Abstract
Scelimeninae is an important subfamily of Tetrigoidea; however, the phylogenetic relationships within Scelimeninae are poorly understood, and its generic classification has remained unstable. In this study, the COI, 16S rRNA and 18S rRNA genes from 24 species in 9 genera within Scelimeninae were amplified and sequenced, the base composition and inter-species genetic distance of the combined sequence of COI, 16S rRNA and 18S rRNA genes were analyzed, and the molecular phylogenetic relationships were reconstructed using Maximum Likelihood (ML) and Bayesian inference (BI) methods. The results of sequence analysis showed that the total length of the combined COI, 16S rRNA and 18S rRNA gene sequence was 3507 bp, including 2345 conservative sites, 1144 variable sites and 901 parsimony-informative sites. The average A+T content was 63.5% and 78.1% in the COI, 16S rRNA sequences, respectively, indicating A+T bias. The average genetic distance between all species was 0.134, and the average genetic distance in the inner group (Scelimeninae) was 0.126. A phylogenetic tree based on the combined sequences of the COI, 16S rRNA and 18S rRNA genes showed that the phylogenetic relationships among 9 Scelimeninae genera were as follows: Criotettix + (((Zhengitettix + Hebarditettix) + (Falconius + (Scelimena + Paragavialidium))) + ((Eucriotettix + Thoradonta) + Loxilobus)). The molecular phylogenetic results generally support the morphological taxonomy; at the genus level, Criotettix, Scelimena, Paragavialidium, Thoradonta and Eucriotettix are monophyletic groups, Scelimena and Paragavialidium form sister groups, and Thoradonta and Eucriotettix also form sister groups, but the relationship between Hebarditettix and Zhengitettix needs further study. At the species level, synonyms may exist between Thoradonta spiculoba and Thoradonta transpicula and Thoradonta nodulosa and Thoradonta obtusilobata, but more studies are required to confirm this inference.
References
Cigliano, M.M., Braun, H., Eades, D.C. & Otte, D. (2018) Orthoptera species file. Version 5.0/5.0. Available from: http://Orthoptera.SpeciesFile.org (accessed 15 April 2018)
Deng, W.A., Zheng, Z.M. & Wei, S.Z. (2007) Fauna of Tetrigoidea from Yunnan and Guangxi. Science and Technology Press, Nanning, 458 pp.
Deng, W.A. (2016) Taxonomic study of Tetrigoidea from China. Ph.D. Dissertation, Huazhong Agricultural University, Wuhan, 341 pp.
Fang, N., Xuan, W.J., Zhang, Y.Y. & Jiang, G.F. (2010) Molecular phylogeny of chinese Tetrigoids (Othoptera, Tetrigoidea) based on the mitochondrial cytochrome CoxidaseI gene. Acta Zootaxonomica Sinica, 35 (04), 696–702.
Flook, P.K. & Rowell, C.H.F. (1997a) The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences. Molecular Phylogenetics and Evolution, 8 (1), 89–103.
Flook, P.K. & Rowell, C.H.F. (1997b) The Effectiveness of Mitochondrial rRNA Gene Sequences for the Reconstruction of the Phylogeny of an Insect Order (Orthoptera). Molecular Phylogenetics and Evolution, 8 (2), 177–192.
Flook, P.K. & Rowell, C.H.F. (1999a) Inferences about Orthopteroid Phylogeny and Molecular Evolution from Small Subunit Nuclear Ribosomal DNA Sequences. Insect Molecular Biology, 7 (2), 163–178.
Flook, P.K. & Rowell, C.H.F. (1999b) Combined Molecular Phylogenetic Analysis of the Orthoptera (Arthropoda, Insecta) and Implications for their Higher Systematics. Systems Biology, 48 (2), 233–253.
Hancock, J.L. (1912) Tetriginae (Acridiinae) in the Agricultural Research Institute, Pusa, Bihar, with descriptions of new species. Memoirs of the Department of Agriculture, India, Entomology Series, 4 (2), 131–160.
Jiang, G.F. (2000) Study on the genetic sequence and system evolution of the mitochondrial cytochrome b gene of Tetrigoidea. Shangxi Normal Universty, Xian, China, 2000, 1–86.
Lehmann, A.W., Devriese, H., Tumbrinck, T., Skejo, J., Lehmann, G.U.C. & Axel Hochkirch, A. (2017) The importance of validated alpha taxonomy for phylogenetic and DNA barcoding studies: a comment on species identification of pygmy grasshoppers (Orthoptera, Tetrigidae). ZooKeys, 679, 139–144.
https://doi.org/10.3897/zookeys.679.12507Li, X.D., Deng, W.A. & Lin, M.P. (2014) First description of male of Systolederus longipennis Zheng et Jiang (Orthoptera: Tetrigoidea). Journal of Huazhong Agricultural University, 33 (04), 50–52.
Leavitt, J.R., Hiatt, K.D., Whiting, M.F. & Song, H. (2013) Searching for the optimal data partitioning strategy in mitochondrial phylogenomics: A phylogeny of Acridoidea (Insecta: Orthoptera: Caelifera) as a case study. Molecular Phylogenetics and Evolution, 67 (2), 494–508.
https://doi.org/10.1016/j.ympev.2013.02.019Lin, L.L. (2014) Study on the mitochondrial genome analysis and phylogenetic relationship of three species of Tetrigoidea. Shangxi Normal Universty, Xian, 2014, 1–140.
Lin, M.P., Li, X.D., Wei, S.Z. & Deng, W.A. (2015) Mitochondrial COI gene partial sequences and phylogenetic relationship of some groups of Tetrigoidea. Journal of Huazhong Agricultural University, 34 (6), 40–48.
Pilar, P.G., Jesus, M. & Mario, G.P. (2012) Taxonomy and morphological characterization of Allotettix simony (Bolívar, 1890) and implications for the systematics of Metrodorinae (Orthoptera: Tetrigidae). Zoological Journal of the Linnean Society, 164, 52–70.
https://doi.org/10.1111/j.1096-3642.2011.00764.xRonquist, F. & Huelsenbeck, J.P. (2003) MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics, 19 (12), 1572–1574.
https://doi.org/10.1093/bioinformatics/btg180Song, H., Moulton, M.J. & Whiting, M.F. (2014) Rampant Nuclear Insertion of mtDNA across Diverse Lineages within Orthoptera (Insecta). PLoS ONE, 9 (10), e110508. https://doi.org/10.1371/journal.pone.0110508
Song, H., Amedegnato, C., Cigliano, M.M., Desutter-Grandcolas, L., Heads, S.W., Huang, Y., Otte, D. & Whiting, M.F. (2015) 300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling. Cladistics, 31, 621–651.
https://doi.org/10.1111/cla.12116Stamatakis, A. (2006) RAxML-VI-HPC: maximum likelihood-based phylogenetic analyses with thousands of taxa and mixed models. Bioinformatics, 22 (21), 2688–2690.
https://doi.org/10.1093/bioinformatics/btl446Tan, K.M., Storozhenko, S.Y., Hwang, W.S. & Meier, R. (2017) Integrative taxonomy reveals two sympatric species of the genus Eucriotettix Hebard, 1930 (Orthoptera: Tetrigidae). Zootaxa, 4268 (3), 377–394.
https://doi.org/10.11646/zootaxa.4268.3.4Thompson, J.D., Gibson, T.J., Plewniak, F., Jeanmougin, F. & Higgins, D.G. (1997) The CLUSTALX windows interface: flexible strategies for multiple sequence alignment aided by quality analysis tools. Nucleic Acids Res, 25 (24), 4876–4882.
https://doi.org/10.1093/nar/25.24.4876Wei, S.Z. (2007) Studies on the whole mitochondrial genomics of four species of Orthoptera. Shangxi Normal University, Xian, 2007, 1–175.
Wang, B. (2015) Cloning and characterization of 18S 28S and 5.8S ribosomal RNA genes and internal transcribed spacers of seven species/strains in genus Prorocentrum. Ocean University of China, 2015, 1–69.
Xiao, B., Feng, X., Miao, W.J. & Jiang, G.F. (2012) The complete mitochondrial genome of grouse locust Tetrix japonica (Insecta: Orthoptera: Tetrigoidea). Mitochondrial DNA, 23 (4), 288–289.
https://doi.org/10.3109/19401736.2012.674123Xiao, B., Chen, W., Hu, C.C. & Jiang, G.F. (2012) Complete mitochondrial genome of the groundhopper Alulatettix yunnanensis (Insecta: Orthoptera: Tetrigoidea). Mitochondrial DNA, 23 (4), 286–287.
https://doi.org/10.3109/19401736.2012.674122Yao, Y.P. (2008) Molecular evolution and systematic study of the 16S rRNA and 18S rRNA gene sequences of Tetrigoidea in China. Journal of Shangxi Normal Universty, Xian, 2008, 1–69.
Zheng, Z.M. (2005) The Tetrigoidea of western China. Science Press. Beijing, 501 pp.
Zhou, F. (2015) Determination of mitochondrial genomes and analysis of orthoptera phylogeny of four species of locusts. Journal of Shangxi Normal Universty, Xian, 2015, 1–103.
Zheng, Z.M. (1996) Three new species of Tetrigidae from China (Orthoptera). Acta Zootaxonomica Sinica, 21 (01), 83–88.
Zhao, L., Lin, L.L. & Zheng, Z.M. (2016) DNA barcoding reveals polymorphism in the pygmy grasshopper Tetrix bolivari (Orthoptera, Tetrigidae). ZooKeys, 582, 111–120.
https://doi.org/10.3897/zookeys.582.6301