Genome size variation in butterflies (Insecta,Lepidotera, Papilionoidea): a thorough phylogenetic comparison

Butterflies have been of great interest to naturalists for centuries, and the study of butterflies has been an integral part of ecology and evolution ever since Darwin proposed his theory of natural selection in 1859. There are > 18 000 butterfly species worldwide, showing great diversity in morphological traits and ecological niches. Compared with butterfly diversity, however, patterns of genome size variation in butterflies remain poorly understood, especially in a phylogenetic context. Here, we sequenced and assembled the mitogenomes of 68 butterflies and measured the genome sizes (C-values) of 67 of them. We also assembled 10 mitogenomes using reads from GenBank. Among the assembled 78 mitogenomes, those from 59 species, 23 genera and one subfamily are reported for the first time. Combining with published data of mitogenomes and genome size, we explored the patterns in genome size variation for 106 butterfly species in a phylogenetic context based on analyses of mitogenomes from 264 species covering six families. Our results show that the genome size of butterflies has a 6.4-fold variation ranging from 0.203 pg (199 Mb) (Nymphalidae: Heliconius xanthocles) to 1.287 pg (1253 Mb) (Papilionidae: Parnassius orleans). Within families, the largest variation was found in Papilionidae (5.9-fold: 0.22–1.29 pg), followed by Nymphalidae (4.8-fold: 0.2–0.95 pg), Pieridae (4.4-fold: 0.22–0.97 pg), Hesperiidae (2.2-fold: 0.3–0.66 pg), Lycaenidae (2.6-fold: 0.39–1.02 pg) and Rioidinidae (1.8-fold: 0.48–0.87 pg). Our data also suggest that butterflies have an ancestral genome size of c. 0.5 pg, and some ancestral genome size increase or decrease events along different subfamilies or tribes produce the diversity of genome size variation in diverse butterflies. Our data provide novel insights into patterns of genome size variation in butterflies and are an important reference for future genome sequencing programmes.     

Mitochondrial genomes of three Bostrichiformia species and phylogenetic analysis of Polyphaga (Insecta,Coleoptera)

Here we determined mitogenomes of three Bostrichiformia species. These data were combined with 51 previously sequenced Polyphaga mitogenomes to explore the higher-level relationships within Polyphaga by using four different mitogenomic datasets and three tree inference approaches. Among Polyphaga mitogenomes we observed heterogeneity in nucleotide composition and evolutionary rates, which may have affected phylogenetic inferences across the different mitogenomic datasets. Elateriformia, Cucujiformia, and Scarabaeiformia were each inferred to be monophyletic by all analyses, as was Bostrichiformia by most analyses based on two datasets with low heterogeneity. The large series Staphyliniformia was never recovered as monophyletic in our analyses. The Bayesian tree using a degenerated nucleotide dataset (P123_Degen) and a site-heterogeneous mixture model in PhyloBayes was supported as the best Polyphaga phylogeny: (Scirtiformia, (Elateriformia, ((Bostrichiformia, Cucujiformia), (Scarabaeiformia + Staphyliniformia)))). For Cucujiformia, the largest series, we inferred a superfamily-level phylogeny: ((Cleroidea, Coccinelloidea), (Tenebrionoidea, (Cucujoidea + Curculionoidea + Chrysomeloidea))).     

Comparative Mitogenomic Analysis of Damsel Bugs Representing Three Tribes in the Family Nabidae (Insecta: Hemiptera)

Background

Nabidae, a family of predatory heteropterans, includes two subfamilies and five tribes. We previously reported the complete mitogenome of Alloeorhynchus bakeri, a representative of the tribe Prostemmatini in the subfamily Prostemmatinae. To gain a better understanding of architecture and evolution of mitogenome in Nabidae, mitogenomes of five species representing two tribes (Gorpini and Nabini) in the subfamily Nabinae were sequenced, and a comparative mitogenomic analysis of three nabid tribes in two subfamilies was carried out.

Methodology/Principal Findings

Nabid mitogenomes share a similar nucleotide composition and base bias, except for the control region, where differences are observed at the subfamily level. In addition, the pattern of codon usage is influenced by the GC content and consistent with the standard invertebrate mitochondrial genetic code and the preference for A+T-rich codons. The comparison among orthologous protein-coding genes shows that different genes have been subject to different rates of molecular evolution correlated with the GC content. The stems and anticodon loops of tRNAs are extremely conserved, and the nucleotide substitutions are largely restricted to TψC and DHU loops and extra arms, with insertion-deletion polymorphisms. Comparative analysis shows similar rates of substitution between the two rRNAs. Long non-coding regions are observed in most Gorpini and Nabini mtDNAs in-between trnI-trnQ and/or trnS2-nad1. The lone exception, Nabis apicalis, however, has lost three tRNAs. Overall, phylogenetic analysis using mitogenomic data is consistent with phylogenies constructed mainly form morphological traits.

Conclusions/Significance

This comparative mitogenomic analysis sheds light on the architecture and evolution of mitogenomes in the family Nabidae. Nucleotide diversity and mitogenomic traits are phylogenetically informative at subfamily level. Furthermore, inclusion of a broader range of samples representing various taxonomic levels is critical for the understanding of mitogenomic evolution in damsel bugs.     

The complete mitochondrial genome of Eterusia aedea (Lepidoptera,Zygaenidae) and comparison with other zygaenid moths

Zygaenidae comprises >1036 species, including many folivorous pests in agriculture. In the present study, the complete mitochondrial genome (mitogenome) of a major pest of tea trees, Eterusia aedea was determined. The 15,196-bp circular genome contained the common set of 37 mitochondrial genes (including 13 protein-coding genes, two rRNA genes, and 22 tRNA genes) and exhibited the similar genomic features to reported Zygaenidae mitogenome. Comparative analyses of Zygaenidae mitogenomes showed a typical evolutionary trend of lepidopteran mitogenomes. In addition, we also investigated the gene order of lepidopteran mitogenomes and proposed that the novel gene order trnA-trnR-trnN-trnE-trnS-trnF from Zygaenidae and Gelechiidae and most other gene rearrangements of this tRNA cluster evolved independently. Finally, the mitogenomic phylogeny of Lepidoptera was reconstructed based on multiple mitochondrial datasets. And all the phylogenetic results revealed the sister relationships of Cossoidea and Zygaenoidea with both BI and ML methods, which is the first stable mitogenomic evidence for this clade.     

The complete mitochondrial genome of <Emphasis Type="Italic">Damora sagana</Emphasis> and phylogenetic analyses of the family Nymphalidae

The monotypic genus Damora (Nymphalidae, Heliconiinae) contains a single species, Damora sagana, which is widely distributed across southern China. Herein, its complete mitogenome was sequenced to further understand lepidopteran mitogenome characteristics, reconstruct the nymphalid family phylogeny, and infer the subdivision of Heliconiinae species. The circular mitogenome was 15,151 bp long, abundant in A and T, and comprised of 13 protein-coding genes (PCGs), 22 transfer RNA (tRNA) genes, 2 ribosomal RNA (rRNA) genes, and one control region with a gene arrangement typical of lepidopteran mitogenomes. ATN codons initiated all PCGs, except cytochrome c oxidase subunit 1 (COX1), which was initiated by a CGA sequence as has been observed in other lepidopterans. Three PCGs (COX1, COX2 and ND4) employed a single T termination signal, whereas others had the typical complete termination codon (TAA). All tRNA genes were folded into the typical cloverleaf structure except for tRNA-Ser (AGN). The A+T-rich region included the conserved motif ‘ATAGA’ followed by a 17 bp poly-T stretch, which was also observed in tribe Argynnini mitogenomes. A phylogenetic tree was constructed via multiple methods using the 13 PCGs data of D. sagana and other available mitogenomes of nymphalid species. All three phylogenetic trees yielded the same topology. These results were consistent with those from previous studies of most major nymphalid groups, except those regarding tribe subdivision in certain subfamilies such as Argynnini + (Acraeini + Heliconiini) for Heliconiine. Furthermore, our analyses identified that the genus Cethosia was grouped with the genus Acraea composing the tribe Acraeini with strong support.     

Structural features of the mitogenome of the leafhopper genus Cladolidia (Hemiptera: Cicadellidae: Coelidiinae) and phylogenetic implications in Cicadellidae

The first two complete mitogenomes of the leafhopper genus Cladolidia (C. biungulata and C. robusta) were sequenced and annotated to further explore the phylogeny of Cladolidia. Both the newly sequenced mitogenomes have a typical circular structure, with lengths of 15,247 and 15,376 bp and A + T contents of 78.2% and 78%, respectively. We identified a highly conserved genome organization in the two Cladolidia spp. through comparative analysis that included the following assessments: genome content, gene order, nucleotide composition, codon usage, amino acid composition, and tRNA secondary structure. Moreover, we detected the base heterogeneity of Cicadellidae mitogenomic data and constructed phylogenetic trees using the nucleotide alignments of 12 subfamilies of 58 leafhopper species. We noted a weak heterogeneity in the base composition among the Cicadellidae mitogenomes. Phylogenetic analyses showed that the monophyly of each subfamily was generally well supported in the family Cicadellidae; the main topology was as follows: (Deltocephalinae + (Treehoppers + ((Megophthalminae + (Macropsinae + (Hylicinae + (Coelidiinae +Iassinae)) + (Idiocerinae + (Cicadellinae + (Typhlocybinae + (Mileewinae + (Evacanthinae +Ledrinae)))))))))). Within Coelidiinae, phylogenetic analyses revealed that C. biungulata and C. robusta belong to Coelidiinae and the monophyly of Cladolidia is well supported. In addition, on the basis of complete mitogenome phylogenetic analysis and the comparison of morphological characteristics, we further confirm the genus Olidiana as a paraphyletic group, suggesting that the genus may need taxonomic revisions.     

Exploring taxonomic diversity and biogeography of the family Nemacheilinae (Cypriniformes)

Nemacheilidae, in the superfamily Cobitoidea, is comprised of many of morphologically similar fish species that occur in Eurasian water bodies. This large group shows inconsistencies between traditional morphological taxonomy and molecular phylogenetic data. We used mitochondrial genomes, recombinase‐activating gene proteins 1 (RAG1) and the mitochondrial cytochrome c oxidase I gene (COI) to study the phylogenetic relationships among Nemacheilidae species using Bayesian inference and maximum likelihood approaches. Phylogenetic analyses based on mitogenomes provided support for two clades (I and II). The mitogenomes, RAG1, and COI results indicated that several species and genera were not consistent with the traditional morphological subdivisions. The two clades inferred from mitogenomes showed clear geographical patterns. The Tibetan Plateau, Hengduan Mountains, and the Iran Plateau may act as a barrier dividing the clades. The estimated timing of clades separation (36.05 million years ago) coincides with the first uplift of the Tibetan Plateau. We conclude that the geological history of the Tibetan Plateau played a role in the diversification and distribution of the Nemacheilidae taxa. These results provided a phylogenetic framework for future studies of this complex group.     

A phylogeny of ranid frogs (Anura: Ranoidea: Ranidae), based on a simultaneous analysis of morphological and molecular data

During the last two decades, major taxonomic rearrangements were instituted in the anuran family Ranidae. Most of these changes were not based on phylogenetic analysis, and many are controversial. Addressing the phylogeny of Ranidae requires broader taxon sampling within the superfamily Ranoidea, the phylogenetic relationships and higher classification of which are also in a state of flux. No comprehensive attempt has yet been made to reconstruct ranid phylogeny using both morphological and molecular data. In the present contribution, data from 178 organismal characters were collated for 74 exemplar species representing the families Arthroleptidae, Hemisotidae, Hyperoliidae, Mantellidae Microhylidae, Petropedetidae, Rhacophoridae, Sooglossidae, and most subfamilies of Ranidae. These were combined with ～1 kb of DNA sequence from the mitochondrial 12S rDNA and 16S rDNA gene regions in a simultaneous parsimony analysis with direct optimization. Results support the classification of Hemisus with the brevicipitine microhylids, confirm that Arthroleptidae (and its two component subfamilies Astylosterninae and Arthroleptinae) are monophyletic, and advocate the recognition of Leptopelidae. Monophyly of Ranidae is compromised by recognition of Petropedetidae, Rhacophoridae and Mantellidae, which should be recognized as subfamilies of Ranidae at present. Furthermore, Petropedetidae was found to be grossly paraphyletic, comprising three clades which are all considered separate subfamilies of Ranidae, i.e., Petropedetinae, Phrynobatrachinae and Cacosterninae. Three well defined subfamilies of Ranidae were consistently retrieved as monophyletic in a sensitivity analysis, i.e., Tomopterninae, Ptychadeninae and Pyxicephalinae. However, Ptychadeninae and Pyxicephalinae were embedded in Raninae and Dicroglossinae, respectively. Ceratobatrachinae is removed from Dicroglossinae. Dicroglossinae is synonymized with Pyxicephalinae. A new subfamily Strongylopinae is proposed. Raninae should be conservatively treated as a “metataxon” (sensu Ford and Cannatella, 1993 ) until such time as it is fully revised. Tomopterninae is removed from synonymy with Cacosterninae. Morphological synapomorphies are reported for major monophyletic clades retrieved in the simultaneous analysis with equal weights. The present study found that many Old World clades appear to contain both African and Asian taxa, contrary to the findings of some recent biogeographical analyses. This study demonstrates the value of broad taxonomic sampling in ranid phylogeny, and highlights the immense contribution that can be made from detailed morphological data. © The Willi Hennig Society 2005.     

Phylogeny in Echinocereus (Cactaceae) based on combined morphological and molecular evidence: taxonomic implications

Echinocereus is a morphologically diverse genus that includes 64 species grouped into eight taxonomic sections based on morphological traits. In previous molecular phylogenetic analyses, the relationships amongst Echinocereus species were not entirely revealed and useful characters to recognize clades were not provided. The inclusion of several sources of evidence in a phylogenetic analysis is likely to produce more supported hypotheses. Therefore, we performed a combined phylogenetic analysis with a set of 44 morphological characters and six chloroplast DNA sequences. Topologies from parsimony and Bayesian analyses were mostly congruent. However, the relationships of E. poselgeri were not consistent between analyses. A second Bayesian analysis using a long-branch extraction test resulted in a topology with the morphological position of E. poselgeri congruent with that in parsimony analysis. Parsimony and Bayesian analyses corroborated the monophyly of Echinocereus, which included eight monophyletic groups. The combined phylogeny integrated into different clades those taxa that were not determined in previous analyses and changed the relationships of some recognized clades. The clades did not recover the recent infrageneric classification. In the present study, a new sectional classification for Echinocereus is proposed based on the eight recovered clades, which is supported by a combination of morphological and molecular characters. An identification key for sections in the genus is included.     

Mitogenomes provide new insights of evolutionary history of Boreheptagyiini and Diamesini (Diptera: Chironomidae: Diamesinae)

Mitogenomes have been widely used for phylogenetic reconstruction of various Dipteran groups, but specifically for chironomid, they have not been carried out to resolve the relationships. Diamesinae (Diptera: Chironomidae) are important bioindicators for freshwater ecosystem monitoring, but its evolutionary history remains uncertain for lack of information. Here, coupled with one previously published and 30 new mitogenomes of Diamesinae, we carried out comparative mitogenomic analysis and phylogenetic analysis. Mitogenomes of Diamesinae were conserved in structure, and all genes arranged in the same order as the ancestral insect mitogenome. All protein‐coding genes in Diamesinae were under stronger purifying selection than those of other nonbiting midge species, which may exhibit signs of adaptation to life at cold living conditions. Phylogenetic analyses strongly supported the monophyly of Diamesinae, with Boreheptagyiini deeply nested within Diamesini. In addition, phylogenetic relationship of selected six genera was resolved, except Sympotthastia remained unstable. Our study revealed that the mitogenomes of Diamesinae are highly conserved, and they are practically useful for phylogenetic inference.     

Reconstructing intraordinal relationships in Lepidoptera using mitochondrial genome data with the description of two newly sequenced lycaenids, Spindasis takanonis and Protantigius superans (Lepidoptera: Lycaenidae)

Lepidoptera is one of the largest insect orders, but the phylogenetic relationships within this order, have yet to be adequately described. Among these unresolved relationships include those regarding the monophyly of the Macrolepidoptera and interfamilial relationships of the true butterflies superfamily Papilionoidea. We present two new mitochondrial genomes (mitogenomes) belonging to the butterfly family Lycaenidae to explore the phylogenetic relationships existing among lepidopteran superfamilies and true butterfly families from a mitogenome perspective, and to evaluate the characteristics of the lepidopteran mitogenomes. Our consensus phylogeny of the Lepidoptera largely supported the superfamilial relationships (((((Bombycoidea + Geometroidea) + Noctuoidea) + Pyraloidea) + Papilionoidea) + Tortricoidea), signifying a lack of support for a traditionally defined Macrolepidoptera. The familial relationships of the true butterflies concordantly recovered the previously proposed phylogenetic hypothesis (((Lycaenidae + Nymphalidae) + Pieridae) + Papilionidae). The test for the effect of optimization schemes (exclusion and inclusion of third codon position of PCGs and two rRNA genes, with and without partitions) on the resolution and relationships within the Lepidoptera have demonstrated that the majority of analyses did not substantially alter the relevant topology and node support, possibly as the result of relatively strong signal in mitogenomes for intraordinal relationships in Lepidoptera.     

Phylogeny, diversity, and classification of the Accipitridae based on DNA sequences of the RAG-1 exon

The avian family Accipitridae has historically been divided into subfamilies or tribes based on features such as general resemblance, feeding ecology, and behavior. Consequently, the monophyly of those groups has been questionable. Recently, three phylogenetic analyses of a majority of the genera have appeared, one based on osteology, one on DNA sequences from a single mitochondrial gene, and the third on mitochondrial plus nuclear DNA sequences, and the resulting phylogenies were in substantial disagreement concerning the composition and basal branching patterns of the clades and hence require further analysis and confirmation. Here we use DNA sequences from the large nuclear RAG-1 exon to investigate the phylogenetic relationships of these birds. Our results largely corroborated the prior study that included nuclear genes. We found strong support for a monophyletic clade comprising the secretarybird Sagittarius serpentarius , the osprey Pandion haliaetus , and the traditional accipitrids. However, every one of the traditionally recognized subfamilies of accipitrids was found to be polyphyletic. The most basal nodes in the phylogeny separate small clades of insectivorous and scavenger species, such as kites and Old World vultures, from the rest of the family. The speciose genera of bird and mammal predators are all relatively derived (terminal) in the phylogeny. Many of the basal clades are cosmopolitan in their distributions, consistent with the great mobility of these raptors. A new classification is proposed that eliminates the problem of polyphyletic intrafamilial taxa.     

The complete mitochondrial genomes of two ghost moths, Thitarodes renzhiensis and Thitarodes yunnanensis: the ancestral gene arrangement in Lepidoptera

ABSTRACT: BACKGROUND: Lepidoptera encompasses more than 160,000 described species that have been classified into 45-48 superfamilies. The previously determined Lepidoptera mitochondrial genomes (mitogenomes) are limited to six superfamilies of the most derived lepidopteran lineage Ditrysia. Compared with the ancestral insect gene order, these mitogenomes all contain a tRNA rearrangement. To gain new insights into Lepidoptera mitogenome evolution, we sequenced the mitogenomes of two ghost moths that belong to primitive lepidopteran lineages and conducted a comparative mitogenomic analysis across Lepidoptera. RESULTS: The mitogenomes of Thitarodes renzhiensis and T. yunnanensis are 16,173 bp and 15,814 bp long with an A+T content of 81.28% and 82.33%, respectively. Different tandem repeats in the A+T-rich region mainly account for the size difference between the two mitogenomes. Both mitogenomes include 13 protein-coding genes, 22 transfer RNA genes, and 2 ribosomal RNA genes. The 1,584-bp sequence from rrnS to nad2 was also determined for Thitarodes sp.QL, which has no repetitive sequence in the A+T-rich region. All three Thitarodes species possess the ancestral gene order with trnI-trnQ-trnM located between the A+T-rich region and nad2, which is different from the gene order trnM-trnI-trnQ in all previously sequenced Lepidoptera species. The formerly identified conserved elements of Lepidoptera mitogenomes (i.e. the motif 'ATAGA' and poly-T stretch in the A+T-rich region and the long intergenic spacer upstream of nad2) are absent in the Thitarodes mitogenomes. The phylogenetic analysis supports that Hepialoidea, represented by T. renzhiensis and T. yunnanensis, occupies a basal position in the currently sampled seven superfamilies. The relationships of the other six superfamilies are (((((Bombycoidea + Geometroidea) + Noctuoidea) + Pyraloidea) + Papilionoidea) + Tortricoidea). CONCLUSION: The mitogenomes of T. renzhiensis and T. yunnanensis exhibit unusual features compared with the previously determined Lepidoptera mitogenomes. Their ancestral gene order indicates that the tRNA rearrangement event occurred after Lepidoptera diverged from other holometabolous insect orders. Phylogenetic analysis based on mitogenome sequences is a power tool for addressing phylogenetic relationships among major Lepidoptera superfamilies. Characterization of the two ghost moth mitogenomes has enriched our knowledge of Lepidoptera mitogenomes and contributed to our understanding of the mechanisms underlying mitogenome evolution, especially gene rearrangements.     

Mitogenomes provide new insights into the evolutionary history of Prodiamesinae (Diptera: Chironomidae)

Evolutionary analysis of Prodiamesinae has long been impeded by lack of information, and its phylogenetic relationship with Orthocladiinae remains questionable. Here, ten complete mitochondrial genomes (mitogenomes) of Orthocladiinae sensu lato were newly sequenced, including three Prodiamesinae species and seven Orthocladiinae species. Coupled with published mitogenomes, a total of 12 mitogenomes of Orthocladiinae sensu lato were selected for a comparative mitogenomic analysis and phylogenetic reconstruction. Mitogenomes of Orthocladiinae sensu lato are conserved in structure, and all genes arrange the same gene order as the ancestral insect mitogenome. Nucleotide composition is highly biased, and the control region displayed the highest A + T content. All protein-coding genes are under purifying selection, and the ATP8 evolves at the fastest rate. In addition, the mitogenomes of Orthocladiinae sensu lato are highly conserved, and they are practically useful for phylogenetic inference, suggesting a re-classification of Orthocladiinae by sinking Prodiamesinae as a subgroup of Orthocladiinae.     

External morphology of adult Libythea celtis (Laicharting [1782]) (Lepidoptera: Nymphalidae)

Libytheinae is a morphologically unusual group of butterflies whose external anatomy has never been described in detail. The subfamily is considered basal within the diverse family Nymphalidae, and libytheines retain many plesiomorphic features, such as a functional female foreleg and smooth larva. In this paper, we describe and illustrate the adult morphology of Libythea celtis Laicharting, make comparisons to other nymphalid subfamilies, and homologize terms whenever possible. We describe major sclerites, sulci, and the presence of cranial and lateral pits on the tibia, the latter which appear to be previously unreported from the lepidopteran literature. When present, sexual dimorphism and geographic variation are noted. It is hoped that the present paper will help stabilize inconsistencies in morphological terms in nymphalid butterflies.     

A molecular phylogeny of anseriformes based on mitochondrial DNA analysis

To study the phylogenetic relationships among Anseriformes, sequences for the complete mitochondrial control region (CR) were determined from 45 waterfowl representing 24 genera, i.e., half of the existing genera. To confirm the results based on CR analysis we also analyzed representative species based on two mitochondrial protein-coding genes, cytochrome b (cytb) and NADH dehydrogenase subunit 2 (ND2). These data allowed us to construct a robust phylogeny of the Anseriformes and to compare it with existing phylogenies based on morphological or molecular data. Chauna and Dendrocygna were identified as early offshoots of the Anseriformes. All the remaining taxa fell into two clades that correspond to the two subfamilies Anatinae and Anserinae. Within Anserinae Branta and Anser cluster together, whereas Coscoroba, Cygnus, and Cereopsis form a relatively weak clade with Cygnus diverging first. Five clades are clearly recognizable among Anatinae: (i) the Anatini with Anas and Lophonetta; (ii) the Aythyini with Aythya and Netta; (iii) the Cairinini with Cairina and Aix; (iv) the Mergini with Mergus, Bucephala, Melanitta, Callonetta, Somateria, and Clangula, and (v) the Tadornini with Tadorna, Chloephaga, and Alopochen. The Tadornini diverged early on from the Anatinae; then the Mergini and a large group that comprises the Anatini, Aythyini, Cairinini, and two isolated genera, Chenonetta and Marmaronetta, diverged. The phylogeny obtained with the control region appears more robust than the one obtained with mitochondrial protein-coding genes such as ND2 and cytb. This suggests that the CR is a powerful tool for bird phylogeny, not only at a small scale (i.e., relationships between species) but also at the family level. Whereas morphological analysis effectively resolved the split between Anatinae and Anserinae and the existence of some of the clades, the precise composition of the clades are different when morphological and molecular data are compared.     

Cytochrome b and Bayesian inference of whale phylogeny

In the mid 1990s cytochrome b and other mitochondrial DNA data reinvigorated cetacean phylogenetics by proposing many novel and provocative hypotheses of cetacean relationships. These results sparked a revision and reanalysis of morphological datasets, and the collection of new nuclear DNA data from numerous loci. Some of the most controversial mitochondrial hypotheses have now become benchmark clades, corroborated with nuclear DNA and morphological data; others have been resolved in favor of more traditional views. That major conflicts in cetacean phylogeny are disappearing is encouraging. However, most recent papers aim specifically to resolve higher-level conflicts by adding characters, at the cost of densely sampling taxa to resolve lower-level relationships. No molecular study to date has included more than 33 cetaceans. More detailed molecular phylogenies will provide better tools for evolutionary studies. Until more genes are available for a high number of taxa, can we rely on readily available single gene mitochondrial data? Here, we estimate the phylogeny of 66 cetacean taxa and 24 outgroups based on Cytb sequences. We judge the reliability of our phylogeny based on the recovery of several deep-level benchmark clades. A Bayesian phylogenetic analysis recovered all benchmark clades and for the first time supported Odontoceti monophyly based exclusively on analysis of a single mitochondrial gene. The results recover the monophyly of all but one family level taxa within Cetacea, and most recently proposed super- and subfamilies. In contrast, parsimony never recovered all benchmark clades and was sensitive to a priori weighting decisions. These results provide the most detailed phylogeny of Cetacea to date and highlight the utility of both Bayesian methodology in general, and of Cytb in cetacean phylogenetics. They furthermore suggest that dense taxon sampling, like dense character sampling, can overcome problems in phylogenetic reconstruction.     

A phylogenetic study of krill (Crustacea: Euphausiacea) reveals new taxa and co-evolution of morphological characters

The first comprehensive phylogenetic study of Euphausiacea (all 86 valid species) is presented. It is based on four molecular markers and 168 morphological characters (including 58 characters of the petasma). Phylogenetic analyses support the monophyly and robustness of the families Bentheuphausidae and Euphausiidae and reveal three major clades for which we erect three new subfamilies: Thysanopodinae, Euphausiinae and Nematoscelinae. All genus-level clades are statistically supported (except Thysanopoda in molecular analyses), deeply nested within the subfamily-level clades, and encompass 14 new species groups. Copulatory structures have a major impact on tree topology in the morphological analysis, the removal of which resulted in only half the number of supported clades and genera. We revealed three groups of morphological characters, which are probably coupled with the same biological role and thus interlinked evolutionarily: (i) antennular peduncle and petasma (copulation); (ii) eyes and anterior thoracopods (feeding); and (iii) shape of carapace and pleon (defence). We analysed the evolutionary pathways of the clades into main oceanic biotopes and compared them with morphological adaptations most likely to be coupled with this process.     

Diet breadth and host plant diversity of tropical- vs. temperate-zone herbivores: South-East Asian and West Palaearctic butterflies as a case study

1. Data on host plant associations of butterflies (Papilionoidea, excluding Hesperiidae) from two biogeographical regions were used to investigate (1) whether tropical herbivores are more narrowly specialized with regard to host plant choice than those of northern temperate zones, and (2) whether tropical butterflies show a greater diversity of host plant affiliations. 2. There was no evidence for a more restricted diet breadth of tropical butterflies, with diet breadth being measured as number of host plant families used per species. In the families Papilionidae, Pieridae, and Nymphalidae, host plant ranges of West Palaearctic and South-East Asian species are similar, whereas in one speciose group within the Lycaenidae, the Polyommatini, tropical species are significantly more polyphagous. 3. Diet breadth also differs among higher butterfly taxa. While Papilionidae, Pieridae, the nymphalid subfamilies Satyrinae, Morphinae, Libytheinae and Apaturinae, as well as the temperate-zone Polyommatini in the Lycaenidae are composed predominantly of host specialists, the degree of polyphagy is higher among the remaining nymphalid subfamilies and in many lycaenids. These results challenge strongly the view that tropical herbivores are generally more specialized in this regard than herbivores of higher latitudes. Rather, chemical constraints and phylogenetic conservatism shape host plant associations in many taxa in such a way that differences between temperate and tropical representatives are slight. 4. Host plant diversity, measured as the number of plant families used per butterfly family and by application of the log-series model, is much higher in South-East Asian Nymphalidae and Lycaenidae (the two largest families) than in their Western Palaearctic relatives. No such differences are observed in the Papilionidae and Pieridae (the two smaller families). Besides effects of sample size, the strong association of papilionid and pierid butterflies with plants characterized by a small set of classes of secondary plant compounds might generally restrict their capability to utilize a broader taxonomic range of host plants. 5. The results indicate that high floral diversity can be reflected by higher diversity of host plant affiliations of herbivores, but taxonomic idiosyncrasies render it difficult to draw generalized conclusions.     

HOST PLANT UTILIZATION,HOST RANGE OSCILLATIONS AND DIVERSIFICATION IN NYMPHALID BUTTERFLIES: A PHYLOGENETIC INVESTIGATION

It has been suggested that phenotypic plasticity is a major factor in the diversification of life, and that variation in host range in phytophagous insects is a good model for investigating this claim. We explore the use of angiosperm plants as hosts for nymphalid butterflies, and in particular the evidence for past oscillations in host range and how they are linked to host shifts and to diversification. At the level of orders of plants, a relatively simple pattern of host use and host shifts emerges, despite the 100 million years of history of the family Nymphalidae. We review the evidence that these host shifts and the accompanying diversifications were associated with transient polyphagous stages, as suggested by the “oscillation hypothesis.” In addition, we investigate all currently polyphagous nymphalid species and demonstrate that the state of polyphagy is rare, has a weak phylogenetic signal, and a very apical distribution in the phylogeny; we argue that these are signs of its transient nature. We contrast our results with data from the bark beetles Dendroctonus, in which a more specialized host use is instead the apical state. We conclude that plasticity in host use is likely to have contributed to diversification in nymphalid butterflies.