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The Phylogenetic Structure of Metriorrhynchini Fauna of Sulawesi

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Zoological Studies 50(5): 645-656 (2011) The Phylogenetic Structure of Metriorrhynchini Fauna of Sulawesi (Coleoptera: Lycidae) with Descriptions of a New Genus, Mangkutanus, and Three New Species of Xylobanus Vaclav Kubecek, Milan Dvorak, and Ladislav Bocak* Department of Zoology, Faculty of Science, Palacky University, Tr. Svobody 26, 771 46 Olomouc, Czech Republic (Accepted May 20, 2011) Vaclav Kubecek, Milan Dvorak, and Ladislav Bocak (2011) The phylogenetic structure of Metriorrhynchini fauna of Sulawesi (Coleoptera: Lycidae) with descriptions of a new genus, Mangkutanus, and three new species of Xylobanus. Zoological Studies 50(5): 645-656. The phylogenetic structure of the Metriorrhynchini fauna from Sulawesi was investigated. We obtained DNA sequences for 3 fragments of the mitochondrial genome: cytochrome oxidase I, tRNA-Leu, and cytochrome oxidase subunit 2; NADH dehydrogenase subunit 5, tRNA- Ser, tRNA-Glu, and tRNA-Phe; and rrnl, tRNA-Leu, and NADH dehydrogenase subunit 1 for a total of 3070 aligned nucleotide positions, and performed a phylogenetic analysis of 12 genera of the Metriorrhynchini from Sulawesi and adjacent regions. Metriorrhynchini formed a monophylum that was split into 2 clades: an Afro- Oriental clade consisting of Cautires Waterhouse, 1879 and Xylobanus Waterhouse, 1879, and an Austral- Oriental clade formed by 10 genera distributed either only in the Australian Region, or with hundreds of species in Australia and New Guinea and a few in the Oriental region. Altogether, 11 genera are known from Sulawesi, four of them endemic to the island, only 2 genera, Xylobanus and Cautires, with a presumed Oriental origin, and 9 belonging to the Australian clade. All 87 metriorrhynchine species except 1 are endemic to Sulawesi, and 79 (91%) of them belong to the Australian lineage. We propose Mangkutanus gen. nov. for 2 species, which were previously placed in Sulabanus Dvorak and Bocak, 2007. The genus is described, diagnostic characters are illustrated, and relationships are discussed. We further redescribe the genus Xylobanus, describe 3 new species, and provide a key to Sulawesi species. http://zoolstud.sinica.edu.tw/Journals/50.5/645.pdf Key words: Phylogeny, mtDNA, Endemism, Dispersal, Taxonomy, Wallacea. Wallace’s line was proposed as a zoogeo- related species in the entire region including graphic border between continental Asia and Sulawesi (Maekawa et al. 2001, Balke et al. 2004 Australia 150 yr ago (Wallace 1860). Although 2009, Campbell et al. 2004, Sikes et al. 2006). permeability of the line is widely acknowledged Herein, we analyzed relationships among genera of (Mayr 1944), the dispersal history of various the Metriorrhynchini to investigate the phylogenetic lineages depends on their biology, and structure of the Sulawesi net-winged beetle fauna. consequently the phylogenetic structure of the The island of Sulawesi began to form about Sulawesi fauna remains poorly studied. Several 23 × 106 yr ago (mya) as a result of collisions authors examined the phylogeny and distribution of between several oceanic landmasses and an Asian animals found in the transitional zone between the continental fragment (Hall 2002), and has never Oriental and Australian regions including Sulawesi been connected to the Asian continent, although in the last 10 yr, but either focused on populations the Makassar Strait was much narrower during of a single species or the distributions of closely cold periods (Voris 2000). The Buton-Tukang Besi *To whom correspondence and reprint requests should be addressed. E-mail:[email protected] 645 646 Zoological Studies 50(5): 645-656 (2011) Platform which forms the southwestern peninsula for DNA extraction were unavailable for these taxa. of Sulawesi is a microcontinent of Australian origin, Sequences representing a further 31 species which collided with Sulawesi about 15 mya. The available from GenBank were added to the dataset Banggai Sula Block drifted to Sulawesi 6 mya (Hall (Bocak et al. 2006 2008). 2002). As a consequence, the Sulawesi fauna may be a mixture of both Asian and Australian DNA extraction, polymerase chain reaction continental faunas, but little is known about the (PCR) amplification, and DNA sequencing balance between them in Sulawesi or about relationships among the faunas of Sulawesi, Total DNA was extracted using the phenol- the Philippines, and Sundaland. Similarities of chloroform protocol, and the extraction yield Sulawesi and Philippine fauna led to the definition was determined using a NanoDrop-1000 of Huxley’s line, which excludes the Philippines spectrophotometer (NanoDrop Technologies, Inc., from the Oriental Region, the significance of which Wilmington, USA). The PCR was performed using was supported by numerous studies (Atkins et al. 0.5-0.6 U Taq polymerase, 1 mM MgCl2, 50 mM of 2001, de Bruyn et al. 2004). each dNTP, 0.2 mM primer, and typically 0.03 mg Although the 1st species of net-winged of a template in a 50-ml reaction volume. Cycle beetles from Sulawesi was described in the 19th conditions were generally 35-40 cycles of 2 min century, study of the fauna was neglected for a at 94°C, 30-60 s at 94°C, 30-60 s at 45-52°C, and long time, and only a few species were described 1-2 min at 72°C, with a final 10 min at 72°C. Three by R. Kleine and M. Pic in 1920s and 1930s (Kleine fragments were included in the study: 1100 bp 1933). Recently, 2 metriorrhynchine genera comprising cox1, tRNA-Leu, and cox2; a 1180-bp and several dozen species were described in a fragment comprising nad5 and the adjacent tRNA- series of taxonomical studies (Bocak 2000a 2001, Phe, tRNA-Glu, and tRNA-Ser; and an 800-bp Bocak and Jass 2004, Bocak et al. 2006, Dvorak fragment comprising rrnl, tRNA-Leu, and nad1 and Bocak 2007 2009). Herein, we conducted a (respectively referred to as cox1, nad5, and rrnl phylogenetic analysis to address relationships and hereafter). The primers used are listed by Bocak the zoogeography of the Metriorrhynchini from and Yagi (2010). Purified PCR products were Sulawesi. The principal question we addressed sequenced on an ABI 3130 automated sequencer was the proportions of Asian and Australian (Applied Biosystems, Foster City, USA) using the lineages based on the phylogenetic hypothesis. Big Dye Terminator Cycle Sequencing Kit vers. 1.1 Additionally, we also addressed the question of (Applied Biosystems, Foster City, USA). the monophyly of morphologically defined genera of previous studies, described a new genus, and Sequence handling and phylogenetic analyses redescribed the genus Xylobanus, describing 3 new species and providing a key to Sulawesi Sequences were edited using the Sequencher species. The role of molecular data has increased 4.6 software package (Gene Codes, Ann Arbor, in taxonomic research (Nalugwa 2010, Chullasorn USA) and aligned with ClustalX 1.8 using the 2011), and we used both sources of information default settings (Thompson et al. 1997). The to delimit the described genera and species. We aligned sequences were inspected with the naked consider the availability of DNA information for eye, and some minor adjustments were made. primary-type specimens as an integral part of A phylogenetic analysis was carried out delimitation of new taxa. under parsimony criteria using PAUP vers. 4.0.10 (Swofford 2003). All DNA fragments were combined for the final analysis. Equal weights MATERIAL AND METHODS were assigned to all positions, and gaps were treated as a 5th character state. For the bootstrap Sampling and geographic origin analysis, 1000 pseudoreplicates were generated and subsequently analyzed. The concatenated In total, 23 species representing 8 genera matrix was further analyzed under the likelihood of the Metriorrhynchini were sequenced, with criterion with RAxML vers. 7.2.3 (Stamatakis et GenBank accession numbers given in table 1. All al. 2008), and the support of the branches was known genera from Sulawesi were included in the assessed by analyzing 100 pseudoreplicates. All analysis, except for Diatrichalus Kleine, 1926 and genes and codon positions in the protein coding Lobatang Bocak, 1998. The properly fixed tissues fragments were partitioned. The model was Kubecek et al. – Metriorrhynchini of Sulawesi 647 Table 1. Specimens included in the study Species Voucher number Geographic origin rrnl cox1 nad5 Cantharis decipiens Baudi 00M14 Czech Republic DQ198685 DQ198608 DQ181417 Lyropaeus sp. 000L11 Malaysia DQ180968 DQ181190 DQ181344 Scarelus nigricornis Malohlava et Bocak 000L15 Malaysia - DQ181194 DQ181348 Lyponia nigrohumeralis Pic 000L17 China DQ180974 DQ181196 DQ181350 Dihammatus sp. 000L12 Malaysia DQ180969 DQ181191 DQ181345 Plateros sp. 000L13 Malaysia DQ180970 DQ181192 DQ181346 Calochromus sp. 000L16 China DQ180973 DQ181195 DQ181349 Macrolycus sp. 000L18 China DQ180975 DQ181197 DQ181351 Broxylus pfeifferi Waterhouse MD0099 Sulawesi HQ456957 HQ456980 HQ457002 B. malinensis Bocak et Jass MD0101 Sulawesi HQ456958 HQ456981 HQ457003 Cautires sp. (A) 000L14 RSA DQ180971 DQ181193 DQ181347 Cautires sp. (B) 000L06 Malaysia DQ180967 DQ181189 DQ181343 Cautires sp. (C) A00021 Taiwan HQ456947 HQ456965 - Cautires sp. (D) A00048 Malaysia HQ456948 HQ456967 HQ456990 Cautires sp. (E) A00080 Cameroon HQ456950 HQ456969 HQ456992 Cautires sp. (F) A00022 Madagascar - HQ456966 HQ456989 Cautires sp. (G) 000132 SE Asia - HQ456987 HQ457009 Cautiromimus sp. 000459 New Guinea - DQ144680 DQ144704 Cladophorus sp. 000451 New Guinea - DQ144681 DQ144705 Leptotrichalus sp. (A) 000208 Sabah - DQ181212 DQ181366 Leptotrichalus sp. (B) A00052
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