Comparative mitochondrial genomic analysis of Macrophthalmus pacificus and insights into the phylogeny of the Ocypodoidea & Grapsoidea

Grapsoidea and Ocypodoidea, two of the most abundant and economically important groups in Brachyura, are of great commercial value to fisheries and aquaculture. However, the taxonomy of Ocypodoidea and Grapsoidea has long been highly disputed. Previous studies have investigated this problem through phylogenetic analysis based on limited taxonomic sampling, with different reports proposing either monophyly or paraphyly, but no definitive conclusion has been reached. In this study, the complete mitogenome of Macrophthalmus pacificus (Ocypodoidea, Macrophthalmidae) is reported on and the relationship between Ocypodoidea and Grapsoidea is further investigated. Sequencing the M. pacificus mitogenome, which is a closed circular molecule containing a typical 37 genes, preliminarily determined the ancestral gene order of Macrophthalmidae, which is consistent with previous studies. Comparative analyses of gene order among Ocypodoidea and Grapsoidea revealed that Varunidae (Grapsoidea) and Macrophthalmidae (Ocypodoidea) have the same rearrangement, which confirms previous research. Larger data analysis revealed that these two families (Varunidae and Macrophthalmidae) cluster into a monophyletic clade as sister groups. Rearrangement and phylogeny lines of evidence is concluded that Varunidae and Macrophthalmidae may be of common origin. Furthermore, the remaining Ocypodoidea and Grapsoidea families mix paraphyletically in the phylogenetic tree. Therefore, both gene rearrangement and phylogenetic analysis support the paraphyly of Ocypodoidea and Grapsoidea, which reinforces this view. These findings provide important information regarding Brachyura's phylogenetic relationships, which demonstrates the advantage of mitogenome sequence data in phylogenetic studies.     

The Entire Mitochondrial Genome of Macrophthalmus abbreviatus Reveals Insights into the Phylogeny and Gene Rearrangements of Brachyura

Brachyuran crabs comprise the most species-rich clades among extant Decapoda and are divided into several major superfamilies. However, the phylogeny of Brachyuran remains controversial, comprehensive analysis of the overall phylogeny is still lacking. Complete mitochondrial genome (mitogenome) can indicate phylogenetic relationships, as well as useful information for gene rearrangement mechanisms and molecular evolution. In this study, we firstly sequenced and annotated the complete mitogenome of Macrophthalmus abbreviatus (Brachyura; Macrophthalmidae). The mitogenome length of M. abbreviatus is 16,322 bp, containing the entire set of 37 genes and a control region typically observed in Brachyuran mitogenomes. The genome composition of M. abbreviatus was highly A+T biased 76.3% showing positive AT-skew (0.033) and negative GC-skew (??0.351). In M. abbreviatus mitogenome, most tRNA genes were folded into the clover-leaf secondary structure except trnH, trnS1 and trnC, which was similar to the other species in Macrophthalmidae. Phylogenetic analysis showed that all families form a monophyletic, and Varunidae and Macrophthalmidae clustered into a monophyletic clade as sister groups. Comparative analyses of rearrangement among Brachyura revealed that Varunidae (Grapsoidea) and Macrophthalmidae (Ocypodoidea) had the same gene order, which reinforced the result of phylogeny. The combined results of two aspects revealed that the polyphyly of Ocypodoidea and Grapsoidea were well supported. In general, the results obtained in this research will contribute to further studies on molecular based for the classification and gene rearrangements of Macrophthalmidae or even Brachyura.

     

Characterization of the complete mitochondrial genome of Uca lacteus and comparison with other Brachyuran crabs

Brachyuran crabs comprise the most species-rich clade among the crustacean order Decapoda and are divided into several major superfamilies. However, the monophyly of the superfamilies Ocypodoidea and Grapsoidea in their current compositions within the Brachyura remains inconclusive. In this study, the complete mitochondrial genome (mitogenome) of Uca lacteus (Ocypodoidea, Ocypodidae) was sequenced, annotated, and compared with those of other Brachyuran crabs. The circular mitogenome of U. lacteus is 15,661 base pairs long and contains the entire set of 37 genes and an A + T-rich region typically observed in decapod mitogenomes. Secondary structures of several tRNAs are partly missing (trnS1), and the number of bases is significantly decreased (trnD and trnF), as discovered in many other metazoans. We compared the gene order of U. lacteus with other species of Ocypodidae and found that they are consistent. The gene rearrangement of Ocypodidae is also identical to that of the ancestor of Brachyura. However, the order of the trnH gene varies from the rearrangement of ancestral Decapoda. Accordingly, we hypothesized that this rearrangement of trnH underwent a translocation during the evolution from Decapoda to Brachyura. The phylogenetic relationship of the 81 Brachyura species and one outgroup was recovered based on 13 protein-coding genes. This analysis confirmed that U. lacteus belongs to the family Ocypodidae and established a paraphyletic relationship between Ocypodoidea and Grapsoidea.     

Subtropical Brazilian mangroves as a refuge of crab (Decapoda: Brachyura) diversity

This study characterized seven mangrove areas of different sizes, located on the southeastern Brazilian coast, with respect to their sediment and vegetation features. We also estimated the richness and composition of the brachyuran assemblages in these mangroves. Crab, vegetation and sediment data were collected during low tide. The larger mangroves showed richer and complex forests, and may be more ecologically stable than smaller mangroves. Twenty-eight species of crabs were recorded, members of nine families: Panopeidae, Pinnotheridae, Gecarcinidae, Grapsidae, Sesarmidae, Varunidae, Ocypodidae, Ucididae, and Portunidae. Ocypodoidea predominated over Grapsoidea in the smaller mangrove areas in pioneer stages of forest establishment, whereas the opposite was recorded for larger and more stable mangroves, where the forest can reach advanced stage of development. Comprido is a small mangrove, but its crab assemblage was similar to the larger ones, probably because of other factors not investigated here (e.g., megalopa settling rate). This study provided basic knowledge for developing conservation strategies for vulnerable mangrove ecosystems.     

Using a multigene phylogenetic analysis to assess generic delineation and character evolution in Verrucariaceae (Verrucariales, Ascomycota)

Verrucariaceae are a family of mostly crustose lichenized ascomycetes colonizing various habitats ranging from marine and fresh water to arid environments. Phylogenetic relationships among members of the Verrucariaceae are mostly unknown and the current morphology-based classification has never been confronted to molecular data. A multilocus phylogeny (nuLSU, nuSSU and RPB1) was reconstructed for 83 taxa representing all main genera of this family to provide a molecular phylogenetic framework necessary to assess the current morphology-based classification. Four main well-supported monophyletic groups were recovered, one of which contains seven robust monophyletic subgroups. Most genera, as traditionally delimited, were not monophyletic. A few taxonomic changes are proposed here to reconcile the morphology-based classification with the molecular phylogeny (Endocarpon diffractellum comb. nov., Heteroplacidium fusculum comb. nov., and Bagliettoa marmorea comb. nov.). Ancestral state reconstructions show that the most recent common ancestor of the Verrucariaceae was most likely crustose with a weakly differentiated upper cortex, simple ascospores, and hymenium free of algae. As shown in this study, the use of symplesiomorphic traits to define Verrucaria, the largest and type genus for the Verrucariaceae, as well as the non monophyly of the genera Polyblastia, Staurothele and Thelidium, explain most of the discrepancies between the current classification based on morphological similarity and a classification using monophyly as a grouping criterion.     

Gastric teeth of some thoracotreme crabs and their contribution to the brachyuran phylogeny

The gastric teeth of three ocypodoid species were investigated using scanning electron microscopy, and the morphological results were discussed with respect to the known food preferences. The species were chosen in particular because of contrasting ideas about their relationships within the Thoracotremata. For the genera Heloecius, Dotilla, Mictyris, and “Uca” (s. str.), we find a specific correlation of the gastric teeth with the suspension feeding. The lateral gastric teeth of Uca have no prominent lateral teeth cusps, and most of their teeth surface consists of transverse comb‐like lamellae. However, this possible food adaptation does not exclude the usability of specific teeth characters to distinguish species of suspension feeders. The closer relationship of the Dotillidae to grapsoid lines of gecarcinid or sesarmid crabs suggested by molecular data is not supported by the gastric teeth. For the genus Ucides, we found several characters that distinguish Ucides from the remaining ocypodoid genera Heloecius, Dotilla, Mictyris, and “Uca.” In particular, the structures of the lateral and the dorsomedian teeth show some similarities to genera of the Gecarcinidae and Sesarmidae. Our results suggest that foregut characters can be used for phylogenetic analyses. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

Reconciling taxonomy and phylogeny in the bristleworm family Eunicidae (polychaete,Annelida)

Eunicid annelids inhabit diverse marine habitats worldwide, have ecological and economic importance and have been pictured in the news as giant predator worms. They compose a traditional stable taxon recently supported as monophyletic but characterized by plesiomorphies. Most genera within the family have been recovered as paraphyletic in previous studies. We present a phylogenetic hypothesis for eunicid based on molecular (COI, 16S rDNA, 18S rDNA) and morphological data (213 characters), including an explicit attempt to account for serial homology. Eunicidae as well as monophyletic genera Marphysa sensu stricto and Lysidice is redefined based on synapomorphies. Nematonereis is synonymized to Lysidice. Leodice and Nicidion are resurrected to name monophyletic groups including species previously included in Eunice and Marphysa sensu lato. Traditional diagnostic characters such as the absence/presence of peristomial cirri, lateral antennae and branchiae are homoplasies and not informative at the generic level. Different coding of traditional characters (i.e. articulation of prostomial appendages) and novel characters of prostomial features and regionalization of the body support the monophyly of the family and genera level clades. Thus, the phylogenetic hypothesis presented here and the evolution of characters provided background information for taxonomic changes yielding evolutionary meaningful classification and diagnoses for the family and genera.     

An assessment of the status of Polycirridae genera (Annelida: Terebelliformia) and evolutionary transformation series of characters within the family

A phylogenetic analysis was performed to determine the monophyly of non‐monotypic genera of the terebelliform family Polycirridae, i.e. Polycirrus, Amaeana, Lysilla, and Hauchiella, and the evolution of characters among members of this clade. The monotypic genera, Enoplobranchus and Biremis, were also included, together with members of both known species in Hauchiella. Representative species were included for remaining genera: 14 species of Polycirrus, six species of Amaeana, and six species of Lysilla. Out‐groups consisted of representatives of Spionidae, Cirratulidae, and Sabellariidae, as well as several species of Telothelepodidae. A total of 40 in‐ and out‐group species were coded for 50 subjects (‘characters’) and 117 subject–predicate relationships (‘states’). Although results are consistent with recent phylogenetic studies within Terebelliformia that suggest Polycirridae monophyly, only Hauchiella was found to be monophyletic, albeit part of the more inclusive clade comprising remaining polycirrid genera. Evolutionary transformation series are discussed for selected characters in relation to the non‐monophyly of Polycirrus, Lysilla, and Amaeana. Implications for the use of supraspecific taxa as ‘taxonomic surrogates’ are highlighted. The definition of Polycirridae is emended. © 2015 The Linnean Society of London     

Reconstruction of phylogenetic relationships within Grapsidae (Crustacea: Brachyura) and comparison of trans-isthmian versus amphi-atlantic gene flow based on mtDNA

Following taxonomic revisions in recent years, the originally large family Grapsidae MacLeay, 1838 has become a relatively small and morphologically homogeneous family in terms of adult and larval morphology. Most available molecular studies including more than one genus of the family have also suggested monophyly of the corresponding taxa. However, no single phylogenetic study has ever included all constituent genera of the Grapsidae. In the current study, a molecular phylogeny based on sequences of the mitochondrial 16S rRNA gene from all eight grapsid genera and 34 species is presented and suggests that up to four genera are not monophyletic. This is mainly due to the polyphyletic nature of the genus Pachygrapsus which can be found in six different lineages of the phylogeny, suggesting that the genus currently does not represent a single evolutionary lineage and is in need of taxonomic revision. Amphi-atlantic and trans-isthmian species pairs or populations in four genera are compared and reveal relatively constant and pronounced divergences across the Panama Isthmus as opposed to moderate divergences across the Atlantic Ocean, thereby suggesting occurrence of gene flow across the Atlantic Ocean during the past three million years.     

白纹方蟹线粒体基因组全序列测定及方蟹总科比较分析

为探讨该总科内部亲缘关系及其与线粒体基因排序之间的相关性,研究以方蟹科(Grapsidae)白纹方蟹(Grapsus albolineatus)为代表种,测定其线粒体基因组全序列。其全长为15577 bp,包含13个蛋白编码基因,22个tRNA基因, 2个rRNA基因和1个控制区。基因组碱基组成为33.4%A、12.0%G、20.6%C和34.0%T,具有明显的AT偏向性(67.4%)。除ATP8和ND1以GTG作为起始密码子外,其余蛋白编码基因均以ATN作为起始密码子;除COⅡ和Cyt b以T作为不完全终止密码子外,其余基因均以TAN作为终止密码子。亮氨酸(Leu)和半胱氨酸(Cys)分别是使用频率最高(15.28%)和最低(0.81%)的两种密码子。除tRNA-Ser1缺少DHU臂外,其余tRNA均能形成典型的三叶草结构。基于13个蛋白编码基因的核苷酸序列同时构建了方蟹总科的贝叶斯树(BI)和最大似然树(ML),两种方法构建的系统发育树扑拓结构一致,均显示所有方蟹科(Grapsidae)种类聚在一起,其中白纹方蟹与同属的细纹方蟹(G. tenuicrustatus)的亲缘关系最近;...     

On the marine sister groups of the freshwater crabs (Crustacea: Decapoda: Brachyura)

Freshwater crab sister group relationships with marine eubrachyuran families were investigated. A morphology-based cladistic analysis was conducted on representatives of the freshwater crab families Deckeniidae, Gecarcinucidae, Parathelphusidae, Potamidae, Potamonautidae, Pseudothelphusidae, and Trichodactylidae using a disparate assemblage of marine heterotreme and thoracotreme brachyurans as possible sister groups. The monophyly of the freshwater crabs sensu lato is falsified. The family Trichodactylidae and the marine portunid subfamily Carcininae form basal groups within the superfamily Portunoidea. The monophyly of the Pseudothelphusidae and the Paleotropical freshwater crab families is supported, and this clade is the sister group of the Thoracotremata (Gecarcinidae, Grapsidae s.l., and Ocypodoidea). The origin, groundplan, and diversification of freshwater crabs are discussed in the context of previously published scenarios of their evolution.     

A revision of the higher taxonomy of the Afrotropical freshwater crabs (Decapoda: Brachyura) with a discussion of their biogeography

The higher taxonomy of the 20 known genera of Afrotropical freshwater crabs is revised to reflect the evolutionary relationships revealed by the consensus of a series of recent morphological and molecular phylogenetic studies of the group. The Afrotropical freshwater crab genera fall into two monophyletic groups, one from Socotra with two genera (Potamidae) and another that includes the remaining 18 genera. The latter group, which includes the bulk of the region's freshwater crab fauna, forms a well-supported monophyletic clade. We recognize two monophyletic sister groups (subfamilies) within the Potamonautidae, one for seven genera from Africa (the Potamonautinae) and one for 11 genera from Africa, the Seychelles, and Madagascar (the Deckeniinae). The Deckeniinae includes two monophyletic groups (tribes), one with seven genera from Madagascar (the Hydrothelphusini), and one with four genera from Africa and the Seychelles (the Deckeniini). The Deckeniini is further divided here into two subtribes, the Deckeniina and the Globonautina. The Platythelphusidae is not recognized, and the Deckeniidae and Globonautinae are lowered in rank. There is no phylogenetic support for the continued inclusion of any genus from the Afrotropical region in the Gecarcinucidae which is treated here as an exclusively Oriental family. The Afrotropical freshwater crabs (excluding those from Socotra) form a monophyletic assemblage that has no representatives outside of the region. The wider biogeographical implications of the taxonomic revision are discussed.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 399–413.     

Molecular phylogeny, taxonomy, and evolution of nonmarine lineages within the American grapsoid crabs (Crustacea: brachyura)

Grapsoid crabs are best known from the marine intertidal and supratidal. However, some species also inhabit shallow subtidal and freshwater habitats. In the tropics and subtropics, their distribution even includes mountain streams and tree tops. At present, the Grapsoidea consists of the families Grapsidae, Gecarcinidae, and Mictyridae, the first being subdivided into four subfamilies (Grapsinae, Plagusiinae, Sesarminae, and Varuninae). To help resolve phylogenetic relationships among these highly adaptive crabs, portions of the mitochondrial genome corresponding to the 16S rRNA gene were sequenced for all grapsoid genera occurring in America. The resulting phylogeny confirms most of the present grapsid subfamilies but suggests reclassification of some of the genera and recognition of new taxonomic units. The two American gecarcinid genera might not represent a sister group to the Grapsidae but rather appear to have evolved within the latter. Colonization of inland habitats evolved in several lineages of the grapsoids, resulting in various forms of nonmarine life and different degrees of independence from the sea.     

Morphology of the first zoeal stages of eleven Sesarmidae (Crustacea,Brachyura, Thoracotremata) from the Indo-West Pacific,with a summary of familial larval characters

Summary

The first zoeal stages of eleven species of Sesarmidae from the Indo-West Pacific were obtained from ovigerous females. Those of Labuanium scandens, L. rotundatum, L. trapezoideum, L. politum, Metasesarma aubryi, Pseudosesarma crassimanum, Stelgistra stormi, and Sesarmops impressum, are described for the first time, while the first zoeal stages of Clistocoeloma merguiense, Metasesarma obesum and Sesarmops intermedium are re-described. Larval characters of all these species are compared with previously described ones for the family and morphological features are re-evaluated. Minute spines on the telson of the zoeae are described as a new larval character in Sesarmidae and their presence or absence in other grapsoid groups is discussed. The results demonstrate that a recurrent combination of reliable larval characters distinguishes zoeae and megalopae of the examined sesarmids from the rest of the Grapsoidea. This appears to be consistent with recent studies that redefine the Sesarmidae.     

Reconstructing the phylogeny of the Sipuncula

Sipunculans are marine spiralian worms with possible close affinities to the Mollusca or Annelida. Currently 147 species, 17 genera, 6 families, 4 orders and 2 classes are recognized. In this paper we review sipunculan morphology, anatomy, paleontological data and historical affiliations. We have conducted cladistic analyses for two data sets to elucidate the phylogenetic relationships among sipunculan species. We first analyzed the relationships among the 45 species of Phascolosomatidea with representatives of the Sipunculidea as outgroups, using 35 morphological characters. The resulting consensus tree has low resolution and branch support is low for most branches. The second analysis was based on DNA sequence data from two nuclear ribosomal genes (18S rRNA and 28S rRNA) and one nuclear protein-coding gene, histone H3. Outgroups were chosen among representative spiralians. In a third analysis, the molecular data were combined with the morphological data. Data were analyzed using parsimony as the optimality criterion and branch support evaluated with jackknifing and Bremer support values. Branch support for outgroup relationships is low but the monophyly of the Sipuncula is well supported. Within Sipuncula, the monophyly of the two major groups, Phascolosomatidea and Sipunculidea is not confirmed. Of the currently recognized families, only Themistidae appears monophyletic. The Aspidosiphonidae, Phascolosomatidae and Golfingiidae would be monophyletic with some adjustments in their definition. The Sipunculidae is clearly polyphyletic, with Sipunculus nudus as the sister group to the remaining Sipuncula, Siphonosoma cumanense the sister group to a clade containing Siphonosoma vastumand the Phascolosomatidea, and Phascolopsis gouldi grouping within the Golfingiiformes, as suggested previously by some authors. Of the genera with multiple representatives, only Phascolosoma and Themiste are monophyletic as currently defined. We are aiming to expand our current dataset with more species in our molecular database and more detailed morphological studies.     

Higher‐level phylogeny of the insect order Hemiptera: is Auchenorrhyncha really paraphyletic?

The higher‐level phylogeny of the order Hemiptera remains a contentious topic in insect systematics. The controversy is chiefly centred on the unresolved question of whether or not the hemipteran suborder Auchenorrhyncha (including the extant superfamilies Fulgoroidea, Membracoidea, Cicadoidea and Cercopoidea) is a monophyletic lineage. Presented here are the results of a multilocus molecular phylogenetic investigation of relationships among the major hemipteran lineages, designed specifically to address the question of Auchenorrhyncha monophyly in the context of broad taxonomic sampling across Hemiptera. Phylogenetic analyses (maximum parsimony, maximum likelihood and Bayesian inference) were based on DNA nucleotide sequence data from seven gene regions (18S rDNA, 28S rDNA, histone H3, histone 2A, wingless, cytochrome c oxidase I and NADH dehydrogenase subunit 4) generated from 86 in‐group exemplars representing all major lineages of Hemiptera (plus seven out‐group taxa). All combined analyses of these data recover the monophyly of Auchenorrhyncha, and also support the monophyly of each of the following lineages: Hemiptera, Sternorrhyncha, Heteropterodea, Heteroptera, Fulgoroidea, Cicadomorpha, Membracoidea, Cercopoidea and Cicadoidea. Also presented is a review of the major lines of morphological and molecular evidence for and against the monophyly of Auchenorrhyncha.     

Phylogeny and taxonomy of casebearer moths (Lepidoptera,Coleophoridae) based on morphological and molecular genetic data. 1. Reconstruction of phylogeny of coleophoridae using analysis of <Emphasis Type="Italic">COI</Emphasis> gene variability

A phylogenetic study of representatives of the family Coleophoridae was conducted using a comprehensive approach, including methods of morphological and molecular genetic analyses. The existent data on the family system were compared with the results of phylogenetic analysis of the COI mitochondrial gene sequences. Four of the five studied subfamilies (Coleophorinae, Ischnophaninae, Augasminae, and Tolleophorinae) corresponded to their location on the phylogram; representatives of Metriotinae were part of Coleophorinae. According to the aggregate data from molecular phylogeny and morphology, the most numerous subfamily of casebearers, Coleophorinae, is polyphyletic within its current boundaries. The results of our analysis of COI molecular divergence does not refute the monophyly of the tribes Casignetellini, Carpochenini, Klinzigedini, Goniodomini, Casasini, and Atractulini from the subfamily Coleophorinae. The allocation of the tribes Aporipturini and Sistrophoecini within the subfamily does not correspond to the molecular data. Monophyly of the genera Ecebalia, Perygra, and Casignetella was confirmed. These genera are well isolated, which reflects the evolutionary significance of the morphological characters chosen for their taxonomic division. The boundaries of the cluster containing these genera correspond to those of the tribe Casignetellini, justifying the allocation of this tribe within the subfamily. The existence of monophyletic tribes Goniodomini (genus Goniodoma) and Carpochenini (genera Ionescumia, Carpochena, and Falkmisa) was also supported. The exceptions were the genera Kasyfia, Tollsia, and Agapalsa, whose monophyly was not confirmed by our results. The distribution of the sequences of species of these genera indicated a paraphyletic origin of Kasyfia and Tollsia and a polyphyletic origin of Agapalsa.     

Sensitive phylogenetics of Clematis and its position in Ranunculaceae

Ranunculaceae are a nearly cosmopolitan plant family with the highest diversity in northern temperate regions and with relatively few representatives in the tropics. As a result of their position among the early diverging eudicots and their horticultural value, the family is of great phylogenetic and taxonomic interest. Despite this, many genera remain poorly sampled in phylogenetic studies and taxonomic problems persist. In this study, we aim to clarify the infrageneric relationships of Clematis by greatly improving taxon sampling and including most of the relevant subgeneric and sectional types in a simultaneous dynamic optimization of phenotypic and molecular data. We also investigate how well the available data support the hypothesis of phylogenetic relationships in the family. At the family level, all five currently accepted subfamilies are resolved as monophyletic. Our analyses strongly imply that Anemone s.l. is a grade with respect to the Anemoclema + Clematis clade. This questions the recent sinking of well‐established genera, including Hepatica, Knowltonia and Pulsatilla, into Anemone. In Clematis, 12 clades conceptually matching the proposed sectional division of the genus were found. The taxonomic composition of these clades often disagrees with previous classifications. Phylogenetic relationships between the section‐level clades remain highly unstable and poorly supported and, although some patterns are emerging, none of the proposed subgenera is in evidence. The traditionally recognized and horticulturally significant section Viorna is both nomenclaturally invalid and phylogenetically unsupported. Several other commonly used sections are likewise unjustified. Our results provide a phylogenetic background for a natural section‐level classification of Clematis.     

A morphological cladistic analysis of Lecythidoideae with emphasis on Bertholletia, Corythophora, Eschweilera, and Lecythis

A cladistic analysis was conducted to test the monophyly of Eschweilera and Lecythis as well as to examine the relationships of these two genera and their close relatives Bertholletia and Corythophora. The study included 86 species, representing all four genera and covering the range of taxonomic and morphological variation in the genera. The data matrix included 49 parsimony-informative characters derived from vegetative, floral, fruit, and seed morphology and anatomy. The results based on the consensus of all most parsimonious trees indicate that Bertholletia, Corythophora, Eschweilera, and Lecythis form a clade supported by brachyparacytic stomata, the absence of pedicels (with subsequent reversals in several clades), a two or four-locular ovary, the presence of an aril, and the absence of cotyledons. Within the clade, the monophyly of Corythophora is supported by the presence of inflorescence scales and the absence of nectar. Eschweilera is monophyletic only if E. congestiflora and E. simiorum are excluded. The monophyly of Eschweilera is supported by the presence of a two-locular ovary. Lecythis is not monophyletic, but sections Corrugata, Pisonis, and Poiteaui are monophyletic. Three species of section Lecythis are more closely related to Eschweilera, and other species of section Lecythis along with Bertholletia excelsa remain as unresolved.     

Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta)

The phylogeny of the basal hymenopteran lineages, including representatives of all ‘symphytan’ families, is anal; In total, 236 morphological characters were scored for 44 exemplars, including six outgroup, two xyelic tenthredinoid, five pamphilioid, three cephoid, five ‘siricoid’, one orussid, and six apocritan taxa. The datas analysed with parsimony under equal weights and under implied weights. The monophyly of the Hymenopte strongly supported but the sistergroup of the Hymenoptera cannot be identified with confidence. The relations of the ‘symphytan’ lineages are Xyeloidea +(Tenthredinoidea+ (Pamphilioidea + (Cephoidea + (Ariaxyelic (Siricidae + (Xiphydriidae +(Orussoidea+Apocrita))))))). Many of the relationships between the superfamilies, especially in the basal branching pattern, are only weakly corroborated. The monophyly of most superfamilies is supported, and all may be monophyletic except the ‘Siricoidea’, which is clearly paraphyletic. It is difficult to di whether the Siricidae or the Anaxyelidae are the closest relatives of Xiphydriidae + (Orussoidea + Apocrita). support for the sistergroup relationship between the Orussoidea and the Apocrita is substantial, putative apomorphies being provided by most character systems. There is also good evidence in favour of the monophj the Apocrita. The internal phylogeny of the Tenthredinoidea differs considerably from the results of earlier anal The Blasticotomidae are the sistergroup of the Tenthredinoidea s.s. Relationships at the base of the Tenthredini s.s. are weakly supported. It is uncertain whether the Tenthredinidae are monophyletic or comprise a 1 paraphyletic grade within the Tenthredinoidea s.s. The Diprionidae may be the sistergroup to Cimbicidae +(Argidae+ Pergidae). Most relationships within the Cimbicidae + (Argidae + Pergidae) clade are corroborated, with the exception of the monophyly of the Argidae. It is proposed to elevate the Anaxyelidae the Xiphydriida both to superfamily status. The family‐level classification of the Tenthredinoidea will probably have to be changed, but this must await further clarification of the phylogeny of this superfamily.