Corresponding author: Joannes Petrus Duffels (
Academic editor: Mike Webb
The type species of
The genus
In his classic work “Cicadas of Malaysia”,
In the last two decades, cicada inventories in the Malaysian and Indonesian parts of Borneo and in Peninsular Malaysia have been strongly intensified. Zaidi and co-workers of Universiti Kebangsaan Malaysia recorded
This study aims to contribute to a better taxonomic knowledge of the cicadas of Borneo anticipating a larger publication on the Bornean cicada fauna. The descriptions of the three new species of
The institutions listed below are the depositories of the material studied. The abbreviations have been used in the lists of material and throughout the text.
BMNH Natural History Museum, London (former British Museum (Natural History))
NHRS Naturhistoriska Riksmuseet, Stockholm
NMWC National Museum of Wales, Cardiff
RMNH Nationaal Natuurhistorisch Museum (former Rijksmuseum voor Natuurlijke Historie), Leiden
ROM Royal Ontario Museum, Toronto
ZMAN Zoölogisch Museum, Universiteit van Amsterdam, Amsterdam
Data on the distribution of the species were derived from the author’s “Biodiversity Database of the Cicadas of South East Asia and the West Pacific”, and plotted on maps of ADC-Worldmap version 2.0 vol. 4 Southern Asia & Australia with the program MapInfo for Power Mac, version 4.03. The localities and other data from the specimen labels in the database are filed in the program File-Maker Pro 4.0. The information about geographical co-ordinates has been retrieved from the following sources: “Atlas van Tropisch Nederland” (
The terminologyadopted in this paper for external features of the body and the male genitalia follows that of Duffels (
The genus
Small cicadas: body length male: 8.1–11.7 mm, female: 9.0–12.4 mm. Head slightly wider than pronotum collar. Vertex black, or reddish brown with a pair of paramedian, squarish, black markings or a pair of spots next to the paired ocelli. Postclypeus protruding weakly to fairly strongly with glabrous nose. Rostrum reaching beyond posterior margin of middle coxae or to anterior margin of hind coxae or beyond. Pronotum slightly wider than mesonotum. Mesonotum with a pair of paramedian, juxtaposed, black, obconical spots and a pair of black to black-brown lateral sigillae. Fore femora with four spines along lower ridge. Tegmina and wings hyaline, wings with four apical areas. Male operculum sickle-shaped and with adjacent setae, margins and apical third with long setae. Timbal with 6–7 ribs and faint intercalary ribs. Pygofer with about equitriangular dorsal beak with long and narrow apex; basal pygofer lobes very long reaching either to about apex of anal segment; upper pygofer lobes mostly distinctly separated from, but sometimes fused with, basal pygofer lobes. Uncus absent. Claspers juxtaposed and different in shape. Theca either with one strong, chitinized appendage, apically divided in two stems, or a pair of more or less similar appendages. Aedeagal basal plates in ventral view triangular to oval. Ventral rib of aedeagal basal plate rod-like and suspended with attachments only at ends.
.
1. | Anterior and ventral parts of postclypeus yellowish to reddish brown. Philippines |
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– | Anterior and ventral parts of postclypeus black with either a yellowish to reddish brown glabrous nose or an oblong area of the same colour reaching from nose toward clypeal suture. Borneo | 2 |
2. | Head width: 3.8–4.0 mm. Abdominal tergite 2 with a black transverse fascia along anterior margin, a reddish brown fascia at about half-length and a yellow-brown fascia along posterior margin. Borneo |
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– | Head width: 2.8–3.3 mm. Abdominal tergite 2 black with a pair of paramedian yellow-brown spots at posterior margin. Borneo | 3 |
3. | Male genitalia as in |
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– | Male genitalia as in |
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The identity of
Dr Gunvi Lindberg, curator of the Swedish Museum of Natural History, Stockholm, kindly sent me the type specimens of
The lectotype is very fragile and partly damaged: the apical half of the wings is missing and the right tegmen is somewhat glued together; sternite 8 is partly damaged; the abdomen is glued to the head and thorax. The pygofer is taken out for the greater part. It is most likely that this damage to the lectotype was caused by an earlier student attempting to pull out the pygofer. The most characteristic feature of the genus
The two paralectotypes are in good condition. One paralectotype bears the following labels: “Ins. / Philipp”, “Semper”, “Tibicen / tener Stål”, “Allotypus” [printed in black cadre; red paper], “NHRS-HEMI 000000010”. The other paralectotype bears the labels: “Ins. / Philipp”, “Semper”, “Tibicen / tener Stål / ♀”, “Paratypus” [printed in black cadre; red paper], “NHRS-HEMI 000000011”
Ground colour reddish brown.
Head. Vertex reddish brown with a pair of comma-shaped dark brown markings next to paired ocelli, a faint brownish ring around each of the ocelli, and dark brown colouration along mediodistal margin of eye. Postclypeus protruding weakly, yellowish to light reddish brown, dorsally with a pair of lateral dark brown spots, anterior and ventral parts of postclypeus with two paramedian series of 7 dark reddish brown transverse grooves. Anteclypeus light reddish brown without marking. Rostrum yellowish brown with brown apex reaching beyond posterior margin of middle coxae. Lorum black but anterior third reddish brown. Gena light reddish brown with a black line along posterior eye margin. Antenna, supra-antennal plate and vertex lobe yellowish brown to reddish brown.
Thorax. Pronotum with a pair of large, rounded rectangular, dark reddish brown to black brown markings that are enclosed by the reddish brown anterior margin of pronotum, the light reddish brown pronotum collar, and a broad median, light reddish brown fascia that strongly widens to anterior margin of pronotum and to pronotum collar. A lanceolate black-brown marking connects the broadly black-brown anterior and posterior oblique fissures.
Mesonotum with a pair of paramedian, black-brown, obconical spots, those are fused at anterior margin of mesonotum and reach to one fourth of mesonotum disk. Scutal depressions in front of cruciform elevation with light brownish suffusion. Lateral sigillae clouded with dark brown and anteriorly slightly wider than anterior part of paramedian obconical spots, gradually narrow to their distal ends near anterior angles of cruciform elevation. Cruciform elevation yellowish.
Legs. Yellow-brown to brownish. Fore femora with four yellow-brown spines with brown apices: a long spine at proximal end of lower ridge of femur, a second spine, half as long as proximal spine, at half-length of lower ridge, a third spine, one third as long as proximal spine, at three fifths of lower ridge, and a tiny spine near distal end of lower ridge.
Tegmina and wings. Hyaline. Venation of tegmina and wings brownish to reddish brown variegated with dark brown.
Operculum. Sickle-shaped with narrowly rounded apex reaching to almost anterior margin of abdominal segment 2, strongly narrowed from base to one fourth of its length, and gradually narrowing from one fourth of length to apex. Apical half with sparse long setae, especially along operculum margins.
Abdomen. Timbal with 7 evenly spaced long ribs and very faint intercalary ribs. Tergite 1 dark reddish brown, tergites 2–7 with a laterally widening, dark reddish brown fascia along anterior margin, medially reaching to half-length or two thirds of tergite, and a laterally narrowing reddish brown fascia; tergite 3 also with a narrow yellowish fascia along posterior margin. Tergite 8 with a laterally narrowing dark reddish brown fascia along anterior segment margin and a laterally narrowing reddish brown fascia, both two fifths as long as tergite and a yellowish fascia along posterior margin, one fifth as high as the tergite. Sternite 2 yellowish, sternites 3 to 6 and anterior half of sternite 7 reddish brown, posterior half of sternite 7 and whole sternite 8 yellowish.
Genitalia (
There are two female paralectotypes, one fully coloured and one with obsolete marking. A description of the fully coloured female follows here:
Head. As in male lectotype but anteclypeus black-brown with light brownish anterior margin and keel, and vertex with additional dark brown marking between paired ocelli and eyes.
Thorax. Pronotum as in male lectotype. Mesonotum as in male lectotype but paramedian obconical spots reaching to one third of mesonotum disk and lateral sigillae black-brown.
Legs, tegmina and wings. As in male lectotype.
Operculum. Basal half broad, narrowed at half its length to two thirds of basal width; apical part curved mediad with narrowly rounded apex reaching to just beyond anterior margin of sternite 2.
Abdomen. Tergite 1 light reddish brown. Tergite 2 reddish brown with laterally narrowing, black fascia along anterior margin medially reaching to two fifths of segment length. Tergites 3–7 with a laterally widening, black fascia along anterior margin, medially reaching to half or three fifths of segment length, a slightly narrower reddish brown fascia at about half-length of tergite and a narrow yellowish green fascia along posterior margin. Tergite 8 with laterally narrowing, black fascia along anterior margin medially reaching to one fourth of segment length, a broad reddish brown fascia and a fairly narrow, yellowish green fascia along posterior margin. Sternite 2 with dark brown transverse marking, sternites 3 to 6 with dark brown transverse band, which is a little less than half as wide as sternite and reaches from anterior sternite margin to two thirds or three fourths of sternite length. Sternite 7 medially dark brown. Segment 9 dorsally with a pair of oblong, paramedian, black-brown markings reaching from anterior margin of segment to three fourths of its length, and laterally with a pair of, round, black-brown spots.
The female paralectotype with the more obsolete marking has no marking on anteclypeus, no additional brown marking on vertex, very light brown lateral sigillae on mesonotum, light reddish brown abdominal tergites with much narrower black marking along their anterior margins, and only small brown median spots on sternites 5 and 6.
Measurements (in mm; 1♂, 2♀). –Body length ♂: 10.2 ♀: 11.5; tegmen length ♂: 10.9, ♀: 12.9–13.5; head width ♂: 3.2, ♀: 3.7–3.8; pronotum width ♂: 3.1, ♀: 3.6–3.7.
Distribution. The type specimens of
24♂ 23♀. Holotype ♂: Malaysia: Sabah: “RMNH Leiden E SABAH / Lahad Datu, 60 km W of: / Danum Valley Field Centre / at junction Sg Segama and / Sg Palum Tambun, 150 m / 4°58"N 117°48'E”, “At light. Bridge of Segama. / 19 Mar 1987, 18.30–21.30. / Clearing, edge of untouched / evergr. lowl. rainforest / leg. Van Tol & Huisman” (RMNH). Paratypes: Malaysia: Sabah: same data as holotype, 3♀ (RMNH), same data as holotype but: 17 & 18.iii.1987, 18.30–21.00, 1♂ 2♀ (RMNH), 20.iii.1987, 18.20–21.00, 1♂ 2♀ (RMNH), clearing nr E trail, 21.ii.1987, 18.30–20.30, 1♀ (RMNH); 60 km W Lahad Datu, DVFC, nr Segama bridge, 4°58'N 117°43'E, 20.x.1987, 150 m, J. Huisman & R. de Jong, 1♂ (RMNH); 60 km W of Lahad Datu, road Kg Silam – DVFC, km 68.5, 4°58'N 117°48'E, 150 m, 24.iii.1987, ML-light, J. Huisman, 1♂ (RMNH); Danum Valley, 5°01'N 117°47'E, 10.ix.1987, 100 m, A.H. Kirk-Spriggs, nmw Sabah (Borneo) Expedition, nmw. z 1987, 094, light trap sample roadside, secondary forest, 2♂ (NMWC), same data but: 11.ix.1987, 1♂ 2♀ (nmwc), 14.x.1987, 1♀ (NMWC); Danum Valley, 5°01'N 117°47'E, 30.ix.1987, 200 m, A.H. Kirk-Spriggs, nmw Sabah (Borneo) Expedition, nmw., z 1987, 094, lowland mixed dipterocarp forest, Grid EZ, light trap sample, understory forest, 1♂ 2♀ (nmwc); Danum Valley, 70 km W Lahad Datu, Field Centre, Main Trail West 0 North 5, 150 m, 15.xii.1989, sample Sab. 69, secondary vegetation/canopy/primary forest margin, at light, M.J. & J.P. Duffels, 1♂ (ZMAN); Bettotan, nr. Sandakan, 17.viii.1927, C.B.K. & H.M.P. F.M.S. Museums, ex. F.M.S. Museum, B.M. 1955–354, 1♂ (BMNH), same data but 24.vii.1927, 1♂ (BMNH); Sungai Darling, 60 m W Sandakan, 26.xi.1989, sample Sab. 43, secondary forest understorey, at light, M.J. & J.P. Duffels, 1♀ (ZMAN). Sarawak: Foot of Mt. Dullit, junction of rivers Tinjar & Lejok, 25.viii.1932, Light trap, Oxford Univ. Exp. B.M. Hobby & A.W. Moore B.M. 1933–254, 2♂ 3♀ (BMNH), same data but 28.viii.1932, 1♀ (BMNH), 31.viii.1932, 1♂ 1♀ (BMNH), 6.x.1932, 1♂ (BMNH); Gunung Mulu Nat. Park, Site 7, Long Pala (Base), 324450, 50 m, Alluvial/secondary forest, Acl-understorey, J.D. Holloway, RGS Mulu exped., B.M. 1978–206, 4♂ 4♀ (BMNH). Brunei: Temburong District, ridge NE of Kuala Belalong, approx. 300 m, x.1992, J.H. Martin, 125 W mv light, B.M. 1992–172, 3♂ (BMNH), same data but: xi.1992, 1♂ (BMNH).
This species name refers to its distribution in Borneo.
Ground colour yellowish to greenish brown, and reddish brown. Marking black or black variegated with reddish brown. Dorsal side of body silvery pilose, ventral side with longer silvery setae.
Head (
Thorax. Pronotum (
Mesonotum (
Legs. Yellow-brown, fore and middle tarsi and distal part of fore tibia often darker brown. Fore femora with four yellow-brown spines with dark brown apical parts: a long, erect spine at proximal end of lower ridge of femur, a second spine, two-thirds as long as proximal spine, at three-fifths of lower ridge, a third, slightly shorter, spine, at four-fifths of lower ridge and a very short, triangular spine near distal end of lower ridge.
Tegmina and wings. Hyaline. Venation of tegmina and wings yellowish to reddish brown.
Operculum (
Abdomen (
Genitalia (
Head as in male, but postclypeus reddish brown from nose to anterior margin of pronotum; vertex reddish brown with exception of the black lateroproximal parts that more or less enclose the paired ocelli; a black spot is attached to proximal side of median ocellus.
Thorax. Pronotum. Rectangular markings as described for males are not black but reddish brown with broad black marking in the oblique fissures and narrow black marking in lateral part of ambient fissure. Mesonotum as in males.
Legs, tegmina and wings as in males.
Operculum. Basal half broad, narrowed at half its length to two thirds of basal width; apical part curved mediad, medial margin weakly concave, and lateral margin convex to narrowly rounded apex, reaching to anterior margin or one third of sternite 2.
Abdomen. Tergites 2–7 with a laterally widening, black fascia along anterior margin medially reaching to one third or two thirds of segment length, a laterally narrowing, reddish brown fascia at about half-length of tergite and a laterally narrowing yellowish, green fascia along posterior margin. Tergite 8 with narrow black fascia and broader reddish brown and yellowish fasciae. Sternite 2 medially black-brown, sternites 3 to 6 with brown-black transverse band, which is one third to half as wide as sternite and reaches from anterior sternite margin to two-thirds of sternite or at most almost to its posterior margin. Segment 9 with a pair of oblong, paramedian, brown to black-brown markings and a pair of round, lateral, brown spots and brownish colouration along basal two-thirds of lower margin.
(in mm; 6♂, 6♀). Body length ♂: 8.1–9.2 ♀: 9.0–10.3; tegmen length ♂: 8.7–10.2, ♀: 10.0–11.8; head width ♂: 2.8–3.2, ♀: 2.9–3.7; pronotum width ♂: 2.5–2.9, ♀: 2.8–3.4.
(
Distribution of
2♂. Holotype ♂: Malaysia: Sabah: “RMNH Leiden E SABAH / Lahad Datu, 60 km W of: / Danum Valley Field Centre / at junction Sg Segama and / Sg Palum Tambun, 150 m / 4°58"N 117°48'E”, “At light. Bridge of Segama. / 26 Mar 1987, 18.30–21.30. / Clearing, edge of untouched / evergr. lowl. rainforest / leg. Van Tol & Huisman” (RMNH). Paratype: Indonesia: Kalimantan Timur: Long Tua, edge of Bahau River, 3°10'N 115°47'E, 440 m, 5–9.iv.1994, UV light, B. Hubley & D.C. Darling, IIS 940526, 1♂ (ROM).
This species is dedicated to my colleague and friend Dr Jan Tol, odonatologist of the Leiden museum, for his significant contribution to our knowledge of the cicadas of Borneo.
Ground colour yellowish to reddish brown. Marking black or black variegated with reddish brown. Ventral side of body with short silvery setae.
Head. (
Thorax. Pronotum (
Mesonotum (
Legs. Yellow-brown to greenish. Fore femora with four reddish brown spines: a long spine at proximal end of lower ridge, a second spine, two-thirds as long as proximal spine, at two thirds of lower ridge, a third spine, half as long as proximal spine at four fifths and a very short, triangular spine near distal end of lower ridge.
Tegmina and wings. Hyaline. Costa and basal half of radius + subcosta of tegmen reddish brown, radius anterior and distal half of subcosta reddish to dark brown. Remaining venation of tegmina and wings light to dark brown.
Operculum. (
Abdomen. (
Genitalia. (
(in mm; 2♂). Body length 10.4–10.7; tegmen length 10.8–12.0; head width 3.2–3.3; pronotum width 2.9–3.1.
The paratype from Kalimantan Timur is alike to the holotype in the male genitalia, but the marking on the body is more extended and black instead of black-brown. The head is black with exception of the medial triangle at posterior margin of head, the nose of the postclypeus and the supra-antennal plates, which are reddish brown. The pronotum has a pair of black squarish markings, the mesonotum is entirely black, and the abdomen has a similar marking as the holotype, but the marking is black instead of brownish black.
(
Distribution of
Holotype ♂: Malaysia, Borneo: Sarawak: “SARAWAK / Gunong Mulu / Nat. Park”, “Site 23, April / W. Melinau Gorge / 250 m. 430558”, “J.D. Holloway / RGS Mulu exped. / B.M. 1978–206”, “FEG 4. Limestone / forest. MV - / canopy/understorey”(BMNH). Paratypes: Malaysia, Borneo, Sarawak: same data as holotype, 4♂ 3♀ (BMNH).
This species name refers to its occurrence in Gunung Mulu National Park, Sarawak, Borneo.
Ground colour and pilosity as in
Head (
Thorax. Pronotum (
Mesonotum (
Legs. Yellow-brown, fore tarsi and fore tibia darker brown; inner side of fore femur with dark brown marking. Fore femora with four, fairly stout, black-brown spines with light brown apices: a long and strong spine at proximal end of lower ridge of femur, a distinctly shorter spine at half-length of lower ridge, a somewhat shorter spine at four fifths of lower ridge and a very short, triangular spine near distal end of lower ridge.
Tegmina and wings. Hyaline. Venation of tegmina and wings greenish to reddish brown.
Operculum (
Abdomen (
Genitalia (
Head. Vertex reddish brown with a black spot laterally of paired ocelli only or with black marking restricted to lateroproximal part of vertex. Postclypeus reddish brown to black, but medially yellowish from nose to frontoclypeal suture or from frontoclypeal suture to (almost) clypeal suture; lateral parts of postclypeus with 7 pairs of brown to black transverse ridges. Anteclypeus and rostrum as in male. Antenna, supra-antennal plate, vertex lobe and gena yellowish to reddish brown. Lorum reddish brown but posterior two thirds of lorum more or less black in two paratypes.
Thorax. Pronotum. As in male, but the rectangular, reddish brown markings do not enclose any black marking.
Mesonotum with a pair of paramedian, juxtaposed, black to brown, obconical spots at anterior margin, reaching to two fifths of mesonotum disk. Scutal depressions in front of cruciform elevation with round, brown spots. Lateral sigillae black to very faint and reddish brown without black marking. Cruciform elevation yellowish.
Legs, tegmina and wings as in male
Operculum. Basal half broad, operculum narrowed at half its length to two thirds of basal width, medial margin concave, lateral margin of apical part basally parallel to medial margin, laterodistal angle obtusely rounded, apical margin straight, reaching to anterior margin or one third of sternite 2 and making an angle of 60 degrees with medial margin.
Abdomen. One paratype has the following marking: tergite 2 with black-brown fascia, about as broad as cruciform elevation, along anterior tergite margin, tergites 3–7 with transverse, black fasciae along anterior margins, that on tergite 3 reaches medially to two fifths of tergite length, that on tergite 4 to one third and those on tergites 5–8 to one fourth or one fifth of tergite length, the fasciae on tergites 3–8 widen laterally to reach to two thirds of tergite length. Sternite 2 medially black-brown, sternites 3 to 6 with brown-black transverse band, which is two thirds as broad as sternite and reaches from anterior sternite margin to two-thirds of sternite, sternite 7 medially black-brown. Segment 9 dorsally with a pair of oblong, paramedian, dark brown markings, laterally with a pair of round, lateral, brown spots and ventrally with brownish colouration along basal two-thirds of lower margin. The other two paratypes differ in the following features: dorsal marking with much narrower and partly missing fasciae; sternite 2 medially brownish, sternite 3 with black-brown transverse band, half as wide as sternite, sternites 4–6 with dark brownish median marking, one third to half as wide as sternite. Segment 9 similar but missing the lateral spots.
(in mm; 5♂, 2♀). Body length ♂: 9.9–11.7, ♀: 12.0–12.4; tegmen length ♂: 12.7–12.9, ♀: 14.2–15.0; head width ♂: 3.8–4.0, ♀: 4.0–4.3; pronotum width ♂: 3.6–3.8, ♀: 3.8–4.1.
(
Distribution of
I am very much indebted to the following curators of collections for the loan of material: Mr B. Hubley & Dr D.C. Darling (ROM), Mr A.H. Kirk-Spriggs and Dr M.R. Wilson (NMWC), Dr. G. Lindberg (NHRS), Dr J. van Tol (rmnh) and Mr M. D. Webb (bmnh).I thank my late wife Greet Duffels van Egmond for her help in the field. Special thanks go to Dick Langerak (ZMAN) for preparing the figures, to Rob Portegies (ZMAN) for making the distribution maps, and to Gerard Verlaan (ZMAN) for technical assistance.