The Cicada Genus Muda Distant (Hemiptera: Cicadidae) from Sundaland: Species and Relationships J.P
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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 <UN> 132 Tijdschrift voor Entomologie, volume 161, 2018 to the genus Muda: Baeturia obtusa (Walker, 1858) MZB Museum Zoologicum Bogoriense, from Java and Baeturia beccarii Distant, 1888 from Cibinong Sumatra, while he listed Muda concolor as a junior NMWC National Museum of Wales, Cardiff synonym of Muda obtusa. In his ‘Monograph of PSML Slovenian Museum of Natural History, Oriental Cicadidae’ (Distant 1892b: 153) and in his Ljubljana catalogue of 1906, Distant mentioned Cicada vir- RMNH Naturalis Biodiversity Center (former guncula Walker, 1856 as one of the species of which Rijksmuseum voor Natuurlijke Histo- he could not find the type specimen in the collection rie), Leiden of the British Museum (Natural History). ROM Royal Ontario Museum, Toronto In his classic work on the cicadas of Sundaland, SEM Snow Entomological Museum, Law- Moulton (1923) recognized two species in the ge- rence, Kansas nus Muda, M. obtusa and M. virguncula, and had TMB Magyar Természettudományi Múzeum, recovered the type specimen of Cicada virguncula in Budapest the BMNH collection, transferred C. virguncula to UMS University of Malaysia Sabah, Kota Muda and also synonymized Muda beccarii with M. Kinabalu, Malaysia virguncula. Moulton (1923) also followed Distant ZMAN Naturalis Biodiversity Center (former (1906) in regarding Muda concolor as a junior syn- Zoölogisch Museum, Universiteit van onym of Muda obtusa. The two species, M. obtusa Amsterdam), Leiden and M. virguncula, are uniform yellowish brown to ZSM Zoologische Staatssammlung, München brown cicadas lacking dark brown or black markings on the head and thorax and often confused in the Data on the distribution of the species were derived literature. Moulton (1923) distinguished these two from the author’s ‘Biodiversity Database of the Ci- Muda species by two characters: (1) the veins of the cadas of South East Asia and the West Pacific’, and ulnar cells of the tegmina running to the radial cell plotted on maps of ADC-Worldmap version 2.0 are angulated in M. obtusa and nearly straight in vol. 4 Southern Asia & Australia with the program M. virguncula, and (2) the upper vein of the lowest MapInfo Professional 11.0. The localities and other cubital cell, which is not convex in M. obtusa and data from the specimen labels in the database are slightly convex in M. virguncula. The present study filed in the program FileMaker Pro 11. Information reveals that these characters are unreliable. After regarding the geographical coordinates has been re- comparison of the types and other material of the trieved from the following sources: Global Gazetteer, two species, it appeared that the two Muda species Google, Atlas van Tropisch Nederland (Anonymous can be separated by the number of timbal ribs, the 1938) and The Times Comprehensive Atlas of the number of apical areas in the wings and some very World (Anonymous 1999). distinctive features of the male genitalia. The terminology adopted in this paper for exter- In recent years, two species were added to Muda: nal features of the body and for the male genitalia the new species M. tua Duffels from the Malay follows that of Duffels (Duffels 1977, 1983; Duffels Peninsula, described in Schouten et al. (2004), and & Turner 2002) and Moulds (2005). M. kuroiwae from the Ryukyu Islands of southern Japan, which was transferred by De Boer (1995a) from Baeturia to Muda, a taxonomic position that Taxonomy was recently confirmed byHayashi & Saisho (2011). Genus MUDA Distant 1897 Muda Distant 1897: 384. Type-species by monoty- py: Muda concolor Distant, 1897 (= Muda virgun- Material and methods cula (Walker, 1856)) The institutions listed below are the depositories of Muda: Distant 1906: 153, 156; Moulton 1923: 156, the material studied. The abbreviations have been 158; De Boer 1995a: 202–204; Hayashi & Saisho used in the lists of material and throughout the text. 2011: 196. Iwasemia Matsumura, 1927: 57. BMNH Natural History Museum, London (for- Nahasemia Matsumura, 1930: 9. mer British Museum (Natural History)) For further references see: Metcalf (1963), Duffels & BPBM Bernice P. Bishop Museum, Honolulu Van der Laan (1985) and Sanborn (2014). MNHN Muséum National d’Histoire Naturelle, Paris MNKM Muzium Negara Malaysia. Kuala Diagnosis Lumpur Body length males: 11–18 mm, females 12–18 mm. MSNG Museo Civico di Storia Naturale ‘G. Ventral side of post- and anteclypeus with sharp me- Doria’, Genova dian keel, which is triangular in cross-section, and Downloaded from Brill.com10/05/2021 12:52:07PM via free access <UN> Duffels: The cicada genus Muda from Sundaland 133 a distinct median groove running from anterior Muda plus Prasiini forming the sistergroup of the postclypeal margin to clypeal suture. Body yellow- Chlorocystini. According to De Boer (l.c.), Muda ish brown to brown either with, or without, brown has at least one character in common with Prasi- to black marking. Head 0.78–0.96 times as wide ini and part of the Chlorocystini: the postclypeus, as pronotum collar. Pronotum with lateral sides ei- which is triangular in cross-section. In his paper ther parallel or slightly converging to the anterior. on the higher classification of the cicadas, Moulds Tegmina and wings hyaline. Tegmina with 8 apical (2005) mentioned this postclypeus character and areas, wings with 5 or 6 apical areas. Operculum of the ill-defined basal lobes of the pygofer, which are male and female very short. Timbals with 5–9 trans- confluent with the lateral pygofer margins, as syn- verse ribs and short intercalary ribs; timbals absent in apomorphies for Prasiini, Parnisini and Chlorocys- M. kinabaluana, a convex sklerite replacing the tim- tini (= clade 9 in Moulds’ Fig. 61). Moulds (2005) bal. Male abdomen more or less inflated; abdomi- did not study Katoa or Muda, but my present study nal tergites 2–8, distinctly arched medially in both of these genera reveals that the two synapomophies males and females. Male sternite 8 large, 1.5–2 times for Prasiini, Parnisini and Chlorocystini mentioned as long as wide. Male pygofer very slender, parallel- by Moulds (2005) are also present in Muda and sided, 2.5–3 times as long as broad and with a me- Katoa. diodorsal carina along its whole length, but in M. Recently, Lee (2012) transferred Katoa from the tua twice as long as broad and without carina. Basal tribe Tibicinini in the subfamily Tibicininae to the pygofer lobes very long, reaching proximal ends of tribe Cicadettini in the subfamily Cicadettinae be- upper pygofer lobes.