New braconid wasps from French Cretaceous amber (Hymenoptera, Braconidae): synonymization with Eoichneumonidae and implications for the phylogeny of Ichneumonoidea

Two new fossils of Braconidae are described from Albian-Cenomanian amber of south-western France, Protorhyssalodes arnaudi gen. n., sp. n., and Aenigmabracon capdoliensis gen. n., sp. n. The former appears superficially similar to the type genus and species of the extinct sub-family Protorhyssalinae, from Turonian New Jersey amber specimens, and the latter both to Protorhyssalus and to members of the extinct family Eoichneumonidae. However, both new taxa display unique combinations of wing venation characters making confident assignment to sub-family impossible. Indeed, they are the first braconids ever known to possess both vein 2-CU and a distinct trace of vein 2-1A on hindwing. The new fossil taxa are incorporated into a morphological analysis of extinct and extant ichneumonoids. As a result of the analyses we synonymize the Eoichneumonidae with the Braconidae.     

Molecular systematics of caridean shrimps based on five nuclear genes: Implications for superfamily classification

Caridean shrimps are the second most diverse group of Decapoda. Over the years, several different systematic classifications, exclusively based on morphology, have been proposed, but the classification of the infraorder Caridea remains unresolved. In this study, five nuclear genes, 18S rRNA, enolase, histone 3, phosphoenolpyruvate carboxykinase and sodium–potassium ATPase α-subunit, were used to examine the systematic status of caridean families and superfamilies. We constructed gene trees based on a combined dataset of 3819 bp, containing 35 caridean species from 19 families in 11 superfamilies. At the family level, and based on our restricted representation, our molecular data support monophyly of the families Glyphocrangonidae, Crangonidae, Pandalidae, Alpheidae, Rhynchocinetidae, Nematocarcinidae, Pasiphaeidae, Atyidae and Stylodactylidae. In contrast, both the Hippolytidae and Palaemonidae are polyphyletic in our analysis. Two major clades are revealed. The Alpheidae, Hippolytidae, Crangonidae, Glyphocrangonidae, Barbouriidae, Pandalidae, Hymenoceridae, Gnathophyllidae and Palaemonidae make up the first clade, while the second clade comprises the Rhynchocinetidae, Oplophoridae, Nematocarcinidae, Alvinocarididae, Campylonotidae, Pasiphaeidae and Eugonatonotidae. Two families, Bathypalaemonellidae and Stylodactylidae, are shown to be basal groups in our tree. At the superfamily level, our results do not support the currently accepted superfamily classification, although there is support for a superfamily Palaemonoidea, though only three out of its eight families are included. The results suggest that the currently accepted superfamily classification of the Caridea does not reflect their evolutionary relationships. A major revision of the higher systematics of Caridea appears thus to be vital, ideally incorporating both molecular and morphological evidence.     

A Molecular Phylogeny of the Aphidiinae (Hymenoptera: Braconidae)

Phylogenetic relationships within the Aphidiinae, and between this and other subfamilies of Braconidae (Hymenoptera), were investigated using sequence data from three genes: elongation factor-1α, cytochrome b, and the second expansion segment of the 28S ribosomal subunit. Variation in both protein-coding genes was characterized by a high level of homoplasy, but analysis of the expansion segment—robust over a range of alignment methods and parameters—resolved some of the older divergences. Parsimony analysis of the combined data suggests the following tribal relationships: (Ephedrini + (Praini + (Aphidiini + Trioxini))). In addition, the cyclostome subfamilies were found to form a clade separate from the Aphidiinae, but relationships between the Aphidiinae and the noncyclostome braconids could not be resolved. The inferred phylogeny also supported a secondary loss of internal pupation within the Praini and a polyphyletic origin of endoparasitism within the Braconidae.     

Molecular phylogenetics of Meliaceae (Sapindales) based on nuclear and plastid DNA sequences

Phylogenetic analyses of Meliaceae, including representatives of all four currently recognized subfamilies and all but two tribes (32 genera and 35 species, respectively), were carried out using DNA sequence data from three regions: plastid genes rbcL, matK (partial), and nuclear 26S rDNA (partial). Individual and combined phylogenetic analyses were performed for the rbcL, matK, and 26S rDNA data sets. Although the percentage of informative characters is highest in the segment of matK sequenced, rbcL provides the greatest number of informative characters of the three regions, resulting in the best resolved trees. Results of parsimony analyses support the recognition of only two subfamilies (Melioideae and Swietenioideae), which are sister groups. Melieae are the only tribe recognized previously that are strongly supported as monophyletic. The members of the two small monogeneric subfamilies, Quivisianthe and Capuronianthus, fall within Melioideae and Swietenioideae, respectively, supporting their taxonomic inclusion in these groups. Furthermore, the data indicate a close relationship between Aglaieae and Guareeae and a possible monophyletic origin of Cedreleae of Swietenioideae. For Trichilieae (Melioideae) and Swietenieae (Swietenioideae) lack of monophyly is indicated.     

Molecular phylogenetics of the siphonophora (Cnidaria), with implications for the evolution of functional specialization

Siphonophores are a group of pelagic colonial hydrozoans (Cnidaria) that have long been of general interest because of the division of labor between the polyps and medusae that make up these "superorganisms." These polyps and medusae are each homologous to free living animals but are generated by an incomplete asexual budding process that leaves them physiologically integrated. They are functionally specialized for different tasks and are precisely organized within each colony. The number of functional types of polyps and medusae varies across taxa, and different authors have used this character to construct phylogenies polarized in opposite directions, depending on whether they thought siphonophore evolution proceeded by a reduction or an increase in functional specialization. We have collected taxa across all major groups of siphonophores, many of which are found exclusively in the deep sea, using remotely operated underwater vehicles (ROVs) and by SCUBA diving from ships in the open ocean. We have used 52 siphonophores and four outgroup taxa to estimate the siphonophore phylogeny with molecular data from the nuclear small subunit ribosomal RNA gene (18S) and the mitochondrial large subunit ribosomal RNA gene (16S). Parsimony reconstructions indicate that functionally specialized polyps and medusae have been gained and lost across the phylogeny. Maximum likelihood and Bayesian analyses of morphological data suggest that the transition rate for decreased functional specialization is greater than the transition rate for increased functional specialization for three out of the four investigated categories of polyps and medusae. The present analysis also bears on several long-standing questions about siphonophore systematics. It indicates that the cystonects are sister to all other siphonophores, a group that we call the Codonophora. We also find that the Calycophorae are nested within the Physonectae, and that the Brachystelia, a historically recognized grouping of short-stemmed taxa, are polyphyletic. [Cnidaria; colonial animals; deep sea; division of labor; functional specialization; Hydrozoa; phylogenetics; Siphonophores.].     

Molecular phylogeny of the order Euryalida (Echinodermata: Ophiuroidea), based on mitochondrial and nuclear ribosomal genes

The existing taxonomy of Euryalida, one of the two orders of the Ophiuroidea (Echinodermata), is uncertain and characterized by controversial delimitation of taxonomic ranks from genus to family-level. Their phylogeny was not studied in detail until now. We investigated a dataset of sequence from a mitochondrial gene (16S rRNA) and two nucleic genes (18S rRNA and 28S rRNA) for 49 euryalid ophiuroids and four outgroup species from the order Ophiurida.The monophyly of the order Euryalida was supported as was the monophyly of Asteronychidae, Gorgonocephalidae and an Asteroschematidae + Euryalidae clade. However, the group currently known as the Asteroschematidae was paraphyletic with respect to the Euryalidae. The Asteroschematidae + Euryalidae clade, which we recognise as an enlarged Euryalidae, contains three natural groups: the Asteroschematinae (Asteroschema and Ophiocreas), a new subfamily Astrocharinae (Astrocharis) and the Euryalinae with remaining genera. These subfamilies can be distinguished by internal ossicle morphology.     

Molecular phylogeny of the assassin bugs (Hemiptera: Reduviidae), based on mitochondrial and nuclear ribosomal genes

The first comprehensive cladistic analysis of Reduviidae, the assassin bugs, based on molecular data is presented and discussed in the context of a recently-published morphological analysis. Assassin bugs are essential components of ecosystems, but also important in agriculture and medicine. Sampling included 94 taxa (89 Reduviidae, 5 outgroups) in 15 subfamilies and 24 tribes of Reduviidae and is based on 3300 base pairs of mitochondrial (16S) and nuclear (18S, 28SD2, 28SD3-5) ribosomal DNA. Partitions of the dataset were aligned using different algorithms implemented in MAFFT and the combined dataset was analyzed using parsimony, partitioned maximum likelihood and partitioned Bayesian criteria. Clades recovered in all analyses, independent of alignment and analytical method, comprise: Cimicomorpha and Reduviidae; Hammacerinae; Harpactorinae; Apiomerini; Peiratinae; Phymatinae; Salyavatinae; Triatominae; Phymatinae + Holoptilinae; the higher Reduviidae (Reduviidae excluding Hammacerinae and the Phymatine Complex); Ectrichodiinae + Tribelocephalinae; (Triatominae + Zelurus) + Stenopodainae. Hammacerinae are rejected as sister group to all remaining Reduviidae in all analyses, as is the monophyly of Reduviinae, Emesinae and Harpactorini. High support values for Triatominae imply that blood-feeding has evolved only once within Reduviidae. Stenopodainae and part of Reduviinae are discussed as close relatives to Triatominae.     

Molecular identification of Aphidius transcaspicus (Hymenoptera: Braconidae: Aphidiinae), a closely associated species of Aphidius colemani,in Korea

Aphidius colemani (Braconidae: Aphidiinae) is an endoparasitoid and one of the most commercially used parasitoids for the control of various pest aphids. However, species identification of A. colemani is difficult because of the high degree of morphological similarity with closely associated species, such as A. transcaspicus and A. platensis. Recently, we identified a large-scale emergence of parasitoids from mummies of Rhopalosiphum maidis on barley banker plants in Yecheon County, Korea. The morphological characteristics of the forewing, petiole, and antenna indicated that our specimen is A. transcaspicus. The partial nucleotide sequence analysis of the mitochondrial cytochrome oxidase subunit I (COI) gene showed that it is 100% identical to Japanese specimens and 1.69% variable to Greek specimens. Analysis of the phylogenetic tree, haplotypes, and amino acid sequence of the COI gene, as well as the internal transcribed spacer 2 (ITS2) of the nuclear ribosomal DNA region, indicated that A. transcaspicus is closer to A. platensis than A. colemani, and Korean and Japanese populations were mostly diversified groups at the intraspecific level. This study provides important information to elucidate the genetic variation in A. transcaspicus and the relationships between closely related species within the A. colemani group.     

Status of Ichneumonidae (Hymenoptera: Ichneumonoidea) in Cuba

Current understanding of Cuban ichneumonids (Hymenoptera: Ichneumonoidea) is analysed. A new Subfamily (Tersilochinae) including 13 genera and two species are first reports for Cuba. In contrast with Braconidae, the sister group, the ichneumonids are poorly known because of the lack of local taxonomists and bacause nets, rather than interception, light and yellow pan traps have been used. Using Ophioninae as a reference, and comparing the percentage of ophionines in the fauna of Costa Rican ichneumonids, the ratio of species of Costa Rican to Cuban Ophioninae and the increase in the number of described Cuban ophionines, we suggest that actual Cuban biodiversity in this group may be between 400 and 600 species.     

Molecular phylogenetics of the bee-eaters (Aves: Meropidae) based on nuclear and mitochondrial DNA sequence data

The bee-eaters (family Meropidae) comprise a group of brightly colored, but morphologically homogeneous, birds with a wide variety of life history characteristics. A phylogeny of bee-eaters was reconstructed using nuclear and mitochondrial DNA sequence data from 23 of the 25 named bee-eater species. Analysis of the combined data set provided a well-supported phylogenetic hypothesis for the family. Nyctiornis is the sister taxon to all other bee-eaters. Within the genus Merops, we recovered two well-supported clades that can be broadly separated into two groups along geographic and ecological lines, one clade with mostly African resident species and the other clade containing a mixture of African and Asian taxa that are mostly migratory species. The clade containing resident African species can be further split into two groups along ecological lines by habitat preference into lowland forest specialists and montane forest and forest edge species. Intraspecific sampling in several of the taxa revealed moderate to high (3.7-6.5%, ND2) levels of divergence in the resident taxa, whereas the lone migratory taxon showed negligible levels of intraspecific divergence. This robust molecular phylogeny provides the phylogenetic framework for future comparative tests of hypotheses about the evolution of plumage patterns, sociality, migration, and delayed breeding strategies.     

Dendrosoter protuberans (Hymenoptera: Braconidae), An Important Elm Bark Beetle Parasitoid

The most favourable period for Dendrosoter protuberans development occurred when wasps were introduced 16 days after the beginning of Scolytus multistriatus development (the greatest number of emerged wasps per a single parasitoid female with a very positive sexual index and high percentage of parasitised elm bark beetle larvae). No significant differences were observed in parasitoid efficiency between the 11^th and the 21^st day. The most unfavourable period for elm bark beetle parasitisation was six days after the beginning of S. multistriatus development (a low reproduction of D. protuberans, a negative sexual index and a low number of destroyed S. multistriatus larvae). The reproduction of both the elm bark beetle and D. protuberans was greater in the first, than in the second generation. The developmental period of D. protuberans in the first generation was approximately two days longer than in the second generation. Furthermore, the development of females was 1-2 days longer than the development of males. D. protuberans developed most successfully on S. multistriatus (the most abundant parasitoid reproduction with the longest developmental period and a very positive sexual index that resulted in a high percentage of parasitised larvae) and then on S. ensifer, S. pygmaeus and Pteleobius kraatzi. On the other hand, S. scolytus larvae were the poorest host of all elm species - the reproduction of D. protuberans in this species was the scarcest with the shortest period of development and a negative sexual index, hence larval parasitisation was the lowest.     

Molecular phylogenetics of Melastomataceae and Memecylaceae: implications for character evolution

Melastomataceae are among the most abundant and diversified groups of plants throughout the tropics, but their intrafamily relationships and morphological evolution are poorly understood. Here we report the results of parsimony and maximum likelihood (ML) analyses of cpDNA sequences from the rbcL and ndhF genes and the rpl16 intron, generated for eight outgroups (Crypteroniaceae, Alzateaceae, Rhynchocalycaceae, Oliniaceae, Penaeaceae, Myrtaceae, and Onagraceae) and 54 species of melastomes. The sample represents 42 of the family's currently recognized ～150 genera, the 13 traditional tribes, and the three subfamilies, Astronioideae, Melastomatoideae, and Memecyloideae (= Memecylaceae DC.). Parsimony and ML yield congruent topologies that place Memecylaceae as sister to Melastomataceae. Pternandra, a Southeast Asian genus of 15 species of which five were sampled, is the first- branching Melastomataceae. This placement has low bootstrap support (72%), but agrees with morphological treatments that placed Pternandra in Melastomatacaeae because of its acrodromal leaf venation, usually ranked as a tribe or subfamily. The interxylary phloem islands found in Memecylaceae and Pternandra, but not most other Melastomataceae, likely evolved in parallel because Pternandra resembles Melastomataceae in its other wood characters. A newly discovered plesiomorphic character in Pternandra, also present in Memecylaceae, is a fibrous anther endothecium. Higher Melastomataceae lack an endothecium as do the closest relatives of Melastomataceae and Memecylaceae. The next deepest split is between Astronieae, with anthers opening by slits, and all remaining Melastomataceae, which have anthers opening by pores. Within the latter, several generic groups, corresponding to traditional tribes, receive solid statistical support, but relationships among them, with one exception, are different from anything predicted on the basis of morphological data. Thus, Miconieae and Merianieae are sister groups, and both are sister to a trichotomy of Bertolonieae, Microlicieae + Melastomeae, and Dissochaeteae + Blakeeae. Sonerileae/Oxysporeae are nested within Dissochaeteae, Rhexieae within Melastomeae, and African and Asian Melastomeae within neotropical Melastomeae. These findings have profound implications for our understanding of melastome morphological evolution (and biogeography), implying, for example, that berries evolved from capsules minimally four times, stamen connectives went from dorsally enlarged to basal/ventrally enlarged, and loss of an endothecium preceded poricidal dehiscence.     

Phylogeny and reclassification of Cryptini (Hymenoptera,Ichneumonidae, Cryptinae), with implications for ichneumonid higher‐level classification

The first comprehensive phylogenetic study of the wasp tribe Cryptini (Hymenoptera, Ichneumonidae, Cryptinae) is presented, based on 109 morphological characters and molecular data from seven loci. The dataset includes 370 species, 308 of which are from Cryptini, covering 182 of its 250 genera. Results from parsimony and likelihood analyses are generally congruent. The topology has several implications for ichneumonid higher‐level classification. Previous definitions of the Ichneumoniformes clade are supported, though newly including the Microleptinae. The cryptine subtribe Ateleutina is consistently recovered outside of the Cryptini clade and should be treated as a separate subfamily, Ateleutinae stat.n. The tribe Phygadeuontini is shown to be polyphyletic: while most of the sampled taxa were recovered in a single clade, many of its members are more closely related to the Ichneumoninae, Ateleutinae or Cryptini. Pending a more detailed study, the group should be treated as a separate subfamily, Phygadeuontinae stat. rev . The former Hemigastrini are recovered as largely monophyletic but with important exceptions. Hemigaster Brullé is recovered as part of the Phygadeuontini and is transferred to that group. Echthrus Gravenhorst is consistently recovered as part of Cryptini, rendering Aptesini as the correct name for the tribe. The subfamily Cryptinae should be restricted to the tribes Aptesini and Cryptini. Within Cryptini, the results show little support for the current subtribal classification, with most subtribes recovered as polyphyletic. A number of relatively stable clades are identified and discussed, but the relationships among them are weakly supported. Most of these clades are morphologically heterogeneous and building a subtribal classification based on them would be ineffectual; they are therefore treated under the informal designation of genus groups. The results highlight the ubiquity of morphological homoplasy in Cryptini, and provide a framework from which to address further systematic and evolutionary questions on this hyperdiverse group of parasitic wasps.     

Molecular phylogenetics of the tribeInuleae s. str.(Asteraceae), based on ITS sequences of nuclear ribosomal DNA

The sequences of the internal transcribed spacer (ITS) region of 18S–26S nrDNA for a sample of 16 taxa from theInuleae s. str. and two outgroup taxa are analysed cladistically with PAUP. A consensus tree of the four most parsimonious cladograms is presented. Three different tests of cladogram stability are conducted (Bremer support, parsimony jackknifing and bootstrapping); all tests indicate a high degree of support for the basal nodes of the tree. The ITS phylogeny of the tribe is compared with previous hypotheses based on morphological data. The position ofAnisopappus as sister group to the rest of the tribe is supported by the molecular data, but the proposed subdivision ofInuleae s. str. into a paleate grade group and an epaleate clade is not. The interpretation of the character evolution of, e.g. receptacular paleae and pappus features within the tribe is discussed.     

Molecular phylogenetics of the juno irises,Iris subgenus Scorpiris (Iridaceae), based on six plastid markers

Relationships among the roughly 55 species of Iris subgenus Scorpiris have been studied. A matrix of six plastid DNA regions (matK, rpl14‐rps8 spacer, infA‐rpl36 spacer, trnE‐trnT spacer, trnL intron and trnL‐F spacer) was produced from 57 accessions (52 taxa) and analysed with both parsimony and Bayesian methods. Five major clades are identified, of which four have strong geographical correlations, whereas the fifth corresponds to Iris section Physocaulon. In our results, several species are placed with species not previously considered to be related, although, in some cases, there are morphological characters that suggest that these newly indicated relationships are reasonable. For some of the other oddly grouped species, we can only assume that remarkable parallelisms in morphology have occurred or hybridization is involved. Presently, with plastid DNA as our only comprehensive data resource, we are not able to evaluate more thoroughly these more puzzling associations of species. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 167 , 281–300.     

Family planning in a stemborer parasitoid: sex ratio, brood size and size-fitness relationships in Parallorhogas pyralophagus (Hymenoptera: Braconidae), and implications for biological control

Various aspects were studied of the brood size and sex allocation strategies, and of size-fitness relationships in Parallorhogas pyralophagus (Marsh), a gregarious ectoparasitoid of Eoreuma loftini Dyar. Brood size was significantly correlated with host size; larger hosts were allocated larger broods. Brood sex ratios were fixed precisely at 1 male per 4 females, and eggs were likely to be deposited in that order; differential mortality did not contribute to this precise sex ratio. The sex allocation strategy of P. pyralophagus is likely to conform to strict, i.e. single foundress, local mate competition. Adoption of this strategy is probably influenced by a limited insemination capacity of males; a smaller proportion of females (0.09 vs. 0.21) remained virgin in broods with precise or higher sex ratios (> or = 0.20 males) relative to broods with lower than precise sex ratios (< 0.20 males). Moreover, all females were inseminated in most broods (60%) with precise or higher sex ratios, whereas this did not occur in broods with lower than precise sex ratios. The hypothesized occurrence of strict local mate competition in P. pyralophagus was supported also by observations that: (i) offspring brood sex ratios were independent of maternal brood sex ratios and number of parental females concurrently allocating offspring to a group of hosts, and; (ii) the rate of superparasitism under no-choice conditions was low (approximately 20%), suggesting that rates of outbreeding in the field are low. Other results suggested that fitness in P. pyralophagus was correlated with adult size; longevity and reproductive capacity both increased with adult size in males and females. However, adult size may be more important for females than for males because the differences in reproductive capacity between the largest and smallest individuals was up to 7.3 times greater in females versus < 2 times in males.     

Molecular phylogenetics of Paphiopedilum (Cypripedioideae; Orchidaceae) based on nuclear ribosomal ITS and plastid sequences

Phylogenetic relationships in the genus Paphiopedilum were studied using nuclear ribosomal internal transcribed spacer (ITS) and plastid sequence data. The results confirm that the genus Paphiopedilum is monophyletic, and the division of the genus into three subgenera Parvisepalum, Brachypetalum and Paphiopedilum is well supported. Four sections of subgenus Paphiopedilum (Pardalopetalum, Cochlopetalum, Paphiopedilum and Barbata) are recovered as in a recent infrageneric treatment, with strong support. Section Coryopedilum is also recovered, with low bootstrap but high posterior probability values for support of monophyly. Relationships in section Barbata remain unresolved, and short branch lengths and the narrow geographical distribution of many species in the section suggest that it possibly underwent rapid radiation. Mapping chromosome and genome size data (including some new genome size measurements) onto the phylogenetic framework shows that there is no clear trend in increase in chromosome number in the genus. However, the diploid chromosome number of 2n = 26 in subgenera Parvisepalum and Brachypetalum suggests that this is the ancestral condition, and higher chromosome numbers in sections Cochlopetalum and Barbata suggest that centric fission has possibly occurred in parallel in these sections. The trend for genome size evolution is also unclear, although species in section Barbata have larger genome sizes than those in other sections. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 176–196.