The Cicada Genus Muda Distant (Hemiptera: Cicadidae) from Sundaland: Species and Relationships J.P
Tijdschrift voor Entomologie 161 (2018) 131–154
The cicada genus Muda Distant (Hemiptera: Cicadidae) from Sundaland: species and relationships J.P. Duffels
A diagnosis is provided of the cicada genus Muda Distant, 1897, with descriptions, illustrations and distribution maps for the five species found in Sundaland. Three of these, Muda obtusa (Walker, 1858), M. virguncula (Walker, 1856) and M. tua Duffels, 2004, are redescribed. Muda beccarii (Distant, 1888) and M. concolor Distant, 1897 are junior synonyms of M. virguncula. Abroma tahanensis (Moulton, 1923) is transferred to Muda and redescribed. Muda kinabaluana is described as new to science, and is very peculiar, missing the timbals. An identification key is provided. Characters and taxonomic position of Muda kuroiwae (Matsumura, 1913) from the Ryukyu Islands of southern Japan are discussed. Muda is characterized by two supposed synapomorphies, viz., the mediodorsal carina of the male pygofer and the movable upper pygofer lobes; these characters are also found in species of Katoa. Further comparative study of Muda and Katoa is needed to reveal their relationships. J.P. Duffels, Naturalis Biodiversity Center, PO Box 9517, 2300 RA Leiden, The Netherlands. [email protected]
Introduction Muda kinabaluana from the Mount Kinabalu massif This paper presents a revision of the species of the of Borneo, in which the sound organs are lost. genus Muda Distant, 1897 occurring in Sundaland: The taxonomic position and characters of Muda Java, Malay Peninsula, Sumatra and Borneo, and kuroiwae (Matsumura, 1913) from the Ryukyu Is- nearby smaller islands. The taxonomic position of lands of southern Japan are discussed separately in a the genus is yet to be established, but the (in cross- remark following the key to the Sundaland species; section) triangular postclypeus is a supposed synapo- the species was transferred from Baeturia to Muda by morphy for a group consisting of the genus Muda, de Boer (1995a). the tribe Chlorocystini and the tribe Prasiini. The The present study provides the characters to genus Muda is either sister to the latter two tribes separate two Muda species with a uniform yellowish or sister to the tribe Prasiini only (De Boer 1995a; brown to light brown body that have been confused Moulds 2005). for about 150 years, viz., M. virguncula and M. ob According to the literature (Moulton 1923; tusa. Muda virguncula is widespread in Sundaland Schouten et al. 2004), three species of Muda are (see Fig. 25 below) and may also occur in the south- found in Sundaland: M. obtusa (Walker, 1858), eastern mainland of South East Asia. Muda obtusa is M. virguncula (Walker, 1856), and M. tua Duffels, presumably endemic to Java. 2004. A recent study of the Sundaland cicadas (Duf- fels 2011) of the genus Abroma Stål, 1866 suggested History of the genus that Abroma tahanensis Moulton, 1923, from the The new genus Muda was proposed by Distant Malay Peninsula, should be transferred to Muda. The (1897) for a new species, Muda concolor, from the present paper confirms this nomenclatorial change Mentawai Islands, west of Sumatra. In his synonym- and also presents the description of the new species ic catalogue of 1906, Distant attributed two species
Tijdschrift voor Entomologie 161: 131–154, Figs 1–26. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 16 September 2019. DOI 10.1163/22119434-20192077 Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Fig. 1. Muda virguncula, male in dorsal view, Brunei, Lamunin. Photo J. van Arkel.
The other Muda species have a fold or an articula- with long spines and smaller spines, described in tion that at least suggests the movability of the upper this paper for M. obtusa and M. kinabaluana, is very pygofer lobe. similar to that of K. taibanensis and K. neokanagana, The two supposed synapomorphies for Muda while the long parallel-sided pygofers of M. obtusa mentioned above are also found in species of Katoa. and M. virguncula are not only found in these spe- The specimens of K. chlorotica examined also have cies but also in two species of Katoa, viz., K. chlo the male pygofer with a mediodorsal carina, speci- rotica (Chou & Lu, 1997 in Chou et al. 1997: 94, mens of other Katoa species have not been studied. Figs 8–29) and K. paura (Chou & Lu, 1997 in Chou The distinct articulations at the bases of the upper et al. 1997: 96, Figs 8–30). pygofer lobes in Katoa taibanensis (Lei & Chou, The species of Muda from Sundaland. Five species of 1995: 100, Fig. 1), K. paucispina (Lei & Chou, Muda recorded from Sundaland are described and 1995: 101, Fig. 2) and K. neokanagana (Chou & Lu, keyed below. Two species, M. virguncula and M. ob 1997 in Chou et al. 1997: 91, Figs 8–26), suggest tusa, have an ochraceous to yellowish brown body that these lobes are movable. Further comparative without brown or black marking, a fairly narrow study of the species of Muda and especially Katoa is head (0.78–0.85 times as wide as pronotum col- needed to establish whether these features are syn lar), a pronotum that narrows to the anterior, and a apomorphies for the two genera or perhaps sustain male pygofer that extends from the tip of the abdo- the synonymy of the two genera. men as a short tail. Two species, M. tahanensis and Other characters found in species of both genera M. tua, have a brown body with a darker brown may help to unravel the relationships in Muda and or black marking, a fairly broad head (0.90–0.96 Katoa. Katoa chlorotica (body length male: 11.9– times as wide as pronotum collar), a parallel-sided 14.3 mm) and K. paura (14.0 mm) are small green pronotum and membranous aedeagal appendages. species, just like Muda obtusa (13.2–13.9 mm) and Muda kinabaluana has brown markings on its body, M. virguncula (14.8–18.0 mm), though specimens a fairly narrow head (0.79–0.81 times as wide as of the latter two species in collections are often yel- pronotum collar) and a parallel-sided pronotum. In lowish brown to brown. It is very remarkable that the both M. obtusa and M. kinabaluana the aedeagus other species of Muda (body length male: 12.2–14.4 has a very interesting feature, viz., a recurved flat, mm) are also small in size, while the other species of chitinized apical appendage with 10–12 long spines Katoa (24.6–26.5 mm) are much larger. The more and a few smaller spines, all along one side; the ae- or less chitinized apical part of the male theca armed deagus of M. virguncula has no appendages. The Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 2–4. Muda virguncula, males, Malaysia, Sabah, Gunung Kinabalu, Sayap, Gua Melayu trail. – 2, Body in dorsal view; 3, pygofer in ventral view with straight unfolded upper pygofer lobes; 4, left timbal in laterodorsal view. bpl, basal pygofer lobe; upl, upper pygofer lobe. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 5–8. Muda virguncula, males. – 5, Basal part of abdomen with operculum in ventrolateral view, Malaysia, Sabah, Tawau Hill, jungle lodge; 6, pygofer in lateral view with straight unfolded upper pygofer lobes, Malaysia, Sabah, Tawau Hill, jungle lodge; 7, pygofer in ventral view with folded upper pygofer lobes, Java, Malang; 8, pygo- fer in lateral view with folded upper pygofer lobes, Java, Malang. bpl, basal pygofer lobe; upl, upper pygofer lobe.
allocation of M. tua in the genus is questionable 1. Body uniform yellowish brown to brown without since it differs from the other species in the male dark brown or black marking ���������������������������� 2 pygofer. – Body brown with dark brown or black marking ������������������������������������������������������������ 3 2. Wings with five apical areas. Timbals with 7–8 Key to the Sundaland species of Muda transverse ribs. Aedeagus with a long apical ap- This key is usable principally for the identification pendage armed with 10–12 long spines. Java of males and may not be applicable for the identi- ������������������������������������������������������������ M. obtusa fication of the females as the females of two species, – Wings with six apical areas. Timbals with five M. obtusa and M. tahanensis, were not available. transverse ribs. Aedeagus without appendage Downloaded from Brill.com10/05/2021 12:52:07PM via free access
armed with spines. Java, Sumatra, Mentawai Is- several publications. By far the most species of cica- lands, Borneo ��������������������������������M. virguncula das in this area belong to two groups: the tribe Chlo- 3. Timbals absent, a convex sklerite replaces the rocystini, including 147 species (De Boer 1995a), timbal (Fig. 20). Borneo �������������M. kinabaluana and the subtribe Cosmopsaltriina including 134 Timbals present. Malay Peninsula ��������������������� 4 species (Beuk 2002, Duffels & Turner 2002, Duffels 4. Timbals with six transverse ribs. Postclypeus in 2018). Taxon cladograms and taxon-area cladograms male 0.66–0.70 times as long as broad. Append- of these groups demonstrate a high degree of ende- ages of aedeagus as in Fig. 13. Malay Peninsula mism at the species and genus level and also provided �����������������������������������������������������������������M. tua the basis for the reconstruction of their biogeograph- – Timbals with nine transverse ribs. Postclypeus in ical history. male 1.27–1.35 times as long as broad. Append- De Boer (1995a) regarded Muda as the sisterge- ages of aedeagus as in Fig. 17. Malay Peninsula nus of the Chlorocystini and therefore South East ������������������������������������������������������ M. tahanensis Asia and the Ryukyu Islands, where Muda occurs, as the most likely source area of the Chlorocystini of New Guinea and the West Pacific. Another group Remark on Muda kuroiwae (Matsumura, 1913) of cicadas, the Cosmopsaltriina, demonstrates a sis- Muda kuroiwae was described by Matsumura (1913: tergroup relationship between the genus Meimuna 85, 86, pl. x, Fig. 3) and is known from the Ryukyu from northeastern Asia and a group of genera oc- Islands of southern Japan (Metcalf 1963). I have curring in the West Pacific Duffels( & Turner 2002). seen a male specimen of this species from Okinawa, Though the area cladograms of Chlorocystini and Ryukyu Islands in the BMNH collection, identified Cosmopsaltriina show considerable congruence and by Dr Masami Hayashi. M. kuroiwae can be distin- though both groups originated in Asia, it was also guished from the other Muda species by its large clear that the Chlorocystini originate from South body size (body length ♂: 20 mm) and the high East Asia (Sundaland and adjacent mainland of number of 13 timbal ribs. The upper pygofer lobes southeastern Asia) while the Cosmopsaltriina origi- are similar to those of M. obtusa and M. virguncula; nate from northeastern Asia. A close relationship or Dr. Masami Hayashi (pers. comm., 19.xii.2010) synonymy of Muda and Katoa, as presented in this kindly informed me that the upper pygofer lobes paper, suggests that the source area of the Chloro- of M. kuroiwae can be folded inwards as described cystini might include not only South East Asia but above in the diagnosis of the genus. The mediodorsal also more northern areas in the provinces of Szech- carina of the pygofer is weakly developed. The apex uan and Shaanxi in Central China, where species of of the aedeagus of M. kuroiwae bears two relatively Katoa have been found. long membranous appendages ending in very sharp, chitinized apices. Descriptions of Sundaland species of Muda Biogeography Distribution of Sundaland species of Muda Muda virguncula (Walker, 1856) One species of Muda, M. virguncula, is widely dis- Figs 1–8, 25 tributed in Java, Sumatra and northern Borneo. The other Sundaland species show more or less endemic Cicada virguncula Walker, 1856: 84. Holotype ♂: ‘Wallace distributions. Muda obtusa is recorded from Djam- / Sing’ [Wallace printed between two lines; Sing hand- pang Tengah, West Java and is probably endemic to written], ‘virguncula n .s.’ [handwritten], ‘Type’ [round West Java. Muda tua is only recorded from Endau label with red circle; print] (BMNH) [examined]. Rompin N.P. in the province of Johor, West Malay- Cicada virguncula: Distant 1892b: 153; Distant 1906: 188. sia, while M. tahanensis is only known from the prov- Muda virguncula: Moulton 1923: 158, 159, 170 (equals ince of Pahang, West Malaysia. Muda kinabaluana Baeturia beccarii); Moulton 1929: 121–122; Metcalf seems to be endemic to the Mount Kinabalu massif 1963: 244, 245; Duffels & Van der Laan: 255. in Sabah, North Borneo. Sanborn et al. (2007) and Baeturia beccarii Distant, 1888: 524. Lectotype ♂: ‘Suma- Boulard (2008) recorded two undescribed species of tra / Mte Singalang / Luglio 1878 / O. Beccari’, ‘ Bae- Muda from Thailand, and I have also seen unidenti- turia / beccarii / Dist [Distant’s handwriting] / typus !’, fied specimens ofMuda from Thailand. ‘Beccarii Dist.’. ‘Typus’ [in red and red cadre], ‘Synty- pus ♂ / Baeturia / Beccarii / W.L. Distant 1888’ [print Historical biogeography in black cadre on red paper], ‘Museo Civico / di Genova’ The origin of West-Pacific cicadas and the routes of (MSNG) [examined]. dispersal of cicadas from Asia into Sulawesi, New Baeturia beccarii: Distant 1892b: 149. Guinea and the West Pacific have been the subject of Muda beccarii; Distant 1906: 156; Blöte 1958: 266. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Muda concolor Distant, 1897: 384. Lectotype: ‘Mentawai / J.M.A. v. Groenendael, 1♂ (ZMAN); Soekaboemi, coll. Si Oban IV-VIII / Modigliani 94’ [print in black cadre], Ouwens 1♂ 1♀ (MZB); Surrounding of Soekaboemi, ‘Muda / concolor Dist. [Distant’s handwriting]’, ‘ Syn- 2000’, 24.ii.1940, J.M.A. v. Groenendael, 1♂ (ZMAN); typus ♀ / Muda / concolor / W.L. Distant 1897’ [print Surrounding of Soekaboemi, 22.ii.1940, J.M.A. v. in black cadre on red paper], ‘Museo Civico / di Genova’ Groenendael, 1♀ (ZMAN); W. Java, Soekanegara, 400– (MSNG) [examined]. 1000 m, ii.1940, native coll., 1♀ (MZB); Soekanangli, x. Muda concolor: Distant 1906: 156; Moulton 1923: 158 (in [19]13, 1♂ (ZMAN); W. Priangan, Tjimerang, 1400– synonymy of M. obtusa); Moulton 1929: 121 (in syn- 1800’, 20–25.xi. 1939, J.M.A. v. Groenendael, 1♀ onymy of Muda virguncula). (ZMAN); Tjimerang, 15.viii. 1937, J.M.A. v. Groenen- Muda obtusa [nec Walker]: Zaidi & Ruslan, 1995a: 219; dael, 1♂ (ZMAN); Tjimerang, 1800 m, 2.iv. 1939, J.M.A. Zaidi & Ruslan, 1995b: 200–203; Zaidi, Ruslan & Ma- v. Groenendael, 1♂ (ZMAN); Off W. Java, P. Panaitan, hadir 1996: 61; Zaidi & Ruslan 1998a: 3, 6; Zaidi & ix.1951, native coll., 2♂ 2♂ (MZB). Kalimantan: East Ruslan 1998b: 370; Zaidi, Ruslan & Azman 1999: 319; Kalimantan, Bulungan, Kayan Mentarang, Nature Re- Zaidi, Noramly & Ruslan 2000: 331; Zaidi, Ruslan & serve, Lalu Birai Res. Stn., 02°51’ N, 115°28’ E, 355 m, Azman 2000: 218; Zaidi, Azman & Ruslan 2001: 112, 20.ii-4.iii.1993, IIS 930000, primary rainforest, mixed 116, 122; Zaidi & Nordin 2001: 185, 191, 204; Zaidi dipterocarp, D.C. Darling, 1♀ (ROM); East Kalimantan, & Azman 2003: 106. Kayan-Mentarang Nature Reserve, IIS 940507, 2°52’ N, For further references before 1980 see: Metcalf (1963). 115°49’ E, 378 m, 28.iii–16.iv.1994, lowland diptero. for- Note: Baeturia beccarii: Distant 1892a: pl. xiv, Figs 27, 27a, b est, WWF station Lalut Birai, B. Hubley, D.C. Darling, (these Figures depict Baeturia famulus Distant, 1906 which UV light, 1♂ 2♀ (ROM); E. Borneo, Tabang, Bengeh is a junior synonym of Papuapsaltria nana (Jacobi, 1903)). River, 2.ix.1956, S.M.R. Wegner, 1♂ (MZB). Sumatra: See paragraph on synonymy. Aceh, Mt. Leuser Nat. Pk., Ketambe Res. Stn., beside Alas River, 350 m, 2–6.xi.1989, ROM 893020, B. Hubley, Material examined. 80♂, 67♀. Indonesia: Java: E [East] D.C.Darling, UV/Hg vapour light, 1♂ (ROM); Benkoel- Java, Blitar, ix.1912–iv.1913, coll. E.H.J. v.d. Beek, acq. en, Marang-Liwa, coll. Noualhier 1898, 2♂ 1♀ (MNHN), vi.1945, 1♂ (ZMAN); Depok, 100 m, 15.xii.1935, M.A. same data but with: Muda obtusa Walk., 1♂ (MNHN); Lieftinck, 1♂ (MZB); W. Java, Djampang Tenggah, 10. North Sumatra, near Brastagi, Gunung Sibayak, 03°14’ N, xii.1939, J.M.A. v. Groenendael, 1♂ 1♀ (ZMAN); Djam- 098°29’ E, 1650 m, 02.iii.2002, M. Fiebiger & K. Larsen, pang Tenggah, 1000’, x.1941, J.M.A. v. Groenendael, 1♂ 1♂ (ZSM); Medan, Soengai Krio, iv.1928, J.C. v.d. Meer (ZMAN); Djampang Tenggah, 1500’, 15.i.1940, J.M.A. v. Mohr, 1♂ (MZB); Lampung Krui, Pahmungan, 5°11’ S, Groenendael, 1♀ (ZMAN), same data but 24.i.1940, 1♂ 103°57’ E, Damar Garden, at light, 23.viii.2001, K. Smets, (ZMAN), 27.i.1940, 1♂ (ZMAN), 2.ii.1940, 1♀ 1♂ (ZMAN), same data but 01.ix.2001, under leaf Lansi (ZMAN), x.1941, 1♀ (ZMAN); Djampang Tenggah, um domesticum (Meliaceae), K. Smets, 1♀ (ZMAN); 1800’, 10.iv.1939, J.M.A. v. Groenendael, 1♂ (ZMAN), Lampung Krui, Pahmungan, 5°11’ S, 103°57’ E, Damar same data but 16.v.1939, 1♂ 1♀ (ZMAN), 18.v.1939, 1♂ Garden, river bank, under leaf Shorea javanica (Dipt.), (ZMAN), 28.v.1939, 1♀ (ZMAN), 25.xi.1939, 1♀ 14.xi.2001, K. Smets, 1♀ (ZMAN); North Sumatra, near (ZMAN); Djampang Tenggah, Tjiajunau, 2600’, ix.1941, Prapat, 02°45’52” N 099°58’20” E, 28.ii.2002, 1050 m, J.M.A. v. Groenendael, 1♂ (ZMAN); W. Java, Djampang T. Kothe, 3♂ 5♀ (ZSM); North Sumatra, Umg. [sur- Wetan, 1800’, 10.xii.1939, J.M.A. v. Groenendael, 1♂ roundings] Prapat, 31.x.1999, U. Buchsbaum, 1♂ (ZSM); (ZMAN), same data but 10.iv.1940, 1♂ (ZMAN); Mont Sumatra’s O. K. [East Coast], Sibolangit, 27.x.1921, 550 Gedeh (J.B. Ledru), R. Oberthür, 1♂ 2♀ (MNHN); Ge- m, 1♀ (ZMAN); N. Sumatra, Sidikalang, 02°41’51’ N, deh, Tjisaroea, Panggerango, 1000 m, vi.[19]32, M.A. 98°18’18” E, 1250 m, 15–16.ii.2002, T. Kothe, 1♂ Lieftinck, 1♀ (MZB); Idjen, 1400 m, Kendeng III, (ZSM); Simarjarunjung, near Lake Toba, 02°50’11” N, vi.1924, Dammerman, Muda virguncula det. Karny, 1♂ 098°46’20” E, 1700 m, 18.ii.1999, U. Buchsbaum, 1♂ 1♀ (MZB); Malang, 1♂ (ZMAN); Malang, Prasia, 1♂ (ZSM); North Sumatra, near Siprok, Danua Marsaput, (TMB); Noesa Kambangan, xi.1911, F.C. Drescher, coll. 01°37’50” N, 099°20’30” E, 1435 m, 27.ii.2002, T. Kothe, F.C. Drescher, 1♂ (ZMAN), same data but x.1913, 1♂ 2♀ (ZSM); Sumatra’s O. K. [East Coast], Tandjong Merah, (ZMAN); W. Java, Palaboeanratoe, xi.[19]35, F. Dupont, 22 m, xi.1917, J.B. Corporaal, 1♂ (ZMAN); S. Sumatra, 1♀ (MZB); Priangan, 1935–1939, J.M.A. v. Groenendael, Lemongs, Wai Lima, xi–xii.1921, nog. 73, Karny & 2♂ 2♀ (ZMAN), same data but vii.1935, 1♂ (ZMAN), Siebers, 1♀ ( MZB); S. Sumatra, Mt. Tanggamoes, 600 m, same data but vii.1937, 1♂ (ZMAN); ZW Priangan, 19–31.iii.1940, M.A. Lieftinck, 1♂ ( MZB); S. Sumatra, 1800’, 4–12.xi.1941, J.M.A. v. Groenendael, 2♂ 1♀ S. W. Lampongs, Mt. Tanggamoes, Gisting, 600–700 m, (ZMAN); W. Java, Selabintanah, xi.1933, Walsh, 1♀ ult. xii. 1939, M.A. Lieftinck, 1♀ (MZB); S. Sumatra, S. (MZB); Soekaboemi, 1600–2000’, 8.iii.1940, J.M.A. v. W. Lampongs dist., Mt. Tanggamoes, Gisting, 1000 m, Groenendael, 2♂ 1♀ (ZMAN); Soekadoemi [= Soekabo- 29.xii.1934, Lieftinck /Toxopeus, 1♀ (MZB). Mentawai emi], 29.iii.1926, Mrs M.E. Walsh, 4♂ 2♀ (SEM); Soeka- Islands: Mentawai, Si Oban iv–viii.[18]94, Modigliani, ♀ boemi, Djampang Kidoel, 1000–1500’, 15.xii.1939, Lectotype of Muda concolor, W.L. Distant 1897’ (MSNG). Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Malaysia, Sabah: 105 km S of Beaufort, Long Pa Sia area, Camp, 2–20 m, 6.xi.[19]57, J.L. Gressitt, light trap, 2♀ Payakalaba, 4°25’ N, 115°44’ E, 1000 m, 13.iv.1987, (BPBM), same data but 3.xi.[19]57 (BMNH); 60 km W somewhat disturbed kerangas vegetation, at light, 04.00– Sandakan, Sungai Darling, 26.xi.1989, sample Sab. 43, 06.15 h, Van Tol & Huisman, 1♀ (RMNH); 105 km S of secondary forest understorey, at light, M.J. & J.P. Duffels, Beaufort, Long Pa Sia area, airstrip Long Pa Sia, 4°24’ N, 2♂ 1♀ (ZMAN); Tawau Hill, jungle lodge, 300 m, gar- 115°43’ E, 1000 m, 15.iv.1987, semicultivated area, near den/secondary growth, at light, 25.iii.2001, J.P. & M.J. disturbed evergreen trop. rainforest, at light, 18.20–20.30 Duffels, 1♂ (ZMAN), same data but 26.iii.2001, 1♂ 1♀ h, Van Tol & Huisman, 1♀ (RMNH); Bettotan, nr Sanda- (ZMAN), 27–29.iii.2001, 1♂ 1♀ (UMS), 3♀ (ZMAN); kan, 25.vii.1927, C.B.K. & H.M.P. F.M.S. museums, ex Tawau, Quoin Hill, Cocoa Res. Sta., 225 m, 14.ix.1962, F.M.S. Museum, B.M. 1955–354, 1♂ (BMNH), same malaise trap, Y. Hirashima, 1♀ (BPBM); Tawau, Quoin data but 31.vii.1927, 1♀ (BMNH), 1.viii.1927, 1♂ Hill, Cocoa Res. Sta., 225 m, 25.ix.1962, K.J. Kuncheria, (BMNH), 1.viii.1927, 1♂ (BMNH); Danum Valley, 5°01’ 1♀ (BPBM). Malaysia: Sarawak: Gunung Mulu Nat. N, 117°47’ E, 120 m, 1.x.1987, lowland mixed diptero- Park, Site 16, Long Pala (Base), 324450, 70 m, Allu./sec- carp forest, light trap sample understory forest edge, NMW ond. forest, MV-on batu-Canopy, March, J.D. Holloway, Sabah (Borneo) Expedition, NMW.Z.1987.094, A.H. RGS Mulu exped., B.M. 1978–206, 2♂ (BMNH); Trail Pa Kirk-Spriggs, l♂ (NMWC); Danum Valley, 5°01’ N, Lungan–Long Rapun, 3°53’ N, 115°35’ E 1200 m, 23. 117°47’ E, 220 m, 26.ix.1987, light trap sample, roadside, ii.1987, J. Huisman, 1♂ (RMNH). Banguey Island: secondary forest, NMW Sabah (Borneo) Expedition, Banguey Island, coll. Noualhier 1898, Distant coll. 1911– NMW.Z.1987.094, A.H. Kirk-Spriggs, 1 ♂ (NMWC), 383, 1♂ (BMNH). Brunei: Temburong District, ridge NE same data but 200 m, 29.ix.1987, 1♀ (NMWC), same of Kuala Belalong, approx. 300 m alt, x.1992, J.H. Martin, data but 11.ix.1987, 1♀ (NMWC); Danum Valley, 70 km coll. B.M. 1992–172, 1♂ (BMNH); Lamunin, 13.i.1996, W Lahad Datu, Field Centre at Sungai Segama above staff- T.W. Harman, 1♀ (ZMAN). quarters, 150 m, 5.xii.1989, sample Sab. 57, at light, M.J. & J.P. Duffels, 1♂ 1♀ (ZMAN); DVFC [Danum Valley Field Station], 60 km W Lahad Datu, buildingsite staff- Diagnosis quarters, 4°58’ N, 117°48’ E, 150 m, 20.x.1987, J. Huis- M. virguncula and M. obtusa can be separated by the man & R. de Jong, 1♀ (RMNH), same data but 28.x.1987, number of apical areas in the wings: six in virguncula 3♀ (RMNH); 19 km N of Kalabakan, Forest Camp, and five in obtusa. The males of M. virguncula and 27.x.1962, Y. Hirashima, light trap, 1♂ (BPBM); M. obtusa are also distinguished by the number of R. Karamvak, Kunantong, 200’, 3.ix.1977, M.E. Bacchus, timbal ribs: five in M. virguncula and 7–8 in M. ob B.M. 1978–48, 2♂ (BMNH); Keningan [= Keningau], tusa. Furthermore, M. virguncula has a very simple, 12–17.i.1959, T.C. Maa, 1♂ 2♀ (BPBM); Gunung long aedeagus without armature, while the aedeagus Kinabalu, Sayap, Gua Melayu trail, 06°10’N, 116°34’ E, of M. obtusa has a long apical appendage armed with 1000 m, clearing in primary forest, 10.iii.2001, at light, 10–11 long spines. Most males of the two species are J.P. Duffels & M.A. Schouten, 1♂ (ZMAN); Gunung separated by the ratio of the distance between the Kinabalu, Sayap, Jalan raya, 116°34’E 06°10’N, 1000 m, two lateral ocelli and the distance between the lat- edge of primary forest, 11.iii.2001, at light, J.P.Duffels & eral ocellus and the eye, which is respectively (0.92) M.A. Schouten, 1♀ (ZMAN); 75 km W Lahad Datu, 1.07–1.46 in males of M. virguncula (n = 14) and confl. S. Sabran, S. Danum, S/N, 4°57’ N, 117°41’ E, 200 0.87–0.92 in males of M. obtusa (n = 3). Moulton m, 23.x.1987, J. Huisman & R. de Jong, 1♀ (RMNH), (1923) distinguished M. virguncula and M. obtusa by same data but 25.x.1987, 1♂ 1♀ (RMNH); 15 km W La- the veins to the radial area and the upper vein of the had Datu, cacao plantation/acacia shadowtrees, sample lower ulnar area, but these features are not reliable to Sab. 61, 7.xii.1989, M.J. & J.P. Duffels, 1♀ (ZMAN); 16 separate these species. km NE Tenom, Agr. Res. Station, resthouse, 5°12’N, 115°59’ E, 270 m, xi.1987, J. Huisman & R. de Jong, 1♀ Lectotype designation for Baeturia beccarii (RMNH); Liawan, 14–19.i.1959, T.C. Maa, 1♂ (BPBM), same data but 14–17.i.1959, 1♀ (BPBM); Poring hot and Muda concolor spring, 21.iv.1999, Trilar leg., 1♂ (PSML); Poring hot Baeturia beccarii Distant, 1888 was probably de- spring, Sg. Montokungon, Ranau–Kg Poring, 450 m, c. scribed after one male and two females from Mt Sin- 6°02’30” N, 116°43’ E, 25.xi.1986, J. Huisman, galang, Sumatra, which are now located in the MSNG 1♂ (RMNH); Poring hot spring, confl S Kepungit × and BMNH collections. At my request, Dr. Fabio S. Langanan, 6°03’ N, 116°42’ E, 450 m, 8.xii.1986, Penati (MSNG) kindly sent me two type specimens Huisman, 1♂ (RMNH); 23 km W Sandakan, Sepilok, tree of B. beccarii, a male and a female, from the MSNG tower, 5°49’ N, 118°06’ E, 1.xi.1987, 0–100 m, ML, collection. The male bears the labels: ‘Sumatra / Mte J. Huisman & R. de Jong, 1♂ (RMNH); Sandakan Bay Singalang / Luglio 1878 / O. Beccari’, ‘Baeturia / (NW), Sepilok For. Res., 1–10 m, 30.x.1957, J.L. Gressitt, beccarii / Dist [Distant’s handwriting] / typus!’, ‘Bec- 1♀ (BPBM); Sandakan Bay (SW), Sapagaya Lumber carii Dist.’. ‘Typus’ [in red and within red cadre], Downloaded from Brill.com10/05/2021 12:52:07PM via free access
‘Syntypus ♂ / Baeturia / Beccarii / W.L. Distant different species and assigned the Figures of Baetu 1888’ [printed on red]. According to Dr Penati, it ria beccarii in his monograph (Distant 1892a: pl. xiv was Dr Raffaello Gestro, vice-director of MSNG in Figs 27, 27a, b) to Baeturia famulus. Since this pub- Distant’s time, who wrote ‘typus!’ at the bottom of lication, Baeturia famulus Distant, 1906 is a valid Distant’s identification label and added the ‘Beccarii name. Half a century later, Blöte (1958) revealed Dist.’ and the ‘Typus’ labels. The syntypus label was that the specimen of Baeturia famulus depicted in added by Dr. Penati. The female type specimen in Distant’s monograph (1892a) is a male from Buja- MSNG bears the following labels: ‘Sumatra / Mte kori, New Guinea, in the BMNH collection and he Singalang / Luglio 1878 / O. Beccari’, ‘Syntypus ♀ regarded this specimen as the holotype of Baeturia / Baeturia / Beccarii / W.L. Distant 1888’, ‘Museo famulus. Metcalf (1963) incorrectly stated that Dis- Civico / di Genova’. Dr. A.J. de Boer (ZMAN), who tant (1906) regarded B. famulus as a synonym of B. studied the so-called ‘Baeturia and related genera beccarii. De Boer (1995b) followed Blöte’s reasoning complex’ (e.g., De Boer 1995a), kindly informed me in establishing the identity of Baeturia famulus, re- that the BMNH has a female specimen of Baeturia vealed that B. famulus is a junior synonym of Thau beccarii with the following labels: ‘Sumatra / Mte mastopsaltria nana (Jacobi, 1903), and allocated Singalang / Luglio 1878 / O. Beccari’, ‘beccarii Dist.’ T. nana to the new genus Papuapsaltria De Boer, [handwritten], ‘Type’ [round label with red margin], 1995. ‘Distant 1911–383’. The round type label was prob- The female lectotype of B. concolor from Men- ably not attached by Distant but by a later curator of tawai [Islands] west of Sumatra, has six apical areas the BMNH. I designate the male syntype of Baeturia in the wings and is therefore synonymized here with beccarii in MSNG as the lectotype of the species; the M. virguncula, which is widespread in Sumatra, Java label ‘paralectotype of Baeturia beccarii’ is added to and Borneo. the female syntypes in MSNG and BMNH. Muda concolor Distant, 1897 was described after two females from Si Oban, Mentawai. Si Oban cor- Description responds to the modern town of Sioban on Sipora Body. Yellowish brown to brown without dark mark- Island in the Mentawai archipelago west of Sumatra. ing, occasionally with bright red specks, parts of the Dr. Fabio Penati (MSNG) kindly sent me on loan a body sometimes tinged with green. Lateral sides of female type specimen of B. concolor from the MSNG mesonotum with scattered silver pilosity, abdomen collection. The labels of this specimen are given dorsally pilose, underside of thorax and abdomen above. I designate this specimen as the lectotype of with more scattered and longer silvery setae. Abdo- B. concolor. I have not seen the other type specimen. men of male and female respectively 1.31–1.7 times (n = 8) and 1.55–1.78 times (n = 8) as long as head and thorax together. Synonymy Head. Head fairly narrow, distinctly narrower than Cicada virguncula Walker, 1856 was described after pronotum at half-length, head in male 0.78–0.82 a male from Singapore in the BMNH collection. times (n = 8) as wide as pronotum collar, in female Distant disregarded Cicada virguncula in his 0.76–0.83 times (n = 8). Postclypeus flat, in male publications because he could not find Walker’s 0.67–0.82 times (n = 8) as long as broad, in female type in the BMNH collection (Distant, 1892b: 153; 0.70–0.82 times (n = 8). Anterior postclypeal margin 1906: 188). triangularly protruding beyond angularly rounded Baeturia beccarii was described by Distant (1888) anterior margins of vertex lobes. Each of the ocelli from Mt. Singalang, Sumatra, and later transferred encircled by a bright reddish ring. Distance between by him to the genus Muda (Distant 1906). After lateral ocelli in male (092) 1.07–1.46 times (n = 14) Moulton (1923) discovered Wallace’s type of Cicada the distance between lateral ocelli and eyes, in female virguncula in a series labelled Muda beccarii in the 1.33–1.55 times (n = 8). Ventral side of postclyp- BMNH, he transferred Cicada virguncula to the ge- eus with two series of 6–7 transverse ridges; in most nus Muda and synonymized Baeturia beccarii with specimens these ridges end laterally in a rasp consist- Muda virguncula. Examination of the holotype of ing of one, two or three, short, longitudinal ridges. Cicada virguncula and the lectotype of Baeturia bec Rostrum reaching just beyond coxae of hind legs. carii, including study of the male genitalia for the Thorax. Pronotum with lateral sides slightly converg- present paper, confirms this synonymy. ing to the anterior. Baeturia famulus, a manuscript name of Stål, was Legs. Ochraceous to light brown, claws either en- listed under Baeturia beccarii in Distant’s ‘Mono- tirely, or only distally, brown. Fore femur with three graph of Oriental Cicadidae’ (1892b: 149). In his spines along underridge: a stout and appressed spine ‘Synonymic Catalogue of Homoptera’ of 1906, at two fifths of length from base, a somewhat shorter, Distant treated Baeturia famulus and B. beccarii as fairly broad, erect spine at two thirds and an again Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 9–11. Muda obtusa, males. – 9, Pygofer in ventral view, W. Java, Djampang Tengah, 1800’, 23.v.1939; 10, basal part of abdomen with operculum in ventrolateral view, W. Java, Djampang Tengah, 1800’, 7.iv.1939; 11, left timbal in laterodorsal view, W. Java, Djampang Tengah, 1800’, 7.iv.1939.
in the reference to the original description. This green. Dorsal side of body with sparse silver pilosity, specimen is designated here as lectotype of Muda ventral side of thorax and abdomen with scattered obtusa. and longer silvery setae. Abdomen 1.08–1.12 times M. obtusa is uncommon in collections and prob- (n = 3) as long as head and thorax together. ably endemic to West Java. Apart from the lecto- Head. Head fairly narrow, distinctly narrower than type from Java, I have found two males and two pronotum at half length and 0.82–0.85 times females of M. obtusa from Djampang Tengah, Java, (n = 3) as wide as pronotum collar. Postclypeus flat, among the unidentifiedMuda ’s available for this 0.56–0.79 times (n = 3) as long as wide. Anterior study. postclypeal margin triangularly protruding beyond angularly rounded anterior margins of vertex lobes. Margins of each of ocelli with more or less distinct Description of male reddish marking. Distance between lateral ocelli Body. Yellowish brown to light brown without dark 0.87–0.92 times (n = 3) as wide as distance between markings, parts of body sometimes tinged with lateral ocellus and eye. Ventral side of postclypeus Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 12–14. Muda tua, males. – 12, Body in dorsal view, paratype Endau Rompin NP, road Staging point to Kuala Jasin, bridge 9, 21.iii.1999; 13, pygofer in ventral view, holotype; 14, basal part of abdomen with operculum in ventrolateral view, holotype.
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reddish tinge. Sternites reddish especially toward pos- terior margins. Tergites 2–8 distinctly arched medi- ally. Lateral margin of abdomen between tergites and epipleurites sharply folded. Sternite 8 large, 1.5 times as long as wide and 1.5 times as long as sternite 7. Male genitalia. Pygofer about twice as long as broad. Basal pygofer lobes fairly narrow, apically rounded and reaching to two thirds of pygofer length. Up- per pygofer lobes very long, tapering to pointed and inwardly curved apex; tips of lobes of both sides crossed. Clasper quadrangular, twice as long as wide, medial margins of claspers more or less parallel. Ae- deagus with two apical appendages: a relatively short Fig. 15. Muda tua, male, right timbal in laterodorsal and narrow one and a very long one, splitting in a view, holotype. very long and fairly narrow, coiled part that widens in its distal half and narrows to the sharp sklerotized apex, and a somewhat shorter, broad part that gradu- parallel to oblique fissures: one pair of fasciae is situ- ally narrows to a sharp apex. ated between anterior and posterior oblique fissures, Female operculum. Similar to that of male. the other pair between posterior fissures and lateral Female abdomen. Light brownish. Posterior margins parts of ambient fissure. Lateral corners of pronotum of tergites somewhat darker brown, sometimes with collar with a pair of dark brown spots. a light reddish tinge. Sternites 3–6 with reddish band Mesonotum light brown with very weakly de- along posterior margin. Tergites sharply folded ven- veloped marking. Two juxtaposed brown triangular trolaterally, and tergites 2–8 arched, as in male. Fe- spots reach from dark brown anterior branches of male pygofer long and slender. Ovipositor very long, cruciform elevation along half-length or two thirds reaching far beyond apex of segment 9. of mesonotum; these two triangles are separated by Measurements in mm. (n = 8♂, 8♀). Body length ♂ a narrow median line. Some specimens have a pair 12.2–14.4, ♀ 14.7–17.5; head width ♂ 3.9–4.2, ♀ of lateral sigillae, hardly darker than the ground co- 4.4–4.9; pronotum width ♂ 4.1–4.5, ♀ 4.6–5.2; lour, but in most specimens the sigillae are reduced tegmen length ♂ 14.2–16.0, ♀ 17.0–19.4. to some spots or even completely lacking. Legs. Ochraceous, often tinged with green, apical part of pretarsus of all legs darkened. Claws brown. Distribution Fore femur with three sharply pointed, brown spines This species is fairly common in Endau Rompin along underridge: a semi-erect, stout spine at two N.P., Johor, West Malaysia, and not recorded from fifths of length from base, a distinctly shorter, broad other localities. spine at two thirds and a broad and curved, distal spine with a very small extra spine more distally. Muda tahanensis (Moulton, 1923) Tegmina and wings. Hyaline. Costa light brownish, remaining venation brown to red–brown or reddish. n. comb. Wings with 5 apical areas. Figs 16–19, 25 Male operculum. Medial two thirds of operculum flat or weakly convex, separated from hollow lateral part Abroma tahanensis Moulton, 1923: 156–157, Lectotype by a distinct longitudinal ridge. Mediodistal corner ♂ [by present designation]: ‘Pahang FMS / G. Tahan of operculum rounded triangular. Distal margin of 5550‘ / March 1921 / E. Seimund’, ‘Abroma tahanensis operculum upcurved and weakly convex up to ridge Moulton / Type ♂ 22.5.23’, ‘1923–352 / Brit. Mus’, separating both operculum parts. Lateral margin dis- ‘Type / H.T’ [round label with red margin; print] tally with convex upcurved protrusion. Meracanthus (BMNH) [examined]. long, reaching across the open space between oper- Abroma tahanensis: Zaidi & Ruslan 1997: 232; Zaidi & culum and abdomen to anterior margin of abdomi- Azman 2005: 50, 58. nal segment 2. For further references see: Sanborn (2014). Timbal organs. Completely visible in lateral view. Timbal with six transverse, slightly curved, parallel Material examined. Malaysia: Pahang, G. Tahan, 5550’, ribs and short intercalary ribs. iii.1921, E. Seimund, male lectotype and male paralecto- Male abdomen. Light brownish. Lateral corners of type of Abroma tahanensis (BMNH); Pahang, King George segment 1 dark brown. Posterior margins of tergites V Nat. Park, Kuala Tahang, 7–14.xii.1958, J.L. Gressitt somewhat darker brown, sometimes with a light collector Bishop Museum, 1♂ (BPBM). Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 16–18. Muda tahanensis. – 16, Body in dorsal view, holotype, male. 17–18, Male, Malaysia, Pahang, King George V Nat. Park, Kuala Tahang: 17, pygofer in ventral view; 18, basal part of abdomen with operculum in ven- trolateral view. bpl, basal pygofer lobe; upl, upper pygofer lobe.
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Description of male The following description is made after the lectotype and a specimen from Kuala Tahang, with exception of the marking on the pronotum that is described after the lectotype, since the marking of the Kuala Tahang specimen is fairly obsolete. Body. Light brown to brown, but abdomen of Kuala Tahang specimen dark brown; markings on head and thorax black to dark brown; posterior mar- gins of abdominal segments often reddish. Head and thorax fairly densely silvery pilose, postclypeus with scattered long setae. Abdomen with scattered pilosity, apical part and ventral side of abdomen also with long, silvery or brownish setae. Abdomen 1.11–1.25 times (n = 2) as long as head and thorax together. Head. Head fairly broad, slightly broader than pro- notum at half-length, head 0.90–0.92 times (n = 2) as wide as pronotum collar. Postclypeus 1.27–1.35 times (n = 2) as long as broad. Anterior postclypeal margin triangularly protruding beyond weakly con- vex anterior margins of vertex lobes. Postclypeus with a pair of oval paramedian spots at frontoclypeal suture. Distance between lateral ocelli 1.0–1.17 times (n = 2) as wide as distance between ocelli and eyes. Ventral side of postclypeus with two series of 6 transverse ridges. Rostrum black-tipped, reaching just beyond coxae of middle legs. Figs 19–20. Muda tahanensis and Muda kinabaluana, Thorax. Pronotum with about parallel lateral sides. tymbal area. – 19, Muda tahanensis, male, left timbal A pair of paramedian, narrow, triangular black spots in laterodorsal view, Malaysia, Pahang, King George V distally of, and about parallel to, anterior oblique fis- Nat. Park, Kuala Tahang; 20, Muda kinabaluana, male sures. A pair of fasciae curve outward from medial paratype 12–15.iii.2001; cs, convex sklerite replacing ends of anterior oblique fissures to the ambient fis- the timbal. sure. A pair of irregularly shaped figures between an- terior and posterior oblique fissures. A pair of fasciae curve from distal ends of posterior oblique fissures in Lectotype designation lateroproximal direction. Lateral corners of prono- Moulton (1923) described this species after 3♂ and tum collar with a pair of light brownish spots. 1♀ from Malay Peninsula, Gunong Tahan, alt. 5.500 Mesonotum light brown with very weakly de- ft, March 1921, coll E. Seimund, in the F.M.S. Mu- veloped marking. A pair of faint, brown, oval spots seums collection. Two specimens of the type series reach from dark brown anterior branches of cruci- were found in the BMNH: one male labelled Type form elevation to half-length of mesonotum. Para- (labels as above in the reference to the original de- median sigillae discernable in the Kuala Tahang scription) and one specimen, also from Gn Tahan, specimen only. missing the abdomen. The specimen labelled ‘Type’ Legs. As in M. tua. is a male and not a female as stated by Moulton Tegmina and wings. Hyaline, venation brown to red- (l.c.) and is designated here as the lectotype of A. brown. Wings with 6 apical areas. tahanensis; the other BMNH specimen has been la- Operculum. Medial two thirds of operculum weakly belled paralectotype. Both specimens were covered convex, separated from hollow lateral part by a weak in mould and were cleaned by us as far as possible. longitudinal ridge. Mediodistal corner of operculum The lectotype is in poor condition: the left tegmen rounded triangular. Distal margin of operculum and wing, the apical part of the right tegmen and slightly upcurved and weakly convex up to ridge the posterior half of the right wing are missing, the separating both operculum parts. Lateral margin dis- mesonotum is partly destroyed around the pin, and tally with convex upcurved protrusion. Meracanthus the apical part of the aedeagus is gone. The other two long, reaching across the open space between oper- specimens of the type series are probably in MNKM, culum and abdomen to anterior margin of abdomi- but this has not been checked. nal segment 2. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Timbal organs. Completely visible in lateral view. appendage and a fairly broad apical appendage, both Timbal with nine transverse, slightly curved, parallel narrow to a sharp sklerotized apex. ribs and short intercalary ribs. Measurements in mm. (n = 2). Body length 10.8; head Abdomen. Tergites 2–8 distinctly arched medially. width 3.3–3.5; pronotum width 3.5–3.8; tegmen Lectotype with light brown abdomen with reddish length 13.8. posterior segment margins and a darkened tergite 8. Kuala Tahang specimen with tergites 1–6 dark brown, but medial area of tergites 3–6 light brown; Distribution tergite 7 light brown but turning laterally to dark This species was originally described from Mount brown; anterior two thirds of tergite 8 with a pair Tahan (5500’) and later recorded from Mount Tahan of dark brown, paramedian squares extending later- (Zaidi & Azman 2005) and from Pahang, Rompin, ally in a curved fascia reaching to the anterior tergite Sg Kinchin base Camp (Zaidi & Ruslan, 1997). For margin; sternites 3–6 dark brown, posterior third this revision I have seen another specimen from Kua- of these sternites reddish brown; sternites 7–8 light la Tahang. All localities are located in the province of brown. Lateral margin of abdomen between tergites Pahang, Peninsular Malaysia. and epipleurites sharply folded. Sternite 8 large, 1.5 times as long as wide. Genitalia. Pygofer about 2.5 times as long as broad, Muda kinabaluana Duffels n. sp. basal part very broad and suddenly narrowing at Figs 20–24, 26 one third of its length from base: pygofer in basal third 1.4 times as wide as at half-length. Basal pygo- This species is very peculiar because the males are fer lobes fairly narrow, apically rounded and reach- missing the timbals. A convex sklerite replaces the ing to 0.8 of pygofer length. Upper pygofer lobe in sound-producing organ. dorso-lateral view 1.5 times as long as broad at base; it strongly narrows from base to half-length, while its Type material. Holotype ♂: Malaysia: Sabah: ‘MA- apical half slightly narrows to the rounded, inwardly LAYSIA, Sabah / Gunong Kinabalu / SAYAP, 1000 m / curved apex that embraces the anal segment. Clasper 116°34’E 06°10’N’, ‘Jalan raya / edge of primary forest quadrangular, twice as long as wide, medial margins / 11.iii.2001’, ‘at light, J.P. Duffels / & M.A. Schouten’ of claspers more or less parallel. Aedeagus ending in (ZMAN). Paratypes: Sabah: Gunong Kinabalu, Sayap, a sharp sklerotized apex and provided with an ap- 1000 m, 06°10’ N, 116°34’ E, ranger house, edge of pri- pendage which is narrow and sklerotized in basal two mary forest, 12–15.iii.2001, at light, J.P. & M.J. Duffels thirds and strongly widened and membranous in the 1♂ (ZMAN); Gunong Kinabalu, Sayap, 1000 m, 06°10’ apical part; this apical part has a fairly narrow basal N, 116°34’ E, Gua Melaya trail, clearing in primary forest,
Fig. 21. Muda kinabaluana, male paratype in dorsal view, Sabah, Gunong Kinabalu, Sayap, Gua Melaya trail, 1000 m, 12.iii.2001. Photo J. van Arkel. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Figs 22–24. Muda kinabaluana, male. – 22, Body in dorsal view, holotype; 23, pygofer in ventral view, paratype 12–15.iii.2001; 24, basal part of abdomen with operculum in ventrolateral view, holotype. bpl, basal pygofer lobe; upl, upper pygofer lobe; as, apical spine of aedeagus. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Fig. 25. Distribution of Muda virguncula (stars) and M. tahanensis (triangles).
10.iii.2001, at light, J.P. Duffels & M.A.Schouten, 1♀ acute-angled with narrowly rounded apex distinctly (ZMAN), same data but, 12.iii.2001, J.P. & M.J. Duf- protruding beyond rounded anterior margins of ver- fels, 1♂ (ZMAN), 13.iii.2001, J.P. & M.J. Duffels, 1♀ tex lobes, in lateral view with knob-shaped apex and (ZMAN). concave lower margin. Ocelli sometimes encircled Other material: Sabah: Gunong Monkobo, 5°48’ N, by reddish ring. Distance between lateral ocelli in 116°56’ E, Dipterocarp. for., 7–13.viii.1987, K.R. Tuck, male 0.69–0.78 times (n = 4) as wide as distance BM 1987–252, 1♂ (BMNH). between lateral ocelli and eyes, in female 0.67–0.78 times (n = 2). Ventral side of postclypeus with two series of 5–6 more or less distinct transverse ridges Description and distinct crests along lateral margins. Rostrum Body. Yellowish brown, sometimes with a light with brown to black apex, just reaching coxae of hind greenish tinge, and with brown marking; one female legs. paratype reddish brown without marking. Head Thorax. Pronotum with parallel lateral sides and a and thorax dorsally with scattered silver pilosity, pair of more or less faint, light brown spots between ventrally golden pilose; abdomen dorsally silvery the two pairs of oblique fissures. Holotype and one to golden pilose, especially along hind margins of paratype with a V-shaped median brown marking at tergites, underside with short and longer golden se- anterior margin of pronotal collar, in the paratype tae. Abdomen of male 0.99–1.17 times (n = 3) as the V-shaped marking is connected to a pair of sub- long as head and thorax, of female 1.01–1.02 times median, low, brown triangles more laterally along (n = 2). anterior margin of pronotum collar. Head. Head as broad as or slightly narrower than Mesonotum. Holotype and one paratype with a pair pronotum at half-length, head in males 0.79–0.81 of broad paramedian fasciae that run parallel from times (n = 3) as wide as pronotum collar, in females anterior margin of mesonotum to two thirds of me- 0.78 times (n = 1). Dorsally light brown to green- sonotum length where they outcurve to anterior ish, ventrally yellowish brown. Postclypeus in male angles of cruciform elevation and extend laterally. 1.12–1.20 times (n = 3) as long as broad, in female Other paratype with brownish paramedian sigillae 1.10–1.14 times (n = 2). Postclypeus in dorsal view instead of parallel paramedian fasciae. Downloaded from Brill.com10/05/2021 12:52:07PM via free access
Fig. 26. Distribution of Muda obtusa (dot), M. tua (triangle) and M. kinabaluana (diamonds).
Legs. Yellowish brown to light green. Fore femur brown marking, reaching from anterior to posterior with three spines along underridge: a stout and segment margin and about twice as wide as long, and semi-erect spine at two fifths of length from base, a a pair of lateral oval spots. Tergites 3–5 with more or somewhat shorter, fairly broad, erect spine at three less quadrangular, brown, median marking, reaching fourths and an again shorter, broad, erect spine more from anterior to posterior segment margin and about distally; apices of all spines reddish. Distal half of 2–3 times as wide as long. Lateral parts of tergites claws brown. 3–6 with brown markings, that are 2–5 times as wide Tegmina and wings. Tegmen with reddish to reddish as long, medially as high as the segments and laterally brown costa, basal third of venation of tegmen red- sometimes narrower. Tergites 6–7 sometimes with dish brown, apical venation of tegmen and venation light brownish median marking. Holotype with red- of wing yellowish brown to green. Wings with 6 api- dish band along posterior margins of sternites 3–6. cal areas. Antero-lateral parts of segment 8 brownish. Lateral Male operculum. Medial two thirds of operculum margin of abdomen between tergites and epipleu- separated from the concave lateral part by a weak rites sharply folded. Sternite 8 large, twice as long as longitudinal ridge. Mediodistal corner of operculum wide and more than twice as long as sternite 7. narrowly rounded. Distal margin weakly upcurved Male genitalia. Pygofer about 2.5 times as long as and convex up to ridge separating both operculum broad, distinctly constricted at three-fifths from parts. Lateral margin distally with strongly convex base, lateral sides otherwise parallel. Basal pygofer upcurved protrusion. Meracanthus long, reaching lobes slightly widening from base to rounded apex across the open space between operculum and ab- and reaching to nine-tenth of pygofer length. Upper domen, and beyond anterior margin of abdominal pygofer lobes almost twice as long as wide at base, ta- segment 2. pering to pointed and upcurved apex; tips of process- Timbal organs. Timbal absent. A convex sklerite re- es of both sides just meeting above and embracing places the timbal organ. the anal segment. Clasper indistinct, unsklerotized, Male abdomen. Tergites distinctly arched medially. basally narrow and widening in apical part. Apex of Tergite 1 with two low markings widening laterad. aedeagus with a strong, apical spine pointing distad Tergite 2 with median trapezoid or quadrangular and a recurved flat, chitinized appendage with about Downloaded from Brill.com10/05/2021 12:52:07PM via free access
12 long and a few small spines pointing proximad (BMNH), Dr. G.A. Samuelson and Mrs K. Kami along one side. (BPBM). Dr M. Wilson and Dr A.H. Kirk Spriggs Female operculum. Similar to that of male. (NMWC). Dr Maryati Mohamed (UMS) greatly Female abdomen. One paratype reddish brown with- facilitated my fieldwork in Sabah in 2001; the very out marking, other paratype yellowish brown with kind assistance received from the staff of UMS is brown marking. Tergites 2–5 as in male, with a me- gratefully acknowledged. I thank Dr Arnold de dian trapezoid or quadrangular marking reaching Boer (ZMAN) for information about the syntype of from anterior to posterior segment margins. Tergites Baeturia beccarii in the BMNH collection and for 3–6 with lateral brown markings, that are 4 times reading an earlier draft of this paper. I thank my late as broad as high on tergites 3–4 and half as broad wife Greet Duffels-van Egmond for her continuous as high on tergites 5–6. Posterior third of sternites encouragement and help in the field, Dick Langerak 6–7 with reddish tinge. Tergites 2–8 arched, as in (ZMAN) for preparing all but two drawings, Erik- male, and weakly folded ventrolaterally. Female py- Jan Bosch (Naturalis) for making two drawings gofer long and slender. Ovipositor long, reaching far (Figs 7, 8), Rob Portegies (ZMAN) for maintaining beyond apex of segment 9. the database of cicada records and for making the Measurements in mm. (n = 3♂, 2♀). Body length ♂ distribution maps, Henk Caspers (RMNH/Natu- 12.2–13.5, ♀ 14.0–14.2; head width ♂ 3.4–3.6, ♀ ralis) for scanning the drawings and Gerard Verlaan 3.6–3.8; pronotum width ♂ 4.3–4.5, ♀ 4.9; tegmen (ZMAN) for technical assistance. length ♂ 17.1–18.1, ♀ 19.2–20.0. References1 Remark Anonymous, 1938. Atlas van tropisch Nederland. – The specimen from Mt Monkobo, mentioned above Koninklijk Nederlands Aardrijkskundig Genootsc- as other material, is not included in the type series hap & Topografische Dienst in Nederlandsch-Indië, because it differs from the type specimens in the fol- Amsterdam/Batavia, The Netherlands/Dutch East lowing features: the median marking on tergites 2–5 Indies, pp. ix, 17, maps 1–31b, legends 1–31b. is missing though tergites 3–4 have a pair of small Anonymous, 1999. The Times Comprehensive Atlas of the paramedian spots, tergites 3–6 have lateral brown World. – Times Books, London, UK, 67 v, 220 pp, markings as in the male types but more faint, the 124 pls. ratio head width:pronotum width is higher (0.92) Beuk, P.L.Th., 2002. Cicadas spreading by island or by spreading the wings? Historic biogeography of dun- than in the males of the type series (0.79–0.81), dubiine cicadas of the Southeast Asian continent and and the body is smaller (body length 11.5 mm, teg- archipelagos. – Ph.D. thesis, University of Amsterdam, men length 14.8 mm; males type series body length The Netherlands, pp. 1–323. 12.2–13.5 mm, tegmen length 17.1–18.1mm). The Blöte, H.C., 1958. On the history of the genus Baeturia male genitalia of the Monkobo specimen are similar Stål (Insecta, Homoptera, Cicadidae). – Archives Néer- to those of the male type specimens however. landaises de Zoologie 13, suppl. 1: 262–269. Boer, A.J. de, 1995a. The phylogeny and taxonomic sta- tus of the Chlorocystini (sensu stricto) (Homoptera, Distribution Tibicinidae). – Contributions to Zoology 65: 201–231. This species is described from Sayap and record- Boer, A.J. de, 1995b. The taxonomy and biogeography of ed from Gunung Monkobo, both in the Mount the genus Papuapsaltria n. gen. (Homoptera, Tibicini- Kinabalu massif of North Borneo. dae). – Tijdschrift voor Entomologie 138: 1–44. Boulard, M., 2008. Les cigales thaies. Liste actualisée in- cluant la description de deux nouveaux genres, de sept Acknowledgements espces nouvelles et les Cartes d’Identité Acoustique I am very grateful to Dr. Cong Wei (Shaanxi, China) (CIA) de Chremistica siamensis Bregman et de Lepto and his student Xu Wang for translating the Chi- psaltria samia (Walker) (Rhynchota, Cicadomorpha, nese text about the genus Katoa and its species from Cicadidae). – Ecole Pratique des Hautes Etudes. Biolo- Chou et al. The Cicadidae of China. I am very much gie et Evolution des Insectes 18: 1–112. indebted to the following curators for the loan and Chou, I., Z. Lei, L. Li, X. Lu & W. Yao, 1997. The Cicadi- dae of China (Homoptera: Cicadoidae). – Ilustrataj gift of material: Mrs Pudji Aswari (MZB), Dr G.V. Insect-Faunoj 2: 1-380, 1-16 pls. Gapud and Dr R.W. Brooks (SEM), Dr M. Go- Distant, W.L., 1888. Descriptions of new species of east- gala and Dr T. Trilar (PMSL), Mr B. Hubley and ern Cicadidae in the collection of the Museo Civico of Dr D.C.Darling (ROM), Mrs T. Kothe (ZSM), Dr Maryati Mohamed (UMS), Dr F. Penati (MSNG), Dr A. Soulier-Perkins (MNHN), Dr J. van Tol 1 * indicates that the publication was not seen and (RMNH), Dr Vásárhelyi (TMB), Mr M.D. Webb that the reference is quoted from Metcalf (1962). Downloaded from Brill.com10/05/2021 12:52:07PM via free access
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Bio- historic biogeography of the subtribe Cosmopsaltriaria diversity, biogeography, and bibliography of the cica- (Homoptera: Cicadidae). – Pacific Insects Monograph das of Thailand (Hemiptera: Cicadoidea: Cicadidae). 39: 1–127. – Zootaxa 1413: 1–46. Duffels, J.P., 2011. New genera and species of the tribe Schouten, M.A., J.P. Duffels & M.I. Zaidi, 2004. A check- Taphurini (Hemiptera, Cicadidae) from Sundaland. – list of the cicada fauna (Homoptera, Cicadoidea) of Deutsche Entomologische Zeitschrift 58: 77–104. Endau Rompin National Park, Malaysia, with descrip- Duffels, J.P., 2018. Revision of the cicada genus Dilobo tion of a new species. – Malayan Nature Journal 56: pyga (Hemiptera, Cicadidae ) from Sulawesi and the 369–386. Moluccas. – Zootaxa 4409: 1–172. Walker, F. 1856. Catalogue of the Homopterous insects Duffels, J.P. & P.A. van der Laan, 1985. Catalogue of the collected at Singapore and Malacca by Mr. A. R. Wal- Cicadoidae (Homoptera, Auchenorhyncha) 1956– lace, with descriptions of new species. – Journal of the 1980. – Series Entomologica 34; Dr. W. Junk, Dor- Proceedings of the Linnean Society London 1: 82–100, drecht, The Netherlands, pp. i–xiv, 1–414. pls 3–4. Duffels, J.P. & H. Turner, 2002. Cladistic analysis and Walker, F., 1858. List of the specimens of Homopterous biogeography of the cicadas of the Indo-Pacific sub- insects in the collection of the British Museum. Sup- tribe Cosmopsaltriina. – Systematic Entomology 27: plement. – Trustees British Museum, London, UK, 235–261. pp. 1–369. GEOnet Names Server of the U.S. Defense Mapping Zaidi, M.I. & S. Azman 2003. Cicada fauna (Homop- Agency (http://geonames.nga.mil/gns/html/). tera: Cicadoidea) of Sabah: with special reference to six Hayashi, M. & Y. Saisho, 2011. The Cicadidae of Japan. – selected protected areas. – Serangga 8: 95–106. Seibundo-shinkosha Co., Tokyo, Japan, 223 pp. (in Zaidi, M.I. & S. Azman, 2005. Cicada fauna of the Na- Japanese). tional Park of Peninsular Malaysia. – Serangga 10: Lee, Y.J., 2012. Descriptions of two genera and species of 49–58. Cicadidae (Hemiptera) from India with some notes Zaidi, M.I., S. Azman & M.Y. Ruslan 2001. Cicada on tribal classification. – Deutsche Entomologische (Homoptera, Cicadoidea) fauna of Langkawi island, Zeitschrift 59: 225–231. Kedah. – Serangga 6: 109–122. Lei, Z. & I. Chou, 1995. Two new species of the genus Zaidi, M.I., B.M. Noramly & M.Y. Ruslan, 2000. Cica- Katoa (Homoptera: Cicadidae). – Entomotaxonomia das (Homoptera: Cicadoidea) from Tibow, Sabah. – 17: 99–102. Serangga 5: 319–333. *Matsumura, S., 1913. A thousand insects of Japan. – Zaidi, M.I. & W. Nordin 2001. Cicadas of Crocker Range Additamenta 1: 1–184, pls i–xv. Park, Sabah (Homoptera, Cicadoidea), an additional Matsumura, S., 1927. New species of Cicadidae from the survey. – Serangga 6: 181–204. Japanese Empire. – Insecta Matsumurana 2: 46–58, Zaidi, M.I. & M.Y. 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