Abstract

Simple SummaryTwenty-seven complete mitochondrial genomes of Phasmatodea have been published in the NCBI. To shed light on the intra-ordinal and inter-ordinal relationships among Phasmatodea, more mitochondrial genomes of stick insects are used to explore mitogenome structures and clarify the disputes regarding the phylogenetic relationships among Phasmatodea. We sequence and annotate the first acquired complete mitochondrial genome from the family Pseudophasmatidae (Peruphasma schultei), the first reported mitochondrial genome from the genus Phryganistria of Phasmatidae (P. guangxiensis), and the complete mitochondrial genome of Orestes guangxiensis belonging to the family Heteropterygidae. We analyze the gene composition and the structure of the three mitochondrial genomes. We recover the monophyly of Phasmatodea and show the sister-group relationship between Phasmatodea and Mantophasmatodea after removal of the Embioptera and Zoraptera species. We recover the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae.Insects of the order Phasmatodea are mainly distributed in the tropics and subtropics and are best known for their remarkable camouflage as plants. In this study, we sequenced three complete mitochondrial genomes from three different families: Orestes guangxiensis, Peruphasma schultei, and Phryganistria guangxiensis. The lengths of the three mitochondrial genomes were 15,896 bp, 16,869 bp, and 17,005 bp, respectively, and the gene composition and structure of the three stick insects were identical to those of the most recent common ancestor of insects. The phylogenetic relationships among stick insects have been chaotic for a long time. In order to discuss the intra- and inter-ordinal relationship of Phasmatodea, we used the 13 protein-coding genes (PCGs) of 85 species for maximum likelihood (ML) and Bayesian inference (BI) analyses. Results showed that the internal topological structure of Phasmatodea had a few differences in both ML and BI trees and long-branch attraction (LBA) appeared between Embioptera and Zoraptera, which led to a non-monophyletic Phasmatodea. Consequently, after removal of the Embioptera and Zoraptera species, we re-performed ML and BI analyses with the remaining 81 species, which showed identical topology except for the position of Tectarchus ovobessus (Phasmatodea). We recovered the monophyly of Phasmatodea and the sister-group relationship between Phasmatodea and Mantophasmatodea. Our analyses also recovered the monophyly of Heteropterygidae and the paraphyly of Diapheromeridae, Phasmatidae, Lonchodidae, Lonchodinae, and Clitumninae. In this study, Peruphasma schultei (Pseudophasmatidae), Phraortes sp. YW-2014 (Lonchodidae), and species of Diapheromeridae clustered into the clade of Phasmatidae. Within Heteropterygidae, O. guangxiensis was the sister clade to O. mouhotii belonging to Dataminae, and the relationship of (Heteropteryginae + (Dataminae + Obriminae)) was recovered.

Highlights

  • Stick and leaf insects (Phasmatodea) belong to an order of polyneopteran insects, which includes over 3000 recognized species subdivided into approximately 500 genera, distributed across major landmasses [1]

  • To illuminate the phylogenetic relationships of Phasmatodea, we first performed maximum likelihood (ML) and Bayesian inference (BI) analyses based on data from 85 species, including the three newly determined sequences, sixty-five previously sequenced mitochondrial genomes, and 13 protein-coding genes (PCGs) of seventeen species of Phasmatodea assembled from transcriptome data [14,33,34,36,55,56,57,58,59,60,61,62,63,64,65,66,67,68,69,70,71,72,73,74,75,76,77] (Table 1, Table 2 and Table S2)

  • According to the previously published complete mitochondrial genomes of stick and leaf insects, we found that the differing lengths of the Phasmatodea genomes (15,590–18,248 bp) were caused mainly by the size of the

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Summary

Introduction

Stick and leaf insects (Phasmatodea) belong to an order of polyneopteran insects, which includes over 3000 recognized species subdivided into approximately 500 genera, distributed across major landmasses [1]. Polyneoptera includes ten insect orders of Blattodea, Dermaptera, Embioptera, Grylloblattodea, Mantodea, Mantophasmatodea, Orthoptera, Phasmatodea, Plecoptera, and Zoraptera Both morphological and molecular data highly support Embioptera as the sister group of Phasmatodea [7,8,9,10,11,12]. Data on mitochondrial genomes considered that Timema did not belong to the Euphasmatodea, but grouped with other orders (e.g., Orthoptera and Embioptera) [6,34]. This result coincided with the study about the morphology of Timema species in egg that indicated that Timema was a separate lineage [35]. We analyzed the gene composition and the structure of the three mitochondrial genomes

Sampling Collection and DNA Extraction
PCR Amplification and Sequencing
Mitochondrial Genome Annotation and Sequence Analyses
Phylogenetic Analyses
Mitochondrial Genome Organization and Composition
Protein-Coding Genes and Codon Usages
Ribosomal RNAs and Transfer RNAs
Intergenic and Overlap Regions
Intergenic and Overlapgenomes
Findings
Conclusions
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