ZooKeys 1092: 195-2 | 9) (2022) A peer-rev iewed open-access journal I doi: 10.3897/zookeys.1092.80993 RESEARCH ARTICLE #7Z,00Ke y https:/ / ZOO keys. pensoft.net Launched to accelerate biodiversity research The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species and phylogenetic implications Zhonghua Wei! | The Key Laboratory of Southwest China Wildlife Resources Conservation of the Ministry of Education, Col- lege of Life Sciences, China West Normal University, 637009, Nanchong, Sichuan Province, China Corresponding author: Zhonghua Wei (wzh1164@126.com) Academic editor: Dmitry Telnov | Received 21 January 2022 | Accepted 18 March 2022 | Published 6 April 2022 http://z00bank.org/F8957AFF-24AE-44E5-9577-43E3160A778B Citation: Wei Z (2022) The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species and phylogenetic implications. ZooKeys 1092: 195-212. https://doi.org/10.3897/zookeys.1092.80993 Abstract Five complete mitochondrial genomes of five species from the subfamily Agrilinae were sequenced and annotated, including Coraebus diminutus Gebhardt, 1928 (15,499 bp), Coraebus cloueti Théry, 1893 (15,514 bp), Meliboeus sinae Obenberger, 1935 (16,108 bp), Agrilus sichuanus Jendek, 2011 (16,521 bp), and Sambus femoralis Kerremans, 1892 (15,367 bp). These mitogenomes ranged from 15,367 to 16,521 bp in length and encoded 37 typical mitochondrial genes: 13 protein-coding genes (13 PCGs), 2 ribosomal RNA genes (2 rRNAs), 22 transfer RNA genes (22 tRNAs), and a control region (CR). Most of PCGs had typical ATN start codons and terminated with TAR or an incomplete stop codon T—. Among these five mitogenomes, Leu2, Ile, Phe, Ser2, Gly, Met, and Val were the seven most frequently encoded amino acids. Interestingly, in A. sichuanus, a 774 bp insertion was present at trnW and trnC junction, which is unusual in Buprestidae. Additionally, phylogenetic analyses were performed based on three kinds of nu- cleotide matrixes (13 PCGs, 2 rRNAs, and 13 PCGs + 2 rRNAs) using Bayesian inference and maximum- likelihood methods. The results showed that the clade of Buprestidae was well separated from outgroups and all Agrilinae species formed to a single highly supported clade. The tribe Coraebini was polyphyletic, as the genus Meliboeus (Coraebini) clustered with the genus Trachys (Tracheini). The rRNA genes had important impact for the tree topology of Agrilinae. Compared to the tribes Tracheini and Agrilini, the tribe Coraebini is a younger group. Keywords Comparative analysis, mitogenome, phylogenetic analysis Copyright Zhonghua Wei. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. 196 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Introduction The superfamily Buprestoidea, which contains the families Buprestidae and Schizo- podidae, differs from other groups of the Elateriformia by their serrate antennae, hy- pognathous head, transverse suture of metaventrite present, and two connate basal abdominal ventrites (Bellamy and Volkovitsh 2016). The buprestid beetles are a large group containing six subfamilies, 521 genera, and more than 15,000 species widely distributed in the world (Bellamy 2008; Kuban et al. 2016). The adults exhibit a broad range of host utilization in leaves, flowers, and stems, whereas larvae are mostly in- ternal feeders on roots and stems, or feed on foliage of woody or herbaceous plants (Bellamy and Volkovitsh 2016). Only adults of the Australian Xyroscelis crocata were reported to feed on the sap of the host plant Macrozamia communis (Bellamy 1997). Although taxonomists have made important contributions to the buprestid classi- fication of subfamilies and tribes based on several morphological characteristics (Cobos 1980, 1986; Téyama 1987; Holtynski 1988, 1993, 2009; Bellamy 2003), the problems of the overall classification in Buprestoidea remain unsettled. In the past two decades, molecular systematic approaches have been used to resolve unsettled classification and phylogenetic relationships in Insecta (Short and Fika¢ek 2013; Cline et al. 2014; Robertson et al. 2015; Kundrata et al. 2017; Gimmel et al. 2019; Lee et al. 2020). As to Buprestidae, Bernhard et al. (2005) first used mo- lecular phylogenetic methods based on three mitochondrial markers (mad1, 12S, and 16S) and confirmed that the Agrilus viridis complex, which is widely distributed across Eurasia, is monophyletic. Pentinsaari et al. (2014) and Pellegrino et al. (2017) used mitochondrial markers to evaluate the diversity of A. viridis complex, their results sug- gest that different feeding forms of A. viridis represent distinct species. Subsequently, Evans et al. (2015) performed the first large-scale phylogenetic trees combing nuclear and mitochondrial data from 141 species to understand the higher-level relationships in Buprestidae. In that study, the monophyly of the family Schizopodidae and sub- families Agrilinae, Julodinae, and Galbellinae were strongly supported, while the in- terrelationships of Chrysochroinae and Buprestinae remained uncertain. Hansen et al. (2016) used molecular systematic methods based on nuclear and mitochondrial data (coi and ak) to investigate the relationships within Chrysobothris femorata species group, and their results showed that some morphological species were not well sepa- rated. Kelnarova et al. (2019) provided a molecular phylogeny of Agrilus species from the Northern Hemisphere and their results suggest that DNA barcoding is a powerful species identification to Agrilus. During this time, the mitogenome emerged as a valuable source for higher-level phy- logenetic analyses, evolutionary strategies, and genetic diversity analyses (Saccone et al. 1999; Krzywinski et al. 2011; Cameron 2014; Qin et al. 2015; Song et al. 2019; Wang et al. 2019). Several buprestid mitogenomes have been sequenced and reported, such as the mitogenome of Chrysochroa fulgidissima (Schénherr, 1817) by Hong et al. (2009); the mitogenome of Agrilus planipennis Fairmaire, 1888 by Duan et al. (2017), who also performed phylogenetic analyses based on 13 PCGs of 45 mitogenomes of coleopterans; The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 197 Table |. Information on the mitogenomes of Buprestidae and two outgroups used in this study. size (bp) 1 Coraebus diminutus OK189521 15,499 38.34 68.42 0.12 -0.25 This study 2 — Coraebus cloueti OK189520 15,514 38.53 69.27 0.11 -0.25 This study 3 Meliboeus sinae OK189522 16,108 40.18 72.42 0.11 -0.22 This study 4 Sambus femoralis OK349489 15,367 40.98 73.23 0.12 -0.18 This study 5 Agrilus sichuanus OK189519 16,521 40.19 71.73 0.12 -0.21 This study 6 = Agrilus planipennis KT363854 15,942 40.25 71.90 0.12 -0.24 Duan et al. 2017 7 ~~ Agrilus mali MN894890 16,204 40.34 74.46 0.08 -0.18 Sun et al. 2020 8 Coraebus cavifrons MK913589 15,686 38.94 69.79 0.12 -0.18 Cao and Wang 2019b 9 Trachys auricollis MHG638286 16,429 38.94 71.05 0.10 -0.20 Xiao et al. 2019 10 Trachys troglodytiformis KX087357 16,316 41.03 74.62 0.10 -0.19 Unpublished 11 = Trachys variolaris MN178497 16,771 39.92 72.11 0.11 -0.21 Cao and Wang 2019a 12. Melanophila acuminata MW287594 15,853 38.74 75.66 0.02 -0.25 Peng et al. 2021 13. Anthaxia chinensis MW929326 15,881 40.12 73.61 0.09 -0.29 Chen et al. 2021 14. Chrysochroa fulgidissima EU826485 15,592 40.31 69.92 0.15 -0.24 Hong et al. 2009 15 Acmaeodera sp. FJ613420 16,217 38.11 68.41 0.11 -0.25 Sheffield et al. 2009 16 = Heterocerus parallelus (outgroup) KX087297 15,845 41.90 74.03 0.13 -0.24 Unpublished 17 Dryops ernesti (outgroup) KX035147 15,672 39.04 72.98 0.07 -0.23 Unpublished the mitogenome of Tiachys variolaris Saunders, 1873 by Cao and Wang (2019a); and the mitogenome of Coraebus cavifrons Descarpentries & Villiers, 1967 by Cao and Wang (2019b). More detailed information of buprestid mitogenomes is presented in Table 1. Currently, the subfamily Agrilinae contains four tribes (Agrilini, Coraebini, Aphanisticini, and Tracheini); however, the phylogenetic placement of several genera of this subfamily remains unstable. The genera in the tribes Coraebini and Agrilini were revised by Kuban et al. (2000). In that study, the genus Sambus in the tribe Coraebini was transferred to Agrilini based on the female behavior of ovipositing on rather smooth surfaces of living plants. Later, Kuban (2016) placed the genera Sambus, Parasambus, and Pseudagrilus in incertae sedis. In order to solve these problems, we contribute mitogenomic data of five species of buprestids, Coraebus diminutus Geb- hardt, 1928, Coraebus cloueti Théry, 1893, Meliboeus sinae Obenberger, 1935, Agrilus sichuanus Jendek, 2011, and Sambus femoralis Kerremans, 1892, and perform a mo- lecular phylogenetic analysis in this study. The phylogenetic trees of 15 species from nine genera belonging to four subfamilies of Buprestidae were constructed based on the newly sequenced and previously reported mitogenomes (Table 1). Material and methods Sampling and DNA extraction Specimens of five species were collected using an entomological net. Among them, C. diminutus, C. cloueti, M. sinae, and A. sichuanus were collected in the Dayaoshan Mountains in Guangxi Zhuang Autonomous Region, and S. femoralis was collected at 198 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Yingjiang County in Yunnan Province, China. Specimens were immediately preserved in 95% ethanol in the field after collected and then stored at —24 °C in the laboratory. The specimens were identified based on morphological characteristics under a Leica M205 FA stereomicroscope. Total DNA was extracted from muscle tissues using the Ezup Column Animal Genomic DNA Purification Kit (Shanghai, China) following the manufacturer's instructions. Sequencing, sequence assembly, annotation, and heterogeneity DNA sequencing and de novo assembly of each mitogenome were performed by Bei- jing Aoweisen Gene Technology Co. Ltd (Beijing, China). 22 tRNA genes were identi- fied using the MITOS webserver, with the parameters of the Invertebrate Mito genetic code (Bernt et al. 2013). Their secondary structures were plotted manually from the MITOS predictions using Adobe Illustrator. Every sequence of tRNA genes was manu- ally checked separately. The PCGs were identified as open reading frames correspond- ing to the 13 PCGs. The rRNAs and control regions were identified by the boundaries of the tRNA genes. The tRNA secondary structures were identified using tR NAscan-SE (Lowe and Chan 2016). Mitogenome maps (Suppl. material 1: Fig. S1) were produced using Organellar Genome DRAW (OGDRAW) (Greiner et al. 2019). The Base com- position and relative synonymous codon usage values were determined using MEGA 6.0 (Kumar 2016). Strand asymmetry was calculated using the formulae AT-skew = (A—T) / (A+ T), and GC-skew = (G — C) / (G + C) (Perna and Kocher 1995). In the control region (CR), tandem repeat elements were detected by Tandem Repeats Finder (Benson 1999). The heterogeneous analysis of the 13 PCGs and two rRNAs datasets were performed using AliGROOVE 1.06 (Kiick et al. 2014), and the nucleotide di- versity (Pi) and the ratio of Ka/Ks of PCGS were calculated with DnaSP v. 5 (Librado and Rozas 2009). Phylogenetic analyses Phylogenetic trees for A. sichuanus, C. diminutus, C. cloueti, M. sinae, S. femoralis, and 10 other buprestid species belonging to four subfamilies were reconstructed by three separate datasets (13 PCGs, 2 rRNAs, and 13 PCGs + 2 rRNAs) using different best- fit models (Table 4). The mitogenomes of Heterocerus parallelus (Heteroceridae) and Dryops ernesti (Dryopidae) were used as outgroups, as they are phylogenetically dis- tant from Buprestidae in the suborder Polyphaga (Xiao et al. 2019). The phylogenetic analyses were performed using PhyloSuite v. 1.2.2 (Zhang et al. 2020). Nucleotide sequences of the 13 PCGs and 2 rRNAs of all 17 mitogenomes were aligned using ClustalW (Thompson et al. 1994) and trimmed using trimAl v. 1.2 (Capella-Gutiérrez et al. 2009). The best-fit model for three datasets was determined by ModelFinder based on Bayesian information criterion. The maximum-likelihood (ML) and Bayesian inference (BI) methods were used to reconstruct the phylogenetic trees by IQ-tree v. 1.6.8 (Guindon et al. 2010) and MrBayes v. 3.2.6 program respectively (Ronquist et al. The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 199 2012). Bayesian analyses were run with two independent chains spanning 2,000,000 generations, four Markov chains, sampling at every 100 generations, and a burn-in period of 0.25 for each chain. The phylogenetic trees were edited and visualized by Figtree v. 1.4.3. Results and discussion Genome organization and base composition The complete mitogenomes of the buprestids A. sichuanus, C. diminutus, C. cloueti, M. sinae, and S. femoralis have the following GenBank accession numbers attributed to them: OK189519, OK189521, OK189520, OK189522, OK349489. The mitog- enomes of these five species contained the 37 typical mitochondrial genes (13 PCGs, 22 tRNAs, and 2 rRNAs) and a control region (CR) (Table 2). The composition and arrangement of the mitochondrial genes in these species (Table 2) were highly similar as those in most other buprestid species (Duan et al. 2017; Cao and Wang 2019a, 2019b; Xiao et al. 2019; Chen et al. 2021; Peng et al. 2021). Four of the 13 PCGs (nad1, nad4L, nad4, and nad5), eight tRNAs (trnQ, trnV, trnL1, trnP, trnH, trnF, trnY, and trnC), and two rRNAS (rrnL and rrnS) are encoded on the N-strand, whereas the other 23 genes (9 PCGs and 14 tRNAs) are encoded on the J-strand. The mitogenome sequence of these five buprestid species ranged in size from 15,367 to 16,521 bp. The mean A + T nucleotide contents of five complete mitogenomes were similar: 68.42% in C. diminutus, 69.27% in C. cloueti, 72.42% in M. sinae, 71.73% in A. si- chuanus, and 73.23% in S. femoralis. The entire mitogenomes had a higher A + T con- tents of 68.42—73.23% (66.05—72.50% for PCGs, 70.95-74.03% for tRNA genes, 75.20-77.33% for rRNA genes, and 74.17—78.38% for the CR) than G + C contents, which is consistent with the typical base of buprestid mitogenomes. The overall AT skews in these five complete mitogenomes were 0.12, 0.11, 0.11, 0.12, and 0.12, re- spectively. These five species showed a positive TA skew, suggesting that a slight AT bias which are similar to those observed in other buprestid species (Duan et al. 2017; Cao and Wang 2019a, 2019b; Xiao et al. 2019; Chen et al. 2021; Peng et al. 2021). Protein-coding regions, codon usage, and nucleotide diversity The total lengths of PCGs in these five buprestid species ranged from 11,090 to 11,158 bp, accounting for 67.54—72.17% of the entire mitogenomes. Similar to the other buprestid mitogenomes, mad5 and atp8 were found to be the largest (1708- 1723 bp) and smallest (156-159 bp) genes, respectively. The majority of PCGs strictly started with an AT'N (ATA/ATT/ATC/ATG) start codon, except for the ad/ starting with TTG. All PCGs strictly terminated with TAR (TAG/TAA) or an incomplete stop codon T-. 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N I/T/T1/T/6 VVO L8ET/6GLVI/LOVI/VOVI/S6IT LTETLILSEL/GEEL/THEL/TELT N Aud 0/0/C/C/0 VOO ITELIOSET/IEET/GEE L/OETT 99NTI/I6TI/9LTI/6LT1/8907 N Dud 8-/EL/ET-/8-/PLZL VOV ELTI/EOET/E8TI/I8TI/C6T1 GOTI/TETI/BITL/TCCL/PCTI [ Mud C-/0/C-/ T/T VWVL/VVL/VVL/DVL/VVL LIV/O.LV/.LIV/LLV/OLV OITI/TETI/GITL/61CIL/CCTI L61/907/L61/Z61/007 [ Epue 0/0/0/0/0 VVO 961/S07/961/961/661 TEL/8ET/6TI/6TL/TET [ WU T-/0/0/0/1- OVV TEL/8ET/67T1/671/TET €9/0£/19/19/€9 N ous €-S/€-/€-/€- IVS 69/99/€9/€9/69 T/T/T/T/T [ [444 saphospnu s1ue8ia3u, += uopoonuy suopuos doyg SUOPOd 11815 OL WoL] UOTISOg puens auary “pouTUJaIap 10U — ‘s77MLOWMaf snquiYS pur ‘avuls snaoqyay “Manos 200 SNGAVAO) ‘SNINUIULIP SNGAVLO) “SNUPYNYAIIS SNILABY :SMOT[OF Sk St 3]GeI BA UT saTads day say} JO JOpIO dT, ‘soMOUdSONW seprsordng poeiouur A[MoU dAy SUT, *7 VIQUL The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 201 Table 3. Summarized mitogenomic characteristics of the five buprestid species in this study. Species PCGs rRNAs tRNA CR Size A+T ATskew _ Size A+T ATskew _ Size A+T AT skew Size A+T AT skew (bp) content (bp) content (bp) content (bp) content A. sichuanus 11,158 70.08 -0.15 1976 75.96 -0.13 1444 74.03 -0.0009 1142 74.17 0.06 C. diminutus 11,093 66.05 -0.14 1984 75.20 -0.11 1477 70.95 0.03 1019 77.72 0.02 C. cloueti 11,093 67.09 -0.15 1983 75.39 -0.11 1414 71.22 0.019 1031 78.27 0.02 M. sinae 11,135 70.70 -0.15 1967 77.33 -0.11 1435 72.13 0.007 1577 78.38 0.13 S. femoralis 11,090 = 72.50 -0.16 1954 75.69 -0.13 1430 73.85 0.03 910 75.82 0.18 Table 4. Best-fit models of three datasets used for phylogeny. ML method BI method 13 PCGs GTR+F+I+G4 GTR+F4+1+G4 2 rRNAs TVM+F+1+G4 GTR+F+1+G4 13 PCGs +2 rRNAs GTR+F+I+G4 GTR+F+1+G4 (0.11—0.12) of these five PCGs (Table 3) were similar among the 15 buprestid species. Summaries of the numbers of amino acids in the annotated PCGs and relative syn- onymous codon usage are presented in Figs 1 and 2. Overall codon usage among the sequenced buprestid mitogenomes was found to be similar, with Leu2, Ile, Phe, Ser2, Gly, Met, and Val being the seven most frequently coded amino acids. The nucleotide diversity (Pi) of the 13 PCGs among five species of Agrilinae is provided (Fig. 3), which ranged from 0.202 to 0.375. In these genes, nad2 (Pi = 0.375) presented the highest variability, followed by nad6 (Pi = 0.346), nad4 (Pi = 0.300), and nad5 (Pi = 0.290); coxI (Pi = 0.20) exhibited the lowest variability. The ratio of Ka/Ks (Fig. 4) for each gene of the 13 PCGs was calculated. The values of nad4 and nad4L are distinctly higher than others, which suggests that the genes nad4 and nad4L have a relatively higher evolutionary rate. 3500 3000 - 3.66% 2500 15.48% 15.65% 10.57% 4.42% 6.13% 4.20% 6.05% 2000 10.63% 1500 1000 15.88% 10.16% BF elLigl2e| Mev wSsieP BT BA BY 4.75% 5.81% BH @SQG@N OK @O BE BC OWOeR 8S28G Figure |. Numbers of different amino acids in the mitogenomes of the five buprestid species; the stop codon is not included. AS: Agrilus sichuanus, CC: Coraebus cloueti, CD: Coraebus diminutus, MS: Meli- boeus sinae, and SF: Sambus femoralis. 202 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Agrilus sichuanus RSCU Coraebus cloueti RSCU Coraebus diminutus RSCU Meliboeus sinae RSCU Sambus femoralis RSCU nm wo = Ala Arg Asn Asp Cys Gin Glu Gly His lle Leul Leu2 Lys Met Phe Pro Serl Ser2 Thr Trp Tyr Val Kefefe] FXefel Miele) EXetel Figure 2. RSCU (relative synonymous codon usage) of the mitogenomes of the five buprestid species; the stop codons are not included. The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 203 -) | a Pa t ND2 col cou ZaATP6 com & NDS Np4 &ND6 CYTB NDI 0.375 0.202 0.240 = 0.280 0.250 = 0.290 0.300 20.346 0.254 0.24 ee) ‘Oo Nn et oo wn 0 2500 5000 7500 10000 12500 Nucleotide Position Figure 3. Nucleotide diversity (Pi) of 13 PCGs among five newly sequenced Agrilinae mitogenomes. : ; J i | | , F | D3 ATP6 ATP8 COI COM CYTB NDI N ND4 ND4L ND5 oo bo 1.5 — 0. wn uKs mKa = Ka/Ks Figure 4. The ratio of Ka/Ks of 13 PCGs among the 15 reported Buprestidae mitogenomes. tRNA, rRNA genes, and heterogeneity The length of rrnZ genes ranged from 1258 bp (S. femoralis) to 1271 bp (A. si- chuanus), whereas rrnS ranged from 696 bp (S. femoralis) to 718 bp (C. diminutus). The A + T content of the rRNA genes ranged from 75.20% (C. diminutus) to 77.33% (M. sinae) (Table 3). Compared with those in other sequenced buprestid mitogenom- 204 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Ay ee cS A ‘ Coraebus diminutus Meliboeus sinae Sambus femoralis Figure 5. The predicted secondary structures of the tRNA-Ser in the mitogenomes of the five buprestid species. es, the rRNA genes in these five newly sequenced buprestid mitogenomes are highly conserved (Hong et al. 2009; Duan et al. 2017; Cao and Wang 2019a, 2019b; Xiao et al. 2019; Sun et al. 2020; Chen et al. 2021; Peng et al. 2021). These rRNAs were located between the CR and ¢trnL1, and separated by zrnV. The total lengths of the 22 tRNA genes ranged from 1414 bp (C. cloueti) to 1444 bp (C. diminutus), whereas individual tRNA genes typically ranged in size from 58 to 70 bp, among which, eight tRNAs were encoded on the N-strand and the remaining 14 encoded on the J-strand. The secondary structures of tRNAs showed a standard clover-leaf structure (Suppl. material 1: Figs S2-S6), except for tRNA-Ser (Fig. 5) which lacks or has an unusual dihydrouridine arm, which forms a loop commonly found in other insects (Xiao et al. 2011; Park et al. 2012; Yu et al. 2016; Yan et al. 2017; Yu and Liang 2018; Li et al. 2019). In A. sichuanus, the longest intergenic nucleotide (774 bp) was located between trnW and trnC, which is an interesting and specific phenomenon in Buprestidae. The degree of heterogeneity of the 13 PCGs dataset was higher than that of the two rRNAs dataset (Suppl. material 1: Fig. S7). Additionally, the heterogeneity in sequence divergences was slightly stronger for Coraebus than for other buprestid genera (Suppl. material 1: Fig. S7). The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 205 Control region The CR, also known as the A + T-rich region (Wolstenholme 1992), was the larg- est non-coding region and located between trn/ and rrnS. The length of CR ranged from 910 bp (S. femoralis) to 1577 bp (M. sinae). The A + T content (74.17—78.38%) of the CR of these five species was found to be higher than that of the whole ge- nome (68.42—73.23%), PCGs (66.05—72.50%), rRNAs (75.20—77.33%), and tRNAs (70.95-73.85%) (Table 3). Moreover, the compositional analysis revealed that the mi- togenomes of the five buprestid species had a positive AT skew (0.02—0.18) in the CR. In these five species, only C. cloueti and C. diminutus had no tandem repeat element detected; however, those of A. sichuanus (20 and 40 bp), M. sinae (53 bp), and S. femo- ralis (265 bp) had different lengths. Phylogenetic analyses Both ML and BI trees using three datasets produced identical topologies (Figs 6-8), (Buprestinae + ((Chrysochroniae + Polycestinae) + Agrilinae)), in terms of subfamily- level relationship. The monophyly of Buprestidae is corroborated again, as all the bu- prestid species converged together as an independent clade, and two outgroup taxa obviously separated from the buprestid clade. The target species C. diminutus, C. clou- eti, Meliboeus sinae, Agrilus sichuanus, and Sambus femoralis, as well as other species of Agrilinae, converged together as an independent clade. And the target species, M. sinae, 1 Agrilus mali > l : ; P we 70 Agrilus planipennis as 1 Ky Nos ~ Agrilus sichuanus . fe Sambus femoralis ] ie a > = 1 ~~ Trachys auricollis § 99 100 sees alee [Dee 1 SG ~ Trachys roglodytiformis |= 2 -- Meliboeus sinae 1 Coraebus cloueti is 0.64 : ae 2 1 100 - Coraebus diminutus & MT tae Coraebus cavifrons E 37 77 Se ee a ee Chrysochroa fulgidissima | Chrysochroinae Buprestinae ~~ Heterocerus parallelus Outgroups ~ Dryops emesti 0.07 Figure 6. Phylogenetic relationships of 15 selected buprestid species using both BI and ML analyses based on 13 PCGs of mitogenomes. The numbers on the branches show posterior probability (BI tree), whereas the values under branches are bootstrap (ML tree). 206 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) i Agrilus mali > 1 [SSS meee Agrilus plaipennis a 0 ~ Agrilus sichuanus : 1oocee6- Coraebus cloueti S 100 Coraebus diminutus S |e 1 100 Coraebus cavifrons E E 1 Trachys auricollis as 1 109 a ve Trachys troglodytitormis z 98 3] » Trachys variolaris = 0.6 vows Maliboeus sinae I 0.99 Acmaeodera sp | Polycestinae 50 eA Chrysochroa fulgidissima _|Chrysochroinae I nn Al mthaxia chinensis Biiprettinas 94 aint, Hy MR ARIAZRL Wed ARSANE REAR 127 on ARH NO! A OA GRE OAR Melanophila acuminata a ae eee sénpedsinapecenystfajebpjottbocfe Heterocerus parallelus sinenk Outgroups 0.05 Figure 7. Phylogenetic relationships of 15 selected buprestid species using both BI and ML analyses based on 2 rRNAs of mitogenomes. The numbers on the branches show posterior probability (BI tree), whereas the values under branches are bootstrap (ML tree). 1 Agrilus mali > L 92 Agrilus planipennis = 0.76 mk Agnilus sichuanus 3 1 Coraebus cloueti |S 100 Coraebus diminutus |&. | d= 0? ae Coraebus cavitrons |B. | = 1 Trachys auricollis 3 & 1 ; - 85 Trachys variolaris e 55 5 Trachys troglodytiformis 5. 0.76 Meliboeus sinae 140 Sambus femoralis ; L Acmaeodera sp | Polycestinae a he Chrysochroa fulgidissima | Chrysochroinae 1 Anthaxia chinensis 100 Melanophila acuminata Buprestinae Heterocerus parallelus Dryops emesti Outgroups 0:07 - Figure 8. Phylogenetic relationships of 15 selected buprestid species using both BI and ML analyses based on 13 PCGs + 2 rRNAs of mitogenomes. The numbers on the branches show posterior probability (BI tree), whereas the values under branches are bootstrap (ML tree). The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 207 was most closely related to the genus Trachys with high value support (Figs 6-8) which is inconsistent with the previous studies (Kuban et al. 2000; Evans et al. 2015). The relationship of Agrilinae clades obtained from 2 rRNAs and 13 PCGs + 2 rRNAs data- sets are identical but with different topology from the 13 PCGs dataset. In the topol- ogy generated from the 13 PCGs dataset, S. femoralis and Agrilus were clustered into a single branch with high support value (Fig. 6, ML: 77, BI: 1) whereas, in the topology generated from the 2 rRNAs and 13 PCGs + 2 rRNAs datasets, S. femoralis split from base of the Agrilinae clades (Figs 7, 8). Based on these results the position of the genus Sambus in the tribe Agrilini was not suitable and suspect. ‘The different tree topologies suggested that the rRNA genes were extremely valuable for the phylogenetic analysis of Agrilinae. Coraebini is the most diverse tribe in Agrilinae, and 10 subtribes are defined (Kuban et al. 2000). The genus Meliboeus (Meliboeina) and Coraebus (Coraebina) in different clades suggested that the tribe Coraebini was polyphyletic, which is consistent with the previous study of Evans et al. (2015). The samples used in this study might be too limited for a comprehensive phylogeny of Buprestidae which still needs a deep study in the future. Conclusions In this study, five mitogenomes (15,367—16,521 bp) were newly sequenced and an- notated, including representatives from the tribes Coraebini and Agrilini in subfamily Agriinae. The mitogenomes of the genera Sambus and Meliboeus are reported for the first time. These five sequences showed a positive AT skew, and the amino acids Leu, Ile, Phe, Ser2, Gly, Met, and Val were most frequently used. The secondary structures of tRNA-Ser are absent the D-arm, which is similar to other orders of Insecta. The rRNA genes are valuable for phylogenetic analyses of Agrilinae as they could affect the tree topologies. The results show that Coaebini is polyphyletic, and the genus Sambus belongs to neither Coaebini nor Agrilini. However, more mitogenome samplings are needed to resolve the phylogeny of the Buprestidae in the future to better understand the phylogenetics of jewel beetles. Acknowledgements I am sincerely grateful to Lanrui Wang (Qingyang, Gansu, China) and Yingqi Liu (China Agricultural University, Beijing, China) for their guidance in using the soft- ware. I also thank Dr Hui-Feng Zhao (Langfang Normal University, Hebei, China) and Menglin Wang (China West Normal University, Sichuan, China) for revising the manuscript. This work was supported by the Doctoral Scientific Research Foundation of China West Normal University (20E054). 208 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) References Bellamy CL (1997) Phylogenetic relationships of Xyroscelis (Coleoptera: Buprestidae). Inverte- brate Systematics 11(4): 569-574. https://doi.org/10.1071/IT94026 Bellamy CL (2003) An illustrated summary of the higher classification of the superfamily Bu- prestoidea (Coleoptera). Folia Heyrovskyana (Supplementum 10): 1-197. Bellamy CL (2008) A World Catalogue and Bibliography of the Jewel Beetles (Coleoptera: Bu- prestoidea). Volumes 1—4. Pensoft series faunistica No. 76-79, Sofia/ Moscow, [8] + 2684 pp. Bellamy CL, Volkovitsh M (2016) 18 Buprestoidea Crowson, 1955. In: Beutel RG, Leschen RAB (Eds) Handbook of Zoology, Arthropoda: Insecta, Volume 1: Morphology and Sys- tematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim), 2"¢ edn. Walter de Gruyter, Berlin/Boston, 543-552. https://doi.org/10.1515/9783110373929-021 Benson G (1999) ‘Tandem repeats finder: A program to analyze DNA sequences. Nucleic Acids Research 27(2): 573-580. https://doi.org/10.1093/nar/27.2.573 Bernhard D, Fritzsch G, Glockner P, Wurst C (2005) Molecular insights into speciation in the Agrilus viridis-complex and the genus Trachys (Coleoptera: Buprestidae). European Journal of Entomology 102(4): 599-605. https://doi.org/10.1441 1/eje.2005.083 Bernt M, Donath A, Juhling E Externbrink F, Florentz C, Fritzsch G, Stadler PF (2013) MITOS: Improved de novo metazoan mitochondrial genome annotation. Molecular Phy- logenetics and Evolution 69(2): 313-319. https://doi.org/10.1016/j.ympev.2012.08.023 Cameron SL (2014) Insect mitochondrial genomics: Implications for evolution and phylog- eny. Annual Review of Entomology 59(1): 95-117. https://doi.org/10.1146/annurev-en- to-011613-162007 Cao LM, Wang XY (2019a) The complete mitochondrial genome of the jewel beetle Trachys variolaris (Coleoptera: Buprestidae). Mitochondrial DNA, Part B, Resources 4(2): 3042— 3043. https://doi.org/10.1080/23802359.2019.1666053 Cao LM, Wang XY (2019b) The complete mitochondrial genome of the jewel beetle Coraebus cavifrons (Coleoptera: Buprestidae). Mitochondrial DNA, Part B, Resources 4(2): 2407— 2408. https://doi.org/10.1080/23802359.2019.1636730 Capella-Gutiérrez S, Silla-martinez JM, Gabaldén T (2009) TrimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics (Oxford, Eng- land) 25(15): 1972-1973. https://doi.org/10.1093/bioinformatics/btp348 Chen B, Wei ZH, Shi AM (2021) The complete mitochondrial genome of the jewel beetle, Anthaxia chinensis (Coleoptera: Buprestidae). Mitochondrial DNA, Part B, Resources 6(10): 2962-2963. https://doi.org/10.1080/23802359.2021.1973920 Cline AR, Smith TR, Miller K, Moulton M, Whiting M, Audisio P (2014) Molecular phy- logeny of Nitidulidae: assessment of subfamilial and tribal classification and formalization of the family Cybocephalidae (Coleoptera: Cucujoidea). Systematic Entomology 39(4): 758-772. https://doi.org/10.1111/syen.12084 Cobos A (1980) Ensayo sobre los géneros de la subfamilia Polycestinae (Coleoptera, Bupresti- dae) (Parte I). EOS. Revista Espafola de Entomologia 54: 15-94. Cobos A (1986) Fauna Iberica de Coleopteros Buprestidae. Consejo Superior de Invertiga- ciones Cientificas, Madrid, 364 pp. The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 209 Duan J, Quan GX, Mittapalli O, Cusson M, Krell PJ, Doucet D (2017) The complete mitog- enome of the Emerald Ash Borer (EAB), Agrilus planipennis (Insecta: Coleoptera: Bupresti- dae). Mitochondrial DNA, Part B, Resources 2(1): 134-135. https://doi.org/10.1080/23 802359.2017.1292476 Evans AM, Mckenna DD, Bellamy CL, Farrell BD (2015) Large-scale molecular phylogeny of metallic wood-boring beetles (Coleoptera: Buprestoidea) provides new insights into rela- tionships and reveals multiple evolutionary origins of the larval leaf-mining habit. System- atic Entomology 40(2): 385-400. https://doi.org/10.1111/syen.12108 Gimmel ML, Bocakova M, Gunter NL, Leschenm RAB (2019) Comprehensive phylogeny of the Cleroidea (Coleoptera: Cucujiformia). Systematic Entomology 44(3): 527-558. https://doi.org/10.1111/syen.12338 Greiner S, Lehwark P, Bock R (2019) OrganellarGenomeDRAW (OGDRAW) version 1.3.1: Expanded toolkit for the graphical visualization of organellar genomes. Nucleic Acids Re- search 47(W1): W59-W64. https://doi.org/10.1093/nar/gkz238 Guindon S, Dufayard J, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New al- gorithms and methods to estimate maximum-likelihood phylogenies: Assessing the per- formance of PhyML 3.0. Systematic Biology 59(3): 307-321. https://doi.org/10.1093/ sysbio/syq0 10 Hansen JA, Moulton JK, Klingeman WE, Oliver JB, Windham MT, Trigiano RN, Reding ME (2016) Molecular systematics of the Chrysobothris femorata species group (Coleoptera: Buprestidae). Annals of the Entomological Society of America 108(5): 950-963. https:// doi.org/10.1093/aesa/sav080 Holyriski RB (1988) Remarks on the general classification of Buprestidae Leach as applied to Maoraxiina. Folia Entomologica Hungarica 49(1): 49-54. Hotynski RB (1993) A reassessment of the internal classification of the Buprestidae Leach (Co- leoptera). Crystal. Series Zoologica (G6éd) 1: 1-42. Hotynski RB (2009) Taxonomic structure of the subtribe Chrysochroina Cast. with review of the genus Chrysochroa Dej. Gondwana, Warszawa, 391 pp. Hong MY, Jeong HC, Kim MJ, Jeong HU, Lee SH, Kim I (2009) Complete mitogenome sequence of the jewel beetle, Chrysochroa fulgidissima (Coleoptera: Buprestidae). Mi- tochondrial DNA Mapping, Sequencing, and Analysis 20(2—3): 46-60. https://doi. org/10.1080/19401730802644978 Kelnarova I, Jendek E, Grebennikov VV, Bocak L (2019) First molecular phylogeny of Agrilus (Coleoptera: Buprestidae), the largest genus on Earth, with DNA barcode database for forestry pest diagnostics. Bulletin of Entomological Research 109(2): 200-211. https:// doi.org/10.1017/S00074853 18000330 Krzywinski J, Li C, Morris M, Conn JE, Lima JB, Povoa MM, Wilkerson RC (2011) Analy- sis of the evolutionary forces shaping mitochondrial genomes of a Neotropical malaria vector complex. Molecular Phylogenetics and Evolution 58(3): 469-477. https://doi. org/10.1016/j.ympev.2011.01.003 Kuban V (2016) Tribe Agrilini, genera incertae sedis. In: Lob! I, Lobl D (Eds) Catalogue of Palaearctic Coleoptera. Volume 3, Scarabaeoidea, Scirtoidea, Dascilloidea, Buprestoidea, Byrrhoidea. Revised and updated edition; Leiden, Boston, 549 pp. 210 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Kuban V, Majer K, Koliba¢ J (2000) Classification of the tribe Coraebini Bedel, 1921 (Coleop- tera, Buprestidae, Agrilinae). Acta Musei Moraviae. Scientiae Biologicae (Brno) 85: 185-287. Kuban V, Volkovitsh MG, Kalashian MJ, Jendek E (2016) Family Buprestidae Leach, 1815. In: Lobl I, Lobl D (Eds) Catalogue of Palaearctic Coleoptera. Volume 3, Scarabaeoidea, Scirtoidea, Dascilloidea, Buprestoidea, Byrrhoidea. Revised and Updated Edition. Apollo Books, Stenstrup, 432-574. Kiick P, Meid SA, Grof C, Wagele JW, Misof B (2014) AliGROOVE-visualization of het- erogeneous sequence divergence within multiple sequence alignments and detection of inflated branch support. Bioinformatics (Oxford, England) 15: e294. https://doi. org/10.1186/1471-2105-15-294 Kumar S, Stecher G, Tamura K (2016) MEGA7: Molecular Evolutionary Genetics Analy- sis version 7.0 for bigger datasets. Molecular Biology and Evolution 33(7): 1870-1874. https://doi.org/10.1093/molbev/msw054 Kundrata R, Jach MA, Bocak L (2017) Molecular phylogeny of the Byrrhoidea—Buprestoidea complex (Coleoptera, Elateriformia). Zoologica Scripta 46(2): 150-164. https://doi. org/10.1111/zsc.12196 Lee MH, Lee S, Leschen RAB, Lee S (2020) Evolution of feeding habits of sap beetles (Co- leoptera: Nitidulidae) and placement of Calonecrinae. Systematic Entomology 45(4): 911-923. https://doi.org/10.1111/syen.12441 Li R, Shu XH, Li XD, Meng L, Li BP (2019) Comparative mitogenome analysis of three species and monophyletic inference of Catantopinae (Orthoptera: Acridoidea). Genomics 111(6): 1728-1735. https://doi.org/10.1016/j.ygeno.2018.11.027 Librado P, Rozas J (2009) DnaSP v5: A software for comprehensive analysis of DNA poly- morphism data. Bioinformatics (Oxford, England) 25(11): 1451-1452. https://doi. org/10.1093/bioinformatics/btp187 Lowe TM, Chan PP (2016) tRNAscan-SE On-line: Integrating search and context for analy- sis of transfer RNA genes. Nucleic Acids Research 33(W1): W686—W689. https://doi. org/10.1093/nar/gkw413 Park JS, Cho Y, Kim MJ, Nam SH, Kim I (2012) Description of complete mitochondrial ge- nome of the black-veined white, Aporia crataegi (Lepidoptera: Papilionoidea), and compar- ison to papilionoid species. Journal of Asia-Pacific Entomology 15(3): 331-341. https:// doi.org/10.1016/j.aspen.2012.01.002 Pellegrino I, Curletti G, Liberatore K Cucco M (2017) Cryptic diversity of the jewel beetles Agrilus viridis (Coleoptera: Buprestidae) hosted on hazelnut. The European Zoological Journal 84(1): 465-472. https://doi.org/10.1080/24750263.2017.1362050 Peng XJ, Liu J, Wang Z, Zhan QZ (2021) The complete mitochondrial genome of the pyroph- ilous jewel beetle Melanophila acuminata (Coleoptera: Buprestidae). Mitochondrial DNA. Part B, Resources 6(3): 1059-1060. https://doi.org/10.1080/23802359.2021.1899079 Pentinsaari M, Mutanen M, Kaila L (2014) Cryptic diversity and signs of mitochondrial intro- gression in the Agrilus viridisspecies complex (coleoptera: Buprestidae). European Journal of Entomology 111(4): 475-486. https://doi.org/10.1441 1/eje.2014.072 Perna NT, Kocher TD (1995) Patterns of nucleotide composition at fourfold degenerate sites of animal mitochondrial genomes. Journal of Molecular Evolution 3(3): 353-358. https:// doi.org/10.1007/BF01215182 The complete mitochondrial genomes of five Agrilinae (Coleoptera, Buprestidae) species... 211 Qin J, Zhang YZ, Zhou X, Kong XB, Wei SJ, Ward RD, Zhang AB (2015) Mitochondrial phy- logenomics and genetic relationships of closely related pine moth (Lasiocampidae: Den- drolimus) species in China, using whole mitochondrial genomes. Genomics 16(1): e428. https://doi.org/10.1186/s12864-015-1566-5 Robertson JA, Slipitski A, Moulton M, Shockley FW, Gorgi A, Lord NP, McKenna DD, Tomaszewska W, Forrester J, Miller KB, Whiting ME, McHugh JV (2015) Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinel- loidea (Coleoptera: Cucujiformia). Systematic Entomology 40(4): 745-778. https://doi. org/10.1111/syen.12138 Ronquist F, Teslenko M, Der Mark PV, Ayres DL, Darling AE, Hohna S, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539-542. https://doi.org/10.1093/sysbio/ sys029 Saccone C, De Giorgi C, Gissi C, Pesole G, Reyes A (1999) Evolutionary genomics in Meta- zoa: The mitochondrial DNA as a model system. Gene 238(1): 195-209. https://doi. org/10.1016/S0378-1119(99)00270-X Sheffield NC, Song H, Cameron SL, Whiting MF (2009) Nonstationary evolution and com- positional heterogeneity in beetle mitochondrial phylogenomics. Systematic Biology 58(4): 381-394. https://doi.org/10.1093/sysbio/syp037 Short AEZ, Fika¢ek M (2013) Molecular phylogeny, evolution, and classification of the Hydro- philidae (Coleoptera). Systematic Entomology 38(4): 723-752. https://doi.org/10.1111/ syen.12024 Song F, Li H, Liu GH, Wang W, James P, Colwell DD, Tran A, Gong S, Cai WZ, Shao R (2019) Mitochondrial genome fragmentation unites the parasitic lice of eutherian mam- mals. Systematic Biology 68(3): 430-440. https://doi.org/10.1093/sysbio/syy062 Sun HQ, Zhao WX, Lin RZ, Zhou ZF, Huai WX, Yao YX (2020) The conserved mitochondri- al genome of the jewel beetle (Coleoptera: Buprestidae) and its phylogenetic implications for the suborder Polyphaga. Genomics 112(5): 3713-3721. https://doi.org/10.1016/j. ygeno.2020.04.026 Thompson JD, Higgins DG, Gibson TJ (1994) CLUSTAL W: Improving the sensitivity of progressive multiple sequence alignment through sequence weighting, position-specific gap penalties and weight matrix choice. Nucleic Acids Research 22(22): 4673-4680. https://doi.org/10.1093/nar/22.22.4673 Toyama M (1987) The systematic positions of some buprestid genera (Coleoptera, Bupresti- daé)Elyrrat ho l—1 1. Wang WQ, Huang YX, Bartlett CR, Zhou FM, Meng H, Qin DZ (2019) Characterization of the complete mitochondrial genomes of two species of the genus Aphaena Guérin-Meé- neville (Hemiptera: Fulgoridae) and its phylogenetic implications. International Journal of Biological Macromolecules 141: 29-40. https://doi.org/10.1016/j.ijbiomac.2019.08.222 Wolstenholme DR (1992) Animal mitochondrial DNA: Structure and evolution. International Review of Cytology 141: 173-216. https://doi.org/10.1016/S0074-7696(08)62066-5 Xiao JH, Jia JG, Murphy RW, Huang DW (2011) Rapid evolution of the mitochondrial ge- nome in chalcidoid wasps (Hymenoptera: Chalcidoidea) driven by parasitic lifestyles. PLoS ONE 6(11): e26645. https://doi.org/10.137 1/journal.pone.0026645 212 Zhonghua Wei / ZooKeys 1092: 195-212 (2022) Xiao LE Zhang SD, Long CP, Guo QY, Xu JS, Dai XH, Wang JG (2019) Complete mitog- enome of a leaf-mining buprestid Beetle, Trachys auricollis, and its phylogenetic implica- tions. Genes 10(12): e992. https://doi.org/10.3390/genes10120992 Yan L, Zhang M, Gao Y, Pape T, Zhang D (2017) First mitogenome for the subfamily Milto- gramminae (Diptera: Sarcophagidae) and its phylogenetic implications. European Journal of Entomology 114: 422-429. https://doi.org/10.14411/eje.2017.054 Yu F, Liang AP (2018) The complete mitochondrial genome of Ugyops sp. (Hemiptera: Delpha- cidae). Journal of Insect Science 18(3): e25. https://doi.org/10.1093/jisesa/iey063 Yu P, Cheng X, Ma Y, Yu D, Zhang J (2016) The complete mitochondrial genome of Brachythemis contaminata (Odonata: Libellulidae). Mitochondrial DNA A DNA Mapping Sequencing Analysis 27: 2272-2273. https://doi.org/10.3109/19401736.2014.984176 Zhang D, Gao F, Jakovli¢ I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An inte- erated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348-355. https://doi.org/10.1111/1755-0998.13096 Supplementary material | Figures S1—S7 Authors: Zhonghua Wei Data type: Images (pdf file) Explanation note: Figure $1. The mitogenome maps of Agrilus sichuanus, Coraebus cloueti, Coraebus diminutus, Meliboeus sinae, and Sambus femoralis. Figure S2. The secondary cloverleaf structure for the tRNAs of Agrilus sichuanus. Figure S3. The secondary cloverleaf structure for the tRNAs of Coraebus cloueti. Figure S4. The secondary cloverleaf structure for the tRNAs of Coraebus diminutus. Figure S5. The secondary cloverleaf structure for the tRNAs of Meliboeus sinae. Figure SO. The secondary cloverleaf structure for the tRNAs of Sambus femoralis. Figure S7. Heterogeneous sequence divergence within datasets 13 PCGs and 2 rRNAs of Bu- prestidae species. Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODDbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://doi.org/10.3897/zookeys.1092.80993.suppl1