Mitochondrial DNA sequences of five squamates: phylogenetic affiliation of snakes.

Complete or nearly complete mitochondrial DNA sequences were determined from four lizards (Western fence lizard, Warren's spinytail lizard, Terrestrial arboreal alligator lizard, and Chinese crocodile lizard) and a snake (Texas blind snake). These genomes had a typical gene organization found in those of most mammals and fishes, except for a translocation of the glutamine tRNA gene in the blind snake and a tandem duplication of the threonine and proline tRNA genes in the spinytail lizard. Although previous work showed the existence of duplicate control regions in mitochondrial DNAs of several snakes, the blind snake did not have this characteristic. Phylogenetic analyses based on different tree-building methods consistently supported that the blind snake and a colubrid snake (akamata) make a sister clade relative to all the lizard taxa from six different families. An alternative hypothesis that snakes evolved from a lineage of varanoids was not favored and nearly statistically rejected by the Kishino-Hasegawa test. It is therefore likely that the apparent similarity of the tongue structure between snakes and varanoids independently evolved and that the duplication of the control region occurred on a snake lineage after divergence of the blind snake.     

鞘翅目昆虫线粒体基因组研究进展

鞘翅目（Coleoptera）是世界上最具多样性的类群,具有很高的生态和形态多样性,这些多样性吸引了很多进化生物学家和分类学家的关注。随着分子生物学的发展,分子生物学技术广泛应用于鞘翅目系统学的研究,但随着研究的深入,简单的分子片段已经不能满足研究的需求,需要发掘更新的分子标记。近年来,线粒体全基因组已经成为鞘翅目分子系统学研究中很重要的分子标记之一,并广泛地应用于鞘翅目昆虫各个阶元的研究中。本文就鞘翅目线粒体全基因组的概况、研究进展及存在问题进行了总结和讨论。目前,鞘翅目线粒体基因组的研究主要包括物种线粒体基因组组成与结构、分子系统学和分子进化等方面。线粒体基因组在解决系统发育和进化方面表现出了很多的优越性,然而也存在着一些缺点,如序列难获得、基因类型单一、各基因进化速率不同、应用较局限等。     

Discovery of Aspidytidae,a new family of aquatic Coleoptera

The six extant aquatic families of Hydradephaga (Coleoptera) known so far represent a diverse group of beetles morphologically highly modified for life in the water. We report the discovery of a new genus with two species from South Africa and China, which differ greatly from all extant families, but resemble the Jurassic-Cretaceous dagger Liadytidae (the dagger symbol indicates that the taxa are known only as fossils). Based on a combined phylogenetic analysis of molecular and morphological data we erect a new family, Aspidytidae, which is the sister group of Dytiscidae plus Hygrobiidae. We propose a new scenario for the evolution of swimming behaviour in adephagan beetles, in which the transition into the aquatic environment is followed by complex and repeated changes in lifestyles, including the secondary complete loss of swimming ability in Aspidytidae.     

The Complete Mitochondrial Genomes of Three Bristletails (Insecta: Archaeognatha): The Paraphyly of Machilidae and Insights into Archaeognathan Phylogeny

The order Archaeognatha was an ancient group of Hexapoda and was considered as the most primitive of living insects. Two extant families (Meinertellidae and Machilidae) consisted of approximately 500 species. This study determined 3 complete mitochondrial genomes and 2 nearly complete mitochondrial genome sequences of the bristletail. The size of the 5 mitochondrial genome sequences of bristletail were relatively modest, containing 13 protein-coding genes (PCGs), 2 ribosomal RNA (rRNA) genes, 22 transfer RNA (tRNA) genes and one control region. The gene orders were identical to that of Drosophila yakuba and most bristletail species suggesting a conserved genome evolution within the Archaeognatha. In order to estimate archaeognathan evolutionary relationships, phylogenetic analyses were conducted using concatenated nucleotide sequences of 13 protein-coding genes, with four different computational algorithms (NJ, MP, ML and BI). Based on the results, the monophyly of the family Machilidae was challenged by both datasets (W12 and G12 datasets). The relationships among archaeognathan subfamilies seemed to be tangled and the subfamily Machilinae was also believed to be a paraphyletic group in our study.     

A comparison of the mitochondrial genomes from two families of Solifugae (Arthropoda: Chelicerata): Eremobatidae and Ammotrechidae

Arachnids are an ancient and diverse group of arthropods, yet few representative mitochondrial genomes have been published for most of the 11 orders. Here, we present and compare sequence and genomic data from two complete mitochondrial genomes from the arachnid order Solifugae (the camel spiders or wind scorpions), representing two families, Ammotrechidae and Eremobatidae. We also make genome-level and sequence comparisons between these taxa and the horseshoe crab, a chelicerate from the sister group to arachnids. In their organization, the two solifuge mitochondrial genomes are similar to that of the horseshoe crab, although both of the solifuges possess a region of repeated sequence. All 13 protein-coding genes and the two ribosomal RNA genes are of similar sizes to those found in the horseshoe crab. The ammotrechid and the eremobatid each have one tRNA gene that differs in location from those of other chelicerates, suggesting that these translocations occurred after the divergence of Solifugae from other arachnid lineages. All 22 tRNA genes in both solifuges are inferred to form secondary structures that are typical of those found in other metazoan mt genomes. However, in the eremobatid, the tRNA(Ser(UCN)) gene in the repeat region appears to have undergone partial duplication and loss of function, and a new tRNA(Ser(UCN)) gene has been created de novo. Our divergence data, in conjunction with the fossil record, indicate that these two solifuge families diverged more than 230 million years ago. Thus, despite several gene rearrangements and duplications, these data indicate a remarkable degree of evolutionary stasis.     

Genealogy of families of SINEs in cetaceans and artiodactyls: the presence of a huge superfamily of tRNA(Glu)-derived families of SINEs.

Several novel (sub)families of SINEs were isolated from the genomes of cetaceans and artiodactyls, and their sequences were determined. From comparisons of diagnostic nucleotides among the short interspersed repetitive elements (SINEs) in these (sub)families, we were able to draw the following conclusions. (1) After the divergence of the suborder Tylopoda (camels), the CHRS family of SINEs was newly created from tRNA(Glu) in a common ancestor of the lineages of the Suina (pigs and peccaries), Ruminantia (cows and deer), and Cetacea (whales and dolphins). (2) After divergence of the Suina lineage, the CHR-1 SINE and the CHR-2 SINE were generated successively in a common ancestor of ruminants, hippopotamuses, and cetaceans. (3) In the Ruminantia lineage, the Bov-tA SINE was generated by recombination between the CHR-2 SINE and Bov-A. (4) In the Suina lineage, the CHRS-S SINE was generated from the CHRS SINE. (5) In this latter lineage, the PRE-1 family of SINEs was created by insertion of part of the gene for tRNA(Arg) into the 5' region of the CHRS-S family. The distribution of a particular family of SINEs among species of artiodactyls and cetaceans confirmed the most recent conclusion for paraphyly of the order Artiodactyla. The present study also revealed that a newly created tRNA(Glu)-derived family of SINEs was subjected both to recombination with different units and to duplication of an internal sequence within a SINE unit to generate, during evolution, a huge superfamily of tRNA(Glu)-related families of SINEs that are now found in the genomes of artiodactyls and cetaceans.     

Phylogenetic and genomic analysis of the complete mitochondrial DNA sequence of the spotted asparagus beetle Crioceris duodecimpunctata

We report the complete mitochondrial DNA sequence of the spotted asparagus beetle, Crioceris duodecimpunctata. The genome complement, gene order, and nucleotide composition of this beetle's mitochondrial genome were found to be typical of those reported for other insects. Unusual features of this genome include the substitution of UCU for GCU as the anticodon for tRNA(Ser), an unusual TpsiC loop for the tRNA(Ile) gene, and the identification of a putative ATT start codon for cox1. The utility of complete mitochondrial genome data for phylogenetic inference of the insect orders was tested, and compared to that of cox1 and combined mitochondrial ribosomal DNA sequences. Even though the number of insect orders represented by complete mitochondrial genomes is still limited, several well-established relationships are evident in the phylogenetic analysis of the complete sequences. Monophyly of the orders Diptera, Lepidoptera, and Coleoptera were consistently recovered. Monophyly of the Holometabola was also observed in some (though not all) analyses. The accumulation of complete mitochondrial sequences from a broader array of insect orders holds the promise of clarifying the early diversification of insects.     

Complex pattern of coalescence and fast evolution of a mitochondrial rRNA pseudogene in a recent radiation of tiger beetles

Transposed copies of mitochondrial DNA into the nucleus (numts) are widespread, but to date they have not been described from the Coleoptera (beetles). Here we report the discovery of a numt derived from a mitochondrial ribosomal RNA gene in Australian tiger beetles (genus Rivacindela). The loss of function of the numt was confirmed by high proportion of transversions, numerous noncompensatory substitutions in stem regions, and large deletions in functionally important sequences. Phylogenetic analysis of orthologous numt sequences was performed together with the corresponding mtDNA lineage for a study of origination and establishment of the transposed copies in closely related populations and species. All numt sequences were strongly supported to be monophyletic, indicating a single origin of this element. However, populations were polymorphic for the presence of the numt, and phylogenetic trees based on the numt sequences showed inconsistencies with the corresponding mtDNA phylogeny, suggesting slower processes of fixation compared to the mtDNA sequences. In a side-by-side comparison with their mtDNA sister lineage, the nucleotide substitution rate of 1.66 x 10(-8) substitutions/site/year in the numts was approximately equal to the average rate of mtDNA in this group but substantially higher than previous estimates of neutral nuclear rates in vertebrates. The numt clade was affected by several deletions but no insertions, with estimates of nucleotide loss exceeding the rate of nucleotide substitutions by approximately five times. The young age of the Rivacindela numt clade, their absence in species outside of a narrow lineage of related individuals, and the high rate of deletions suggest that insertions do not persist in this group, which is consistent with the view that comparatively small genomes as those of Coleoptera harbor fewer mitochondrial and other nuclear pseudogenes.     

The conserved mitochondrial genome of the jewel beetle (Coleoptera: Buprestidae) and its phylogenetic implications for the suborder Polyphaga

In this study, we sequenced the mitochondrial (mt) genome of Agrilus mali (Coleoptera: Buprestidae) using next-generation sequencing, and accordingly annotated 13 protein-coding, 22 tRNA, and 2 rRNA genes and a 1458-bp non-coding region. Comparative analysis indicated that the mt genome of A. mali is relatively conserved, with a typical gene content and order identical to those of other coleopterans. However, the newly sequenced mt genome is characterized by a relatively higher A + T content compared with that of other species within the family Buprestidae. Phylogenetic analysis based on Bayesian inference revealed that the evolutionary relationship among the six infraorders of the suborder Polyphaga is (Scirtiformia + (Elateriformia + ((Scarabaeiformia + Staphyliniformia) + (Bostrichiformia + (Cucujiformia))))). However, the topology indicated that the family Buprestidae is a sister group to other Polyphaga infraorders, excluding Scirtiformia as a monophyly, and thus the monophyly of Elateriformia was not supported. This study not only presents the mt genome of a species in the family Buprestidae and a comparative analysis of jewel beetles but also examines the contribution of mt genomes in elucidating phylogenetic relationships within the suborder Polyphaga of Coleoptera.     

Resolving phylogeny at the family level by mitochondrial cytochrome oxidase sequences: phylogeny of carrion beetles (Coleoptera, Silphidae)

We investigated the phylogenetic relationships of carrion beetles (Coleoptera, Silphidae) using 2094 bp of their mitochondrial cytochrome oxidase subunit I and II and tRNA leucine gene sequences. Shorter fragments of this gene region previously have been used to establish generic relationships in insects. In this study, they provided more than sufficient resolution, although the third positions of the protein-coding sequences reached saturation for the deeper divergences. This first published phylogeny for the Silphidae comprises 23 species from 13 genera sampled across the geographic range of the family. In addition, we included species from three related families as outgroups. One of these families, the Agyrtidae, was, until recently, included in the Silphidae, but its resolution here justifies its current position as a separate family. The silphid subfamilies Nicrophorinae and Silphinae are monophyletic in all analyses. All genera for which several species were sampled are supported as monophyletic groups, with the exception of the genus Silpha. European and North American representatives of two Nicrophorus species described from both continents are supported as each others' closest relatives. The lineage that colonized Gondwanaland and that most likely originated in the Palearctic is the most basal within the Silphinae.     

Gene recruitment--a common mechanism in the evolution of transfer RNA gene families

The evolution of alloacceptor transfer RNAs (tRNAs) has been traditionally thought to occur vertically and reflect the evolution of the genetic code. Yet there have been several indications that a tRNA gene could evolve horizontally, from a copy of an alloacceptor tRNA gene in the same genome. Earlier, we provided the first unambiguous evidence for the occurrence of such "tRNA gene recruitment" in nature--in the mitochondrial (mt) genome of the demosponge Axinella corrugata. Yet the extent and the pattern of this process in the evolution of tRNA gene families remained unclear. Here we analyzed tRNA genes from 21 mt genomes of demosponges as well as nuclear genomes of rhesus macaque, chimpanzee and human. We found four new cases of alloacceptor tRNA gene recruitment in mt genomes and eleven cases in the nuclear genomes. In most of these cases we observed a single nucleotide substitution at the middle position of the anticodon, which resulted in the change of not only the tRNA's amino-acid identity but also the class of the amino-acyl tRNA synthetases (aaRSs) involved in amino-acylation. We hypothesize that the switch to a different class of aaRSs may have prevented the conflict between anticodon and amino-acid identities of recruited tRNAs. Overall our results suggest that gene recruitment is a common phenomenon in tRNA multigene family evolution and should be taken into consideration when tRNA evolutionary history is reconstructed.     

Wolbachia distribution in selected beetle taxa characterized by PCR screens and MLST data

Wolbachia (Alphaproteobacteria) is an inherited endosymbiont of arthropods and filarial nematodes and was reported to be widespread across insect taxa. While Wolbachia's effects on host biology are not understood from most of these hosts, known Wolbachia‐induced phenotypes cover a spectrum from obligate beneficial mutualism to reproductive manipulations and pathogenicity. Interestingly, data on Wolbachia within the most species‐rich order of arthropods, the Coleoptera (beetles), are scarce. Therefore, we screened 128 species from seven beetle families (Buprestidae, Hydraenidae, Dytiscidae, Hydrophilidae, Gyrinidae, Haliplidae, and Noteridae) for the presence of Wolbachia. Our data show that, contrary to previous estimations, Wolbachia frequencies in beetles (31% overall) are comparable to the ones in other insects. In addition, we used Wolbachia MLST data and host phylogeny to explore the evolutionary history of Wolbachia strains from Hydraenidae, an aquatic lineage of beetles. Our data suggest that Wolbachia from Hydraenidae might be largely host genus specific and that Wolbachia strain phylogeny is not independent to that of its hosts. As this contrasts with most terrestrial Wolbachia–arthropod systems, one potential conclusion is that aquatic lifestyle of hosts may result in Wolbachia distribution patterns distinct from those of terrestrial hosts. Our data thus provide both insights into Wolbachia distribution among beetles in general and a first glimpse of Wolbachia distribution patterns among aquatic host lineages.     

Gene rearrangements in snake mitochondrial genomes: highly concerted evolution of control-region-like sequences duplicated and inserted into a tRNA gene cluster

Mitochondrial DNA (mtDNA) regions corresponding to two major tRNA gene clusters were amplified and sequenced for the Japanese pit viper, himehabu. In one of these clusters, which in most vertebrates characterized to date contains three tightly connected genes for tRNA(Ile), and tRNA(Gln), and tRNA(Met), a sequence of approximately 1.3 kb was found to be inserted between the genes for tRNA(Ile) and tRNA(Gln). The insert consists of a control-region-like sequence possessing some conserved sequence blocks, and short flanking sequences which may be folded into tRNA(Pro), tRNA(Phe), and tRNA(Leu) genes. Several other snakes belonging to different families were also found to possess a control-region-like sequence and tRNA(Leu) gene between the tRNA(Ile)and tRNA(Gln) genes. We also sequenced a region surrounded by genes for cytochrome b and 12S rRNA, where the control region and genes for tRNA(Pro) and tRNA(Phe) are normally located in the mtDNAs of most vertebrates. In this region of three examined snakes, a control-region- like sequence exists that is almost completely identical to the one found between the tRNA(Ile) and tRNA(Gln) genes. The mtDNAs of these snakes thus possess two nearly identical control-region-like sequences which are otherwise divergent to a large extent between the species. These results suggest that the duplicate state of the control-region- like sequences has long persisted in snake mtDNAs, possibly since the original insertion of the control-region-like sequence and tRNA(Leu) gene into the tRNA gene cluster, which occurred in the early stage of the divergence of snakes. It is also suggested that the duplicated control-region-like sequences at two distant locations of mtDNA have evolved concertedly by a mechanism such as frequent gene conversion. The secondary structures of the determined tRNA genes point to the operation of simplification pressure on the T psi C arm of snake mitochondrial tRNAs.      

Diversity patterns of soil macro-Coleoptera in Mexican shaded and unshaded coffee agroecosystems: an indication of habitat perturbation

The soil Coleoptera community was sampled with pitfall traps in shaded and unshaded coffee agroecosystems in Veracruz, Mexico. The insect collection resulted in a total of 31 species which belong to nine families. The most frequent families collected in this study (in terms of species and number of organisms) were Scarabaeidae and Carabidae.Species diversity was measured using the Hill's family of diversity numbers. The Coleoptera community was more diverse in shaded than in unshaded coffee. Also, the collected organisms were more evenly distributed between species in shaded coffee.The diversity of the scarab beetles was shown to be strongly affected by the degree of forest perturbation. The richness and evenness of scarab beetles was correlated with the diversity of shade trees present in coffee agro-ecosystems; scarab richness drops from 19 species collected in virgin tropical rain-forests, to five species in polyspecific shade (more than ten species of shade trees) and six species in monogeneric shade (three species of shade trees) coffee agro-ecosystems, and to three scarab species in unshaded coffee. Evenness in scarab beetles follows a similar pattern: a single species tends to gradually become dominant as more shade trees are removed from the agro-ecosystem. It is suggested that some shade trees should be preserved within the agro-ecosystem during the intensification of coffee production in Mexico.     

The complete mitochondrial genomes of two schizothoracine fishes (Teleostei,Cypriniformes): A novel minisatellite in fish mitochondrial genomes

Minisatellites, a class of variable number tandem repeats (VNTRs), are abundant throughout the control region in animal mitochondrial DNA (mtDNA) but rare in other regions of animal mtDNA. Here, we reported a novel minisatellite in fish mitochondrial genomes. We first determined the complete mitochondrial genomes of two schizothoracine fishes (Herzensteinia microcephalus and Schizopygopsis pylzovi) and found a type of minisatellites in a novel region between the tRNA‐Thr and tRNA‐Pro genes in their mtDNA. To explore the origin and evolution of the minisatellites in different schizothoracine and closely related fishes, we analyzed the available 80 fish mitogenomes which represent five closely related tribes of cyprinine fishes. The results from the phylogenetic analyses show that the schizothoracine fishes sensu stricto is not a monophyletic group and is divided into two clades (Schizothoracini and Schizopygopsini); and the minisatellite is only present in Schizopygopsini distributed in the region between the two tRNA genes (tRNA‐Thr and tRNA‐Pro) of the mtDNA. This is the first record of a minisatellite in a non‐control region of fish mitogenome.     

Sequence and gene organization of the chicken mitochondrial genome. A novel gene order in higher vertebrates

The 16,775 base-pair mitochondrial genome of the white Leghorn chicken has been cloned and sequenced. The avian genome encodes the same set of genes (13 proteins, 2 rRNAs and 22 tRNAs) as do other vertebrate mitochondrial DNAs and is organized in a very similar economical fashion. There are very few intergenic nucleotides and several instances of overlaps between protein or tRNA genes. The protein genes are highly similar to their mammalian and amphibian counterparts and are translated according to the same variant genetic code. Despite these highly conserved features, the chicken mitochondrial genome displays two distinctive characteristics. First, it exhibits a novel gene order, the contiguous tRNA(Glu) and ND6 genes are located immediately adjacent to the displacement loop region of the molecule, just ahead of the contiguous tRNA(Pro), tRNA(Thr) and cytochrome b genes, which border the displacement loop region in other vertebrate mitochondrial genomes. This unusual gene order is conserved among the galliform birds. Second, a light-strand replication origin, equivalent to the conserved sequence found between the tRNA(Cys) and tRNA(Asn) genes in all vertebrate mitochondrial genomes sequenced thus far, is absent in the chicken genome. These observations indicate that galliform mitochondrial genomes departed from their mammalian and amphibian counterparts during the course of evolution of vertebrate species. These unexpected characteristics represent useful markers for investigating phylogenetic relationships at a higher taxonomic level.     

Diversity of aquatic and riparian Coleoptera of the Philippines: checklist,state of knowledge,priorities for future research and conservation

The diversity of the aquatic and strictly riparian beetles of the Philippines is reviewed. A checklist (with 317 species and subspecies) of the aquatic and riparian Coleoptera of the Philippines is provided for the first time herein. This checklist covers the following families (number of species and subspecies in parentheses): Hydroscaphidae (1), Gyrinidae (15), Dytiscidae (65), Noteridae (4), Haliplidae (2), Hydraenidae (15), Hydrochidae (2), Spercheidae (1), Hydrophilidae (72), Scirtidae (49), Elmidae (23), Dryopidae (2), Psephenidae (16), Eulichadidae (1), Limnichidae (20), Heteroceridae (5), Malachiidae (5), Lampyridae (15), Nitidulidae (4). A total of 201 species and subspecies (63%) are considered to be endemic to the Philippines. Aquatic/riparian beetle genera and families that are still unrecorded but likely to occur in this country are briefly discussed. Based on estimates at the genus level, it is concluded that about 850 aquatic and riparian species can be expected to occur in the Philippines. The implication of the Philippine biogeographic history for this diversity is discussed. The potential of aquatic/riparian beetles as bioindicators is briefly examined. The dilemma of aquatic invertebrate conservation is analysed and priorities for future research are outlined.     

tRNA Signatures Reveal a Polyphyletic Origin of SAR11 Strains among Alphaproteobacteria

Molecular phylogenetics and phylogenomics are subject to noise from horizontal gene transfer (HGT) and bias from convergence in macromolecular compositions. Extensive variation in size, structure and base composition of alphaproteobacterial genomes has complicated their phylogenomics, sparking controversy over the origins and closest relatives of the SAR11 strains. SAR11 are highly abundant, cosmopolitan aquatic Alphaproteobacteria with streamlined, A+T-biased genomes. A dominant view holds that SAR11 are monophyletic and related to both Rickettsiales and the ancestor of mitochondria. Other studies dispute this, finding evidence of a polyphyletic origin of SAR11 with most strains distantly related to Rickettsiales. Although careful evolutionary modeling can reduce bias and noise in phylogenomic inference, entirely different approaches may be useful to extract robust phylogenetic signals from genomes. Here we develop simple phyloclassifiers from bioinformatically derived tRNA Class-Informative Features (CIFs), features predicted to target tRNAs for specific interactions within the tRNA interaction network. Our tRNA CIF-based model robustly and accurately classifies alphaproteobacterial genomes into one of seven undisputed monophyletic orders or families, despite great variability in tRNA gene complement sizes and base compositions. Our model robustly rejects monophyly of SAR11, classifying all but one strain as Rhizobiales with strong statistical support. Yet remarkably, conventional phylogenetic analysis of tRNAs classifies all SAR11 strains identically as Rickettsiales. We attribute this discrepancy to convergence of SAR11 and Rickettsiales tRNA base compositions. Thus, tRNA CIFs appear more robust to compositional convergence than tRNA sequences generally. Our results suggest that tRNA-CIF-based phyloclassification is robust to HGT of components of the tRNA interaction network, such as aminoacyl-tRNA synthetases. We explain why tRNAs are especially advantageous for prediction of traits governing macromolecular interactions from genomic data, and why such traits may be advantageous in the search for robust signals to address difficult problems in classification and phylogeny.