Characterization of the complete mitochondrial genome of an endemic species in China, Aulocera merlina (Lepidoptera: Nymphalidae: Satyrinae) and phylogenetic analysis within Satyrinae.

The mitochondrial genome (mitogenome) has been extensively used as molecular markers in determining the insect phylogenetic relationships. In order to resolve the relationships among tribes and subtribes of Satyrinae at the mitochondrial genomic level, we obtained the complete mitogenome of Aulocera merlina (Oberthür, 1890) (Lepidoptera: Nymphalidae: Satyrinae) with a size of 15,259 bp. The mitogenome consisted of 37 typical genes, including 13 protein-coding genes (PCGs), 2 ribosomal RNA genes (rRNAs), 22 transfer RNA genes (tRNAs), and an A + T-rich region. The gene organization and arrangement were similar to those of all other known Satyrinae mitogenomes. All PCGs were initiated with the canonical codon pattern ATN, except for the cox1 gene, which used an atypical CGA codon. Nine PCGs used the complete stop codon TAA, while the remaining PCGs (cox1, cox2, nad4, and nad5) were terminated with a single T nucleotide. The canonical cloverleaf secondary structures were found in all tRNAs, except for trnS1 which lacked a dihydrouridine arm. The 448 bp A + T-rich region was located between rrnS and trnM, and it included the motif ATAGA followed by a 19-bp poly-T stretch and a microsatellite-like (TA)6 element preceded by the ATTTA motif. The phylogenetic tree, inferred using Bayesian inference and maximum likelihood methods, generated similar tree topologies, revealing well-supported monophyletic groups at the tribe level and recovering the relationship ((Satyrini + Melanitini) + ((Amathusiini + Elymniini) + Zetherini)). The close relationship between Satyrina and Melanargiina within the Satyrini was widely accepted. Additionally, Lethina, Parargina, and Mycalesina were closely related and collectively formed a sister group to Coenonymphina. Moreover, A. merlina was closely related to Oeneis buddha within the Satyrina. These findings will provide valuable information for future studies aiming to elucidate the phylogenetic relationships of Satyrinae.

The complete mitochondrial genome of Stibochiona nicea (Gray, 1846) (Lepidoptera: Nymphalidae) and phylogenetic analysis.

In this study, the complete mitochondrial genome (mitogenome) of Stibochiona nicea (Gray, 1846) (Lepidoptera: Nymphalidae) was first reported with 15,298 bp in size, containing 13 protein-coding genes (PCGs), 22 tRNA genes, two rRNA genes (rrnL and rrnS), and one control region. The nucleotide composition of the entire mitogenome is highly A + T biased (81.5%). The gene content and arrangement of the newly sequenced mitogenome are identical to those of the other available mitogenomes of Nymphalidae. All PCGs start with the conventional ATN codons, except for cox1 initiating with atypical CGA(R). Nine PCGs (atp8, atp6, cox3, nad1, nad2, nad3, nad4l, nad6, and cob) utilize a typical stop codon TAA, whereas the remaining PCGs (cox1, cox2, nad4, and nad5) end with an incomplete stop codon T-. Phylogenetic analysis revealed that S. nicea is closely related to Dichorragia nesimachus within Pseudergolinae, which further forms the sister group to the grouping of (Nymphalinae + (Cyrestinae + (Biblidinae + Apaturinae))). The complete mitogenome of S. nicea will provide useful genetic information for improving the taxonomic system and phylogenetics of Nymphalidae.

Complete mitochondrial genome of the Woodland Brown, Lopinga achine Scopoli, 1763 (Nymphalidae: Satyrinae) and its phylogenetic analysis.

In this study, the complete mitochondrial genome (mitogenome) of the Woodland Brown, Lopinga achine Scopoli, 1763 (Nymphalidae: Satyrinae) was determined to be 15,284 bp in size, including 37 typical mitochondrial genes and a control region. The gene content and arrangement of the mitogenome are identical to that of the majority of other sequenced nymphalids. All protein-coding genes (PCGs) are started with the conventional ATN codons, except for cox1 gene which is initiated by atypical CGA(R) codon. Nine PCGs use a typical stop codon of TAA, whereas the remaining PCGs (cox1, cox2, nad4, nad5) end with an incomplete T. The length of rrnL and rrnS are 1333 and 755 bp, respectively, separated by trnV. The phylogenetic tree inferred with Bayesian inference method reveals the phylogenetic relationships among the four tribes of Satyrinae analyzed as ((Satyrini + Melanitini) + (Elymniini + Amathusiini)). The newly sequenced species L. achine was clustered together with other two species of Parargina and formed a sister group with two species of the genus Lethe within Satyrini.

The complete mitochondrial genome sequence of Dodona eugenes (Lepidoptera: Riodinidae).

The complete mitochondrial genome (mitogenome) sequence of Dodona eugenes (Lepidoptera: Riodinidae) was determined and analyzed. The mitogenome is 15,680 bp in length with consisting of 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, two ribosomal RNA genes (rrnL and rrnS), and one AT-rich region. The gene content, orientation, and order are identical to that of the majority of other lepidopteran insects. The D. eugenes mitogenome includes a cox1 gene with an atypical CGA(R) start codon and three genes (cox1, nad5, and nad4) exhibiting incomplete stop codons. All tRNAs have a typical secondary cloverleaf structure, except for trnS1 which lacks the dihydrouridine (DHU) arm. The 825-bp long AT-rich region is the longest among sequenced riodinids, which range from 349 to 423 bp. The conclusion of phylogenetic analysis highly supported the monophyly of Riodinidae, which is standing as the sister of the family Lycaenidae.

Dated phylogeny and dispersal history of the butterfly subfamily Nymphalinae (Lepidoptera: Nymphalidae).

The origin and dispersal history of the large butterfly subfamily Nymphalinae are not fully understood, due to internal phylogenetic and time calibration issues. We conducted phylogenetic and dating analyses using mitochondrial and nuclear genes of biogeographically diverse groups of the Nymphalinae in order to resolve some controversial relationships and the paleobiogeographic pattern of the subfamily. Our results support the sister relationship of Vanessa (Tribe Nymphalini) and the Nymphalis-group, and the grouping of the three old-world genera (Rhinopalpa, Kallimoides and Vanessula) within Tribe Victorinini. Molecular dating analyses invoking two additional calibrations under the butterfly-host plant coevolutionary scenarios result in a relatively deeper divergence of the subfamily's two major clades (Nymphalini and the Kallimoids), compatible with the Cretaceous floral turnover scenario during the so-called Cretaceous Terrestrial Revolution. Phylobiogeographic analyses reveal that the Oriental region is probably the center of early divergences for Nymphalinae after the Cretaceous-Paleogene (K-Pg) mass extinction, followed by repeated dispersals into the rest of the Old World and the New World during various periods beginning in Eocene. The biogeographic history indicates that temperature changes and host-plant diversification may have facilitated the dispersals of this butterfly subfamily, with accelerated global colonization during the middle to late Miocene.

The complete mitochondrial genome of comma, polygonia c-aureum (Lepidoptera: Nymphalidae: Nymphalinae).

The mitochondrial genome (mitogenome) of Polygonia c-aureum (Lepidoptera: Nymphalidae: Nymphalinae) is determined to be 15,209 bp in length and shows AT bias (80.6%). Similar to other butterflies, it contains 37 typical mitochondrial genes and one AT-rich region (D-loop). All protein-coding genes (PCGs) started with ATN, except for cox1 gene with CGA(R), which is often found in other butterflies, and seven PCGs harbour the typical stop codon TAA, whereas cox1, cox2, nad3, nad5, nad4 and nad1 end with a single T. The rrnL and rrnS genes are 1332 bp and 773 bp in length, respectively. The 342 bp AT-rich region contains non-repetitive sequences, but harbour several features common to the lepidopterans, including the motif ATAGA followed by a 19-bp poly-T stretch and a microsatellite-like (TA)8 element preceded by the ATTTA motif. The complete mitogenome sequence provided here would be useful for further understanding the taxonomy and phylogeny of Nymphalinae.

The complete mitochondrial genome of Callerebia suroia (Lepidoptera: Nymphalidae: Satyrinae).

The complete mitochondrial genome (mitogenome) of Callerebia suroia (Lepidoptera: Nymphalidae: Satyrinae) was determined and analyzed in this paper. The circular genome is 15,208 bp long, including 37 typical mitochondrial genes and one non-coding AT-rich region. All protein-coding genes (PCGs) started with ATN, except for COI gene with CGA(R), which is often found in other butterflies; nine PCGs harbor the typical stop codon TAA, whereas COI, COII, ND5 and ND4 end with a single T. All tRNA genes display typical secondary clover-leaf structures, except for tRNA(Ser)(AGN), whose dihydrouridine (DHU) arm is replaced by a simple loop. The lrRNA and srRNA genes are 1,347 bp and 753 bp in length, with their AT contents of 84.4% and 85.4%, respectively. The 417 bp AT-rich region contains non repetitive sequences, but harbor several features common to the lepidopterans, including the motif ATAGA followed by a 19-bp poly-T stretch and a microsatellite-like (TA)8 element preceded by the ATTTA motif.

Morphological characters are compatible with mitogenomic data in resolving the phylogeny of nymphalid butterflies (lepidoptera: papilionoidea: nymphalidae).

Nymphalidae is the largest family of butterflies with their phylogenetic relationships not adequately approached to date. The mitochondrial genomes (mitogenomes) of 11 new nymphalid species were reported and a comparative mitogenomic analysis was conducted together with other 22 available nymphalid mitogenomes. A phylogenetic analysis of the 33 species from all 13 currently recognized nymphalid subfamilies was done based on the mitogenomic data set with three Lycaenidae species as the outgroups. The mitogenome comparison showed that the eleven new mitogenomes were similar with those of other butterflies in gene content and order. The reconstructed phylogenetic trees reveal that the nymphalids are made up of five major clades (the nymphaline, heliconiine, satyrine, danaine and libytheine clades), with sister relationship between subfamilies Cyrestinae and Biblidinae, and most likely between subfamilies Morphinae and Satyrinae. This whole mitogenome-based phylogeny is generally congruent with those of former studies based on nuclear-gene and mitogenomic analyses, but differs considerably from the result of morphological cladistic analysis, such as the basal position of Libytheinae in morpho-phylogeny is not confirmed in molecular studies. However, we found that the mitogenomic phylogeny established herein is compatible with selected morphological characters (including developmental and adult morpho-characters).

The complete mitochondrial genome of Blue Pansy, Junonia orithya (Lepidoptera: Nymphalidae: Nymphalinae).

The complete mitochondrial genome (mitogenome) of Junonia orithya Linnaeus (Lepidoptera: Nymphalidae: Nymphalinae) is determined to be 15,214 bp in length, including 37 typical mitochondrial genes and an AT-rich region. Its gene order and orientation are identical to those of other butterfly species. All PCGs are initiated by typical ATN codons, except for cox1 gene which is started by CGA codon. Nine genes use complete termination codon (TAA), whereas the cox1. cox2. nad1 and nad4 genes end with single T. Except for trnS1(AGN), all tRNA genes display typical secondary cloverleaf structures as those of other insects. The 331 bp long AT-rich region contains several features common to the other lepidopterans, such as the ATAGA motif followed by a 18 bp poly-T stretch, two microsatellite-like (TA)9 elements, a 5 bp poly-A stretch immediately upstream of trnM gene.

The complete mitochondrial genome of Gonepteryx rhamni (Lepidoptera: Pieridae: Coliadinae).

The complete mitochondrial genome (mitogenome) of Linnaeus Gonepteryx rhamni (Lepidoptera: Pieridae: Coliadinae) is a circular molecule of 15,023 bp in length, containing 37 typical coding genes and one non-coding AT-rich region. Its gene order and content are identical to the common type found in most insect mitogenomes. All protein coding genes (PCGs) start with a typical ATN initiation codon, except for the cox1, which use CGA as its start codon. Nine genes use standard complete termination codon (TAA), whereas the cox1, cox2, nad4 and nad5 genes end with single T. Except for trnS1(AGN), all tRNA genes display typical secondary cloverleaf structures as those of other insects. Additionally, the 371 bp long AT-rich region contains a few structures common to the other lepidopterons, such as the motif ATAGA followed by a 17 bp poly-T stretch, a microsatellite-like (AT)10 element preceded by the ATTTA motif, and a 10 bp poly-A presented immediately upstream of trnM gene.

The first mitochondrial genome for the butterfly family Riodinidae (Abisara fylloides) and its systematic implications.

The Riodinidae is one of the lepidopteran butterfly families. This study describes the complete mitochondrial genome of the butterfly species Abisara fylloides, the first mitochondrial genome of the Riodinidae family. The results show that the entire mitochondrial genome of A. fylloides is 15 301 bp in length, and contains 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNA genes and a 423 bp A+T-rich region. The gene content, orientation and order are identical to the majority of other lepidopteran insects. Phylogenetic reconstruction was conducted using the concatenated 13 protein-coding gene (PCG) sequences of 19 available butterfly species covering all the five butterfly families (Papilionidae, Nymphalidae, Peridae, Lycaenidae and Riodinidae). Both maximum likelihood and Bayesian inference analyses highly supported the monophyly of Lycaenidae+Riodinidae, which was standing as the sister of Nymphalidae. In addition, we propose that the riodinids be categorized into the family Lycaenidae as a subfamilial taxon. The Riodinidae is one of the lepidopteran butterfly families. This study describes the complete mitochondrial genome of the butterfly species Abisara fylloides, the first mitochondrial genome of the Riodinidae family. The results show that the entire mitochondrial genome of A. fylloides is 15 301 bp in length, and contains 13 protein-coding genes, 2 ribosomal RNA genes, 22 transfer RNA genes and a 423 bp A+T-rich region. The gene content, orientation and order are identical to the majority of other lepidopteran insects. Phylogenetic reconstruction was conducted using the concatenated 13 protein-coding gene (PCG) sequences of 19 available butterfly species covering all the five butterfly families (Papilionidae, Nymphalidae, Peridae, Lycaenidae and Riodinidae). Both maximum likelihood and Bayesian inference analyses highly supported the monophyly of Lycaenidae+Riodinidae, which was standing as the sister of Nymphalidae. In addition, we propose that the riodinids be categorized into the family Lycaenidae as a subfamilial taxon.

Complete mitochondrial genome of the Common Evening Brown, Melanitis leda Linnaeus (Lepidoptera: Nymphalidae: Satyrinae).

The complete mitochondrial genome (mitogenome) of Melanitis leda (Lepidoptera: Nymphalidae: Satyrinae) is a circular molecule of 15,122 bp in length, containing 37 typical animal mitochondrial genes and 1 control region, known in insects as the AT-rich region. Its gene content and order are identical to all other available butterfly mitogenomes. All protein-coding genes (PCGs) start with a typical ATN initiation codon, except for COI, which is initiated by the CGA codon as observed in other butterfly species. A total of 97 bp of intergenic spacers are interspersed in 11 regions, ranging in size from 1 to 45 bp. The 314-bp-long AT-rich region is the smallest of all the butterfly corresponding regions available and contains some conserved structures similar to those found in other butterfly mitogenomes, including the motif ATAGA followed by a 19-bp poly-T stretch and a microsatellite-like (AT)6 element preceded by the ATTTA motif.

Complete mitogenome of the Painted Jezebel, Delias hyparete Linnaeus (Lepidoptera: Pieridae) and its comparison with other butterfly species.

In the present study, we report the first complete mitochondrial genome (mitogenome) of the Painted Jezebel, Delias hyparete. The mitogenome of Delias hyparete is 15 186 bp in length, and has typical sets of 37 genes: 13 protein-coding genes (PCGs), 2 ribosomal RNAs, 22 transfer RNAs and a non-coding A+T-rich region. All protein-coding genes are initiated by ATN codons, except for COI, which is tentatively designated by the CGA codon, as observed in other butterfly species. A total of 10 PCGs harbored the complete termination codon TAA or TAG, while the COI, COII and ND5 genes ended at a single T residue. All 22 tRNA genes show typical clover structures, with the exception of the tRNA(Ser(AGN)) which lacks the dihydrouridine (DHU) stem and is instead replaced by a simple loop. Thirteen intergenic spacers totaling 153 bp, and 13 overlapping regions totaling 46 bp are scattered throughout the whole genome. The 377 bp long of D. hyparete A+T-rich region is not comprised of large repetitive sequences, but harbors several features characteristic of the lepidopteran insects, including the motif ATAGA followed by an 18 bp poly-T stretch, a microsatellite-like (AT)(5) element preceded by the ATTTA motif, an 10 bp polyA-like stretch (AAAAATAAAA) present immediately upstream tRNA(Met).

Complete mitogenome of the Lesser Purple Emperor Apatura ilia (Lepidoptera: Nymphalidae: Apaturinae) and comparison with other nymphalid butterflies.

The complete mitochondrial genome of Apatura ilia (GenBank accession no. JF437925) was determined as a circular DNA molecule of 15 242 bp, with common genes of 13 putative proteins, 2 rRNAs, and 22 tRNAs and of the same gene arrangement as in other sequenced lepidopterans. All protein-coding genes had the typical start codon ATN, except for the COI's using CGA as its start codon as previously demonstrated in other lepidopteran species. The comparison of the nucleotide sequences of the A. ilia mitogenome with ten other Nymphalidae species showed nearly identical gene orientation and arrangement, with only a few alterations in non-coding fragments. The nucleotide composition and codon frequency all fell into the range estimated for the order Lepidoptera. The A. ilia mitochondrial genome had the canonical set of 22 tRNA genes folded in the typical cloverleaf structure, with an unique exception of tRNA(Ser) (AGN). The mitochondrial genes from A. ilia were overlapped in a total of 33 bp at 9 locations, as well as interleaved with a total of 155 bp intergenic spacers, spread over 12 regions with the size ranging from 1 to 49 bp. Furthermore, the spacer between ND6 and Cyt b harbored a microsatellite-like repeat (TA)(23) not found in other completely sequenced nymphalid genomes. The 403 bp AT-rich region harbored two conserved motifs (ATAGA, ATTTA), a 21 bp polyT stretch, a 10 bp poly-A region, along with two microsatellite-like repeats ( (TA)(10) and (TA)(7)), as detected in other nymphalid butterflies.