Phylogenetic relationships among superfamilies of Hymenoptera

The first comprehensive analysis of higher‐level phylogeny of the order Hymenoptera is presented. The analysis includes representatives of all extant superfamilies, scored for 392 morphological characters, and sequence data for four loci (18S, 28S, COI and EF‐1α). Including three outgroup taxa, 111 terminals were analyzed. Relationships within symphytans (sawflies) and Apocrita are mostly resolved. Well supported relationships include: Xyeloidea is monophyletic, Cephoidea is the sister group of Siricoidea + [Xiphydrioidea + (Orussoidea + Apocrita)]; Anaxyelidae is included in the Siricoidea, and together they are the sister group of Xiphydrioidea + (Orussoidea + Apocrita); Orussoidea is the sister group of Apocrita, Apocrita is monophyletic; Evanioidea is monophyletic; Aculeata is the sister group of Evanioidea; Proctotrupomorpha is monophyletic; Ichneumonoidea is the sister group of Proctotrupomorpha; Platygastroidea is sister group to Cynipoidea, and together they are sister group to the remaining Proctotrupomorpha; Proctotrupoidea s. str. is monophyletic; Mymarommatoidea is the sister group of Chalcidoidea; Mymarommatoidea + Chalcidoidea + Diaprioidea is monophyletic. Weakly supported relationships include: Stephanoidea is the sister group of the remaining Apocrita; Diaprioidea is monophyletic; Ceraphronoidea is the sister group of Megalyroidea, which together form the sister group of [Trigonaloidea (Aculeata + Evanioidea)]. Aside from paraphyly of Vespoidea within Aculeata, all currently recognized superfamilies are supported as monophyletic. The diapriid subfamily Ismarinae is raised to family status, Ismaridae stat. nov. © The Will Henning Society 2011.     

Molecular phylogeny of the apocritan wasps: the Proctotrupomorpha and Evaniomorpha

Phylogenetic relationships among the apocritan wasps were investigated using comparative sequence data from the mitochondrial 16S rRNA gene. The gene fragment contained three length polymorphic regions. Relationships that were not sensitive to the alignment procedure are discussed, but should be considered as preliminary only. Maximum parsimony analysis recovered the Evaniomorpha ( sensu Rasnitsyn, 1988) as a monophyletic group, with the pattern of relationships Ceraphronoidea + (Evanioidea + [Megalyroidea + Trigonalyoidea]). The Proctotrupomorpha ( sensu Rasnitsyn, 1988) were also recovered as a monophyletic group, while the Proctotrupoidea were recovered as paraphyletic. The Proctotrupoidea were only monophyletic if the Platygastroidea and Chalcidoidea were included (i.e. Proctotrupomorpha). Relationships of the Proctotrupomorpha that were supported by this analysis included Diapriidae + (Platygastroidea + Chalcidoidea), and the Vanhorniidae falling inside the Proctotrupidae. However, resolution of other relationships within the Proctotrupomorpha was less reliable, as indicated by low decay indices and low bootstrap proportions. Similarly, the placement of the Cynipoidea relative to the other apocritan lineages was also not recovered confidently.     

Evolution of the hymenopteran megaradiation

The Hymenoptera--ants, bees and wasps--represent one of the most successful but least understood insect radiations. We present the first comprehensive molecular study spanning the entire order Hymenoptera. It is based on approximately 7 kb of DNA sequence from 4 gene regions (18S, 28S, COI and EF-1α) for 116 species representing all superfamilies and 23 outgroup taxa from eight orders of Holometabola. Results are drawn from both parsimony and statistical (Bayesian and likelihood) analyses, and from both by-eye and secondary-structure alignments. Our analyses provide the first firm molecular evidence for monophyly of the Vespina (Orussoidea+Apocrita). Within Vespina, our results indicate a sister-group relationship between Ichneumonoidea and Proctotrupomorpha, while the stinging wasps (Aculeata) are monophyletic and nested inside Evaniomorpha. In Proctotrupomorpha, our results provide evidence for a novel core clade of proctotrupoids, and support for the recently proposed Diaprioidea. An unexpected result is the support for monophyly of a clade of wood-boring sawflies (Xiphydrioidea+Siricoidea). As in previous molecular studies, Orussidae remain difficult to place and are either sister group to a monophyletic Apocrita, or the sister group of Stephanidae within Apocrita. Both results support a single origin of parasitism, but the latter would propose a controversial reversal in the evolution of the wasp-waist. Generally our results support earlier hypotheses, primarily based on morphology, for a basal grade of phytophagous families giving rise to a single clade of parasitic Hymenoptera, the Vespina, from which predatory, pollen-feeding, gall-forming and eusocial forms evolved.     

Simultaneous analysis of 16S, 28S, COI and morphology in the Hymenoptera: Apocrita -evolutionary transitions among parasitic wasps

Simultaneous analysis of morphological and molecular characters from the 16S rDNA, 28S rDNA and cytochrome oxidase 1 genes was employed to resolve phylogenetic relationships among the apocritan (Insecta: Hymenoptera: Apocrita) wasps. Parsimony analyses, employing a broad range of models, consistently recovered the Proctotrupomorpha as a natural group, the Megalyridae and Trigonalidae as sister groups, a clade comprising the Monomachidae, Diapriidae, and Maamingidae, the Vanhorniidae and Proctotrupidae as sister groups, the Procto-trupoidea as polyphyletic, and the Evaniomorpha as a grade (but including the Ichneumonoidea, Aculeata, and Stephanidae). The Proctotrupomorpha, containing virtually all of the wholly endoparasitic lineages, was consistently recovered as an apical clade, with the remaining groups forming a parapbyletic grade below them. Although the relative placement of the groups forming this basal grade varied among analyses, the most commonly recovered arrangement is consistent with the ancestral biology being ectoparasitism of coleopteran, wood-boring larvae. Furthermore, the recovery of the ectoparasitic-containing proctotrupomorphs (Chalcidoidea and, in some analyses, Ceraphronoidea) as apical lineages argues that these biologies are reversals.     

Grooming behaviours in the Hymenoptera (Insecta): potential phylogenetic significance

Grooming behaviours from representatives of 36 families of Hymenoptera were video recorded and analysed. Thirty-three distinct types of grooming movements were recognized. The evolutionary pattern of each behaviour is discussed. Some behaviours displayed consistent variation between taxa, and appear to be informative regarding higher level relationships within the order. Putative synapomorphies are reported that support the monophyly of the Apocrita, Cynipoidea, Platygastroidea, Ichneumonoidea, and Chalcidoidea including the Mymaridae.     

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.     

Per arborem ad astra: morphological adaptations to exploiting the woody habitat in the early evolution of Hymenoptera

We survey morphological features of larval and adult wasps that undergo their entire larval development inside wood and interpret them in view of the lifestyle. The evolution of some of the characters is explored by mapping them on a recently published phylogeny of Hymenoptera. Based on this phylogeny, it is reasonable to assume that wood-living wasps evolved from a xylophagous/mycetophagous stage as displayed by woodwasps to a carnivorous/parasitoid lifestyle, preying on woodboring insect larvae. The latter mode of life is probably ancestral to the Apocrita which comprise the majority of the order; they share this lifestyle with their sister group, the Orussidae. However, most apocritan wasps have radiated into other habitats, the Orussidae and Stephanidae apparently being the only taxa that have retained the ancestral lifestyle of carnivorous wasps. Other apocritan lineages associated with wood (e.g., Aulacidae, Megalyridae, basal Cynipoidea and some Ichneumonoidea and Chalcidoidea) possibly entered this habitat secondarily and independently acquired morphological traits associated with it. The woody habitat was occupied by Hymenoptera during a crucial stage in their evolution where the transition from the phytophagous to carnivorous lifestyle took place. The anatomy of both larva and adults was extensively transformed in the process.     

The phylogeny of lower Hymenoptera (Insecta), with a summary of the early evolutionary history of the order

A cladistic analysis of the lower Hymenoptera, including all the ‘symphytan’ families and the apocritan families Stephanidae, Megalyridae, Trigonalyidae, Ibaliidae, Vespidae and Gasteruptiidae, has been undertaken. A total of 98 characters were scored for 21 taxa. Twenty equally parsimonious minimum-length trees were obtained. The phylogenetic status of the Xyelidae is uncertain: they might be monophyletic. or the Xyelinae might be the sistergroup of the rest of the Hymenoptera. The non-xyelid Hymenoptera are probably monophyletic; the phylogeny Tenthredinoidea + (Megalodontoidea + (Cephidae + (Anaxyelidea + (Siricidae + (Xiphydriidae + (Orussidae + Apocrita)))))) is proposed for this clade. The Blasticotomidae are probably the sistergroup of all othe Tenthredinoidea, but tenthredinoid phylogeny is otherwise uncertain. Substantial homoplasy occurs within the ‘siricoid’ families, making the relative positions of the Anaxyelidae and Siricidae uncertain. The Stephanidae might be the sistergroup of the rest of the Apocrita; the phylogeny of the remaining apocritan taxa included is insufficiently elucidated. The phylogeny proposed here supports the hypothesis that the appearance of parasitism in the Hymenoptera took place in the common ancestor of Orussidae + Apocrita, the host of which was probably wood boring insect larvae. The exact larval mode of feeding of the ancestral hymenopteran cannot be determined due ot the diversity of lifestyles in the basal lineages of the order.     

Higher and lower‐level relationships of the deep‐sea fish order Alepocephaliformes (Teleostei: Otocephala) inferred from whole mitogenome sequences

Fishes of the order Alepocephaliformes, slickheads and tubeshoulders, constitute a group of deep‐sea fishes poorly known in respect to most areas of their biology and systematics. Morphological studies have found alepocephaliform fishes to display a mosaic of synapomorphic and symplesiomorphic characters, resulting in great difficulties when attempting to resolve intra‐ and interrelationships. Molecular data recently added to the confusion by removing Alepocephaliformes from the Euteleostei and placed them as incertae sedis within the Otocephala. In the present study we attempt to further clarify relationships of Alepocephaliformes by adding newly determined whole mitogenome sequences from 19 alepocephaliforms in order to address 1) phylogenetic position of Alepocephaliformes within the Otocephala; and 2) intrarelationships of Alepocephaliformes. The present study includes 96 taxa of which 30 are alepocephaliforms and unambiguously aligned sequences were subjected to partitioned maximum likelihood and Bayesian analyses. Results from the present study support Alepocephaliformes as a genetically distinct otocephalan order as sister clade to Ostariophysi (mostly freshwater fishes comprising Gonorynchiformes, Cypriniformes, Characiformes, Siluriformes and Gymnotiformes). The disputed family Bathylaconidae was found to be an artificial assemblage of the two genera Bathylaco and Herwigia, with the former as the sister group of the family Alepocephalidae and the latter nested within Alepocephalidae. Platytroctidae was found to be monophyletic as sister clade to the rest of Alepocephaliformes. Previously unrecognized clades within the family Alepocephalidae are presented and a clade comprising Alepocephalus, Conocara and Leptoderma was recovered as the most derived. As long as the current classification is being followed, the genera Alepocephalus, Bathytroctes, Conocara and Narcetes were all found non‐monophyletic. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 923–936.     

Molecular phylogeny of the Byrrhoidea–Buprestoidea complex (Coleoptera,Elateriformia)

The superfamilies of Elateriformia have been in a state of flux since their establishment. The recent classifications recognize Dascilloidea, Buprestoidea, Byrrhoidea and Elateroidea. The most problematic part of the elateriform phylogeny is the monophyly of Byrrhoidea and the relationships of its families. To investigate these issues, we merged more than 500 newly produced sequences of 18S rRNA, 28S rRNA, rrnL mtDNA and cox1 mtDNA for 140 elateriform taxa with data from GenBank. We assembled an all‐taxa (488 terminals) and a pruned data set, which included taxa with full fragment representation (251 terminals); both were aligned in various programs and analysed using maximum‐likelihood criterion and Bayesian inference. Most analyses recovered monophyletic superfamilies and broadly similar relationships; however, we obtained limited statistical support for the backbone of trees. Dascilloidea were sister to the remaining Elateriformia, and Elateroidea were sister to the clade of byrrhoid lineages including Buprestoidea. This clade mostly consisted of four major lineages, that is (i) Byrrhidae, (ii) Dryopidae + Lutrochidae, (iii) Buprestoidea (Schizopodidae sister to Buprestidae) and (iv) a clade formed by the remaining byrrhoid families. Buprestoidea and byrrhoid lineages, with the exception of Byrrhidae and Dryopidae + Lutrochidae, were usually merged into a single clade. Most byrrhoid families were recovered as monophyletic. Callirhipidae and Eulichadidae formed independent terminal lineages within the Byrrhoidea–Buprestoidea clade. Paraphyletic Limnichidae were found in a clade with Heteroceridae and often also with Chelonariidae. Psephenidae, represented by Eubriinae and Eubrianacinae, never formed a monophylum. Ptilodactylidae were monophyletic only when Paralichas (Cladotominae) was excluded. Elmidae regularly formed a clade with a bulk of Ptilodactylidae; however, elmid subfamilies (Elminae and Larainae) were not recovered. Despite the densest sampling of Byrrhoidea diversity up to date, the results are not statistically supported and resolved only a limited number of relationships. Furthermore, questions arose which should be considered in the future studies on byrrhoid phylogeny.     

Phylogeny and classification of Cucujoidea and the recognition of a new superfamily Coccinelloidea (Coleoptera: Cucujiformia)

A large‐scale phylogenetic study is presented for Cucujoidea (Coleoptera), a diverse superfamily of beetles that historically has been taxonomically difficult. This study is the most comprehensive analysis of cucujoid taxa to date, with DNA sequence data sampled from eight genes (four nuclear, four mitochondrial) for 384 coleopteran taxa, including exemplars of 35 (of 37) families and 289 genera of Cucujoidea. Maximum‐likelihood analyses of these data present many significant relationships, some proposed previously and some novel. Tenebrionoidea and Lymexyloidea are recovered together and Cleroidea forms the sister group to this clade. Chrysomeloidea and Curculionoidea are recovered as sister taxa and this clade (Phytophaga) forms the sister group to the core Cucujoidea (Cucujoidea s.n .). The nitidulid series is recovered as the earliest‐diverging core cucujoid lineage, although the earliest divergences among core Cucujoidea are only weakly supported. The cerylonid series (CS) is recovered as monophyletic and is supported as a major Cucujiform clade, sister group to the remaining superfamilies of Cucujiformia. Currently recognized taxa that were not recovered as monophyletic include Cucujoidea, Endomychidae, Cerylonidae and Bothrideridae. Biphyllidae and Byturidae were recovered in Cleroidea. The remaining Cucujoidea were recovered in two disparate major clades: one comprising the nitidulid series + erotylid series + Boganiidae and Hobartiidae + cucujid series, and the other comprising the cerylonid series. Propalticidae are recovered within Laemophloeidae. The cerylonid series includes two major clades, the bothriderid group and the coccinellid group. Akalyptoischiidae are recovered as a separate clade from Latridiidae. Eupsilobiinae are recovered as the sister taxon to Coccinellidae. In light of these findings, many formal changes to cucujiform beetle classification are proposed. Biphyllidae and Byturidae are transferred to Cleroidea. The cerylonid series is formally recognized as a new superfamily, Coccinelloidea stat.n. Current subfamilies elevated (or re‐elevated) to family status include: Murmidiidae stat.n. , Teredidae stat.n. , Euxestidae stat.n. , Anamorphidae stat.rev. , Eupsilobiidae stat.n. , and Mycetaeidae stat.n. The following taxa are redefined and characterized: Cleroidea s.n. , Cucujoidea s.n. , Cerylonidae s.n. , Bothrideridae s.n. , Endomychidae s.n. A new subfamily, Cyclotominae stat.n. , is described. Stenotarsinae syn.n. is formally subsumed within a new concept of Endomychinae s.n.     

基于18S rDNA序列的蝽次目(半翅目：异翅亚目)

利用18SrDNA分子约1 912 bp的序列对蝽次目21个科53个种进行系统发育分析。运用MP法、ML法和NJ法分析后的结果表明:蝽次目的单系性得到很高的支持;扁蝽总科成为毛点类的姐妹群;毛点类基本确定为两大分支:一支包含蝽总科和红蝽总科;另一支主要由长蝽总科、缘蝽总科和南蝽总科组成;长蝽总科和缘蝽总科都是多系;长蝽总科中,跷蝽科和皮蝽科的关系最近,构成姐妹群,位于整个毛点类的基部;与长蝽总科中另外两个科长蝽科和地长蝽科的关系很远。说明利用18SrDNA分子对研究蝽次目的系统发育关系是适合的,能够重建蝽次目;扁蝽总科和蝽总科单系性的结果与形态学的研究以及Li et al (2005)的研究一致;但较Li et al(2005)的研究更进一步把红蝽总科从广义的缘蝽总科中分出来;并建议皮蝽科作为一个独立的总科更合适。     

Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.     

A review of the lower actinopterygian phylogeny

A review of lower actinopterygian phylogeny has led us to the conclusion that the Cladistia are the sister group of Recent actinopterygians (Actinopteri) and that the extinct Palaeonisciformes are a paraphyletic group, comprising stem-group actinopterygians (e.g. Cheirolepis ), stem-group actinopterans (e.g. Moythomasia ) and relatives of higher actinopterans such as Pteronisculus . Our analysis further concluded that the Acipenseriformes formed a clade together with Saurichthys and Birgeria , which was most parsimoniously resolved when the Acipenseriformes and Saurichthys were sister groups.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 144 , 511−525.     

Global interrelationships of Plesiosauria (Reptilia,Sauropterygia) and the pivotal role of taxon sampling in determining the outcome of phylogenetic analyses

Previous attempts to resolve plesiosaurian phylogeny are reviewed and a new phylogenetic data set of 66 taxa (67% of ingroup taxa examined directly) and 178 characters (eight new) is presented. We recover two key novel results: a monophyletic Plesiosauridae comprising Plesiosaurus dolichodeirus, Hydrorion brachypterygius, Microcleidus homalospondylus, Occitanosaurus tournemirensis and Seeleyosaurus guilelmiimperatoris; and five plesiosaurian taxa recovered outside the split between Plesiosauroidea and Pliosauroidea. These taxa are Attenborosaurus conybeari, ‘Plesiosaurus’macrocephalus and a clade comprising Archaeonectrus rostratus, Macroplata tenuiceps and BMNH 49202. Based on this result, a new name, Neoplesiosauria, is erected for the clade comprising Plesiosauroidea and Pliosauroidea. Taxon subsamples of the new dataset are used to simulate previous investigations of global plesiosaurian relationships. Based on these simulations, most major differences between previous global phylogenetic hypotheses can be attributed to differences in taxon sampling. These include the position of Leptocleididae and Polycotylidae and the monophyly or paraphyly of Rhomaleosauridae. On this basis we favour the results recovered by our, larger analysis. Leptocleididae and Polycotylidae are sister taxa, forming a monophyletic clade within Plesiosauroidea, indicating that the large‐headed, short‐necked ‘pliosauromorph’ body plan evolved twice within Plesiosauria. Rhomaleosauridae forms the monophyletic sister taxon of Pliosauridae within Pliosauroidea. Problems are identified with previous phylogenetic definitions of plesiosaurian clades and new, stem‐based definitions are presented that should maintain their integrity over a range of phylogenetic hypotheses. New, rank‐free clade names Cryptoclidia and Leptocleidia are erected to replace the superfamilies Cryptoclidoidea and Leptocleidoidea. These were problematic as they were nested within the superfamily Plesiosauroidea. The incongruence length difference test indicates no significant difference in levels of homoplasy between cranial and postcranial characters.     

First molecular phylogeny of the major clades of Pseudoscorpiones (Arthropoda: Chelicerata)

The phylogenetic relationships of the major lineages of the arachnid order Pseudoscorpiones are investigated for the first time using molecular sequence data from two nuclear ribosomal genes and one mitochondrial protein-encoding gene. The data were analyzed using a dynamic homology approach with the new program POY v.4 under parsimony as the optimality criterion. The data show monophyly of Pseudoscorpiones as well as many of its superfamilies (Feaelloidea, Chthonioidea, Cheiridioidea and Sternophoroidea), but not for Neobisiodea or Garypoidea. Cheliferoidea was not monophyletic either due to the position of Neochelanops, which grouped with some garypoids. In all the analyses, Feaelloidea constituted the sister group to all other pseudoscorpions; Chthonioidea is the sister group to the remaining families, which constitute the group Iocheirata sensu Harvey--a clade including pseudoscorpions with venom glands within the pedipalpal fingers. This phylogenetic pattern suggests that venom glands evolved just once within this order of arachnids.     

A revision and phylogenetic analysis of Stapeliopsis (Apocynaceae)

A survey of morphological characters is carried out for Stapeliopsis . The information obtained from this is combined with molecular data from the plastid trn L-F DNA region and ITS1 of the nuclear encoded 18S−26S rRNA cistron, to obtain a hypothesis of the evolutionary relationships among the species. It is shown that Stapeliopsis is monophyletic in a combined molecular and morphological analysis. Stapeliopsis is sister to a clade containing Huernia , Orbea and Tromotriche . The species of Stapeliopsis group into two clades. One contains S. khamiesbergensis , S. neronis and S. urniflora , and this is highly supported. The remaining species fall into an unsupported clade in which S. exasperata is sister to the others. The genera Hermanschwartzia Plowes and Neopectinaria Plowes are rejected. It is shown that a synapomorphy for Stapeliopsis is the laterally flattened inner corona-lobes, which touch the anthers only at their bases. Eight species of Stapeliopsis are recognized, with no subgeneric divisions.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 125–155.     

Phylogenetic relationships of the north-eastern Atlantic and Mediterranean blenniids

The phylogenetic relationships of 27 north-eastern Atlantic and Mediterranean blennioids are analysed based on a total of 1001 bp from a combined fragment of the 12S and 16S mitochondrial rDNA. The most relevant results with implications in current blenniid taxonomy are: (1) Lipophrys pholis and Lipophrys (=  Paralipophrys ) trigloides are included in a well-supported clade that by the rule of precedence must be named Lipophrys ; (2) the sister species of this clade are not the remaining species of the genus Lipophrys but instead a monotypic genus comprising Coryphoblennius galerita ; (3) the smaller species of Lipophrys were recovered in another well-supported and independent clade, which we propose to be recognized as Microlipophrys ; (4) although some authors included the genera Salaria and Lipophrys in a single group we have never recovered such a relationship. Instead, Salaria is more closely related to the genera Scartella and Parablennius ; (5) the genus Parablennius , which was never recovered as a monophyletic clade, is very diverse and may include several distinct lineages; (6) the relative position of Aidablennius sphynx casts some doubts on the currently recognized relationships between the different blenniid tribes. Meristic, morphological, behavioural and ecological characters support our results and are also discussed. The possible roles of the tropical West African coast and the Mediterranean in the diversification of blenniids are discussed.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 283–295.     

Morphological appraisal of Collembola phylogeny with special emphasis on Poduromorpha and a test of the aquatic origin hypothesis

Phylogenetic relationships within Collembola were determined through the cladistic analysis of 131 morphological characters and 67 exemplar taxa representing the major families of the group, with special emphasis on Poduromorpha. The results show that the order Poduromorpha is monophyletic and the sister group to the remaining Collembola, with Entomobryomorpha monophyletic and the sister group to the clade Neelipleona + Symphypleona. In Entomobryomorpha, Actaletidae is the sister group of the remaining families. In Poduromorpha, Tullbergiinae is monophyletic as well as Onychiurinae and the group Tetrodontophorinae + Onychiurinae which is the sister group of the remaining Poduromorpha; Tetrodontophorinae is paraphyletic; Onychiuridae is polyphyletic; Isotogastruridae is not an intermediate between Poduromorpha and Entomobryomorpha, it is the sister group of Tullbergiinae; Odontellidae is monophyletic and the sister group to the clade Neanuridae + Brachystomellidae; in Neanuridae, Frieseinae and the group Pseudachorutinae + Morulinae + Neanurinae are monophyletic; Morulinae is the sister group of Neanurinae; Pseudachorutinae is paraphyletic; Hypogastruridae is polyphyletic; Podura aquatica (Poduridae) is not 'primitive', it clusters with the genera Xenylla and Paraxenylla in Hypogastruridae. On the basis of these relationships and the position of the aquatic species, the most parsimonious hypothesis is a terrestrial edaphic origin for the springtails.     

Towards simultaneous analysis of morphological and molecular data in Hymenoptera

Principles and methods of simultaneous analysis in cladistics are reviewed, and the first, preliminary, analysis of combined molecular and morphological data on higher level relationships in Hymenoptera is presented to exemplify these principles. The morphological data from Ronquist et al . (1999) matrix, derived from the character diagnoses of the phylogenetic tree of Rasnitsyn (1988) , are combined with new molecular data for representatives of 10 superfamilies of Hymenoptera by means of optimization alignment. The resulting cladogram supports Apocrita and Aculeata as groups, and the superfamly Chrysidoidea, but not Chalcidoidea, Evanioidea, Vespoidea and Apoidea.