Mitogenomes Of Species Research Articles

Mitogenomic Characterization and Phylogenetic Placement of African Hind, Cephalopholis taeniops: Shedding Light on the Evolution of Groupers (Serranidae: Epinephelinae).

The global exploration of evolutionary trends in groupers, based on mitogenomes, is currently underway. This research extensively investigates the structure of and variations in Cephalopholis species mitogenomes, along with their phylogenetic relationships, focusing specifically on Cephalopholis taeniops from the Eastern Atlantic Ocean. The generated mitogenome spans 16,572 base pairs and exhibits a gene order analogous to that of the ancestral teleost's, featuring 13 protein-coding genes (PCGs), two ribosomal RNA genes (rRNAs), 22 transfer RNA genes (tRNAs), and an AT-rich control region. The mitogenome of C. taeniops displays an AT bias (54.99%), aligning with related species. The majority of PCGs in the mitogenome initiate with the start codon ATG, with the exceptions being COI (GTG) and atp6 (TTG). The relative synonymous codon usage analysis revealed the maximum abundance of leucine, proline, serine, and threonine. The nonsynonymous/synonymous ratios were <1, which indicates a strong negative selection among all PCGs of the Cephalopholis species. In C. taeniops, the prevalent transfer RNAs display conventional cloverleaf secondary structures, except for tRNA-serine (GCT), which lacks a dihydrouracil (DHU) stem. A comparative examination of conserved domains and sequence blocks across various Cephalopholis species indicates noteworthy variations in length and nucleotide diversity. Maximum likelihood, neighbor-joining, and Bayesian phylogenetic analyses, employing the concatenated PCGs and a combination of PCGs + rRNAs, distinctly separate all Cephalopholis species, including C. taeniops. Overall, these findings deepen our understanding of evolutionary relationships among serranid groupers, emphasizing the significance of structural considerations in mitogenomic analyses.

The First Mitochondrial Genome of an Odonata Endemic to South America, Chalcopteryx rutilans (Rambur, 1842) (Odonata: Polythoridae), and Its Implications for the Phylogeny of the Zygoptera

Chalcopteryx rutilans Rambur, 1842 (Polythoridae, Odonata) is a species widely distributed in central Amazonia. Due to its sensitivity to environmental changes, it is a bioindicator species used to evaluate the environmental conditions of streams in federally protected areas. By sequencing C. rutilans mitogenome, we report the first whole mitogenome from the Polythoridae family and the first from an Odonata species endemic to South America. The entire mitogenome has 15,653 bp and contains 13 protein-coding, 22 tRNA, and two rRNA genes. The nucleotide composition of the mitogenome is 42.7%, T: 25.5%, C: 19.4%, and G: 12.4%. The AT and GC skews of the mitogenome sequence were 0.249 and −0.220, respectively. C. rutilans was recovered as a sister to Pseudolestes mirabilis Kirby, 1900 (Pseudolestidae), demonstrating the absence of mitogenomes of species from multiple families in the current literature. Mitogenome data from this study will provide useful information for further studies on the phylogeny and conservation of Polythoridae.

Affordable de novo generation of fish mitogenomes using amplification-free enrichment of mitochondrial DNA and deep sequencing of long fragments.

Biomonitoring surveys make use of metabarcoding tools to describe the community composition. These studies match their sequencing results against public genomic databases to identify the species. However, mitochondrial genomic reference data are yet incomplete, only a few genes may be available, or the suitability of existing sequence data is suboptimal for species level resolution. Here, we present a dedicated and cost-effective workflow with no DNA amplification for generating complete fish mitogenomes for the purpose of strengthening fish mitochondrial databases. Two different strategies using long fragment sequencing with Oxford Nanopore technology coupled with mitochondrial DNA enrichment were used. One where the enrichment is achieved by preferential isolation of mitochondria followed by DNA extraction and nuclear DNA depletion ("mitoenrichment"). A second enrichment approach takes advantage of the CRISPR Cas9 targeted scission on previously dephosphorylated DNA ("targeted mitosequencing"). The sequencing results varied between tissue, species, and integrity of the DNA. The mitoenrichment method yielded 0.17%-12.33% of sequences on target and a mean coverage ranging from 74.9 to 805-fold. The targeted mitosequencing experiment from native genomic DNA yielded 1.83%-55% of sequences on target and a 38 to 2123-fold mean coverage. These produced complete mitogenomes of species with homopolymeric regions, tandem repeats, and gene rearrangements. We demonstrate that deep sequencing of long fragments of native fish DNA can be achieved with low computational resources in a cost-effective manner, opening the discovery of mitogenomes of nonmodel or understudied fish taxa to a broad range of laboratories worldwide.

Phylogenetic relationships of the Amblyomma cajennense complex (Acari: Ixodidae) at mitogenomic resolution

The genus Amblyomma is the third most diverse in the number of species within the Ixodidae, with practically half of its species distributed in the Americas, though there are also species occurring in Africa, Asia, and Australia. Within the genus, there are several species complexes with veterinary and public health importance. The Amblyomma cajennense complex, in the Americas, is represented by six species with a wide distribution, from Texas to northern Argentina. We combined two sequencing techniques to generate complete mitogenomes of species belonging to the Amblyomma cajennense complex: genome skimming and long-range PCRs sequencing methods. Thus, we generated seven new mitochondrial genomes for all species of the Amblyomma cajennense complex, except for Amblyomma interandinum. Genetic distances between the mitogenomes corroborate the clear differentiation between the five species of the Amblyomma cajennense complex. The phylogenetic relationships of these species had previously been evaluated by combining partial nuclear and mitochondrial genes and here these relationships are corroborated with a more robust framework of data, which demonstrates that the conjunction of mitochondrial and nuclear partial genes can resolve close relationships when entire genes or genomes are unavailable. The gene order, structure, composition, and length are stable across these mitogenomes, and they share the general characteristics of Metastriata. Future studies should increase the number of available mitogenomes for this genus, especially for those species from the Indo-Pacific region and Africa, by means of a better understanding of their relationships and evolutionary process.

The complete mitochondrial genome of the freshwater snail Cipangopaludina ampullacea (Küster, 1852) (Gastropoda: Viviparidae)

Cipangopaludina ampullacea (Küster, 1852) is a freshwater snail endemic to China. In this study, the complete mitochondrial genome of C. ampullacea was sequenced using next-generation sequencing. The mitogenome is 16,892 bp long and comprises a total of 37 genes, including 13 protein-coding genes, two rRNA genes, and 22 tRNA genes. It is consistent with the basic characteristics of other known viviparid mitochondrial genomes. Phylogenetic analysis using related species mitogenomes showed that Cipangopaludina and Margarya are mutually non monophyletic. Our study provides valuable information to reconstruct the taxonomy and evolution of viviparid snails more comprehensively.

Complex Physical Structure of Complete Mitochondrial Genome of Quercus acutissima (Fagaceae): A Significant Energy Plant

Quercus acutissima Carruth. is a Chinese important energy plant with high ecological and economic values. While the species chloroplast genome has been reported, its mitochondrial genome (mitogenome) is still unexplored. Here, we assembled and annotated the Q. acutissima mitogenome, and we compared its characteristic differences with several closely related species. The Q. acutissima mitogenome’s main structure is branched with three distinguished contigs (linear molecule 1, circular molecule 2, and circular molecule 3) with 448,982 bp total length and 45.72% GC content. The mitogenome contained 51 genes, including 32 protein-coding, 16 tRNA and 3 rRNA genes. We examined codon usage, repeated sequences, genome recombination, chloroplast to mitochondrion DNA transformation, RNA editing, and synteny in the Q. acutissima mitogenome. Phylogenetic trees based on 29 species mitogenomes clarified the species classification. Our results provided comprehensive information of Q. acutissima mitogenome, and they are expected to provide valuable information for Fagaceae evolutionary biology and to promote the species germplasm utilization.

Complete mitochondrial genome of the abalone shell-boring Polydora hoplura (Polychaeta, Spionidae)

Polydora hoplura Claparede 1868 (Polychaeta, Spionidae) is a mollusk shell-boring species which has been reported as a harmful species. In the present study, we sequenced and reported the mitochondrial genome (mitogenome) of P. hoplura from Korea. Its mitogenome was determined by long range PCR and primer walking. The mitogenome was found to be 17,707 bp in length and contained two ribosomal RNAs (rRNAs), 23 transfer RNAs (tRNAs), and 12 protein-coding genes which showed different gene arrangement with that in related mitogenomes of species of Sedentaria (Annelida). Interestingly, the atp8 gene was not found in the mitogenome of P. hoplura, whereas it is present in the mitogenomes of related species P. brevipalpa and P. websteri. Moreover, nad2 (3’ end) and cox1 (5’ end) genes overlapped in 23 bases. The mitogenome structure and gene contents of P. hoplura provide useful information on the evolution and phylogenetic relationship among polychaete species.

Characterization of the mitogenomes of long-tailed giant rat, Leopoldamys sabanus and a comparative analysis with other Leopoldamys species

Two mitogenomes of long-tailed giant rat, Leopoldamys sabanus (Thomas, 1887), which belongs to the family Muridae were sequenced and assembled in this study. Both mitogenomes have a length of 15,973 bp and encode 13 protein-coding genes (PCGs), 22 transfer RNA genes, two ribosomal RNA genes and one control region. The circular molecule of L. sabanus has a typical vertebrate gene arrangement. Phylogenetic and BLASTn analysis using 10 Leopoldamys species mitogenomes revealed sequence variation occurred within species from different time zones. Along with the taxonomic issues, this suggests a landscape change might influence genetic connectivity.

Unique Mitochondrial Single Nucleotide Polymorphisms Demonstrate Resolution Potential to Discriminate Theileria parva Vaccine and Buffalo-Derived Strains.

Distinct pathogenic and epidemiological features underlie different Theileria parva strains resulting in different clinical manifestations of East Coast Fever and Corridor Disease in susceptible cattle. Unclear delineation of these strains limits the control of these diseases in endemic areas. Hence, an accurate characterization of strains can improve the treatment and prevention approaches as well as investigate their origin. Here, we describe a set of single nucleotide polymorphisms (SNPs) based on 13 near-complete mitogenomes of T. parva strains originating from East and Southern Africa, including the live vaccine stock strains. We identified 11 SNPs that are non-preferentially distributed within the coding and non-coding regions, all of which are synonymous except for two within the cytochrome b gene of buffalo-derived strains. Our analysis ascertains haplotype-specific mutations that segregate the different vaccine and the buffalo-derived strains except T. parva-Muguga and Serengeti-transformed strains suggesting a shared lineage between the latter two vaccine strains. Phylogenetic analyses including the mitogenomes of other Theileria species: T. annulata, T. taurotragi, and T. lestoquardi, with the latter two sequenced in this study for the first time, were congruent with nuclear-encoded genes. Importantly, we describe seven T. parva haplotypes characterized by synonymous SNPs and parsimony-informative characters with the other three transforming species mitogenomes. We anticipate that tracking T. parva mitochondrial haplotypes from this study will provide insight into the parasite’s epidemiological dynamics and underpin current control efforts.

Sequence comparison of the mitochondrial genomes in two species of the genus Nerita (Gastropoda: Neritimorpha: Neritidae): phylogenetic implications and divergence time estimation for Neritimorpha

Many Nerita species live in warm-water environments, and they are some of the few organisms from the intertidal zone that can live in both freshwater and seawater. Previous comparative studies of the mitogenomes of Nerita species suggest that the genome rearrangements are very conservative. Generally, the species possess a set of similar mitochondrial gene arrangements, but nucleotide sequences can be used to elucidate phylogenetic relationships at various levels of divergence. Here, the mitogenomes of Nerita undata and Nerita balteata were sequenced and found to be 15,583bp and 15,571bp, respectively. The mitogenomes of both species contain 13 protein-coding genes, 22 tRNA genes, and two rRNA genes. The nucleotides of the two mitogenomes are highly similar, with the same gene composition and genomic organization as those present in other Nerita species. The tRNA secondary structures were different from those of other gastropods: trnS2 is not folded into typical secondary structures, and the dihydrouridine (DHU) arm simply forms a loop. The phylogenetic analysis showed that Neritimorpha is a sister group of Vetigastropoda and Caenogastropoda. Nerita balteata is a sister group of Nerita versicolor and Nerita undata, and all three species belong to Neritimorpha. This study contributes towards the comparative mitogenomic analysis of Neritidae and phylogenetic considerations among Neritimorpha species. The estimation of divergence time revealed that the two Nerita species were differentiated in the late Paleogene of the Cenozoic Era, and their evolution may be related to environmental changes.

Comparative analysis of mitogenomes among three species of Haplotropidini grasshoppers and a new species of the genus Sulcotropis (Orthoptera: Acridoidea: Pamphagidae) from China.

The complete mitochondrial genomes of three species of Haplotropidini were sequenced, annotated and analyzed. Then, combined with 18 species mitogenomes of Acridoidea and 1 species of Tridactyloidea, the phylogenetic relationships were reconstructed by maximum likelihood (ML) and Bayesian (BI) methods based on PCGs. The phylogenetic relationship tree showing that Sulcotropis Yin et Chou is a valid genus and not a synonym of Haplotropis Saussure, 1888. A new species Sulcotropis xiaowutaiensis sp. nov. is described in this paper from China, it is allied to Sulcotropis cyanipes Yin et Chou, 1979, but differs from latter by median carina of pronotum cut by posterior transverse sulcus slightly, epiphallus with middle part equal both sides in high, cercus of male gradually widened at base, interspace of mesosternum narrowed in the base slightly and subgenital plate of female oblong, hind margin with small acute angle in the middle. Type specimens are deposited in the College of Life Sciences, Hebei University, Baoding, Hebei, China.

Mitochondrial genome of Tamarixia radiata (Hymenoptera: Chalcidoidea: Eulophidae) and phylogenetic analysis

Tamarixia radiata plays an important role in biological control of the psyllid Diaphorina citri Kuwayama, vector of the huanglongbing (HLB). In this study, we sequenced and analyzed the mitochondrial genome (mitogenome) of T. radiata, the first mitogenome of species in the family Eulophidae. This mitogenome was 14,752 bp long and encoded 13 protein-coding genes (PCGs), 22 transfer RNA genes (tRNAs) and two ribosomal RNA unit genes (rRNAs). All 13 PCGs were initiated by the ATN (ATG, ATT, and ATA) codon. Twelve PCGs terminate with the stop codon TAA or TAG except for nad1 which end with the incomplete codon T−. Phylogenetic analysis showed that T. radiata got together with three Pteromalidae species, indicating the close relationship of Eulophidae and Pteromalidae.

Convergent herbivory on conifers by Choristoneura moths after boreal forest formation

Mitogenomes are useful markers for phylogenetic studies across a range of taxonomic levels. Here, we focus on mitogenome variation across the tortricid moth genus Choristoneura and particularly the spruce budworm (Choristoneura fumiferana) species complex, a notorious pest group of North American conifer forests. Phylogenetic relationships of Tortricidae, representing two subfamilies, four tribes and nine genera, were analyzed using 21 mitogenomes. These included six newly-sequenced mitogenomes for species in the spruce budworm complex plus three additional Choristoneura species and 12 previously published mitogenomes from other tortricids and one from the Cossidae. We evaluated the phylogenetic informativeness of the mitogenomes and reconstructed a time-calibrated tree with fossil and secondary calibrations. We found that tortricid mitogenomes had conserved protein and ribosomal regions, and analysis of all protein-coding plus ribosomal genes together provided an efficient marker at any taxonomic rank. The time-calibrated phylogeny showed evolutionary convergence of conifer feeding within Choristoneura, with two independent lineages, the Nearctic spruce budworm complex and the Palearctic species Choristoneura murinana, both shifting onto conifers about 11 million years ago from angiosperms. These two host-plant shifts both occurred after the formation of boreal forest in the late Miocene. Haplotype diversification within the spruce budworm complex occurred in the last 4 million years, and is probably linked to the initial cooling cycles of the Northern Hemisphere in the Pliocene.

Revisiting the mitogenomic phylogeny of Salmoninae: new insights thanks to recent sequencing advances.

The phylogeny of the Salmonidae family, the only living one of the Order Salmoniformes, remains still unclear because of several reasons. Such reasons include insufficient taxon sampling and/or DNA information. The use of complete mitochondrial genomes (mitogenomics) could provide some light on it, but despite the high number of mitogenomes of species belonging to this family published during last years, an integrative work containing all this information has not been done. In this work, the phylogeny of 46 Salmonidae species was inferred from their mitogenomic sequences. Results include a Bayesian molecular-dated phylogenetic tree with very high statistical support showing Coregoninae and Salmoninae as sister subfamilies, as well as several new phylogenetic relationships among species and genus of the family. All these findings contribute to improve our understanding of the Salmonidae systematics and could have consequences on related evolutionary studies, as well as highlight the importance of revisiting phylogenies with integrative studies.

The Trouble with MEAM2: Implications of Pseudogenes on Species Delimitation in the Globally Invasive Bemisia tabaci (Hemiptera: Aleyrodidae) Cryptic Species Complex.

Molecular species identification using suboptimal PCR primers can over-estimate species diversity due to coamplification of nuclear mitochondrial (NUMT) DNA/pseudogenes. For the agriculturally important whitefly Bemisia tabaci cryptic pest species complex, species identification depends primarily on characterization of the mitochondrial DNA cytochrome oxidase I (mtDNA COI) gene. The lack of robust PCR primers for the mtDNA COI gene can undermine correct species identification which in turn compromises management strategies. This problem is identified in the B. tabaci Africa/Middle East/Asia Minor clade which comprises the globally invasive Mediterranean (MED) and Middle East Asia Minor I (MEAM1) species, Middle East Asia Minor 2 (MEAM2), and the Indian Ocean (IO) species. Initially identified from the Indian Ocean island of Réunion, MEAM2 has since been reported from Japan, Peru, Turkey and Iraq. We identified MEAM2 individuals from a Peruvian population via Sanger sequencing of the mtDNA COI gene. In attempting to characterize the MEAM2 mitogenome, we instead characterized mitogenomes of MEAM1. We also report on the mitogenomes of MED, AUS, and IO thereby increasing genomic resources for members of this complex. Gene synteny (i.e., same gene composition and orientation) was observed with published B. tabaci cryptic species mitogenomes. Pseudogene fragments matching MEAM2 partial mtDNA COI gene exhibited low frequency single nucleotide polymorphisms that matched low copy number DNA fragments (<3%) of MEAM1 genomes, whereas presence of internal stop codons, loss of expected stop codons and poor primer annealing sites, all suggested MEAM2 as a pseudogene artifact and so not a real species.

A novel mitochondrial genome of Arborophila and new insight into Arborophila evolutionary history.

The lineage of the Bar-backed Partridge (Arborophila brunneopectus) was investigated to determine the phylogenetic relationships within Arborophila as the species is centrally distributed within an area covered by the distributions of 22 South-east Asian hill partridge species. The complete mitochondrial genome (mitogenome) of A. brunneopectus was determined and compared with four other hill partridge species mitogenomes. NADH subunit genes are radical in hill partridge mitogenomes and contain the most potential positive selective sites around where variable sites are abundant. Together with 44 other mitogenomes of closely related species, we reconstructed highly resolved phylogenetic trees using maximum likelihood (ML) and Bayesian inference (BI) analyses and calculated the divergence and dispersal history of Arborophila using combined datasets composed of their 13-protein coding sequences. Arborophila is reportedly be the oldest group in Phasianidae whose ancestors probably originated in Asia. A. rufipectus shares a closer relationship with A. ardens and A. brunneopectus compared to A. gingica and A. rufogularis, and such relationships were supported and profiled by NADH dehydrogenase subunit 5 (ND5). The intragenus divergence of all five Arborophila species occurred in the Miocene (16.84~5.69 Mya) when there were periods of climate cooling. We propose that these cooling events in the Miocene forced hill partridges from higher to lower altitudes, which led to geographic isolation and speciation. We demonstrated that the apparently deleterious +1 frameshift mutation in NADH dehydrogenase subunit 3 (ND3) found in all Arborophila is an ancient trait that has been eliminated in some younger lineages, such as Passeriformes. It is unclear of the biological advantages of this elimination for the relevant taxa and this requires further investigation.

The complete mitochondrial genome of Pycnarmon lactiferalis (Lepidoptera: Crambidae)

The complete mitochondrial genome of Pycnarmon lactiferalis (Walker, 1859) has been determined. The entire sequence is 15,219 bp in length which contains 13 protein-coding genes (PCGs), 22 transfer RNA genes, two ribosomal RNA genes, and an A + T-rich region. All PCGs start with the typical ATN codon except for COI with CGA. TAA is used for all the PCGs, while the COI, COII and ND5 possess incomplete termination codons T or TA. The secondary structure of 22 tRNAs have the typical clover-leaf pattern except for tRNASer(AGN) lacking the dihydrouridine (DHU) stem. The A + T-rich region, located between the srRNA and tRNAMet, do not contain the motif “ATAGA” that is conserved across other lepidopteran species mitogenomes. A phylogenetic relationship of Pyraloidea has been reconstructed based on 13 PCGs data using Bayesian inference (BI) method.

Mitochondrial genome of two marine fungal species

Here is a first record for a mitochondrial genome of marine fungus Acremonium fuci and closely related species Emericellopsis sp. One strain for each species, differentiated by morphological features was studied. Complete mitochondrial sequences are 24 565 and 24 951 bp in length. The circular molecule encodes all genes typical for fungal mitochondrial genomes: 15 protein-coding genes, 28 tRNAs, and large and small subunits of RNA. All structural genes are located on one strand and transcribed in one direction. Mitogenomes of species have 99% identity in protein-coding genes, but differ in some structural features: one of them has additional ORF with a similarity to cox1 gene, and another one has an intron in nad5 gene.

The complete mitogenome of the Morton Bay bug Thenus orientalis (Lund, 1793) (Crustacea: Decapoda: Scyllaridae) from a cooked sample and a new mitogenome order for the Decapoda

The mitochondrial genome sequence of the Morton Bay bug, Thenus orientalis, is documented, which makes it the second mitogenome for species of the family Scyllaridae and the ninth for members of the superfamily Palinuroidae. Thenus orientalis has a mitogenome of 16,826 base pairs consisting of 13 protein-coding genes, 2 ribosomal subunit genes, 23 transfer RNAs, and a non-coding AT-rich region. The base composition of the T. orientalis mitogenome is 31.31% for T, 23.77% for C, 31.05% for A, and 13.87% for G, with an AT bias of 62.36%. In addition to a duplicated trnS1 and several other tRNA gene rearrangements, the mitogenome gene order has novel protein coding gene order with the nad6 and cob genes translocated as a block to a location downstream of the nad3 gene.

Mitogenomic sequences effectively recover relationships within brush-footed butterflies (Lepidoptera: Nymphalidae).

BackgroundMitogenomic phylogenies have revealed well-supported relationships for many eukaryote groups. In the order Lepidoptera, 113 species mitogenomes had been sequenced (May 14, 2014). However, these data are restricted to ten of the forty-three recognised superfamilies, while it has been challenging to recover large numbers of mitogenomes due to the time and cost required for primer design and sequencing. Nuclear rather than mitochondrial genes have been preferred to reconstruct deep-level lepidopteran phylogenies, without seriously evaluating the potential of entire mitogenomes. Next-generation sequencing methods remove these limitations by providing efficiently massive amounts of sequence data. In the present study, we simultaneously obtained a large number of nymphalid butterfly mitogenomes to evaluate the utility of mitogenomic phylogenies by comparing reconstructions to the now quite well established phylogeny of Nymphalidae.ResultsWe newly obtained 30 nymphalid mitogenomes via pyrosequencing on the Roche 454 GS Junior system, and combined these sequences with publicly accessible data to provide a 70-taxa dataset covering 37 genes for a 15,495 bp alignment. Polymorphic sites were not homogeneously distributed across the gene. Two gene regions, nad6 and 3’ end of nad5, were most variable, whereas the cox1 and 5’ ends of rrnL were most conserved. Phylogenetic relationships inferred by two likelihood methods were congruent and strongly supported (>0.95 posterior probability; ML bootstrap >85%), across the majority of nodes for multiple partitioning strategies and substitution models. Bayes factor results showed that the most highly partitioned dataset is the preferred strategy among different partitioning schemes. The most striking phylogenetic findings were that the subfamily Danainae not Libytheinae was sister of the remaining brush-footed butterflies and that, within Limenitidini, the genus Athyma was clearly polyphyletic. None of the single-gene phylogenies recovered the highly supported topologies generated on the basis of the whole mitogenomic data.ConclusionsThirty mitogenomes were assembled with 89% completeness from the contigs of pyrosequencing-derived reads. Entire mitogenomes or higher-quality sequences could be obtained by increasing pyrosequencing read coverage or by additional Sanger sequencing. Our mitogenomic phylogenies provide robust nodal support at a range of levels, demonstrating that mitogenomes are both accurate and efficient molecular markers for inferring butterfly phylogeny.Electronic supplementary materialThe online version of this article (doi:10.1186/1471-2164-15-468) contains supplementary material, which is available to authorized users.