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Article

Three Complete Mitochondrial Genomes of Ocellarnaca (Orthoptera, Gryllacrididae) and Their Phylogenies

by
Ting Luo
,
Yanting Qin
,
Xiangyi Lu
,
Siyu Pang
and
Xun Bian
*
Key Laboratory of Ecology of Rare and Endangered Species and Environmental Protection (Guangxi Normal University), Ministry of Education, Guilin 541006, China
*
Author to whom correspondence should be addressed.
Biology 2025, 14(9), 1231; https://doi.org/10.3390/biology14091231
Submission received: 17 June 2025 / Revised: 22 August 2025 / Accepted: 28 August 2025 / Published: 10 September 2025
(This article belongs to the Special Issue Progress in Wildlife Conservation, Management and Biological Research)
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Simple Summary

We performed a systematic analysis of mitochondrial genome features in the family Gryllacrididae (also called raspy cricket). The mitochondrial genomes of three species within the genus Ocellarnaca were sequenced, revealing 111 simple sequence repeats (SSRs) across 18 species. Phylogenetic trees based on maximum likelihood and Bayesian inference demonstrated that all species of Ocellarnaca clustered together, forming a monophyletic group, with all branches showing statistically robust support values. The mitogenomic resources for the genus Ocellarnaca were expanded in this study.

Abstract

The Raspy crickets are an interesting group of nocturnal animals that bear femoral-abdominal stridulation and spin silk from the mouthparts. Gryllacridid classification is the subject of ongoing discussion. Here, we present the first mitogenomic sequences for three Ocellarnaca taxa: O. braueri (15,597 bp), O. fuscotessellata (15,607 bp), and O. emeiensis (16,510 bp). Three mitochondrial genomes exhibited the conventional metazoan gene and conserved the characteristic gene order across Gryllacrididae species. Evolutionary selection analyses showed that atp8 was the least evolutionarily constrained mitochondrial gene, whereas cox1 was the most conserved across lineages. The three Ocellarnaca species harbored 5–8 mitochondrial DNA sequence repeats (mtSSRs), falling within the 1–8 range detected in all analyzed Gryllacrididae species. Magnigryllacris and Ocellarnaca exhibited higher mtSSR counts than related genera sharing analogous male abdominal apex morphology. Phylogenomic analyses of 35 mitogenomes from 21 Gryllacrididae species supported Ocellarnaca as monophyletic and a sister to Magnigryllacris (bootstrap = 100%), with O. fuscotessellata resolved as sister to the clade (O. sp. + (O. emeiensis + O. braueri)). This study expands the mitogenomic resources for Ocellarnaca, which will facilitate further resolution of phylogenetic reconstruction within this genus and across Gryllacrididae genera.

1. Introduction

Members of the family Gryllacrididae (also called raspy cricket) [1] are nocturnal animals that hide by day and come out by night. The family contains 972 extant species and 133 genera, primarily distributed in tropical and subtropical regions worldwide [2]. China has 195 species belonging to 30 genera, mainly distributed in southern regions [1,2,3]. As their name suggests, raspy crickets produce a raspy noise during courtship or when threatened, achieved through femoral-abdominal stridulation, wherein the hind femur rubs against pegs on tergites II and III [4]. These animals are also notable for their ability to produce silk from their mouthparts for nest construction, and their exceptional nest-locating capabilities, which ensure that they can return to their shelters easily [5]. However, behavioral, ecological, and applied research on raspy crickets is limited because of their nocturnal habits.
Gryllacrididae classification is the subject of ongoing discussion [1]. Raspy crickets have been traditionally classified into eight distinct types, based on the presence or absence of a median furrow or split on the male ninth abdominal tergite [6]. However, another classification system divided Gryllacrididae into five types based on tegminal venation [7,8]. While the monophyly of Gryllacrididae has been confirmed [9,10], its internal relationships remain unresolved. Observations of vein variation contributed to the identification of three gryllacridid generic groups from Southeast Asia and New Guinea [11]. One year later, a new classification segregated Gryllacrididae into two distinct subfamilies (namely Gryllacridinae and Hyperbaeninae) based on common generic characteristics and distribution [12]. However, in a taxonomic revision, Gryllacridinae was placed under Stenopelmatidae, whereas Hyperbaeninae was regarded as a new synonym of Gryllacridinae [13].
Recent advances in molecular biology and genomics have been crucial in addressing phylogenetic relationships. Mitochondrial genomes (mitogenomes) are effective tools for studying evolution in humans, animals, and plants [14,15,16,17,18,19]. Phylogenomic analyses have provided resolution of the phylogenetic relationships among Orthoptera: (Rhaphidophoridae (Schizodactylidae ((Gryllacrididae (Stenopelmatidae  +  Anostostomatidae))  +  (Prophalangopsidae  +  Tettigoniidae)))) [20]. A recent study also supported the non-monophyletic relationships between the subfamilies Hyperbaeninae and Gryllacridinae based on 13 protein-coding genes (PCGs) [21], aligning with Brunner’s classification but contradicting that of Cadena-Castañeda [12]. Additionally, the monophyly of Homogryllacris Liu, 2007, Furcilarnaca Gorochov, 2004, Ultragryllacris Gorochov & Dawwrueng, 2015, and Capnogryllacris Karny, 1937 was recovered [22]. A phylogenetic analysis of Chinese Gryllacrididae species revealed that the genera Phryganogryllacris Karny, 1937, Capnogryllacris, and Eugryllacris Karny, 1937 failed to form monophyletic groups, resulting in six new genera being proposed [1]. The genus Ocellarnaca Gorochov, 2004 is defined by a brownish or yellowish-brown color, a median ocellus as large as or slightly wider than the antennal sockets, and the ninth abdominal tergite in males having a pair of lobiform processes, each bearing one spine. Ocellarnaca currently includes 17 valid species, but the complete mitogenome of only one species of unknown name has been defined (Ocellarnaca sp., MT849269) [23].
Mitochondrial DNA sequence repeats (mtSSRs) are polymorphic DNA motifs characterized by tandemly repeated short units and elevated mutation rates, enabling their wide employment as stable and versatile genetic markers in molecular studies [24,25].
We assembled and annotated the mitochondrial genomes of three species in this investigation, O. braueri (Griffini, 1911), O. emeiensis Li, Fang, Liu & Li, 2014, and O. fuscotessellata (Karny, 1926), and compared and characterized their structure and nucleotide composition. MtSSRs were analyzed across the family Gryllacrididae to identify stable molecular markers. Furthermore, phylogenetic reconstruction was conducted to elucidate the taxonomic position of the three species relative to other Gryllacrididae species.

2. Materials and Methods

2.1. Sample Acquisition and DNA Extraction

Samples of three Ocellarnaca species were collected in 2019 in the field: O. braueri (GXNUXZ5) and O. fuscotessellata (GXNUXZ7) from Maoershan, Xing’an (25.882° N, 110.490° E, 12 July 2019, collected by Xun Bian), and O. emeiensis (GXNUXZ34) from Rongshui (25.075° N, 109.283° E, 1 August 2019, collected by Wei Bin) (Figure S1). All locations were located in the Guangxi Zhuang Autonomous Region of China. Field-collected specimens were promptly fixed in 100% ethanol and transferred to the laboratory for storage at −18 °C. Subsequently, DNA extraction was performed on hind femur muscle samples using the TIANamp Genomic DNA Kit (Tiangen, Beijing, China) in accordance with the supplier’s protocol. The voucher specimens were deposited at Guangxi Normal University, Guilin, China.

2.2. Next-Generation Sequencing, Genomic Assembly and Analysis

Total DNA was high-throughput sequenced on an Illumina NovaSeq platform, provided by Beijing Berry Genomics Co., Ltd. (Beijing, China). High-quality clean reads were obtained by filtering raw paired-end data through CLC Genomics Workbench 12 (CLC Bio, Aarhus, Denmark) with default parameters [26]. Processed reads were aligned to the mitogenomic reference sequence with Homogryllacris anelytra Shi, Guo & Bian, 2012 (GenBank Accession number NC033998) [10] as a reference. The sequences were assembled using NOVOPlasty v.4.2.1 [27] and annotated by the MITOS web server (http://mitos.bioinf.uni–leipzig.delindex.py, accessed on 10 March 2023) [28]. Mitogenome circular diagrams were generated using the Proksee online platform (accessed on 4 March 2025) [29] and submitted to GenBank with the accession identifiers NC_069863 (O. fuscotessellata), NC_069865 (O. emeiensis), and NC_069864 (O. braueri). Furthermore, the mitogenomes nucleotide composition and codon utilization frequencies of O. braueri, O. fuscotessellata, and O. emeiensis were proven using MEGA v.11.0 [30]. Relative synonymous codon usage (RSCU) was calculated with PhyloSuite v.1.2.3 [31]. AT-skew and GC-skew were calculated using the following formulas: AT-skew = (A − T)/(A + T) and GC-skew = (G − C)/(G + C). The evolutionary rate of each gene was computed using the DnaSP v.6.12.03 [32].
The detection of short mtSSRs was performed using the MIcroSAtellite tool (https://webblast.ipk-gatersleben.de/misa/, accessed on 5 March 2025) [33] on eighteen complete mitogenomes sequenced from Gryllacrididae species, including O. braueri, O. fuscotessellata, and O. emeiensis. The set minimum repeat lengths were as follows: ≥12 for mononucleotides, ≥6 for dinucleotides, ≥4 for trinucleotides, and ≥3 for tetra-, penta-, and hexanucleotides [34]. An interruption of 0 was assumed between two microsatellites. To standardize the comparative analysis across genomes of varying sizes, SSR distributions were normalized to 1 kilobase (kb) genomic intervals using two metrics: relative abundance (RA): the total number of SSR motifs per kb of genome, and relative density (RD): the total length of SSR sequences per genome nucleotide.

2.3. Phylogenetic Analysis

Phylogenetic analyses were conducted, including the three recently obtained mitogenomes of Ocellarnaca, and 32 mitogenomes belonging to 21 species of Gryllacrididae available in the National Center for Biotechnology Information (NCBI) GenBank database (Table S1). Four additional mitogenomes were employed as outgroups (Table S1): Troglophilus neglectus Krauss, 1879 NC_011306 [35] and Rhaphidophora quadrispina Liu & Bian, 2021 NC_067624 [36] from Rhaphidophoridae, Ammopelmatus fuscus (Haldeman, 1852) NC_028058 [37] from Stenopelmatidae, and Tarragoilus diuturnus Gorochov, 2001 NC_021397 [38] from Prophalangopsidae.
Thirteen PCGs and two rRNAs were aligned using MAFFT v7.505 [39]. Although MAFFT alignment does not account for structural constraints, the FFT-NS-1 (fast) algorithm is effective for most global rRNA alignments, delivering rapid processing and accurate results for related species. The alignments of 35 genes from 21 Gryllacrididae species were concatenated to generate datasets for four taxa: (1) the PCG12 matrix with 7642 bp, including the first and second codon positions of PCGs; (2) the PCG123 matrix with 11,463 bp, including all three codon positions of PCGs; (3) the PCG12+2R matrix with 10,007 bp, including the first and second codon positions of PCGs and two rRNA genes; and (4) the PCG123+2R matrix with 13,828 bp, including all three codon positions of PCGs and two rRNA genes. Heterogeneity analysis was performed using AliGROOVE v.1.08 [40]. PhyloSuite v.1.2.3 [31] was used to construct phylogenetic trees using the Bayesian inference (BI) and maximum likelihood (ML) methods. For ML analysis, IQ-TREE v2.2.0 [41] was used with 1000 replicates, where the optimal nucleotide substitution model was identified as TIM + F + I + G4 [42] (Table S2). Bayesian inference was performed in MrBayes v3.2.7a [43] through two parallel MCMC simulations. The optimal substitution model (GTR + F + I + G4), identified by ModelFinder [42] under the BIC framework (Table S2), involves 2 million generations with sampling every 1000 generations. After discarding the first 25% of generations as burn-in, a consensus topology was reconstructed from the remaining trees, with nodal support assessed via posterior probabilities. The BI analysis was considered to have achieved convergence when the average standard deviation of split frequencies (ASDSF) fell below 0.01 [42]. We employed the iTOL online tool (https://itol.embl.de, accessed 7 May 2025) to enhance topological clarity [44].

3. Results and Discussion

3.1. Genome Structure and Composition of the Three Raspy Cricket Species

The complete mitogenomes of O. braueri, O. emeiensis, and O. fuscotessellata had lengths of 15,597 bp, 16,510 bp, and 15,607 bp, respectively. Three mitogenomes exhibited the typical 37 genes, comprising 13 PCGs, 22 tRNAs, and 2 rRNAs, along with a control region. Gene distribution showed 23 genes encoded on the J-strand and 14 genes on the N-strand (Figure 1).
The three species exhibited a similar nucleotide composition, with a strong bias towards AT base composition, ranging from 71.87% to 74.68% (mean value = 73.22%), consistent with the pattern observed in most orthopteran insects [10,45,46,47]. RSCU analysis identified UUA (Leu2) as the most preferred codon, with RSCU value varying from 12.02 to 12.88. By contrast, UGU (Cys) was the least used, with the highest values utilizing U and A, meaning an overall skew toward A and T in the three mitogenomes (Figure 2).

3.2. Protein-Coding Genes and Evolutionary Rates

Nine PCGs (nad2, cox1, cox2, atp8, atp6, cox3, nad3, nad6, and cytb) were encoded on the J-strand, whereas four PCGs (nad5, nad4, nad4l, and nad1) were encoded on the N-strand (Figure 1). Among all PCGs, nad5 exhibited the maximum length (1735–1738 bp), while atp8 was the least (159 bp). Eight PCGs (nad2, cox1, cox2, atp8, atp6, nad4l, nad3, and cytb) exhibited complete length conservation across the three species. Mitochondrial initiation codons predominantly utilized the ATN, except for nad1, where TTG was identified in O. fuscotessellata (Table S3) and O. braueri (Table S4). Although TAA served as the canonical termination codon for most PCGs, truncated termination signals (T/TA) were observed in cox2, nad4, and nad5 of O. braueri and O. emeiensis (Tables S4 and S5, respectively). This incomplete termination pattern was also conserved in cox2 and nad5 of O. fuscotessellata (Table S3).
We performed Ka/Ks analysis of the 13 PCGs of the mitogenomes to investigate the evolutionary selection constraints of the three Ocellarnaca species (Figure 3). All PCGs exhibited Ka/Ks values less than 1, demonstrating purifying selection across the genomes [48]. Among them, atp8 of O. braueri and O. fuscotessellata exhibited the most elevated Ka/Ks ratios, indicating the relaxed purifying selection. By contrast, cox1 of all three species had the lowest Ka/Ks ratio, suggesting that harmful mutations were eliminated by purifying selection to preserve the integrity of the complex core subunits [49,50].

3.3. RNAs and Control Regions

The total lengths of 22 tRNAs of the three species ranged between 1451 bp (O. fuscotessellata; Table S3) and 1455 bp (O. braueri and O. emeiensis; Tables S4 and S5, respectively), among the 22 tRNA genes spanning from 62 to 74 bp. Fourteen tRNA genes (trnI, trnM, trnW, trnL2, trnK, trnD, trnG, trnA, trnR, trnN, trnS1, trnE, trnT, and trnS2) were J-strand encoded, and eight (trnQ, trnC, trnY, trnF, trnH, trnP, trnL1, and trnV) were N-strand located. Except for trnS1, the tRNAs of the three mitogenomes lacked a dihydrouridine (DHU) arm, forming a typical cloverleaf structure common in metazoans [51,52,53]. Moreover, 20 wobble base pairs (G-U) were found in O. braueri, 24 in O. fuscotessellata, and 23 in O. emeiensis. The rRNAs (12S and 16S) were transcribed from the N-strand, exhibiting size variations from 1299 to 1355 bp, located at a conserved position between trnL1 and trnV. The length of 12S rRNA was 774–776 bp, located between trnV and CR. The length heterogeneity (726–1703 bp) in the regulatory regions located at the 12S rRNA-trnI junction. Thirty-five SSR loci are present in the control region, among which 13 exhibited mutations (Table S6), suggesting that dynamic changes in repetitive sequences may represent one of the key factors contributing to length variation in this region.

3.4. Mitochondrial Microsatellites (mtSSRs)

To conduct mtSSR analysis for the whole mitogenomes, we used the three circular mitogenomes that contained the control regions. The mitogenomes of O. braueri, O. fuscotessellata, and O. emeiensis contained 5–8 mtSSRs, a range that fell within the range of 1–8 mtSSRs observed across all 18 analyzed Gryllacrididae species. O. braueri and O. fuscotessellata had the highest number of mtSSRs and presented 8 mtSSRs each. By contrast, O. emeiensis had only five mtSSRs, of which one was located in trnD and another in the control region (Table 1). A total of 111 mtSSRs were found, representing less than 1% of the mitogenomes analyzed and consistent with nuclear genome microsatellite densities (0.02–3.1%) in insects. A total of 49 SSR mutations were detected in this study, suggesting their potential utility as SSR markers for further investigation (Table S6). Analysis revealed variations in the relative abundance of different SSRs across genera. The average number of mtSSRs in the genera Magnigryllacris Li, Yin & He, 2024 and Ocellarnaca was higher than in other genera with a similarly shaped median ocellus and a male abdominal apex split along the midline (Table 2).
The RA and RD of the SSRs showed marked variation across mitogenomes. RA ranged from 0.062 mtSSR/kb (Furcilarnaca wufengensis Bian, Shi & Guo, 2013, OL519601) to 0.513 mtSSR/kb (Ocellarnaca emeiensis, NC069865), and RD spanned from 0.757 bp/kb (Marthogryllacris erythrocephala maculatis (Liu, Lu & Bian, 2022) OL979481) to 6.575 bp/kb (Dracogryllacris spinosa (Li, Liu & Li, 2014) OL944077) (Table 2). When comparing the number of SSRs within genomes, we found that the percentages of di- (31.5%) and tetranucleotide SSRs (41.4%) were higher than mono- (7.2%) and trinucleotide (19.8%) (Figure 4).
Among the identified mtSSRs, 50.5% were localized within PCGs (with the nad6, nad4, and nad5 exhibiting the highest mtSSR contents), 45% in non-coding regions (including tRNAs, rRNAs, and control regions), and 4.5% in the intergenic spacer region (IR). The highest abundance of microsatellites corresponded to nad6 (19.8%), followed by nad4 (9%) and nad5 (9%) (Figure 5). A substantial proportion of the coding-region mtSSRs in Gryllacrididae were found in Nicotinamide adenine dinucleotide (NADH) dehydrogenase genes involved in the electron transport system. As a critical redox cofactor, NADH mediates fundamental biological processes, including cellular redox homeostasis, bioenergetic metabolism, mitochondrial bioenergetics, transcriptional regulation, and cell signaling cascades, with its redox couples being indispensable for maintaining physiological functions [54,55,56].

3.5. Substitution Saturation and Heterogeneity Analysis

Heterogeneity among the four datasets (PCG12, PCG123, PCG12+2R, and PCG123+2R) indicated that the PCG12 and PCG123 datasets were unsaturated and had stronger phylogenetic signals. Therefore, they were more suitable for phylogenetic relationships analysis (Figure 6).

3.6. Phylogenetic Analysis

Both ML and BI analyses yielded identical tree topologies, supporting the monophyly of Ocellarnaca (ML and BI analyses of PCG123 in Figure 7; PCG12, PCG123+2R, and PCG12+2R analyses in Figures S2, S3, and S4, respectively). Within Ocellarnaca, O. fuscotessellata formed a sister group to the clade comprising O. sp. and the (O. emeiensis + O. braueri) subclade. Dracogryllacris Li, Yin & He, 2024 comprises two clades: (A) Dracogryllacris melanocrania (Karny, 1929) (KX057731), and (B) the remaining sequences within Dracogryllacris. The monophyly of Homogryllacris, Furcilarnaca, Ocellarnaca, Ultragryllacris, and Marthogryllacris in Gryllacrididae was recovered, with the following phylogenetic relationship: Camptonotus + ((((Sericgryllacris + Phryganogryllacris) + Homogryllacris) + Furcilarnaca) + ((Magnigryllacris + Ocellarnaca) + (Ultragryllacris + ((clade A + Marthogryllacris) + clade B)))), similar to that found in former phylogenetic studies [21,22]. The results of this study, consistent with those of Li [1] and Liu [21], which did not recover the monophyly of Hyperbaeninae and Gryllacridinae (Figure 7), in contrast to the classification proposed by Cadena-Castañeda [12]. Cadena-Castañeda divided Gryllacrididae into Hyperbaeninae and Gryllacridinae based on morphological characters, including body size, wing morphology, tarsal structure, and male genitalia/ovipositor features [12]. However, Liu [21] suggested that the presence or absence of a median groove or midline division on the male ninth abdominal tergite- a character that appears to better reflect Gryllacrididae’s true phylogenetic relationships.
However, our results also indicated that taxonomy at the species level remains partially uncertain. Specifically, Furcilarnaca chirurga (ON055390), F. chirurga (OL502168), and F. chirurga (OL470659) were not grouped on the same branch (Figure 7), because the F. chirurga specimen (ON055390) may have been misidentified. Regarding previous studies of Gryllacrididae, few have considered multiple datasets to explore the classification of raspy crickets [21,22]. Molecular or morphological data alone were less likely to provide a complete evolutionary history, even though they could yield unique trees [57,58,59,60]. Future studies could integrate functional morphology with evolutionary models to reveal the evolutionary mechanisms underpinning key adaptive traits in Orthoptera.

3.7. Implications for Conservation and Monitoring

Despite its non-threatened status, Ocellarnaca, as a common gryllacridid, plays a significant ecological role in forest ecosystems by exerting influence on higher trophic levels through its population dynamics. Its population dynamics may influence higher trophic levels. Due to its specific microhabitat humidity requirements, population fluctuations of Ocellarnaca could serve as indicators of forest floor ecosystem health, warranting its inclusion in long-term biodiversity monitoring programs.

4. Conclusions

This study reports the complete mitochondrial genomes of three Ocellarnaca species (O. braueri, O. fuscotessellata, and O. emeiensis), which exhibit conserved genomic characteristics including size uniformity, AT-rich composition, consistent strand asymmetry (AT-skew and GC-skew), and parallel codon usage patterns. All of the tRNAs maintained complete secondary structures, whereas trnS1 lacks a DHU arm. A total of 111 Gryllacrididae mtSSRs were identified. Among the three examined species of Ocellarnaca, mtSSR counts ranged from 5 to 8. Both Ocellarnaca and Magnigryllacris exhibited significantly higher mtSSR numbers than lineages with equivalent male abdominal valve lengths. Over 50% of mtSSRs were localized within PCGs, with NADH dehydrogenase genes harboring 91.1% of these PCG-associated repeats. Phylogenetic analysis of 21 Gryllacrididae species based on mitochondrial data reconstructed Ocellarnaca as monophyletic, within which three species clustered with Ocellarnaca sp., and recovered a sister-group relationship with Magnigryllacris. This study supports the need to reevaluate the traditional boundaries between Hyperbaeninae and Gryllacridinae in raspy cricket taxonomy, providing insights for future taxonomic classifications. This study provides fundamental biological data for Ocellarnaca, a common yet ecologically significant taxon. While facing no extinction risk, its habitat necessitates inclusion in regional biodiversity monitoring frameworks to safeguard against potential future anthropogenic or environmental disturbances.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/biology14091231/s1, Figure S1: Ecological photos of three Ocellarnaca species; Figure S2: Phylogenetic tree of Gryllacrididae inferred from the ML and BI analyses of the PCG12; Figure S3: Phylogenetic tree of Gryllacrididae inferred from the ML and BI analyses of the PCG123+2R; Figure S4: Phylogenetic tree of Gryllacrididae inferred from the ML and BI analyses of the PCG12+2R; Table S1: List of mitogenomes used for phylogenetic analysis. Table S2: The best-fit partition model of ML and BI analyses; Table S3: Mitochondrial genome content, length, and codon information of Ocellarnaca fuscotessellata; Table S4: Mitochondrial genome content, length, and codon information of Ocellarnaca braueri; Table S5: Mitochondrial genome content, length, and codon information of Ocellarnaca emeiensis; Table S6: Detailed information on mtSSR mutations.

Author Contributions

Conceptualization, T.L., Y.Q. and X.L.; investigation, formal analysis, T.L. and S.P.; writing—original draft T.L. and Y.Q.; resources, writing-review and editing, X.B. All authors have read and agreed to the published version of the manuscript.

Funding

This study was supported by the National Natural Science Foundation of China (Grant No. 32360126 and 31802000) for X.B., Innovation Project of Guangxi Graduate Education (Grant No. XYCS2025109) for T.L., and Innovation Project of Guangxi Graduate Education (Grant No. XYCS2025110) for Y.Q.

Institutional Review Board Statement

The retrieval and processing of samples were exempt from permit requirements because the invertebrate species involved are not subject to regulatory protection.

Informed Consent Statement

Informed consent was obtained from all subjects involved in the study.

Data Availability Statement

The original data presented in the study are openly available in the National Center for Biotechnology Information (NCBI, https://www.ncbi.nlm.nih.gov/, accessed on 16 January 2025) genome database.

Conflicts of Interest

The authors declare no conflicts of interest.

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Biology 14 01231 g001
Figure 1. Genome circle map of three Ocellarnaca species. (A): O. braueri; (B): O. fuscotessellata; and (C): O. emeiensis. The various gene regions are represented in distinct hues in the depicted genetic sequence. Twenty-two tRNAs are presented through their respective amino acid abbreviations. The J-strand is shown on the outer ring, while the N-strand is displayed on the inner ring. The GC content is graphically represented in black.
Figure 1. Genome circle map of three Ocellarnaca species. (A): O. braueri; (B): O. fuscotessellata; and (C): O. emeiensis. The various gene regions are represented in distinct hues in the depicted genetic sequence. Twenty-two tRNAs are presented through their respective amino acid abbreviations. The J-strand is shown on the outer ring, while the N-strand is displayed on the inner ring. The GC content is graphically represented in black.
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Biology 14 01231 g002
Figure 2. Relative synonymous codon usage (RSCU) of the three complete Ocellarnaca mitogenomes. Note, RSCU is a relative value calculated from codon usage counts, which labels it as ‘(relative value, no unit)’ to indicate that it is a relative metric.
Figure 2. Relative synonymous codon usage (RSCU) of the three complete Ocellarnaca mitogenomes. Note, RSCU is a relative value calculated from codon usage counts, which labels it as ‘(relative value, no unit)’ to indicate that it is a relative metric.
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Biology 14 01231 g003
Figure 3. The Ka/Ks ratios of 13 PCGs of the three complete Ocellarnaca mitogenomes. Note, Ka: nonsynonymous substitution rate, Ks: the synonymous substitution rate.
Figure 3. The Ka/Ks ratios of 13 PCGs of the three complete Ocellarnaca mitogenomes. Note, Ka: nonsynonymous substitution rate, Ks: the synonymous substitution rate.
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Biology 14 01231 g004
Figure 4. (A): The mitochondrial DNA sequence repeats in 18 Gryllacrididae species. (B): Relative percentage of four sizes of mtSSRs in 18 Gryllacrididae mitogenomes. The percentages of mono-, di-, tri-, and tetranucleotides are shown in different colors. The Gryllacrididae species are shown to the left.
Figure 4. (A): The mitochondrial DNA sequence repeats in 18 Gryllacrididae species. (B): Relative percentage of four sizes of mtSSRs in 18 Gryllacrididae mitogenomes. The percentages of mono-, di-, tri-, and tetranucleotides are shown in different colors. The Gryllacrididae species are shown to the left.
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Figure 5. Location of mtSSRs in the mitogenomes of 18 Gryllacrididae species.
Figure 5. Location of mtSSRs in the mitogenomes of 18 Gryllacrididae species.
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Figure 6. Evolutionary rate heterogeneity among the four Gryllacrididae datasets was assessed using AliGROOVE similarity analysis. The algorithm computes pairwise comparison scores ranging from −1 (red; indicating significant evolutionary rate divergence) to +1 (blue; representing perfect rate congruence).
Figure 6. Evolutionary rate heterogeneity among the four Gryllacrididae datasets was assessed using AliGROOVE similarity analysis. The algorithm computes pairwise comparison scores ranging from −1 (red; indicating significant evolutionary rate divergence) to +1 (blue; representing perfect rate congruence).
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Figure 7. Phylogenetic tree of Gryllacrididae inferred from the maximum likelihood (ML) and Bayesian inference (BI) analyses of the PCG123. Numbers on the branches show bootstrap values. Map the morphological characteristics, the presence or absence of a median groove or midline division on the male ninth abdominal tergite across the two clades onto the phylogenetic tree.
Figure 7. Phylogenetic tree of Gryllacrididae inferred from the maximum likelihood (ML) and Bayesian inference (BI) analyses of the PCG123. Numbers on the branches show bootstrap values. Map the morphological characteristics, the presence or absence of a median groove or midline division on the male ninth abdominal tergite across the two clades onto the phylogenetic tree.
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Table 1. The identified mitogenome sequence repeats (mtSSRs) of three species of Ocellarnaca.
Table 1. The identified mitogenome sequence repeats (mtSSRs) of three species of Ocellarnaca.
SpeciesmtSSRRepeatsSize (bp)StartEndLocation Region
Ocellarnaca braueri
(NC_069864)		(ATT)441238763887IR
(TAA)441299489959nad6
(TTAT)331210,12410,135nad6
(TAAA)331213,84713,858rrnL
(AATA)331214,57714,588rrnS
(TA)881615,23615,251CR
(AT)771415,47615,489CR
(AT)771415,50815,521CR
Ocellarnaca emeiensis
(NC_069865)		(ATTA)331238303841trnD
(T)12121216,00316,014CR
(TA)771416,10316,116CR
(AT)771416,34116,354CR
(AT)661216,37316,384CR
Ocellarnaca fuscotessellata
(NC_069863)		(ATT)441238803891atp8
(AT)881655765591IR
(TTAT)331263216332trnE
(ATT)441264076418nad5
(TAAA)331290209031nad4
(TTAT)331210,14110,152nad6
(CA)771411,67811,691IR
(AT)771415,47215,485CR
Table 2. SSR distribution patterns in 18 Gryllacrididae species, with quantitative characterization of repeat number, total length, relative abundance, and relative density.
Table 2. SSR distribution patterns in 18 Gryllacrididae species, with quantitative characterization of repeat number, total length, relative abundance, and relative density.
SpeciesKbTotal Number of SSRRATotal Length of SSR (bp)RD
Camptonotus carolinensis (NC_028060)15.21130.197225692503.287095
Dracogryllacris melanocrania (KX057731)16.13640.24789291362.231036
Dracogryllacris melanocrania (OL944079)15.71140.254598689241.527592
Dracogryllacris nigromarginat (OL944078)16.63950.300498828362.163592
Dracogryllacris nigromarginata (OL978587)16.21840.246639536503.082994
Dracogryllacris nigromarginata (OL978588)15.77630.190162272120.760649
Dracogryllacris spinosa (OL944076)15.82520.126382306603.791469
Dracogryllacris spinosa (OL944077)15.81720.1264462291046.575204
Dracogryllacris spinosa (OK539822)16.50520.1211754011046.301121
Furcilarnaca armata (NC_067618)15.78730.190029771261.646925
Furcilarnaca armata (OL544941)15.83020.126342388483.032217
Furcilarnaca armata (OL826861)15.67720.127575429120.765453
Furcilarnaca chirurga (NC_067622)15.45440.258832665241.552996
Furcilarnaca chirurga (OL502168)15.44140.25905058120.777152
Furcilarnaca chirurga (ON055390)15.55010.064308682241.543408
Furcilarnaca wufengensis (NC_067623)15.95410.062680206382.381848
Furcilarnaca wufengensis (OL519601)16.02210.062414181261.622769
Furcilarnaca wufengensis (OL826860)15.47810.06460783261.679804
Homogryllacris anelytra (NC_033998)15.70620.12733987603.820196
Homogryllacris yunnana (OM731663)16.20950.308470603482.961318
Magnigryllacris tiga (MZ540210)15.51370.451234449644.125572
Marthogryllacris erythrocephala maculatis (OL876382)16.16330.185609107442.722267
Marthogryllacris erythrocephala maculatis (OL979480)16.17130.185517284865.318162
Marthogryllacris erythrocephala maculatis (OL979481)15.84530.189334175120.757337
Ocellarnaca braueri (NC_069864)15.59780.512919151503.205745
Ocellarnaca emeiensis (NC_069865)16.51050.512590504382.301635
Ocellarnaca fuscotessellata (NC_069863)15.60780.30284676402.562953
Ocellarnaca sp. (MT849269)16.15730.185678034382.351922
Phryganogryllacris superangulata (NC_069838)15.97640.250375563362.253380
Sericgryllacris xiai (NC_033994)15.87630.188964475382.393550
Ultragryllacris rubricapitis (OM731664)15.55850.321378069664.242191
Ultragryllacris rubricapitis (OM683271)16.62530.180451128603.609023
Ultragryllacris rubricapitis (OM683272)15.76630.190282887120.761132
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Luo, T.; Qin, Y.; Lu, X.; Pang, S.; Bian, X. Three Complete Mitochondrial Genomes of Ocellarnaca (Orthoptera, Gryllacrididae) and Their Phylogenies. Biology 2025, 14, 1231. https://doi.org/10.3390/biology14091231

AMA Style

Luo T, Qin Y, Lu X, Pang S, Bian X. Three Complete Mitochondrial Genomes of Ocellarnaca (Orthoptera, Gryllacrididae) and Their Phylogenies. Biology. 2025; 14(9):1231. https://doi.org/10.3390/biology14091231

Chicago/Turabian Style

Luo, Ting, Yanting Qin, Xiangyi Lu, Siyu Pang, and Xun Bian. 2025. "Three Complete Mitochondrial Genomes of Ocellarnaca (Orthoptera, Gryllacrididae) and Their Phylogenies" Biology 14, no. 9: 1231. https://doi.org/10.3390/biology14091231

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Luo, T., Qin, Y., Lu, X., Pang, S., & Bian, X. (2025). Three Complete Mitochondrial Genomes of Ocellarnaca (Orthoptera, Gryllacrididae) and Their Phylogenies. Biology, 14(9), 1231. https://doi.org/10.3390/biology14091231

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