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Complete Mitochondrial Genomes of Two Oriental Dobsonflies

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Complete Mitochondrial Genomes of Two Oriental Dobsonflies Zootaxa 3964 (1): 044–062 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2015 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3964.1.2 http://zoobank.org/urn:lsid:zoobank.org:pub:634BDD12-A3F5-4A81-A84A-FDFB0409191C Complete mitochondrial genomes of two Oriental dobsonflies, Neoneuromus tonkinensis (van der Weele) and Nevromus exterior (Navás) (Megaloptera: Corydalidae), and phylogenetic implications of Corydalinae YUNLAN JIANG1, YAJUN ZHOU1, YIRAN WANG1, LU YUE1, YAN YAN 1, MENGQING WANG2 & XINGYUE LIU1,3 1Department of Entomology, China Agricultural University, Beijing 100193, China 2Key Laboratory of Integrated Pest Management in Crops, Ministry of Agriculture, Institute of Plant Protection, Chinese Academy of Agricultural Sciences, Beijing, 100081, China 3Corresponding author. E-mail: [email protected] Abstract The complete mitochondrial (mt) genomes of two Oriental endemic dobsonfly species, Neoneuromus tonkinensis (van der Weele) and Nevromus exterior (Navás), were determined and analyzed, which represent the first mt genomes of the genera Neoneuromus van der Weele, 1909 and Nevromus Rambur, 1842. The mt genome of N. tonkinensis is a typical circular DNA of 15776 bp with A+T content being 76.3%, while that of N. exterior is 15763 bp with A+T content being 77.5%. Both mt genomes are composed of 37 genes with an ancestral gene arrangement of the insect mt genome. Eleven of the 13 protein coding genes (PCGs) start with codon ATT and ATG, except for cox1 and nad1 respectively having ATC and ATA as the start codons in the mt genome of N. tonkinensis. Complete termination codons TAG and TAA were found in nine PCGs, while the remaining four genes are supposed to end with a single T. Most tRNAs are folded into the typical clover-leaf structure except for the trnS1 whose dihydrouridine arm is a simple loop. The secondary structure of rrnL con- sists of five structural domains and 50 helices, while the rrnS includes three domains and 34 helices. In the phylogenomic analysis, both Bayesian inference (BI) and maximum likelihood (ML) approaches, based on sequence data of all 13 PCGs and two rRNA genes of the mt genomes, suggested that Neoneuromus and Nevromus form a monophyletic group, which is the sister group of the lineage including Corydalus and Acanthacorydalis but not the sister group of Acanthacorydalis van der Weele, 1907 as previously reported based on morphological data. Key word: Corydalinae, Neoneuromus, Nevromus, mitochondrial genome, phylogeny Introduction Corydalinae (dobsonfly) is one of the subfamilies of the megalopteran family Corydalidae. Adult dobsonflies are characterized by the head with well-developed postocular plane (usually bearing a pair of postocular spines), and by the male genitalia having callus cerci not fused with ectoprocts and having a pair of well-developed ninth gonostyli, while the dobsonfly larvae can be easily distinguished by the presence of ventral abdominal tufts as a respiratory structure for adaptation of aquatic habitats (Glorioso 1981; Yang & Liu 2010). Currently, there are ca. 160 dobsonfly species sorted into nine genera which are vicariously distributed in America, eastern and southern Asia, and southern Africa (Yang & Liu 2010). Neoneuromus van der Weele, 1909 and Nevromus Rambur, 1842 are Oriental endemic genera of Corydalinae respectively including nine and six valid described species (Yang & Liu 2010; Liu et al. 2012). Adults of these two genera are relatively large-sized with acute postocular spines and the male genitalia are characterized by the male ninth tergum medially not separated anteriorly with an ovoid internal fossa and by the male ninth sternum which is attenuate and much narrower than ninth tergum (Liu & Yang 2004; Liu et al. 2012). Previous phylogenetic studies on the intergeneric phylogeny of Corydalinae using morphological data suggested that Neoneuromus and Nevromus are sister groups (Glorioso 1981; Penny 1993; Contreras-Ramos 1998, 2011). However, the position of these two genera in Corydalinae has not been resolved. Neoneuromus + 44 Accepted by B. Kondratieff: 28 Apr. 2015; published: 29 May 2015 Nevromus was assigned to be the sister lineage of the clade including Acanthacorydalis van der Weele, 1907, Corydalus Latreille, 1802, Chloronia Banks, 1908, and Platyneuromus van der Weele, 1909 in Glorioso (1981) and Penny (1993). Whereas, Neoneuromus + Nevromus was recovered to be the sister group of Acanthacorydalis in Contreras-Ramos (2011). These hypotheses were based on morphological evidence, but no molecule-based phylogenetic study is currently available to resolve these questions. In this study, we determined the first complete mitochondrial (mt) genomes of two representative species of Neoneuromus and Nevromus, i.e. Neoneuromus tonkinensis (van der Weele, 1907) and Nevromus exterior Navás, 1927. We analyzed the genomic organization, gene arrangement, codon usage, and the secondary structure of tRNAs and rRNAs, and we compared the mt genomes of N. tonkinensis and N. exterior with other sequenced mt genomes of Corydalinae. The present phylogenomic analysis did not support the sister group relationships between Neoneuromus + Nevromus and Acanthacorydalis, but recovered these two genera as the sister lineage of Acanthacorydalis + Corydalus. Material and methods Specimens and DNA extraction. A specimen of N. tonkinensis was collected at Tam Dao National Park, Vinh Phuc Province, Vietnam on May 17, 2012. The specimen of N. exterior was collected at Bac Kan City, Bac Kan Province, Vietnam on May 19, 2012. Both specimens were collected by Xingyue Liu using a light trap. The specimens were preserved in 95% ethanol and stored at -20°C in the Entomological Museum of China Agricultural University (CAU), Beijing, China. Total genomic DNA of N. tonkinensis and N. exterior were extracted by using the TIANamp Genomic DNA Kit (Tiangen Biotech, Beijing, China) from the mesothoracic muscle. PCR amplification and sequencing. The mt genomes of N. tonkinensis and N. exterior were generated by amplification of overlapping PCR fragments. Primers for the present PCR are provided in Tables 1–2. All PCRs used NEB LongAmp Taq DNA polymerase (New England BioLabs, Ipswich, MA, USA) under the following amplification conditions: 30s at 95°C, 40 cycles of 10s at 95°C, 50s at 40–55°C, 1kb/min at 65°C depending on the size of amplicons, and the final elongation step at 65°C for 10 min. The quality of PCR products was evaluated by 1% agarose gel electrophoresis. TABLE 1. Primer sequences of Neoneuromus tonkinensis mt genome used in this study. Number Primer ID Nucleotide sequence (5’-3’) Reference 1 TF210 AATTAAGCTACTAGGTTCATACCC Simon et al. 2006 TR1284 ACARCTTTGAAGGYTAWTAGTTT Simon et al. 2006 2 F20 (SPB-586) CCATTCCATTTYTGATTTCC Simon et al. 2006 R20 (SPB-1738) TTTATTCGTGGAAATGCTATGTC Simon et al. 2006 3 S2F TGGTACCTCAAGGAACACCTC Present study S2R TGCTCCTAAAGCTCCGGTTA Present study 4 F01 (SPA-2756) ACATTTTTTCCTCAACATTT Simon et al. 2006 R01 (SPA-3665) CCACAAATTTCTGAACACTG Simon et al. 2006 5 F02 (SPA-3399) TCTATTGGTCATCAATGGTACTG Simon et al. 2006 R02 (SPA-4061) GAAAATAAATTTGTTATCATTTTCA Simon et al. 2006 6 F03 (SPA-3790) CATTAAGTGACTGAAAGCAAGTA Simon et al. 2006 R03 (SPA-4552) ATGACCTGCAATTATATTAGC Simon et al. 2006 7 TF4463 TTTGCCCATCTWGTWCCNCAAGG Simon et al. 2006 TR5460 TCAACAAAATGTCARTAYCA Simon et al. 2006 8 F05 (SPA-4792) GTAGATGCAAGCCCTTGACC Simon et al. 2006 R05 (SPA-5731) ATTGGATCAAATCCACATTC Simon et al. 2006 9 TF5470 GCAGCTGCYTGATAYTGRCA Simon et al. 2006 ......continued on the next page MITOCHONDRIAL GENOMES OF TWO ORIENTAL DOBSONFLIES Zootaxa 3964 (1) © 2015 Magnolia Press · 45 TABLE 1. (Continued) Number Primer ID Nucleotide sequence (5’-3’) Reference TR5731 TTAGGGTCAAATCCRCAYTC Simon et al. 2006 10 S3F CCATTTTGGATTTGAAGCAG Present study S3R GCTCCTTGATTTCATTCGTGA Present study 11 F06 (SPA-5747) CCATTTGAATGTGGRTTTGATCC Simon et al. 2006 R06 (SPA-6384) AAAATTAAAAGCATAATATTGAAG Simon et al. 2006 12 S4F CACGAATGAAATCAAGGAGCTT Present study S4R GGAATATGGATTGTATGAG Present study 13 F07 (SPA-6172) AGAGGCAATTTATTGTTAATAA Simon et al. 2006 R07 (SPA-7211) TTAAGGCTTTATTATTTATATGTGC Simon et al. 2006 14 S13F GAATAACATACAGTTAATCCTGTAG Present study S13R GATTTGGTTATTTTTATTGAATGGG Present study 15 F08 (SPA-7077) TTAAATCCTTTGAGTAAAATCC Simon et al. 2006 R08 (SPA-7793) TTAGGTTGAGATGGTTTAGG Simon et al. 2006 16 S14F CCCAGAATAAATTTTTCCATGTTGT Present study S14R GCTCCTATTTCCGGGTCTATAATTT Present study 17 TF-J7806 GAMACAARACCTAACCCATCYCA Simon et al. 2006 TR-N8727 AAATCTTTRATTGCTTATTCWTC Simon et al. 2006 18 TF-J8641 CCAGAAGAACATAANCCRTG Simon et al. 2006 TR-N9629 GTTTGTGAGGGWGYTTTRGG Simon et al. 2006 19 TF-J9648 ACCTAAAGCTCCCTCACAWAC Simon et al. 2006 TR-N10608 CCAAGTARTGAWCCAAARTTTCA Simon et al. 2006 20 S5F AACTTTACGAACAACACACCCTCT Present study S5R GCAAACCCTCCTCAAACTCA Present study 21 F12 (SPB-11335) CATATTCAACCAGAATGATA Simon et al. 2006 R12 (SPB-12067) AATCGTTCTCCATTTGATTTTGC Simon et al. 2006 22 F13 (SPB-11876) CGAGGTAAAGTACCACGTACTCA Simon et al. 2006 R13 (SPB-12595) GTTGGATTTCTAACTTTATTRGARCG Simon et al. 2006 23 F14 (SPB-12261) TACCTCATAAGAAATAGTTTGAGC Simon et al. 2006 R14 (SPB-13000) TTACCTTAGGGATAACAGCGTAA Simon et al. 2006 24 F15 (SPB-12888) CCGGTCTGAACTCAGATCATGTA Simon et al. 2006 R15 (SPB-13889) ATTTATTGTACCTTTTGTATCAG Simon et al. 2006 25 F16 (SPB-13342) CCTTTGCACAGTCAAAATACTGC Simon et al. 2006 R16 (SPB-14220) TTATGCACACATCGCCCGTC Simon et al. 2006 26 F17 (SPB-14197) GTAAAYCTACTTTGTTACGACTT Simon
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