1? (ZMAN); SlopeGoen. Gede, 3000’, 23.i.1940, J. M. A. van Groenendael,1? (ZMAN); Djampangtengah, 1500’, 31.i.1940, J. M. A.van Groenendael, 1? (ZMAN); W. Semarang, Teak forest,5.vi.1926, L. G. E. Kalshoven, 1? (RMNH); Semarang,vi.1919, E. Jacobson, 1? (BMNH); Semarang, Edw. Jacob-son, 1/ (RMNH); Java, 1?, 1/ (RMNH); Java, Ledru 1?(RMNH); Java Merid. 1500’, 1891, H. Frühstorfer, 1?(BMNH); Djokjakarta, 21.v.1973, H. Hazewinkel, 1/(ZMAN); Depok bij [near] Batavia, v-vi. 1932, W. C. v.Heurn, 1/ (RMNH); INDONESIA, SUMATRA: Mt. Dempo,4.ix.1987, J. D. Weintraub, 1? (ZMAN).

Purana barbosae (Distant)(figs. 5-7, 11)

Leptopsaltria barbosae Distant, 1889a: 37. – Holotype ?:‘Barbosae’ [handwritten]; ‘Jolo’ [handwritten]; ‘Type’[printed on round label with red margin], ‘Distant coll.1911–383’ (BMNH) [examined].

Leptopsaltria barbosae; Distant 1890: pl. v, figs. 14, 14a-b;Distant 1892: xi; Moulton 1923: 122 (in syn. of Puranacarmente).

Purana barbosae; Distant 1906: 51; Distant 1912: 41;Moulton 1923: 122; Kato 1932:161.

Purana carmente barbosae; Moulton 1923: 122, 123; Met-calf 1963: 465.

The following references probably relate to other species:Purana barbosae; Zaidi et al., 1999: 201-202 (= ?P. obducta,

P. hermes or P. infuscata).Purana barbosae; Zaidi et al., 2000: 303 (= ?P. obducta, P.

hermes or P. infuscata).

Purana barbosae closely resembles P. carmente butdiffers in colour, size and shape of the male opercula.The opercula of P. barbosae are shorter, less attenuated

posteriorly, and not pointed outward, while their api-ces are broader and more rounded, and their inner api-cal margins only shaded with black. The species isknown only from the holotype, which is in fairly poorcondition: the body is very dark and most markings arevery faint, while some markings, like the lateral fasciaeof the mesonotum, are almost completely absent.

Description of maleHead. – Ochraceous to tawny. Genae with dark

brown marking at inner margin of eye. Mandibularplates covered with long silvery setae, lower two thirdsblack. Upper half of postclypeus on both sides with aseries of 4-5 pairs of short, transverse, brown to blackstreaks which are medially connected by an arcuate linethat encloses the upper part of a broad midventral partof the ground colour and continues into a black col-oration covering basal third of postclypeus at clypealsuture. Anteclypeus pubescent, lateral parts black. Ros-trum ochraceous, short, apex black, reaching tohalfway abdominal segment 2. Markings surroundingthe ocelli as in P. carmente but very weakly developed.

Thorax. Pronotum. – Markings as in P. carmente,but with crescent-shaped proximal ends of central fas-ciae connected. Anterolateral corner of pronotal col-lar with a more distinct and sharper lateral tooth.

Mesonotum. – Markings partly very faint, proba-bly due to immaturity of the holotype. Anterior halfof median fascia narrow, posterior half twice as broad,and broadened at cruciform elevation. Paramedian

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Fig. 11. Distribution of Purana barbosae (dots), P. obducta(squares) and P. dimidia (triangles).

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Figs. 12-14. Male abdomen in ventral view. – 12, Purana hermes, holotype, Borneo, Sabah, Danum Valley; 13, P. infuscata,holotype, Malaysia, Sabah, Bettotan; 14, P. sagittata, holotype, Malaysia, Selangor, Ulu Gombak.

12 13

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fasciae narrow and more strongly curved inwardsthan in other species of the carmente group. A pair ofround black spots in front of cruciform elevation.Lateral markings faint and about as wide as, or slight-ly wider than, posterior part of median fascia.

Tegmina and wings. – As in P. carmente.Legs. – As in P. carmente, but proximal spine a lit-

tle longer, and angle between femur and proximalspine more acute.

Operculum (fig. 6). – Short, about 1.4 times as longas broad, reaching to halfway abdominal segment 3,apex broadly rounded, situated medial of mid-line ofoperculum. Lateral margin straight, medial margin al-most straight. Basal part of operculum dark brown.Medial and apical margins of operculum shaded withdark brown. Distance between opercula at apices 2.7times the distance at point of closest approximation.

Abdomen. – Slender, 1.7 times as long as broad.Dorsal side of abdomen without markings. Lateralhalf of timbal cover mainly black. Tubercles on seg-ment 4 smaller, more elongate, and more directedoutwards than the round, distally directed tubercleson segment 3.

Genitalia (fig. 7). – Pygofer ochraceous, narrow atbase, abruptly broadened in the middle, and narrow-ing distally. Basal pygofer lobes short, reachinghalfway the pygofer length, and broadly connected toanterolateral part of pygofer. Uncus brown to blackwith a broad median lobe, which has a sutural linefrom its base to its blunt bicuspidate apex; medianlobe fused with pair of somewhat shorter, flattened,fairly broad and rounded, lateral lobes. Lateralprocesses of pygofer short with recurved, roundedapices, which do not reach beyond anal valves.

Measurements in mm. – Body length: 22.0; tegmenlength: 26.7; head width: 6.5: pronotum width: 7.0.

RemarksA male specimen from the northwestern part of

Mindanao (Silipon, Buk. Mind. P.I. 24.vi.1932, L.Phillips collector, F. H. Wymore collection), depositedin the CAS collection, resembles the holotype in struc-ture of genitalia and opercula. This specimen differsfrom the holotype of P. barbosae in the medianmesonotal fascia, which is very broad along its wholelength. Due to the poor state of the holotype and thelack of sufficient material for comparison this specimenfrom Mindanao has not been attributed to P. barbosae.

Distribution (fig. 11).The holotype is from Jolo (Philippines), a small is-

land east of Sabah. The records of Purana barbosaefrom Sabah, Borneo by Zaidi et al. (1999) and Zaidiet al. (2000) probably should be ascribed to P. obduc-ta, P. hermes or P. infuscata.

Purana obducta sp. n.(figs. 8-11)

Type material. – Holotype ?: ‘ MALAYSIA / Pahang/ J. P. & M. J. Duffels / & M. Y. Ruslan’, ‘Taman Ne-gara NP / Sungai Relau, E of Merapoh / 4°41’N102°03’E / 9-11.iii.1999’, ‘gardens / at light’ (ZMAN).– Paratypes 41? 3/: MALAYSIA, PENINSULAR

MALAYSIA: Pahang, Bukit Ibam, 90 km WNW ofKuala Rompin, ca. 50 m, 4-9.X.1961, K. J. Kuncheriacollector Bishop, 1? (BPBM); Pahang, Merapoh,31.iii.1993, Zaidi, Ruslan & Kundin, 1? (UKM);Merapoh, 30.viii.1996, Ismail & Muzamil, 1?(UKM), same locality, 10-11.iii.1997, 8? (UKM); Pa-hang, Merapoh, Kuala Juram, 5-9.v.1997, Ruslan, Is-mail, Azman & Bidi, 2? (UKM), same locality but 28-29.vii.1995, Zaidi & Ruslan, 1? (UKM), same localitybut 10-11.iii.1997, Ismail & Muzamil, 2? (UKM);Taman Negara NP, Kuala Juram, E of Merapoh,4°39’N 102°08’E, 11.iii.1999, J. P. & M. J. Duffels,M. Zaidi & M. Y. Ruslan, 1? (ZMAN), same data but13.iii.1999, 1? (ZMAN), 15.iii.1999, 2? (ZMAN);Taman Negara, Kuala Kenyam, 29-31.viii.1995, Zai-di, Ruslan & M’dir, 5? (UKM); Pahang, Frazer Hill,1.v.1993, Ruslan & Kundin, 1? (UKM); Selangor,Bangi, 1? (UKM), 21.iii.1985, Vijay, 1? (UKM); Se-langor, Bukit Kutu, 3500’, 13.ix.1929, H. M. Pendle-bury, Ex F.M.S. Museum, B.M. 1955-354, 1?(BMNH); Selangor, Bukit Kutu, 3500’, 19.iii.1931, H.M. Pendlebury, 1? (MNM), same data but 10-11.iv.1931, 1? (MNM); Selangor, Hulu Langat SgCongkak, 27-29.xi.1992, Zaidi & Badrol, 1/ (UKM);Perak, Temenggor, Ekspedisi MNS-Belum, 21.xi-5.xii.1993, 1? (UKM); Perak, Lata Ketabu, 9.xii.1996,Muzamil, 1/ (UKM); Perak, Banding, 29-30.vi.1991,Ismail, Yusof & Zabidi, 1? (UKM); Perak, Taiping, ExF.M.S.Museum B.M. 1955-354, 1? (BMNH); Kelan-tan, Gua Musang, 19-22.ii.1994, Zaidi & Talib, 1?(UKM); Gua Musang, in the town, 10-15.iii.1999, J.P. & M. J. Duffels, M. Zaidi & M. Y. Ruslan, 1?(ZMAN); S. Tengah, 4.xii.1923, 1? (BMNH); KualaLumpur, ex coll. Agr. Dept., Ex F.M.S. MuseumB.M.1955-354, 1? (BMNH); Kuala Lumpur,23.iii.1927, H. M. Pendlebury F.M.S. Museums, ExF.M.S. Museum B.M.1955-354, 1? (BMNH).MALAYSIA, SABAH: Tawau, Maliau Basin, 12-25.v.1996, 1? (UMS). MALAYSIA, SARAWAK: Bidi,1908, C. J. Brooks, 1? (BMNH); Kuching, 11.xii, 1?(BMNH). Other material: without locality, 1/ (ZMAN).

This species can be recognised by the male opercu-la, which cover the first pair of tubercles partly orcompletely, and are merely shaded with black alongtheir inner apical margins. The male genitalia of thenew species are rather similar to P. barbosae, but itslateral uncus lobes are more rounded.

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DescriptionHead. – Ochraceous to tawny and somewhat

lighter than in P. carmente. Pattern of markings onhead quite similar to carmente, but on the whole some-what narrower. Eyes encircled with black except at lev-el of vertex lobes. Genae with transverse fascia betweeneyes and postclypeus just below antenna. Ventral sideof postclypeus with two paramedian series of 4-5 longand 3-4 short, brown to black, transverse streaks.Transverse streaks medially connected by an arcuateline that encloses a midventral part of ground colourand continues in black coloration covering basal onefourth of postclypeus at clypeal suture. Black-tippedrostrum reaching to two thirds of 2nd abdominal seg-ment. Black markings on mandibular plate as in car-mente. Triangular markings around lateral ocellishorter and more rounded than in carmente, not con-nected to anterior margin of pronotum, and some-times enclosing a large ochraceous spot. Markings par-allel to posterior margins of spots around ocelli not asdistinct and more irregular shaped than in carmente.

Thorax. Pronotum. – Ochraceous, lighter than inP. carmente. Anterolateral corner of pronotum collarwith distinct lateral tooth. Markings as in carmente butsomewhat narrower. Two narrow black central fasciaereach from anterior margin of pronotum to pronotalcollar; anterior ends triangular, posterior ends cres-cent-shaped, and not quite connected. Dark markingsof posterior oblique fissures and ambient fissure some-what less distinct than in carmente. Area between ante-rior and posterior oblique fissures without spots.

Mesonotum. – Markings as in P. carmente but nar-rower. Median fascia narrowing from anterior marginto two thirds of mesonotum length, where it broadensto 2-3 times its anterior width, but more distally thefascia narrows again and usually continues on the cru-ciform elevation. Lateral fasciae sometimes interrupt-ed and 0.5-3 times as broad as narrowest part of medi-an fascia. Cruciform elevation with reddish tinge, andwith a narrow black fascia along posterior margin.

Tegmina and wings. – Hyaline, somewhat bronzedtowards apex. Venation ochraceous in basal half,brownish to black in apical half. Tegmina without in-fuscations. Fourth apical area of tegmina 2-2.5 timesas long as broad (n = 10)

Legs. – Proximal spine on fore femur a little longerthan in P. carmente, and angle between femur andproximal spine more acute than in carmente. Apices ofspines always ochraceous. Tibiae with reddish tinge.

Male operculum (fig. 9). – Fairly long, 2 times aslong as broad (n = 19), reaching beyond and partlycovering tubercles 3rd abdominal segment. Triangu-lar, tapering towards rounded or angularly roundedapex. Medial margin convex; basal two thirds of later-al margin convex, apical one third weakly concave (lessdistinct than in P. carmente). Apex of operculum me-

dial of midline of operculum. Ground colour ochra-ceous, basal part black, black band along medial mar-gin relatively broad at base and narrow towards apex.Distance between opercula at apices 2.4-2.8 times thedistance at point of closest approximation (n = 19).

Male abdomen. – About 0.8-1.1 times as long ashead and thorax together (n = 19). Ground colourtawny to ochraceous. Timbal covers with triangularblack markings as in P. carmente. Tergite 2 withoutcentral black marking. Black markings along posteriormargins of tergites as in carmente, but less well devel-oped. Ventral side light ochraceous, but brownish tothe posterior, posterior third of sternite 7 and almostentire sternite 8 dark brown to black. Ventral sidewith golden setae. Tubercles on sternites 3 and 4 as incarmente.

Male genitalia (fig. 10). – Pygofer ochraceous, nar-row at base, abruptly broadening halfway, and nar-rowing distad. Basal pygofer lobes short, brownish,reaching to two thirds of pygofer length, basal twothirds or three fourths of these lobes broadly connect-ed to lateroventral part of pygofer. Uncus brown toblack, consisting of a broad median lobe with bluntbicuspidate apex and two somewhat smaller laterallobes, which are more rounded than in barbosae. Lat-eral processes of pygofer short with recurved roundedapex, reaching just beyond anal valves.

Female operculum. – Short, almost reaching poste-rior margin of abdominal segment 2. Lateral marginconvex, posterior margin straight to slightly convex,laterodistal corner rounded. Ochraceous, medial mar-gin more distinctly shaded with black than in the maleoperculum. Operculum densely covered with silverysetae, and medial half often covered with powderywhite wax.

Female abdomen. – Abdomen 0.8-0.9 times aslong as head and thorax together (n =2). Groundcolour ochraceous. Dorsal side of abdomen coveredwith short silvery hairs. Marking along posterior mar-gins of tergites as in P. carmente. Ventral side of ab-domen with numerous blackish markings. Segment 9with dark coloration along ventral margins and a pairof triangular spots at anterior margin of pygofer.Ovipositor sheath dark brown, ovipositor ochra-ceous. Ventral side of abdomen with long setae, apexof ovipositor sheath with a bundle of similar setae.

Measurements in mm (31?, 2/). – Body length?: 19.2-22.0 (20.7 ± 0.7), /: 17.8-19.3 (18.4 ± 0.7);tegmen length ?: 25.9-28.5 (27.4 ± 0.7), /: 26.0-28.8 (27.6 ± 1.2); head width ?: 6.2-7.0 (6.8 ± 0.2),/: 6.5-6.8 (6.7 ± 0.1); pronotum width ?: 6.8-8.1(7.3 ± 0.3), /: 6.8-7.3 (7.1 ± 0.2).

Distribution (fig. 11).This species is found in Peninsular Malaysia,

Sabah, and Sarawak.

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Etymology The name of this species is derived from the Latin

word ‘obductio’ meaning ‘covering’, referring to thefact that this is the only species of the P. carmente groupin which the opercula cover the first pair of tubercles.

Purana hermes sp. n.(figs. 12, 15, 18)

Type material. – Holotype ?: ‘BORNEO: Sabah , /Danum Valley / 5°01’N 117°47’E / 20.X.1987 120 m/ A. H. Kirk-Spriggs’, ‘NMW Sabah (Borneo) / Expedi-tion, / NMW.Z. 1987.094’, ‘light trap sample / primaryforest edge’ (NMWC) – Paratypes 10? 2/: MALAYSIA,SABAH: same data as holotype, 1/ (NMWC); Keningau,kg. Senoa, 500 m, 7.v.1988, Nordin Wahid leg.,00261, 1/ (UKM); Sandakan, Woodin, 1? (BMNH);Sandakan Dist., 16-30.ix.1973, Rumidi & Labuk, 4?(BMNH) 1? (ZMAN); Tawau, Maliau Basin, 12-25.vi.1996, 1? (UMS); Tawau, Brumas camp,xi.1974, C. Pruett, 1? (BMNH). MALAYSIA, SARAWAK:Mt. Dulit. R. Koyan 2500 ft Primary forest,14.x.1932, B. M. Hobby & A. W. Moore, 1?(BMNH); Foot of Mt. Dulit, Junction of rivers Tinjar& Lejok, 1.ix.1932, B. M. Hobby & A. W. Moore,1? (BMNH).

In external features P. hermes is very similar to P.carmente, but it can be distinguished from this speciesby the presence of a pair of claspers in the male geni-talia. The average body size of P. hermes is slightlylarger than that of other species of the carmente group.

DescriptionHead. – Ochraceous. Eyes encircled with black.

Genae with a broad transverse line between eyes andpostclypeus. Lower two thirds of mandibular plateblack. Transverse streaks on ventral side of post-clypeus as in P. carmente. Lateral sides of anteclypeusand ventral side of postclypeus at clypeal suture black.Anteclypeus pubescent, mandibular plate coveredwith long silvery setae. Black-tipped rostrum reachingposterior margin of second abdominal segment.Black markings around lateral ocelli as in carmente,sometimes with small ochraceous spots lateral of me-dian ocellus. Y-shaped figures slightly curved out-ward, strongly broadened anteriorly; medial leg of Y-shaped figure sometimes interrupted.

Thorax. Pronotum. – Ochraceous, pronotal collarsomewhat paler and with a blunt lateral tooth. Centralfasciae fairly narrow, strongly dilated and triangularshaped at anterior margin, slightly broadened and cre-sent shaped posteriorly. Dark patch on lateroproximalangle of pronotum collar larger than in P. carmente.

Mesonotum. – As in P. carmente but lateral fasciaesometimes interrupted.

Tegmina and wings. – As in P. carmente.Legs. – As in P. carmente.Male operculum (fig. 12). – The shape, length, and

pattern of coloration resembles that of P. carmente.Operculum 1.7-2.0 times as long as broad (n = 11),reaching posterior margin of 3rd abdominal segment.Apex angularly rounded to pointed, situated medialto lateral of midline of operculum. Medial marginconvex, lateral margin strongly convex at base andstrongly concave from two thirds of length to apex.Boundary between black medial half and ochraceouslateral part follows curvature of lateral margin of op-erculum more closely than in carmente. Basal part ofoperculum black. Distance between opercula atapices 2.5-2.9 times the distance at point of closestapproximation (n = 11).

Male abdomen. – Abdomen 1.0-1.1 times as longas head and thorax together (n = 11). Markings onabdomen as in P. carmente.

Male genitalia (fig. 15). – Pygofer ochraceous, longand oval as in P. carmente but without the constrictionhalfway. Basal pygofer lobes long (about as long as incarmente), dark-coloured, and reaching to halfway ortwo thirds of pygofer length. Median uncus lobe darkbrown to black, slightly curved inwards, broad at base,abruptly constricted at one fifth of length, and slight-ly narrowing toward blunt bicuspidate apex. Claspernarrow, outwards curved, and darkened to the apex.Anal valves long and pointed. Lateral process of py-gofer hairy, slightly constricted at basal part and notreaching beyond anal valves.

Female operculum. – Somewhat longer than in P.carmente, reaching towards posterior margin of 2ndabdominal segment. Lateroproximal corner onlypartly black. Operculum densely covered with silverysetae and often covered with powdery white wax.

Female abdomen. – As in P. carmente, but browncoloration along ventral margins of 9th abdominal seg-ment somewhat lighter and narrower than in carmente.

Measurements in mm (11?, 2/). – Body length?: 21.4-23.6 (22.5 ± 0.6), /: 18.8-20.1 (19.4);tegmen length ?: 27.2-31.2 (29.6 ± 1.4), /: 27.6-28.0 (27.8); head width ?: 6.8-7.0 (6.9), /: 6.5-6.8(6.7 ± 0.1); pronotum width ?: 7.8-8.0 (7.9 ± 0.2),/: 7.2-7.5 (7.3).

Distribution (fig. 18)This species is recorded from the Bornean states of

Malaysia: Sabah and Sarawak.

Etymology This new species described above is most closely re-

lated to Purana carmente. Since the name carmentewas probably derived from Carmentis, the goddess ofprophecy, the new species is named after Hermes, sonof Carmentis.

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Figs. 15-17. Male pygofer in ventral view. – 15, Purana hermes, holotype, Borneo, Sabah, Danum Valley; 16, P. infuscata,holotype, Malaysia, Sabah, Bettotan; 17, P. sagittata, holotype, Malaysia, Selangor, Ulu Gombak (cl: clasper; ml: median un-cus lobe).

cl

ml

cl

ml

15 16

17

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Purana infuscata sp. n.(figs. 13, 16, 18)

Type material. –Holotype ?: MALAYSIA, SABAH: ‘N.Borneo / Bettotan / NR. Sandakan / July 24th 1927 /C. B. K. & H. M. P. / F.M.S. / Museums’, ‘ExF.M.S. / Museum / B.M. 1955-354’ (BMNH) –Paratypes 3?: MALAYSIA, SABAH: N. Borneo, San-dakan Dist., 16-30.ix.1973, Rumidi & R. Labuk, C.Pruett, B.M. 1975-590, 2? (BMNH); INDONESIA,KALIMANTAN: Barabei, Z.O. Afd. Borneo, geschenk[gift] A. Pool 1883, 1? (ZMAN).

P. infuscata can be separated from the other speciesof the carmente group by its slightly infuscated basalveins of the 2nd and 3rd apical areas of the tegmina.The male opercula are short, reach to two thirds of ab-dominal segment 2, and have a sparse black coloration.This species is similar to P. obducta in colour andmarkings on head, thorax, and abdomen, but differs inshape and structure of male genitalia and opercula.

Description of male Head. – Ochraceous, markings distinct. Black

markings surrounding the eyes interrupted at level ofthe vertex lobes. Transverse fascia between eyes andpostclypeus interupted or narrowed in the middle.Rostrum short, reaching halfway abdominal segment2. Black triangular markings around lateral ocelli as inP. carmente and P. obducta, connected with anteriormargin of pronotum and enclosing a large ochraceousspot. Y-shaped figures sometimes very broad.

Thorax. Pronotum. – As in P. obducta, but lateraltooth on pronotal collar blunt rather than pointed,and crescent-shaped posterior ends of central fasciaetouching each other.

Mesonotum. – Markings narrow, as in P. obducta.Median fascia continuing on cruciform elevation. Lat-eral fascia either continuous from anterior to nearlyposterior margin of mesonotum or consisting of a fas-cia, reaching from anterior margin to two thirds of me-sonotum length, and a black spot at posterior margin.

Tegmina and wings. – Pale hyaline, tegmina verylightly bronzed toward apices; basal veins of 2nd and3rd apical areas of tegmina weakly infuscated. Vena-tion ochraceous in basal half, brownish to black inapical part.

Legs. – As in P. obducta.Operculum (fig. 13). – Operculum short and tri-

angular, 1.8 times as long as broad (n = 4), reachingto two thirds of abdominal segment 3. Apex sharplypointed, lateral to medial of midline of operculum.Lateral margin straight, medial margin almoststraight and with fairly narrow black band. Basal partof operculum ochraceous. Distance between operculaat apices 2.7 times the distance at point of closest ap-

proximation (n = 4). Timbal cavities clearly visiblefrom lateroventral view.

Abdomen. – Broad, as long as head and thorax to-gether (n = 4). Ground colour of dorsal side ochra-ceous, ventral side light ochraceous. Tubercles casta-neous, tubercles on abdominal segment 4 somewhatlighter than those on segment 3. Timbal covers lighterthan abdomen, triangular markings small and distinct.

Genitalia (fig. 16). – Shape of pygofer broadly oval,slightly broadened in the middle. Basal pygofer lobesfairly long and narrow with darkened rounded apexreaching to three fourths of pygofer length; pygoferlobes connected to lateroventral part of pygofer onlyat base. Median uncus lobe brown to black, broad atits base, rapidly narrowing into a gently inward curv-ing long, tongue-shaped lobe with central fissure andbicuspidate apex. A pair of slender, strongly devel-oped black claspers is attached to basal part of uncus.Apices of claspers point downwards rather than out-wards as in P. hermes. Lateral process of the pygoferrounded, short, not reaching beyond anal valve, anddarkening towards apex.

Measurements in mm (4?). – Body length ?:20.2-21.8 (21.2 ± 0.6); tegmen length ?: 26.8-28.3(27.3 ± 0.6); head width ?: 6.5-6.9 (6.7 ± 0.2);pronotum width ?: 7.1-7.2 (7.1 ± 0.2).

RemarksOne of the paratypes from Sandakan differs from

the other three type specimens in having weakly devel-oped black markings, especially around ocelli and onpostclypeus, and in having no markings on the timbalcovers at all. However, since this specimen is similar tothe other specimens in genitalia, operculum and othercharacters, we have no doubt about its identity.

Distribution (fig. 18).This species is only found in Borneo.

Etymology Purana infuscata is the only species within the car-

mente group with weakly developed infuscations atbasal veins of 2nd and 3rd apical areas of tegmina.

Purana sagittata sp. n.(figs. 14, 17, 18)

Type material. – Holotype ?: ‘MALAYSIA: Selangor /Kuala Lumpur, Ulu Gombak, / Gombak Field Station/ 25.5.1996, 250 m / T. Trilar, M. & M. Gogala leg’(PMSL). – Paratypes 14? 5/: MALAYSIA, PENINSULAR

MALAYSIA: same data as holotype but: 20.v.1996, 1?(PMSL); Johore, 2nd mile Kota Tinggi, 1.iv.1948, J. A.Le Doux, 1? (MNM); Pahang, Taman Negara NP,Kuala Juram E. of Merapoh 4°39 N 102°08’ E,15.iii.1999, J. P. & M. J. Duffels, M. Zaidi & M. Y.

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Ruslan, 1? (ZMAN); Johor, Endau Rompin N.P.,NERC, 02°31’44’’N, 103°24’02’’E, disturbed forest,15.viii.1999, J. P. Duffels, M. A. Schouten & M. Y.Ruslan, 1/ (ZMAN), same locality, 17.viii.1999, M. Z.Hazman, 1? (ZMAN), same locality, 6.xi.1999, M. A.Schouten, 2? (ZMAN) 1/ (UKM), same locality,14.x.1999, M. A. Schouten & F. Cheong, 1? (UKM);Johor, Endau Rompin N.P., NERC, 02°31’44’’N,103°24’02’’E, disturbed forest, 7.xi.1999, M. A.Schouten, 1? (UKM), same data but 8.xi.1999, 1?(UKM), 9.xi.1999, 1?, 1/ (ZMAN); Johor, EndauRompin N.P., 1st river crossing Sungai Jasin, primarylowland rainforest, 12.xi.1999, M. I. Zaidi, M. Y.Ruslan, M. A. Schouten, F. Cheong, 1? (ZMAN); Jo-hor, Endau Rompin N.P., Bridge 9, 12.xi.1999, M.A. Schouten, 1? (BMNH), same data but: 11.xi.1999,1/ (ZMAN). MALAYSIA, SABAH: Sabah, 25-30.viii.1991,L. Danum, 1? (UKM). INDONESIA, SUMATRA: Suma-tra’s O. K. [= Oost Kust (East Coast)], Lau Rakit, 300m, 29.viii.1921, J. B. Corporaal, 1? 1/ (ZMAN).

P. sagittata can be easily separated from the otherspecies of the P. carmente group by its extremely slen-der and pointed median uncus lobe. In the otherspecies of the carmente group the uncus lobes arebroader and have a more or less bicuspidate apex.

Dr Tomi Trilar and Dr Matija Gogala of the Priro-doslovni Muzej Slovenije, Ljubljana describe the songof P. sagittata in this issue (Trilar & Gogala 2002).

DescriptionHead. – Tawny to ochraceous, fairly dark. Pattern

of markings on head quite similar to P. obducta, but

Y-shaped figures fairly broad. Apices of triangularmarkings surrounding ocelli connected to anteriormargin of pronotum. Black-tipped rostrum reachingposterior margin of 2nd abdominal segment.

Thorax. Pronotum. – Tawny to ochraceous, fairlydark. Central fasciae posteriorly ending in two cres-cent-shaped, juxtaposed markings which are fused infront of pronotal collar. Pronotal collar of fresh spec-imens with olivaceous spot at lateroproximal corner.

Mesonotum. – Markings as in P. obducta but para-median fasciae slightly oblique to almost straight,hardly dilated at anterior margin of mesonotum.

Tegmina and wings. – As in P. obducta, but morebronzed.

Legs. – As in P. obducta.Male operculum (fig. 14). – Slender, 1.6-2.0 times

as long as broad (n=11), not reaching beyond twothirds of the 3rd abdominal segment. Apex angularlyrounded, situated medial or lateral of midline of theoperculum. Lateral margin almost straight. Medialmargin slightly convex towards apex, and with distinctblack band from base to apex. Medial half of basal partof operculum black. Opercula of fresh specimens withgreen tinge. Distance between opercula at apices 2.3-2.6 times the distance at point of closest approximation(n=11).

Male abdomen. – As in P. obducta, but triangularmarking on timbal cover fairly small, not reaching tohalfway the length of timbal cover. Tergites coveredwith a dense silvery to golden pubescence, which isparticularly dense on segments 7 and 8, ventral side ofthe abdomen with scattered long setae.

Male genitalia (fig. 17). – Pygofer rather small and

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Fig. 18. Distribution of Purana carmente (dots), P. hermes (squares), P. sagittata (triangles), and P. infuscata (diamonds).

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broadly oval to round, outer margin strongly convex.Basal pygofer lobes black-tipped, short, reachinghalfway the pygofer, and almost entirely separatedfrom lateroventral part of pygofer. Basal pygofer lobesbroader and less far apart than in other species of theP. carmente group. Median uncus lobe very long,strongly curved inward, and abruptly broadened dis-tally to an arrowhead-shaped apex. Uncus dorsallyconnected to basal part of claspers. Claspers broader,and more flattened than in P. infuscata and hermes.Lateral processes of pygofer rounded, darkened to-ward apex, and as long as anal valves.

Female operculum. – As in P. obducta, but some-what longer and with less distinct black markings.Posterior margin straight to slightly convex, laterodis-tal corner flattened to rounded, lateral margin con-vex. Operculum in fresh specimens covered withpowdery white wax.

Female abdomen. – As in P. obducta.Measurements in mm (11?, 2/). – Body length ?:

21.4-23.6 (22.5±0.6), /: 18.8-20.1 (19.4); tegmenlength ?: 27.2-31.2 (29.6±1.4), /: 27.6-28.0 (27.8);head width ?: 6.8-7.0 (6.9), /: 6.5-6.8 (6.7±0.1);pronotum width?:7.8-8.0 (7.9±0.2),/:7.2-7.5 (7.3).

Distribution (fig. 18)The species is known from Peninsular Malaysia,

Borneo (Sabah) and Sumatra.

EtymologyThe species is named after the typically shaped apex

of the male uncus lobe that resembles an arrowhead.

Purana dimidia Chou & Lei (figs. 11, 19, 20)

Purana dimidia Chou & Lei in Chou et al., 1997: 230-231,368-369, pl. X. fig.111.

This species is known from the type specimens,one male from Tonkin and one male from Hekouxi-aonanxi, Yunnan province, which are deposited inthe Institute of Zoology, Academia Sinica in Beijing.

We have not seen any specimen of this species. Thedescription presented below is a copy of the shortcharacterisation of the species as given in Chou et al.(1997) in the English summary. Fig. 19 and 20 arecopies of figs. 9-74: F & K in this book.

‘Body length (?) 20.8-21.0 mm, tegmen length25.2-26.1 mm, width of head 5.9-6.1 mm, width ofbase of mesonotum 5.8-6.2 mm.

This new species is similar to P. carmente, but differsfrom the latter in the following aspects: 1) operculareaching the middle of the third abdomen sternite, lat-ter species reaching the middle of the fourth abdomensternite, 2) there is not spot on the tegmina of P.dimidia; 3) lateral margin of pronotum different inshape.

Holotype ?, Tonkin, 1939-VII, coll. A. de Coo-man; Paratype, 1?, Hekouxiaonanxi, Yunnan Pro-vince, 200m, 1956-VI-8, Huang Keren. (Type speci-mens are deposited in the Institute of Zoology,Academia Sinica).’

RemarksP. carmente has no spots on the tegmina, just like P.

dimidia.

ACKNOWLEDGEMENTS

We are very much indebted to the following cura-tors for the loan of material under their care: Mr K.Arakaki (BPBM), Dr P. H. Arnaud (CAS), Dr M.Gogala and Dr T. Trilar (PMSL), Mr A. H. Kirk-Spriggs (NMCW), Dr Maryati Mohamed (UMS), Mr J.van Tol (RMNH), Mr M. D. Webb (BMNH), Dr ZaidiMohd. Isa and Mr Ruslan Mohd. Yusop (UKM), andDr H. J. Zainal Abidin (MNM). Grants from Nuffic(Jan Tinbergen Scholarship CN 5731) and the Ams-terdamse Universiteits Vereniging (AUV), which en-abled the first author to visit Malaysia from August toNovember 1999, are gratefully acknowledged. A visitof the second author to Peninsular Malaysia in 1999was partially financed by the Netherlands Foundationfor the Advancement of Tropical Research (WR 84-

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Figs. 19-20. Purana dimidia. – 19, male abdomen in ventralview; 20, male pygofer in ventral view. Reproduced fromChou et al., 1997, figs. 9-74, F & K.

19

20

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482). Our special thanks go to Dr Zaidi Mohd. Isaand Mr Ruslan Mohd. Yusop who enabled us to studythe collection of the Centre for Insect Systematics(UKM), established the contact with the Malaysian Na-ture Society, and arranged material and transport todo fieldwork in Endau Rompin National Park. Field-assistance was given by the Endau Rompin team of theMalaysian Nature Society: Francis Cheong, ChewKeng Lin, Sing Yun Chin, Lili Bte Tokiman, HazmanBin Md. Zaki, Ivy S. Abdullah, and Heah Hock Heng.They are thanked for their help in the field and logis-tic support. We thank Dr Matija Gogala and Dr TomiTrilar for their cooperation during the fieldwork. Wewould like to thank Dick Langerak for preparing thefigures, and Gerard Verlaan for technical assistance.We are very grateful to Dr Arnold de Boer for criticalreading of an earlier draft of this paper. The advisesgiven by Mr Paul Beuk and Dr Arnold de Boer werehighly appreciated.

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Received: 19 February 2001Accepted: 24 September 2001

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