Phylogeny of the infraorder Culicomorpha (Diptera: Nematocera) based on 28S RNA gene sequences

Phylogenetic relationships between the families of the infraorder Culicomorpha were investigated by using partial 28S ribosomal RNA gene sequences. All families traditionally placed in this infraorder were investigated and confirmed as clades. On the other hand, some of the morphological relationships between these families were found to be in disagreement with phylogenies based on molecular characters. Our results did not support the generally accepted division of the Culicomorpha into two superfamilies, the Culicoidea (Culicidae + Corethrellidae + Chaoboridae + Dixidae) and the Chironomoidea (Chironomidae + Ceratopogonidae + Simuliidae + Thaumaleidae). Precisely, if the sister-group relationship between Culicidae, Chaoboridae and Corethrellidae was clearly confirmed, the Dixidae, traditionally considered as closely related to these two families, were not placed close to them on our trees. On the other hand, strong evidence was found for grouping together the Simuliidae and the Thaumaleidae, in spite of the cytological and morphological differences between these two families. The position of the Ceratopogonidae was uncertain, and the Chironomidae appeared as a possible sister group to the rest of Culicomorpha. The phylogenetic positions of the groups characterized by feeding on vertebrate blood or insect haemolymph (the Culicidae, Chaoboridae, Ceratopogonidae and Simuliidae) suggest that haematophagy has appeared at least twice in the evolution of Culicomorpha.     

Phylogeny of the subfamilies of Chironomidae (Diptera)

Summary The phylogeny of the subfamilies of Chironomidae are cladistically analysed using parsimony. A data matrix is presented and some characters discussed. Different outgroup taxa, constraints and options are used, characters unordered or ordered, weighted or unweighted, the results reweighted or not and the results discussed. Telmatogetoninae in all cladograms forms the sister group of the remaining subfamilies. Aphroteniinae in some cladograms forms the sister group of all subfamilies except Telmatogetoninae, whereas in other cladograms, including the preferred cladogram, it is part of Tanypodoinae, which otherwise includes Podonominae, Usumbaromyiinae and Tanypodinae. Chilenomyiinae is basal in Tanypodoinae in some cladograms. In most cladograms, including the preferred cladogram, it is basal in Chironomoinae, which also includes Buchonomyiinae, Diamesinae, Prodiamesinae, Orthocladiinae and Chironominae. The preferred cladogram is compared with the relationships between different subfamilies suggested by previous authors.     

Fungal and oomycete parasites of Chironomidae,Ceratopogonidae and Simuliidae (Culicomorpha,Diptera)

Members of the families Chironomidae (chironomids or non-biting midges), Ceratopogonidae (ceratopogonids or biting midges) and Simuliidae (simulids or blackflies) are ubiquitous dipterans of the infraorder Culicomorpha. They are extremely diversified in ecological strategies. Their larvae play major roles in aquatic food webs as detritivores or predators, whereas their adults can be general predators (Chironomidae), hemolymphagous or hematophagous predators (Ceratopogonidae and Simuliidae) or pollinators. Both larval and adult stages are commonly infected by bacteria, viruses, protists, nematodes, true fungi and oomycetes. These phylogenetically diverse assemblages of microorganisms can simultaneously infect multiple species of chironomids, ceratopogonids and simulids, and each host may become trophically interrelated with other hosts by sharing their parasites. Here, we review the information on fungal and oomycete parasites of these dipteran groups with special reference to the natural regulation of host populations, the impact of parasitism in food webs, and the potential of these parasites as biocontrol agents.     

Phylogenetic analysis of the Blephariceromorpha,with special reference to mountain midges (Diptera: Deuterophlebiidae)

Abstract. A cladistic analysis of the Blephariceromorpha (here including the Nymphomyiidae, Deuterophlebiidae and Blephariceridae) and related Diptera provides a test of the phylogenetic hypotheses of Rohdendorf (1964, 1974), Hennig (1973), Wood & Borkent (1989) and Courtney (1990a). In particular, monophyly of the Blephariceroidea and Blephariceromorpha (sensu Wood & Borkent), and their relationship to other Diptera, is tested. Evaluation of larval, pupal and adult characters supports the hypothesis of Wood & Borkent, as modified by Courtney. Four larval features suggest that the Blephariceromorpha + Psychodomorpha form a monophyletic group, although an alternate hypothesis predicting that the Blephariceromorpha is the sister group of the Psychodomorpha + (Ptychopteromorpha + Culicomorpha), is discussed. Monophyly of the Blephariceromorpha (Nymphomyioidea + Blephariceroidea) is supported by one adult and five larval characters. Monophyly of the Blephariceroidea (Deuterophlebiidae + Blephariceridae) is supported by thirteen synapotypies, including features of the larva (six), pupa (three) and adult (four). Nineteen, nineteen and nine hypothesized synapotypies support monophyly of the Nymphomyiidae, Deuterophlebiidae and Blephariceridae, respectively.     

Phylogenetics and temporal diversification of the earliest true flies (Insecta: Diptera) based on multiple nuclear genes

Abstract Relationships among families of the lower Diptera (formerly suborder ‘Nematocera’) have been exceptionally difficult to resolve. Multiple hypotheses based on morphology have been proposed to identify the earliest lineages of flies and place the phylogenetic origin of the higher flies (Brachycera), but convincing support is limited. Here we resolve relationships among the major groups of lower Diptera using sequence data from four nuclear markers, including both ribosomal (28S rDNA) and protein‐coding (CAD, TPI and PGD) genes. Our results support both novel and traditional arrangements. Most unexpectedly, the small, highly‐specialized family Deuterophlebiidae appears to be sister to all remaining Diptera. Other results include the resolution of the traditional infra‐orders Culicomorpha (including a novel superfamily Simulioidea = Thaumaleidae + Simuliidae), Tipulomorpha (Tipulidae sensu lato + Trichoceridae) and Bibionomorpha sensu lato. We find support for a limited Psychodomorpha (Blephariceridae, Tanyderidae and Psychodidae) and Ptychopteromorpha (Ptychopteridae), whereas the placement of several enigmatic families (Nymphomyiidae, Axymyiidae and Perissommatidae) remains ambiguous. According to genetic data, the infra‐order Bibionomorpha is sister to the Brachycera. Much of the phylogenetic signal for major lineages was found in the 28S rDNA gene, whereas protein‐coding genes performed variably at different levels. In addition to elucidating relationships, we also estimate the age of major lower dipteran clades, based on molecular divergence time estimates using relaxed‐clock Bayesian methods and fossil calibration points.     

PHYLOGENETIC RELATIONSHIPS IN SEED PLANTS

Abstract— The phylogenetic relationships of nineteen extant and fossil seed plants are considered. Analysis of 31 characters produced ten topologically similar and equally parsimonious cladograms. A strict consensus tree derived from these cladograms places Lyginopteris as the sister taxon to the other seed plants included. Within this clade all the taxa considered, except medullosans and cycads, form a single monophyletic group defined by the presence of flattened seeds and saccate pollen ("platy-sperms"). Relationships between medullosans, cycads, and "platysperms" were not resolved, but within the "platysperm" clade conifers and cordaites ( Cordaixylon, Mesoxylon ) + Ginkgo form a monophyletic group ("coniferophytes"). The "higher platysperms" (glossopterids, Caytonia , corystosperms, Bennettitales, Pentoxylon , Gnetales, and angiosperms) are also monophyletic, but their relationship to "coniferophytes," peltasperms, and Callistophyton is unresolved. Pentoxylon is placed as sister taxon to the Bennettitales, and together they form the sister group to a clade in which Gnetales and angiosperm are sister taxa. The Bennettitales + Pentoxylon + Gnetales + angiosperms ("anthophytes") form a monophyletic sister group to the corystosperms. This analysis is compared with current classifications of seed plants. It does not support a close relationship between Bennettitales and cycads, it provides no evidence for seed plant polyphyly, and it strongly suggests that the current concept of seed ferns has little value in a phylogenetic context.     

A Cladistic Analysis of the Genus Cyttaria (Fungi-Ascomycotina)*

Abstract—Cyttaria Berkeley (Cyttariaceae, Cyttariales, Class Discomycetes), is a genus of eleven species, seven from South America and four from Australasia (Australia, Tasmania, and New Zealand). Cyttaria is a monophyletic genus defined by the following synapomorphies: fleshy to gelatinous stromata with endostromatic apothecia; and complete lack of fungal chitin in the cell walls, having instead β-1–3-glucan. All Cyttaria are exclusive parasites of Nothofagus species. A cladistic analysis of the genus was performed using 18 characters from macromorphology, micro-morphology, and phenology. Polarity of characters is based on the outgroup comparison method (using the Class Discomycetes as a whole) and on the ontogenetic criterion. A hypothetical outgroup was constructed using all plesiomorphic states. Two equally parsimonious cladograms were produced, each with 40 steps and a consistency index of 0.70. These differed in the position of the South American species C. hookeri and C. johowii. In one cladogram, C. hookeri is the sister group to the rest of the genus, and in the other, both species form a monophyletic group (ascospores ovoid) that is the sister group of the rest of the genus. Our analysis of the two characters causing this difference (position of apothecia on stroma, and shape of ascospores) supports the latter hypothesis. In both cladograms, all of the Australasian species form a monophyletic group (thick ectostroma), and, within that, two subgroups are defined: C. gunnii-C. pallida (papillae present), and C. nigra-C. septentrionalis (ectostroma with black incrustations). The Australasian species form a monophyletic group (spermatangia absent) with C. berteroi , and this last group forms a monophyletic group (conidia absent) with C. espinosae. Cytlaria darwimi and C. exigua form another monophyletic group (very thick ectostroma). Excluding C. hookeri and C. johowii, C. hariotii is the sister group to the remaining eight species.     

Phyletic patterns of early development in gastropod molluscs

Cell lineage data for 30 exemplar gastropod taxa representing all major subclades and the outgroup Polyplacophora were examined for phylogenetic signal using cladistic analysis. Most cell lineages show phyletic trends of acceleration or retardation relative to the outgroup and more basal ingroup taxa, and when coded this variation is phylo-genetically informative. PAUP analyses of a cell lineage data set under three sets of character ordering assumptions produced similar tree topologies. The topologies of the strict consensus trees for both ordered and Dollo (near irreversibility of character transformations) character assumptions were similar, whereas the unordered character assumption recovers the least phyletic information. The cell lineage cladograms are also in agreement with the fossil record of the timing and sequence of gastropod subclade origination. A long branch lies between the Patellogastropoda+Vetigastropoda grade and the Neritopsina+Apogastropoda clade. The geological timing of this long branch is correlated with the first large-scale terrestrially derived eutrophication of the near-shore marine habitat, and one possible explanation for this branch may be a developmental shift associated with the evolution of feeding larvae in response to the more productive conditions in the near-shore water column. Although character transformations are highly ordered in this data set, developmental rate characters (like all other morphological and molecular characters) are also subject to homoplasy. Finally, this study further supports the hypothesis that early development of gastropod molluscs has conserved a strong phyletic signal for about half a billion years.     

Molecular analysis of the biting midges (Diptera: Ceratopogonidae), based on mitochondrial cytochrome oxidase subunit 2

Sequences from the mitochondrial cytochrome oxidase subunit 2 gene (cox2) were determined for 14 species from the family Ceratopogonidae, representing 12 genera and all five subfamilies, along with six representatives of other nematoceran families. The purpose was to develop a molecular phylogeny of the Ceratopogonidae, and interpret the phylogenetic position of the family within the infraorder Culicomorpha. These taxa have been analysed using cladistic methodology which, in combination with an excellent fossil record, provides a well established morphological phylogeny. Sequence analysis of cox2 revealed a high degree of sequence divergence among the species, reflecting in part the antiquity of the family, but also a significant acceleration of sequence evolution in the ceratopogonids compared to other nematoceran Diptera. Phylogenetic reconstruction by neighbor-joining and maximum parsimony gave strong support for an early separation of an ancient lineage that includes the two genera, Austroconops and Leptoconops, from the remainder of the family. The results support the existence of a clade that includes two subfamilies, Dasyheleinae and Forcipomyiinae, and this clade appears as sister to the remaining subfamily, Ceratopogoninae. The molecular phylogeny also supports monophyly of the Ceratopogonidae, and either a sister or paraphyletic relationship of this family with the Chironomidae.     

中国灵猫科的支序系统学分析

选取分布于中国境内的8属9种灵猫科动物的88个骨骼性状、31个外部形态性状和1个行为性状,运用替代外群法分别以赤狐(犬科狐属)和青鼬(鼬科貂属)共同或单独作为外群进行支序分析,得出10个支序图,其步长(TL)为：106～136,一致性指数(CI)为0．581—0．660,保留指数(RI)为0．610—0．714。经合意分析得到4个相似的支序图,其TL为41—136,CI为0．581—0．732,RI为0．610—0．818,结果支持：①斑灵狸、大灵猫、小灵猫构成一个单系群,与传统分类一致(均属灵猫亚科);②椰子狸、花面狸、熊狸构成一个单系群,与传统分类一致(均属长尾狸亚科);③红颊獴、食蟹獴与灵猫科其他种为姐妹群关系(Bootstrap检验支持率100％),建议獴类为一亚科;④长颌带狸拥有较多自近裔性状如三个门齿孔,是一个高度特化的种类,其系统地位有待进一步研究。     

Generic-level relationships within the ant subfamily Dolichoderinae (Hymenoptera: Formicidae)

A cladistic analysis was undertaken to determine relationships among extant genera of the ant subfamily Dolichoderinae. Twenty-one of the twenty-two currently recognized genera within the subfamily were examined using 104 morphological characters. A single fully resolved, most-parsimonious tree was found when a combination of ordered and unordered characters was used. When all characters were coded as unordered, seventy most-parsimonious trees were found. The following results were found with both character coding methods. Leptomyrmex was placed basal to the remainder of the subfamily and the monophyletic sets Dolichoderus + Liometopum + Axinidris + Tapinoma + Technomyrmex, Froggattella + Iridomyrmex + Ochetellus + Papyrius + Philidris + Turneria , and Bothriomyrmex + Dorymyrmex + Forelius + Loweriella were suggested. The genera Linepithema and Doleromyrma showed a small amount of instability in moving between neighbouring sister groups when the character coding method changed. The genera Anillidris and Anonychomyrma were difficult to place as they showed major differences in their positions between the two character coding methods.     

Phytogeny of the nematocerous families of Diptera (insecta)

The relationships of the nematocerous families of Diptera are cladistieally analysed using the parsimony programs PAUP and Hennig86. An extensive review, as well as a data matrix, is presented for 98 almost exclusively morphological characters (larva, 56; pupa, 6; adult, 36). Four infraorders are recognized, viz , Ptychopteromorpha, Culicomorpha, Blephariceromorpha, Bibionomorpha, and a clade containing the 'higher Nematocera' and Brachycera. Traditionally the family Nymphomyiidae or the infraorder Tipulomorpha (=Tipulidae, with or without Trichoceridae) are considered the most basal clade of the extant Diptera. On the basis of our cladistic analysis it is suggested that the Ptychopteromorpha-Culicomorpha clade is the sister-group of all other extant Diptera. We provide evidence that the Axymyiidae are part of a monophyletic Bibionomorpha. The latter infraorder is proposed as the sister-group of the higher Nematocera and Brachycera. We transfer the Tipulidae (Tipulomorpha) to the higher Nematocera, at a position next to Trichoceridae and near the Anisopodidae-Brachycera lineage. Previous hypotheses concerning nematocerous relationships are reviewed.     

Molecular paleobiological insights into the origin of the Brachiopoda

Most studies of brachiopod evolution have been based on their extensive fossil record, but molecular techniques, due to their independence from the rock record, can offer new insights into the evolution of a clade. Previous molecular phylogenetic hypotheses of brachiopod interrelationships place phoronids within the brachiopods as the sister group to the inarticulates, whereas morphological considerations suggest that Brachiopoda is a monophyletic group. Here, these hypotheses were tested with a molecular phylogenetic analysis of seven nuclear housekeeping genes combined with three ribosomal genes. The combined analysis finds brachiopods to be monophyletic, but with relatively weak support, and the craniid as the sister taxon of all other brachiopods. Phylogenetic-signal dissection suggests that the weak support is caused by the instability of the craniid, which is attracted to the phoronids. Analysis of slowly evolving sites results in a robustly supported monophyletic Brachiopoda and Inarticulata (Linguliformea+Craniiformea), which is regarded as the most likely topology for brachiopod interrelationships. The monophyly of Brachiopoda was further tested with microRNA-based phylogenetics, which are small, noncoding RNA genes whose presence and absence can be used to infer phylogenetic relationships. Two novel microRNAs were characterized supporting the monophyly of brachiopods. Congruence of the traditional molecular phylogenetic analysis, microRNAs, and morphological cladograms suggest that Brachiopoda is monophyletic with Phoronida as its likely sister group. Molecular clock analysis suggests that extant phoronids have a Paleozoic divergence despite their conservative morphology, and that the early brachiopod fossil record is robust, and is not affected by taphonomic factors relating to the late-Precambrian/early-Cambrian phosphogenic event.     

The position of Cetacea within mammalia: phylogenetic analysis of morphological data from extinct and extant taxa

Knowledge of the phylogenetic position of the order Cetacea (whales, dolphins, and porpoises) within Mammalia is of central importance to evolutionary biologists studying the transformations of biological form and function that accompanied the shift from fully terrestrial to fully aquatic life in this clade. Phylogenies based on molecular data and those based on morphological data both place cetaceans among ungulates but are incongruent in other respects. Morphologists argue that cetaceans are most closely related to mesonychians, an extinct group of terrestrial ungulates. They have disagreed, however, as to whether Perissodactyla (odd-toed ungulates) or Artiodactyla (even-toed ungulates) is the extant clade most closely related to Cetacea, and have long maintained that each of these orders is monophyletic. The great majority of molecule-based phylogenies show, by contrast, not only that artiodactyls are the closest extant relatives of Cetacea, but also that Artiodactyla is paraphyletic unless cetaceans are nested within it, often as the sister group of hippopotamids. We tested morphological evidence for several hypotheses concerning the sister taxon relationships of Cetacea in a maximum parsimony analysis of 123 morphological characters from 10 extant and 30 extinct taxa. We advocate treating certain multistate characters as ordered because such a procedure incorporates information about hierarchical morphological transformation. In all most-parsimonious trees, whether multistate characters are ordered or unordered, Artiodactyla is the extant sister taxon of Cetacea. With certain multistate characters ordered, the extinct clade Mesonychia (Mesonychidae + Hapalodectidae) is the sister taxon of Cetacea, and Artiodactyla is monophyletic. When all fossils are removed from the analysis, Artiodactyla is paraphyletic with Cetacea nested inside, indicating that inclusion of mesonychians and other extinct stem taxa in a phylogenetic analysis of the ungulate clade is integral to the recovery of artiodactyl monophyly. Phylogenies derived from molecular data alone may risk recovering inconsistent branches because of an inability to sample extinct clades, which by a conservative estimate, amount to 89% of the ingroup. Addition of data from recently described astragali attributed to cetaceans does not overturn artiodactyl monophyly.     

Ants of the genus Myrmecia Fabricius: A review of the species groups and their phylogenetic relationships (Hymenoptera: Formicidae: Myrmeciinae)

Abstract. Myrmecia Fabricius is revised at species-group level. Nine groups are recognized: those of M.aberrans, M.cephalotes, M.gulosa, M.mandibularis, M.nigrocincta, M.picta, M.pilosula, M.tepperi and M. urens. A key to the species groups is provided, and worker diagnoses, illustrations and species lists are given for each. Eight groups are constituted much as in the previous classification of John Clark, but defined using new characters. Phylogenetic relationships are investigated, with six cladograms derived from four sets of data, each with a different outgroup. The most plausible cladograms suggest that: (1) the aberrans group is the sister group to the others; (2) the pilosula, tepperi and mandibularis groups constitute a monophyletic assemblage, though monophyly of the first two is not confirmed; (3) the gulosa, nigrocincta, urens and picta groups constitute a monophyletic assemblage, though monophyly of the picta group is not confirmed; (4) the phylogenetic position of the cephalotes group is unclear.     

A preliminary cladistic study of the families of the superorder Lamiiflorae

LU AN-MING, 1990. A preliminary cladistic study of the families of the superorder Lamiiflorae. A preliminary cladistic analysis was undertaken to evaluate the relationships between families of the superorder Lamiiflorae sensu Dahlgren. Several character complexes were surveyed, and ultimately 29 informative characters were used for the study. Three families, Clethraceae, Oleaceae and Solanaceae were selected for outgroup comparison and polarization of the characters. A data matrix was constructed for the 23 ingroup families. The data matrix was analysed with the cladistic parsimony program Hennig86. Three equally parsimonious cladograms were found. Many family interrelationships could not be resolved, although several groups were common to all three cladograms, as shown by a strict consensus tree. The Retziaceae emerged as the sister group to the remaining families. About half of those appeared in a large polytomy in the consensus tree. There was also one possibly monophyletic complex of families involving the Lamiales with the families Verbenaceae, Lamiaceae, Phrymaceae and Callitrichaceae as well as the three isolated families Trapellaceae, Hippuridaceae, and Hydrostachyaceae. Within this complex, Verbenaceae and Lamiaceae came out as sister groups, as did Callitrichaceae and Hydrostachyaceae, with Hippuridaceae as sister group to them. However, the results must be regarded as tentative.     

The phylogeny of land plants: A cladistic analysis based on male gametogenesis

A cladistic analysis was carried out to resolve phylogenetic pattern among bryophytes and other land plants. The analysis used 22 taxa of land plants and 90 characters relating to male gametogenesis.Coleochaete orChara/Nitella were the outgroups in various analyses using HENNIG86, PAUP, and MacClade, and the land plant phylogeny was unchanged regardless of outgroup utilized. The most parsimonious cladograms from HENNIG86 (7 trees) have treelengths of 243 (C.I. = 0.58, R.I. = 0.82). Bryophytes are monophyletic as are hornworts, liverworts, and mosses, with hornworts identified as the sister group of a liverwort/moss assemblage. In vascular plants, lycophytes are polyphyletic andSelaginella is close to the bryophytes.Lycopodium is the sister group of the remaining vascular plants (minusSelaginella). Longer treelengths (over 250) are required to produce tree topologies in which either lycophytes are monophyletic or to reconstruct the paraphyletic bryophyte phylogeny of recent authors. This analysis challenges existing concepts of bryophyte phylogeny based on more classical data and interpretations, and provides new insight into land plant evolution.     

Trichomycete insect symbionts in Great Smoky Mountains National Park and vicinity

Collections of trichomycete symbionts of larval aquatic insects in Great Smoky Mountains National Park and vicinity in the southern Appalachian region of the USA resulted in finding many taxa of Harpellales, including an unusual new monotypic genus, Barbatospora ambicaudata in Simuliidae, and five new species in Thaumaleidae or Chironomidae, Harpellomyces montanus, Smittium lentaquaticum, Sm. minutisporum, Stachylina gravicaudata and St. stenospora. In addition a new species of Amoebidium (Amoebidiales), A. appalachense, attached to the anal tubules of bloodworms (Chironomidae) is described. Axenic cultures of three of the new taxa were obtained, plus Sm. culisetae. Fourteen identified species representing 13 genera of previously known Harpellales are recorded from Plecoptera, Ephemeroptera and Diptera, as well as a new Dipteran host record for an unidentified harpellid that was found in a Blephariceridae. Also identified were Paramoebidium corpulentum and many undetermined species of Paramoebidium (Amoebidiales) from four orders of aquatic insect larvae. The occurrence of an Enterobryus species in Diplopoda and another Eccrinales from an aquatic beetle is noted. Amoebidiales,     

Two Requirements for Obtaining Valid Common Patterns under Different Assumptions in Vicariance Biogeography

In vicariance biogeography, widespread or sympatric taxa can be dealt with under assumptions 0, 1, and 2. Data from cladogenetic relationships among taxa of a monophyletic group and their distribution over areas are assumed, in the order 0 → 1 → 2, to represent decreasing information about vicariance events. A less strict assumption carries a larger solution set, i.e., the number of possible area cladograms increases with the decrease in strictness of the assumption applied. We formulate two requirements for obtaining valid general area cladograms from data of several monophyletic groups of taxa. First, the assumptions, and with them the sets of area cladograms derived under these assumptions, should be inclusive. Second, sets of single group area cladograms should be compared for different monophyletic groups under a single assumption. When these two requirements are met, area cladograms become consistent with respect to the processes (vicariance, extinction, and dispersal) that are a priori assumed. The explanatory power increases for any particular monophyletic group of taxa when the set of valid general area cladograms contains a subset of area cladograms derived under a more strict assumption. We discuss examples from literature of how violation of these two requirements affects the results.