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Discovery of Eutheiini (Coleoptera: Staphylinidae: Scydmaeninae) in Australia, with Implications for Phylogeny and Biogeography of Paraneseuthia

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Discovery of Eutheiini (Coleoptera: Staphylinidae: Scydmaeninae) in Australia, with Implications for Phylogeny and Biogeography of Paraneseuthia Eur. J. Entomol. 108: 687–696, 2011 http://www.eje.cz/scripts/viewabstract.php?abstract=1668 ISSN 1210-5759 (print), 1802-8829 (online) Discovery of Eutheiini (Coleoptera: Staphylinidae: Scydmaeninae) in Australia, with implications for phylogeny and biogeography of Paraneseuthia PAWEà JAàOSZYēSKI Museum of Natural History, Wrocáaw University, Sienkiewicza 21, 50-335 Wrocáaw, Poland; e-mail: [email protected] Key words. Coleoptera, Staphylinidae, Scydmaeninae, Eutheiini, Paraneseuthia, Oriental, Australia, taxonomy, phylogeny, biogeography Abstract. The scydmaenine tribe Eutheiini is recorded from Australia for the first time. Paraneseuthia carltoni sp. n. and P. booloumba sp. n. are described and illustrated, both from Queensland. In a parsimony-based phylogenetic analysis using adult mor- phological characters including genital features, the Australian species together with the Melanesian type species of Paraneseuthia Franz, P. peckorum Franz, were found to be more closely related to East Palearctic congeners than to most of the Paraneseuthia in the Sunda-Papuan area. The topology of the tree and biogeographic data suggest a Sundaland origin of this genus, with three major dispersal routes from a center located in present-day Sumatra: (i) north-eastern colonization of the Palearctic Far East, via a conti- nental or island-arc route; (ii) south-eastern dispersal to East Australia; and (iii) eastern dispersal to Melanesia, possibly via the Qua- ternary Outer-Melanesian Arc. The important role of dispersal in the evolution of Paraneseuthia is supported by the presence of this genus on isolated volcanic islands, such as the southern Moluccas and Fiji, which were never connected to larger land masses. INTRODUCTION known from females only and therefore undescribed. A Paraneseuthia Franz, 1986 is one of the most enigmatic survey of museum collections revealed two highly intri- genera within the tribe Eutheiini. Originally not explicitly guing, relatively flat and elongate specimens of Eutheiini placed in any tribe (Franz, 1986), later treated as a from Queensland, Australia. The tribe Eutheiini was pre- member of the Cephenniini (Newton & Franz, 1998) and viously known predominantly from the Palearctic and Nearctic regions, with only a few species reaching SE finally transferred to the Eutheiini (JaáoszyĔski & Hosh- ina, 2004), this genus initially was known only from Asia, Melanesia and Mesoamerica (e.g., Franz & Löbl, Melanesia and the Russian Far East (Franz, 1986; Kurba- 1990; Newton & Franz, 1998; O’Keefe, 1999; JaáoszyĔ- tov, 1990, 1991). This disjunct distribution turned out to ski, 2003). The discovery of this tribe on the Australian be a result of an inadequate study of Scydmaeninae and a mainland significantly fills a gap in the general distribu- general rareness of Paraneseuthia. More detailed recent tion of Eutheiini. Detailed examination revealed that both work has resulted in transferring some previously mis- Australian specimens not only belong to Paraneseuthia, but also their aedeagi show surprising similarities, not placed Japanese species to this genus (JaáoszyĔski & Hoshina, 2004) and in descriptions of new species from with those of their Sunda-Papuan congeners, but rather Sumatra, Borneo, West Papua and East New Guinea the Palearctic Far Eastern species. This observation prompted the present study, in which results of a prelimi- (JaáoszyĔski, 2008a, 2009, 2010). Access to new material made it possible to describe the morphology of Parane- nary phylogenetic analysis of species of Paraneseuthia seuthia in detail and confirm that north-eastern species and biogeographic data are combined to propose a hypothesis concerning the evolution and dispersal routes are truly congeneric with Sunda-Papuan taxa (JaáoszyĔ- ski, 2010). The most unusual characters of this genus are of this unusual genus. the maxillary palpus composed of only three palpomeres MATERIAL AND METHODS and a relatively complicated aedeagus in some species. Specimen handling, imaging and measurements Also characteristic of Paraneseuthia are the relatively stout and strongly convex body, elytra more rounded than Specimens used in this study (Table 1) are deposited in the truncated and exposing only a part of the pygidium, rudi- following collections: BM – Bishop Museum, Honolulu, Hawai; MHNG – Muséum d’Histoire Naturelle, Geneva, Switzerland; mentary basal elytral foveae, very high (i.e., strongly pro- NHMW – Naturhistorisches Museum Wien, Vienna, Austria; jecting ventrad) keel on the mesoventrite and lack of an PCPJ – private collection of the author, Wrocáaw, Poland; occipital constriction. Some of these characters, very PCSK – private collection of Sergei Kurbatov, Moscow, Russia; unusual for the Eutheiini, are shared only with a single SEMC – University of Kansas, Natural History Museum and genus of this tribe, Euthiconus Reitter, 1881. Biodiversity Research Center (Snow Entomological Collec- Recent studies yielded a number of specimens from tions), USA; SMNS – Staatliches Museum für Naturkunde, islands located east of the Wallace’s line, including the Stuttgart, Germany. Moluku Archipelago and New Guinea (JaáoszyĔski, Dry-mounted specimens were relaxed in warm water and dis- 2010). Unfortunately, most species from that area are sected; details of the morphology of insects permanently mounted in Canada balsam were studied under light stereo- 687 TABLE 1. List of taxa examined and depositories of the specimens. Species Origin Depository Eutheia scydmaenoides Stephens, 1830 Poland PCPJ Euthiconus conicicollis (Fairmaire et Laboulbène, 1855) Poland PCPJ Paraneseuthia bicolor JaáoszyĔski, 2010 Indonesia: Sumatra MHNG Paraneseuthia booloumba sp. n. Australia: Queensland SEMC Paraneseuthia carltoni sp. n. Australia: Queensland SEMC Paraneseuthia devia JaáoszyĔski, 2008 Indonesia, West Papua BM Paraneseuthia guineana JaáoszyĔski, 2009 Indonesia, West Papua SMNS Paraneseuthia holzneri (Franz, 1976) Japan: Honshu NHMW Paraneseuthia inexpectata JaáoszyĔski, 2006 Japan: Honshu PCPJ Paraneseuthia levigata JaáoszyĔski, 2010 Papua New Guinea: Madang SEMC Paraneseuthia paradoxa (K. Sawada, 1962) Japan: Honshu PCPJ Paraneseuthia peckorum Franz, 1986 Fiji NHMW Paraneseuthia quadrifoveata JaáoszyĔski, 2010 E Malaysia: Sabah MHNG Paraneseuthia saga Kurbatov, 1991 Russia: Kunashir PCSK Paraneseuthia spinosa JaáoszyĔski, 2010 Indonesia: Sumatra MHNG Paraneseuthia trepida Kurbatov, 1990 Russia: Primorsky Krai PCSK Paraneseuthia sp. 1 Japan: Iriomote Is. PCPJ Paraneseuthia sp. 2 Papua New Guinea: Morobe SEMC Paraneseuthia sp. 3 Papua New Guinea: Morobe SEMC Paraneseuthia sp. 4 E Malaysia: Sabah MHNG Paraneseuthia sp. 5 Indonesia: Moluccas (Yamdena) MHNG Paraneseuthia sp. 6 Indonesia: Moluccas (Kai Besar) MHNG Paraneseuthia sp. 7 Indonesia: Moluccas (Kai Besar) MHNG Veraphis japonicus (K. Sawada, 1962) Japan: Honshu PCPJ scopic and compound microscopes. Habitus images were taken genera of Eutheiini in detail – Eutheimorphus Franz & Löbl, using a Nikon Coolpix 4500 camera mounted on a Nikon 1990 and Paeneutheia JaáoszyĔski, 2003 (both from Borneo) are Eclipse 1500 stereoscopic microscope; image stacks were pro- known from minute and fragile types only and the Italian cessed using Combine ZP (Hadley, 2010). The measurements endemic Euthiopsis Müller, 1925 is very rare and no specimens and abbreviations used in the text are as follows: body length were available for dissection. (BL) is the sum of the lengths of the head, pronotum and elytra Phylogenetic analysis was based on 40 parsimony- measured separately; length of head (LH) was measured from a informative, non-additive adult morphological characters; inap- hypothetical line joining posterior margins of eyes to anterior plicable characters were assigned a gap value (“–”) and treated margin of the clypeus; width of head (HW) includes eyes; equivalent to missing data (“?”). Data matrix was assembled in length of antennae (AnL) was measured in ventral view; length Nexus Data Editor for Windows v. 0.5.0 (Page, 2001); parsi- of pronotum (PL) was measured along midline; width of pro- mony analysis was conducted in TNT (Goloboff et al., 2008) notum (PW) is its maximum width; length of elytra (EL) was under equal weights using the implicit enumeration strategy; the measured along suture; width of elytra (EW) is its maximum analysis was rooted to Eutheia. The standard bootstrapping width, combined; elytral index (EI) is length divided by com- (1,000 replicates) was conducted also in TNT. Trees were dis- bined width; length of aedeagus (AeL) was measured in ventral played in TreeView (Page, 1996) and annotated in Corel Photo view, from the base of median lobe to apices of apical projec- Paint. The data matrix is presented in Table 2. tions. All measurements are given in millimeters. List of adult morphological characters and character states Terminology and palaeogeographic background used in phylogenetic analysis Terminology of morphological structures follows that of 1. Body shape: slender and flattened (0); moderately slender and JaáoszyĔski (2010). Biogeographic terms and palaeogeographic moderately convex (1); stout and strongly convex (2). data are based on Keast & Miller (1996), Hall & Holloway 2. Body length: large, distinctly more than 1 mm (0); small, (1998) and Metcalfe et al. (2001). The term “Australian” is used 1 mm and less (1). strictly in its geographical sense, i.e. referring only to the Aus- 3. Vestiture on dorsum: short, dense and recumbent to suberect tralian continent and not to the islands east of
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