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1 Zootaxa 3936 (2): Copyright 2015 Magnolia Press Article ISSN (print edition) ZOOTAXA ISSN (online edition) Notes on Australian Laius Guérin-Méneville, Dicranolaius Champion and Intybia Pascoe with description of new species related to Dicranolaius c-purpureus (Lea) (Coleoptera: Melyridae: Malachiinae) ZHENHUA LIU 1, ADAM ŚLIPIŃSKI 2 & HONG PANG 1,3 1 State Key Laboratory of Biocontrol, Key Laboratory of Biodiversity Dynamics and Conservation of Guangdong Higher Education Institute, College of Ecology and Evolution, Sun Yat-Sen University, Guangzhou , China. F-mail: liuzhh22@mail2.sysu.edu.cn; Lsshpang@mail.sysu.edu.cn 2 Australian National Insect Collection, CSIRO National Research Collections Australia, GPO Box 1700, Canberra, ACT 2601, Australia. Adam.Slipinski@csiro.au 3 Corresponding author Abstract Remarks on Laius Guérin-Méneville and Dicranolaius Champion are provided. Laius falcifer Champion, 1921 is designated as the type species of Dicranolaius Champion, Dicranolaius bellulus is attributed to Boisduval (1835) not to Guérin-Méneville (1830). Dicranolaius c-purpureus (Lea) is redescribed and two closely related new species are described: D. weiri sp. n., and D. similis sp. n. Intybia Pascoe, 1866 is recorded from Australia for the first time and Laius filamentarius Lea, 1917 is regarded as member of Intybia comb. n.. Key words: Coleoptera, Melyridae, Malachiinae, Laius, Dicranolaius, Intybia, Australia, taxonomy, new species Introduction The flower beetles of the subfamily Malachiinae are usually easily recognised among other Australian beetles due to their brightly coloured surfaces and lateral eversible vesicles that become inflated and brightly red when the beetle is disturbed (Lawrence and Ślipiński, 2013). The Malachiinae have been popular subject of taxonomic research in the last 100 years, mostly due to the concerted efforts of Walter Wittmer and Alfons Evers who described a large number of genera and species from all over the world. The Australian Melyridae were extensively studied only at the beginning of the 20 th Century by Arthur M. Lea who described a large number of new species and reviewed the taxa described earlier by Thomas Blackburn and others. He and his British contemporary, George Champion provided detailed illustrations of diagnostic characters located on male antennomeres in Laius Guérin-Méneville and related taxa. Champion (1921a, b) reviewed Laius species at the British Museum and established the subgenus Dicranolaius to include four Asian species with males bearing modified basal antennomeres and fore tarsomere 2 projecting and bearing apical comb. Wittmer (1952) reviewed Australian Laius species and concluded that only L. filamentarius Lea (and possibly L. minutus Lea) belonged to that genus. He described two new genera (Troglolaius and Flabellolaius) for a few Laius species bearing peculiar characters on male pronotum and antennae but transferred the bulk of the species to elevated to the genus level Dicranolaius Champion based on the observation by Lea that the male protarsomere 2 was of a peculiar shape and tipped with black (Lea, 1909: 151). Champion (1921a, b) included Intybia Pascoe (1866) as a synonym or subgenus of Laius, which was followed until Evers (1994) revalidated Intybia as independent genus. Evers concepts of Laius, Intybia and Dicranolaius have been accepted by Wittmer (1995) and others, e.g., Yoshitomi (2008, 2014), Asano & Kawashima (2010), Yoshitomi and Lee (2010), Asano (2012) and Plonski (2013), resulting in transfer of most Laius species to Intybia. Laius contains 35 species of uniformly bluish melyrids with modified male protibia confined to the rocky 272 Accepted by R. Gerstmeier: 19 Feb. 2015; published: 19 Mar. 2015

2 seashores of the Pacific Ocean, but the distinctiveness of Intybia and Dicranolaius is less certain as it is based solely on the presence of the tarsal comb in Dicranolaius that could have been reduced independently many times. The present paper is the first of a planned series of papers revising Australian Malachiinae and provides nomenclatorial framework for further research on the Australian taxa related to Laius and currently placed in the tribe Apalochrini (Mayor, 2007). In this paper we are still following the generic concepts outlined by Evers (1994) but are developing a molecular dataset to test the generic limits of the Australian malachiines in a near future. Material and methods The terms used in morphological descriptions follow Lawrence at al. (2011) and Lawrence & Ślipiński (2013). Measurements were made as follows: (TL) body length apical edge of clypeus to apex of elytra; (BW) body width (also as elytral width) elytra at widest point; (PL) pronotal length median line from anterior margin to posterior margin; (PW) pronotal width across widest point; (EL) elytral length base of scutellum to elytral apex along suture; (PL) penis length from anterior to the posterior margin of penis. Habitus and individual structures in glycerol were photographed using a Zeiss AxioCam HRc mounted on a Zeiss Discovery V20 microscope. Layers were then aligned and stacked in Zeiss software to obtain a fully sharpened image. Entire beetles or abdomens of selected males and females were cleared in 10% KOH, dissected, examined and photographed in open glycerol slides. The following acronyms have been used for institutions, which have provided specimens for study: ANIC: Australian National Insect Collection, CSIRO, Canberra, Australia; BMNH: The Natural History Museum, London, United Kingdom; QMB: Queensland Museum, Brisbane, Australia; SAMA: South Australian Museum, Adelaide, Australia. Taxonomy Laius Guérin-Méneville Laius Guérin-Méneville, 1830: Plate 2, fig. 10. Type species, by monotypy: Laius cyaneus Guérin-Méneville, Guérin- Méneville, 1838: 78. Remarks. The Zoology part of the Coquille voyage book series was printed in numerous fascicles (livraisons) over many years and in spite of the 1830 date printed on the front page, the text of Insecta by Guérin-Méneville was printed only in 1838 (Cretella 2010). However, the livraison No. 19 with plate 2 illustrating this species was published in 1830 (Cretella 2010) validating the joint description of genus and species Laius cyaneus. Included species. Four Australian species can be classified in Laius: Laius alleni Lea, 1909: 161 (Queensland, Cairns; SAMA, ANIC); Laius purpureipennis Lea, 1916: 321 (Northern Territory; SAMA, ANIC); Laius tibialis Gahan, 1900: 102 (Christmas Island; BMNH, ANIC); Laius triangulatus Wittmer, 1954: 305 (Western Australia, Baudin Point; BMNH). Intybia Pascoe Intybia Pascoe, 1866: 448. Type species, by monotypy: Intybia guttata Pascoe, Remarks. In addition to the species included below, some species currently classified in Dicranolaius will certainly be transferred to Intybia after detailed examinations of the type specimens and additional material. Laius filamentarius Lea was thought to be a true Laius by Wittmer (1952) however, according to the current generic concepts (Evers 1994) this species should be classified in Intybia based on the modified basal antennomeres in male, but with fore femur, tibia and tarsi simple without secondary sexual characters. ON AUSTRALIAN MALACHIINAE Zootaxa 3936 (2) 2015 Magnolia Press 273

3 Intybia filamentaria (Lea) comb. n. Laius filamentarius Lea, 1917: 136. Wittmer, 1952: 187. Remarks. This is a very distinctive species with male antennomere 3 strongly expanded and bearing two filamentous processes anteriorly. Numerous syntypes from Queensland, Emerald (SAMA, ANIC) have been examined. Dicranolaius Champion Dicranolaius Champion, 1921b: 195. Type species, here designated, Laius falcifer Champion, 1921b: 204. Remarks. Champion (1921b) included four species in the subgenus Dicranolaius (L. falcifer Champion, 1921b, L. tetrastictus Champion, 1921a, L. flavonotatus Champion, 1921b and L. erythrocephalus Champion 1921b. Of these four L. falcifer appeared to be the best candidate to be designated as the type species of the genus. The other three have been thought to be unfit because: L. tetrastictus was originally based on female only, L. flavonotatus is a junior homonym (see below) and the holotype of L. erythrocephalus is a teneral specimen. Dicranolaius bellulus (Boisduval) Malachius bellulus Boisduval, 1835: 135. Malachius bellulus Guérin-Méneville, 1838: 78. Remarks. This common Australian species has been credited to Guérin-Méneville (1830) but the description by Boisduval (1835) should be given precedence due to the printing delay of the Insecta part by Guérin-Méneville to 1838 (Cretella 2010). Interestingly Boisduval cites Guérin-Méneville as the author of the species with correct page (78). Dicranolaius c-pupureus (Lea) (Figures 1E, 1F, 2B, 2H, 2I, 3C, 3I) Laius c-purpureus Lea, 1914: 220. Dicranolaius c-purpureus: Wittmer, 1952: 188. Description. Length mm, PL/PW , EL/EW , HW/PW , EL/PL Male. Head bicoloured, labrum, clypeus and anterior part of frons yellow or orange, remaining part of frons from middle of eyes and vertex black; pronotum entirely yellow or orange; scutellum black; elytra yellow or orange with dark blue slightly iridescent humeral area and C-shaped preapical marking not reaching suture (Figs 1E, 1F); antennomeres 1 3 yellow (Fig. 2H), remaining segments dark orange and brown; fore femur yellow or orange, protibia yellow or brown, tarsi orange; mid and hind legs brown or dark blue. Ventral side: prothorax yellow or orange, meso- and metathorax dark; abdomen yellow with the apex of terminal segment dark. Vestiture double of short and dense yellow or orange setae and sparse and twice longer bristles, which are pale on yellow or orange parts of elytra and darker on blue spots. Head concave on vertex, covered with dense white setae and sparse yellow and dark bristles, very finely punctured. Antennal scape expanded, flat; antennomere 3 with two dorsal pointed processes on outer margin (Fig. 2I). Pronotum broader than head, widest at anterior third. Lateral margins and base with narrow bead; disk shiny, not distinctly punctured. Scutellum truncate apically with dense silver adpressed setae. Elytra widest at apical third; sides gradually expanded posterolaterally; surfaces densely punctured, punctures about twice as large as eye facets, 0.5 diameter apart. Fore femur with hump-like projection in middle of upper surface (Fig. 2B), second segment of fore tarsi prominent with dense black teeth along anterior edge. 274 Zootaxa 3936 (2) 2015 Magnolia Press LIU ET AL.

4 FIGURE 1. Dicranolaius species, dorsal and lateral habitus. (A, B) Dicranolaius weiri sp. n.; (C, D) Dicranolaius similis sp. n.; (E, F) Dicranolaius c-pupureus (Lea). ON AUSTRALIAN MALACHIINAE Zootaxa 3936 (2) 2015 Magnolia Press 275

5 FIGURE 2. Dicranolaius, males; A C Foreleg; D I antenna and antennomeres 1 4. (A, D E) Dicranolaius weiri sp. n.; (C, F G) Dicranolaius similis sp. n.; (B, H I) Dicranolaius c-pupureus (Lea). 276 Zootaxa 3936 (2) 2015 Magnolia Press LIU ET AL.

6 FIGURE 3. Dicranolaius, males; A C Penis; D F Tergite VIII, dorsal; G I Sternite VIII, ventral (A, D, G) Dicranolaius weiri sp. n.; (B, E, H) Dicranolaius similis sp. n.; (C, F, I) Dicranolaius c-pupureus (Lea). Tergite VIII with deep incision apically and a long median spur (Fig. 3F); sternite VIII as in Figure 3I. Aedeagus (Fig. 3C) with strong longitudinal sclerite and numerous spines in endophallus. Female. Similar to male in colour. Antennae not modified; frons and vertex flat; fore femora simple, second segment of fore tarsi normal. ON AUSTRALIAN MALACHIINAE Zootaxa 3936 (2) 2015 Magnolia Press 277

7 Types. Laius C-purpureus Lea Queensland Cotype/ Carins N. Queensland Hacker/ Paratype/ On permanent loan from Macleay Museum University of Sydney (2, ANIC). Other specimens examined. Queensland: Burdekin R., Selheim, 14.i.1964, G. Monteith (5 males; QMB); Dotswood Stn, 'E 19 45'S Reelborrom Creek, 10.v R Parrerson. A198/ on foliage of Myrtecea melaleuca (1 male; ANIC; dissection). Northern Territory: McArthur River 16.47S E, 14km S by W of Cape Crawford, 25.x.1975, M.S. Upton (1 male, 1 female; ANIC); 14.30S E, 3km SSW of Katherine,12.xi.1979, T. Weir (2 females; ANIC); McArthur River, 16.05S E, 2km SSE of Borroloola, iv.1976, J.E. Feehan (1 male; ANIC; dissection); Delamere, v.1968, M. Mendum (2 females; ANIC); 15.58S E 12km NNE of Borroloola, 1.xi.1975, M.S. Upton (1 male; ANIC); Goose Lagoon 16.10S E 11km SW by S of Borroloola, 31.x. 1975, M.S. Upton (1 male; ANIC); no locality, J.G. Brooks Bequest, 1976 (1 male; ANIC). Distribution. North Queensland and Northern Territory. Dicranolaius weiri sp. n. (Figures 1A, 1B, 2A, 2D, 2E, 3A, 3D, 3G) Etymology. Dedicated to Tom Weir of the Australian National Insect Collection, who has collected the holotype. Diagnosis. This species is externally identical to D. c-purpureus and can be reliably distinguished only by male genitalia and the sexual characters on antennae and fore femora. Females of D. weiri can usually be separated from those of D. c-purpureus in having the C-shaped apical elytral mark occupying longer stretch of lateral edge (Fig. 1A), while in most specimens of D. c-purpureus the mark only touches the edge posteriorly (Fig. 1C). Description. Length mm. PL/PW , EL/EW , HW/PW , EL/PL Male. Head bicoloured with labrum, clypeus and anterior part of frons yellow or orange, remaining part of frons from middle of eyes and vertex bluish or black; prothorax uniformly yellow or orange; elytra orange or yellow with dark blue slightly iridescent humeral area and C-shaped preapical marking (Figs 1A, 1B); antennomeres 1 4 yellow, remaining segments dark brown or black; fore leg yellowish with dark brown tibia, mid and hind legs dark blue. Meso- and metaventrite dark, abdomen yellow with two terminal segments dark blue or black. Vestiture double, consisting of short and moderately dense yellow erect or semierect setae (adpressed on scutellum and part of head) and sparser and distinctly longer bristles, pale on yellow parts of elytra and usually darker on darker spots. Head finely punctured with vertex weakly concave, covered with dense white adpressed setae and sparser bristles. Antenna with strongly modified basal segments (Figs 2D, 2E), scape slightly expanded; antennomere 3 expanded with dorsal pointed process on outer margin (Fig. 2E). Pronotum broader than head, widest at anterior third; lateral margins and pronotal base with narrow bead; disk smooth and shiny, sparsely setose. Scutellum truncate apically, covered with dense whitish adpressed setae. Elytra widest at apical third, sides gradually expanded posterolaterally; surfaces densely punctured, punctures about twice as large as eye facets, puncture diameter apart. Fore femur with deep excavation on dorsal outer surface as in Figure 2A. Fore tarsus with tarsomere 2 projecting over 3, bearing black comb composed of dense teeth. Tergite VIII (Fig. 3D) with deep incision apically and short median spur; sternite VIII as in Figure 3G. Aedeagus (Fig. 3A); endophallus with numerous smaller sclerites and spines. Female. Identical to male but with frons flat or weakly convex, antennomeres 1 and 3 not modified and fore leg with femur simple and tarsomere 2 without comb. Types. Holotype male, Northern Territory: 14.04S E, Fergusson R., 31 km SE by S of Pine Creek, N.T. 14 Nov. 1979, T. Weir (ANIC). Paratypes: same data as the holotype (2, males, 5, females; 5, ANIC; 2, SYSU; 1 dissection); Tindal, N. T S E, 1 20 Dec.1967, light trap, W.J.M. Vestjens (1 male, ANIC, dissection); 12.06S E, Cooper Creek, N.T. 19 km E. by S of Mt. Borradaile 31.v.73 at light, E.G. Matthews (1 male, ANIC); Katherine, 20 Nov. 1974, on Eucalyptus sp., E.M. Exley & R.I. Storey (1 male, QMB). Distribution. Northern Territory. 278 Zootaxa 3936 (2) 2015 Magnolia Press LIU ET AL.

8 Dicranolaius similis sp. n. (Figures 1C, 1D, 2C, 2F, 2G, 3B, 3E, 3H) Etymology. Species name has been derived from the Latin adjective similis, as this new species is very similar to both D. c-purpureus and D. weiri. Diagnosis. This species differs from D. c-purpureus and D. weiri in being somewhat shorter with black head, the dorsal bristles longer and the elytral markings more extensive at base. However, the male dimorphic characters and the male terminalia are the best distinguishing features. Description. Length mm, PL/PW , EL/EW , HW/PW , EL/PL Male. Head and scutellum black, prothorax uniformly yellow or orange, elytra orange or yellow with dark blue slightly iridescent humeral area and C-shaped preapical marking with slightly lighter centre (Figs 1C, 1D). Antennomeres 1 3 orange, remaining segments dark brown or black (Fig. 2F); fore leg black with dark brown tarsus, mid and hind legs black. Meso- and metaventrites dark, abdomen yellow with terminal segment dark. Vestiture double, consisting of short and dense yellow setae and sparse, longer and thicker black bristles. Head with dense white setae and sparse black bristles, very finely punctured. Antennal scape expanded and triangular (Fig. 2F); antennomere 3 expanded and flat (Fig. 2G). Pronotum broader than head, widest at anterior third; lateral margins and pronotal base with narrow bead, posterior edge more or less emarginate medially; pronotal disk without apparent punctures, shiny. Scutellum truncate, apically with dense silver adpressed setae. Elytra widest at apical third; sides gradually expanded posterolaterally; surface densely and weakly punctured, punctures about the same size as eye facets, separated by 1 diameter. Fore leg with dense white and yellow adpressed setae and few black bristles on tibiae, femur with deep excavation medially on upper surface as in Figure 2C, tarsomere 2 elongate with dense comb along the anterior edge; mid and hind legs with dense white and yellow adpressed setae and few black bristles on femora and tibiae. Tergite VIII with deep incision medially without spine (Fig. 3E); sternite VIII divided (Fig. 3H). Aedeagus (Fig. 3B) with very dense spines in endophallus. Female. Unknown. Types. Holotype male, Queensland: Camooweal, Q S E 16.v.68. K. Armstrong (ANIC). Paratype: Katherine, N.T., , E. Matthews (1, male; ANIC, dissected). Distribution. Queensland and Northern Territory. Acknowledgments The authors acknowledge CSIRO, Australia and Sun Yat-sen University, China for support of research. This project was also supported by the National Natural Science Foundation of China Grants No , ) and key project of Science-technology basic condition platform from The Ministry of Science and Technology of the People s Republic of China (Grant No. 2005DKA21402). We are grateful to the following curators and the institutions for providing us with a material under their care, Geoff Monteith and Susan Wright (QMB); Roger Booth (BMNH) and Peter Hudson (SAMA). We thank John F. Lawrence, Adriean Mayor and Roland Gerstmeier for critical review of the manuscript. Cate Lemann (CSIRO) is sincerely acknowledged for technical assistance. References Asano, M. (2012) Taxonomic notes on the genera Dicranolaius and Intybia of the Borneo Island, with description of new species (Coleoptera, Malachidae). Japanese Journal of Systematic Entomology, 18 (2), Asano, M. & Kawashima, I. (2010) Discovery of the genus Dicranolaius (Coleoptera, Malachidae) from Japan with description of a new species. Japanese Journal of Systematic Entomology, 12 (2), Boisduval, J.B. (1835) Faune entomologique de l'océan pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage. Deuxième partie. Coléoptères et autres ordres. In: Voyage de découvertes de L'Astrolabe exécuté par ordre du Roi, pendant les années , sous le commandement de M.J. Dumont d'urville. J. Tastu, Paris, pp. 1 vii ON AUSTRALIAN MALACHIINAE Zootaxa 3936 (2) 2015 Magnolia Press 279

9 Cretella, M. (2010) The complete collation and dating of the section Zoologie of the Coquille voyage. Bollettino Malacologico, 46 (2), Gahan, C.J. (1900) Order 4. - Coleoptera. New species descriptions. In: Waterhouse, C.O., Gahan, C.J. & Arrow, G.J. (Eds.), A Monograph of Christmas Island (Indian Ocean): physical features and geology; with descriptions of the fauna and flora by numerous contributors. Printed by order of the Trusteespp, London, pp. i xiii Guérin-Méneville, F.É. ([1830]1838) Insectes. In: Duperrey, L.I. (Ed.), Voyage autour du monde, Exécuté par Ordre du Roi, sur La Corvette de Sa Majesté, LaCoquille, pendant les années 1822, 1823, 1824 et 1825, Sous le Ministère et conformément aux Instructions de S.E.M. le Marquis de Clermont-Tonnerre, Ministre de la Marine; Et publié sous les auspices de son Excellence Mgr le Cte de Chabrol, Ministre de la Marine et des Colonies, par M.L.I. Duperrey, Capitaine de Frégate, Chevalier de Saint-Louis et Membre de la Légion d Honneur, Commandant de l Expedition. Zoologie, par M. Lesson. Tome Second. 2e Partie. Arthus Bertrand, Libraire-Editeur, Imprimerie de Firmin Didot, In-4, Paris, pp. i-xii, 9-319, [plates 1 and 2 of Insectes published in 1830; text in 1838] Lawrence, J.F. & Ślipiński, A. (2013) Australian Beetles. Vol. 1: Morphology, Classification and Keys. CSIRO Publishing, Collingwood, Victoria, viii pp. Lea, A.M. (1909) Revision of the Australian and Tasmanian Malacodemidae. Transactions of the Entomological Society of London, 1909, , pls. II VI. Lea, A.M. (1914) On Australian and Tasmanian Coleoptera with descriptions of new species. Part II. Proceedings of the Royal Society of Victoria, New Series, 26, Lea, A.M. (1916) Notes on some miscellaneous Coleoptera with descriptions of new species. Part II. Transactions of the Royal Society of South Australia, 40, Lea, A.M. (1917) Notes on some miscellaneous Coleoptera, with descriptions of new species. Part III. Transactions of the Royal Society of South Australia, 41, pls. XII XV. Mayor, A. (2007) Malachiidae. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 4. Elateroidea Derodontoidea Bostrichoidea Lymexyloidea Cleroidea Cucujoidea. Apollo Books, Stenstrup, pp , [total page number 935 pp.] Plonski, I. (2013) Studies on the genus Intybia Pascoe (Coleoptera: Malachiidae). I. Some nomenclatorial acts and faunistic records. Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen, 65, Wittmer, W. (1952) Remarks on some Australian Laius Guer. (Col: Malachiidae). Proceedings of The Linnaean Society of New South Wales, 76, Wittmer, W. (1954) Zur Kenntnis der Malachiidae (Col.) Australiens. Annals and Magazine of Natural History, Series 12, 7, Wittmer, W. (1957) Eine neue Gattung aus der familie Malachiidae (Col.) aus Australien. Annals and Magazine of Natural History, Series 12, 10, Wittmer, W. (1995) Zur Kenntnis der Familie Malachiidae (Coleoptera) 2. Beitrag. Entomologica Basiliensia, 18, Yoshitomi, H. (2008) Contribution to the taxonomy of the genus Laius Guérin-Méneville in Indonesia, with description of new species (Coleoptera: Malachiidae). Koleopterologische Rundschau, 78, Yoshitomi, H. (2014) Comparative morphology of the endophallic structures of the genus Laius (Coleoptera, Melyridae), with the descriptions of three new species. European Journal of Taxonomy, 97, Yoshitomi, H. & Lee, C. (2010) Revision of the Taiwanese and Japanese Species of the Genus Laius (Insecta: Coleoptera: Malachiidae). Zoological Studies, 49 (4), Zootaxa 3936 (2) 2015 Magnolia Press LIU ET AL.