Molecular phylogeny of the Myzorhynchella Section of Anopheles (Nyssorhynchus) (Diptera: Culicidae): genetic support for recently described and resurrected species

Phylogenetic relationships among species of the Myzorhynchella Section of Anopheles (Nyssorhynchus) were investigated using the nuclear ribosomal DNA second internal transcribed spacer (ITS2), the nuclear whitegene and mitochondrial cytochrome oxidase subunit I (COI) regions. The recently described Anopheles pristinus and resurrected Anopheles guarani were also included in the study. Bayesian phylogenetic analyses found Anopheles parvus to be the most distantly related species within the Section, a finding that is consistent with morphology. An. pristinus and An. guarani were clearly resolved from Anopheles antunesi and Anopheles lutzii, respectively. An. lutzii collected in the same mountain range as the type locality were found within a strongly supported clade, whereas individuals from the southern state of Rio Grande do Sul, tentatively identified as An. lutzii based on adult female external morphology, were distinct from An. lutzii, An. antunesi and from each other, and may therefore represent two new sympatric species. A more detailed examination of An. lutzii sensu latoalong its known geographic range is recommended to resolve these anomalous relationships.     

Systematic studies on Anopheles galvaoi Causey,Deane & Deane from the subgenus Nysssorhynchus blanchard (Diptera: Culicidae)

Anopheles galvaoi, a member of the subgenus Nyssorhynchus, is redescribed based on morphological characters of the adults male and female, fourth-instar larva and pupa. Female, male genitalia, larval and pupal stages are illustrated. Data about medical importance, bionomics, and distribution are given based on literature records. Adult female of An. galvaoi can be easily misidentified as An. benarrochi Gabald n and An. aquasalis Curry. A few characters are indicated for identifying female and immatures of An. galvaoi. Phylogenetic relationships among An. galvaoi and six other species of the Oswaldoi Subgroup are estimated using COII mtDNA and ITS2 rDNA gene sequences. Lectotype of An. galvaoi, an adult female from Rio Branco, State of Acre, is invalidated.     

Phylogenetic relationships among Heliconiinae genera based on morphology (Lepidoptera: Nymphalidae)

Abstract.  Phylogenetic relationships among Heliconiinae genera are proposed based on early-stage and adult morphology. Parsimony analyses of forty-nine species in twenty-nine genera indicate that Heliconiinae can be divided into four main groups: (1) Pardopsis , Acraea and Actinote ; (2) Cethosia plus Neotropical genera; (3) Oriental genera and (4) fritillaries. Analyses of adult characters suggested that Oriental genera form a monophyletic group, whereas those of adult plus early-stage characters artificially split this group into three separate lineages. Our character set does not contain enough phylogenetic information to resolve relationships among fritillary genera with confidence, and further studies of this group are needed. The classification of the Heliconiinae is revised based on our results.     

Phylogeny of the Pyretophorus Series of Anopheles subgenus Cellia (Diptera: Culicidae)

A cladistic analysis based on morphological characters is presented for the Pyretophorus Series of subgenus Cellia of Anopheles. The cibarial armature was investigated for the first time since the pioneering work of Christophers (1933) , which is still used to define taxonomic series within Cellia, and for the first time ever using scanning electron microscopy. Christophers’ observations of characters of the cibarial armature were corroborated and several novel characters discovered. Despite difficulties in delimiting and encoding morphological characters at the species level, the analysis provided insights into relationships within the Pyretophorus Series. The data support the monophyly of the Series and establish the Oriental species as a monophyletic group. The Afrotropical species are basal and paraphyletic relative to the Oriental species, with Anopheles christyi placed in the most basal position. The pattern of relationships suggests that the capacity to transmit malaria is an ancestral condition subsequently lost independently in several lineages.     

林氏按蚊线粒体全基因组序列的测定及基于线粒体基因组的按蚊属系统发育分析(英文)

【目的】对林氏按蚊Anopheles lindesayi完整的线粒体基因组进行测序及分析,依据已知的线粒体基因组构建并讨论按蚊属蚊虫的分子系统发育关系。【方法】对林氏按蚊线粒体基因组进行测序、注释,并对其基本特征和基本组成进行分析。基于串联的13个蛋白质编码基因的核苷酸序列和氨基酸序列,用ML法和贝叶斯法构建林氏按蚊和按蚊属其他32种蚊虫的系统发育树,据此探讨按蚊属蚊虫的系统发育关系和系统分类。【结果】林氏按蚊线粒体基因组全长为15 366 bp,包含13个蛋白质编码基因,22个tRNA基因,2个rRNA基因和一段控制区。林氏按蚊线粒体基因组呈现明显的AT偏斜和GC偏斜,AT偏斜为正,GC偏斜为负。除了COX1使用TCG和ND5使用GTG作为起始密码子以外,其他蛋白质编码基因的起始密码子均遵循ATN原则;终止密码子为TAA或者T。除了tRNA^Ser(AGN)以外,其他的tRNA基因均呈现典型的三叶草二级结构。控制区AT含量最高,为94.54%。滑窗分析显示蛋白质编码基因是用于构建亚属或属水平系统发育关系的最佳分子标记。系统发育树强烈支持塞蚊亚属Cellia、按蚊亚属Anopheles、徕蚊亚属Nyssorhynchus和柯特蚊亚属Kerteszia均为单系群。小五斑按蚊An. atroparvus和四斑按蚊An. quadrimaculatus A这两个种聚到一起,从传统的形态分类上讲,它们和林氏按蚊均属于按蚊亚属按蚊系蚊虫。但本研究构建的4个系统发育树均显示,(小五斑按蚊An. atroparvus+四斑按蚊An. quadrimaculatus A)和林氏按蚊被属于迈蚊系的中华按蚊分开,这为两个系的分类提供了新的论点。【结论】本研究获得了林氏按蚊的完整的线粒体基因组,探析了按蚊属的线粒体基因组特征和系统发育关系,为进一步研究蚊科线粒体基因组和系统发育关系提供了依据。     

Higher-level systematics of rodents (Mammalia,Rodentia): Evidence from the mitochondrial 12S rRNA gene

Phylogenetic relationships among major rodent superfamilies traditionally have been difficult to establish because of the apparent high level of convergence and parallelism seen among morphological characters and/or rapid differentiation of rodent groups in the Paleocene/Eocene. Nucleotide sequence data from the mitochondrial 12S rRNA gene were used to clarify phylogenetic relationships among the major groups of rodents as defined by Brandt (1855) and Tullberg (1899). Based on the approximately 800 bp analyzed for the 12S rRNA gene in 59 mammalian species, including 25 of the 32 extant rodent families, the major rodent groups that could be defined as monophyletic clades were the Hystricognathi, the Muroidea, and the Geomyoidea. In addition, support for superfamilial sister-group relationships was found for Aplodontoidea with Sciuroidea and Dipodoidea with Muroidea.     

The classification of genus Anopheles (Diptera: Culicidae): a working hypothesis of phylogenetic relationships

The internal classification of genus Anopheles is updated to reflect taxonomic actions published since the classification was last reviewed in 1994. Both formal and informal taxa are included. The classification is intended to aid researchers and students who are interested in analysing species relationships, making group comparisons and testing phylogenetic hypotheses. The genus includes 444 formally named and 40 provisionally designated extant species divided between six subgenera: Anopheles, Cellia, Kerteszia, Lophopodomyia, Nyssorhynchus and Stethomyia. Subgenera Anopheles, Cellia and Nyssorhynchus are subdivided hierarchically into nested informal groups of morphologically similar species that are believed to represent monophyletic lineages based on morphological similarity. Changes to the classification include additional species, eliminated species and changes to the hierarchical organization and composition of supraspecific groups, some as a result of molecular studies.     

Evolution of mitochondrial and ribosomal gene sequences in anophelinae (Diptera: Culicidae): implications for phylogeny reconstruction

In this study, two mitochondrial genes, cyt b and ND5, and the D2 expansion segment of the 28S nuclear ribosomal gene were used to reconstruct a phylogeny of the mosquito subfamily Anophelinae. The ingroup consisted of all three genera of Anophelinae and five of six subgenera of Anopheles. Six genera of Culicinae were used as the outgroup. Extreme conservation at the protein level coupled with rapid saturation of synonymous positions probably accounted for the lack of meaningful phylogenetic signal in the cyt b gene. In contrast, abundant variation at all codon positions of the ND5 gene allowed recovery of the basal and most of the recent relationships. Phylogenetic analysis of D2 produced results consistent with those of ND5. Combined analysis indicated well-supported monophyletic Anophelinae (with Chagasia basal), Anopheles + Bironella, and subgeneric clades within the genus Anopheles. Moreover, subgenera Nyssorhynchus and Kerteszia were supported as a monophyletic lineage. The Kishino-Hasegawa test could not reject the monophyly of Anopheles, whereas the recently proposed hypothesis of close affinity of Bironella to the subgenus Anopheles was rejected by the analyses of ND5 and combined data sets. The lack of resolution of Bironella and Anopheles clades, or basal relationships among subgeneric clades within Anopheles, suggests their rapid diversification. Recovery of relationships consistent with morphology and previous molecular studies provides evidence of substantial phylogenetic signal in D2 and ND5 genes at levels of divergence from closely related species to subfamily in mosquitoes.     

Phylogenetic relationships in Ephedra (Ephedraceae) inferred from chloroplast and nuclear DNA sequences

Sequences of the nuclear ribosomal DNA internal transcribed spacer region 1 and the chloroplast-encoded genes maturase K and ribulose-1,5 biphosphate carboxylase large subunit were obtained from species of Ephedra (Ephedraceae) representing the geographic range and morphological diversity of the genus. Phylogenetic analyses of the DNA data indicate that relationships within the genus are better predicted by geographic region of origin than by ovulate cone characters. The sampled species with dry, winged (versus fleshy) ovulate cone bracts or single-seeded cones do not form monophyletic groups and therefore the previous classification systems of Ephedra based on these aspects of bract morphology appear to be largely unnatural. Three groups were identified among the Old World species studied, one comprising European and Mediterranean species and two including only Asian species. The sequence data suggest a possible early divergence of a New World clade of Ephedra from among the Old World groups. The South American species form a distinct clade apparently related to one of two groups of North American species, which accords with a frequent floristic pattern of close relationships between species groups in western South America and southwestern North America.     

熊蜂族系统发育关系的研究(膜翅目:蜜蜂总科)(英文)

本文选取熊蜂雌雄外部形态特征、行为学特征,对熊蜂28个亚属之间的系统发育关系利用支序分析的方法进行了探讨。研究结果表明传统的熊蜂属与拟熊蜂属是两个单系群,Mendacibombus亚属在支序图中的位置与传统看法有所不同。另外,根据能蜂各亚属间的亲缘关系及它们的地理分布建议熊蜂属中分两个亚属群：欧亚—亚洲群与欧亚—美洲群。     

Phylogeny of Dendroctonus bark beetles (Coleoptera: Curculionidae: Scolytinae) inferred from morphological and molecular data

Bark beetles in the genus Dendroctonus may attack and kill several species of coniferous trees, some of them causing major economic losses in temperate forests throughout North and Central America. For this reason, they have been widely studied. However, various aspects of the taxonomy and evolutionary history of the group remain contentious. The genus has been subdivided in species groups according to morphological, biological, karyological or molecular attributes, but the evolutionary affinities among species and species groups within the genus remain uncertain. In this study, phylogenetic relationships among Dendroctonus species were reassessed through parsimony‐based cladistic analysis of morphological and DNA sequence data. Phylogenetic inference was based on 36 morphological characters and on mitochondrial DNA sequences of the cytochrome oxidase I (COI) gene. Analyses were carried out for each dataset, as well as for the combined data analysed simultaneously, under equal and implied weights. According to the combined analysis, the genus Dendroctonus is a monophyletic group defined by at least three synapomorphic characters and there are four main lineages of varied composition and diversity within the genus. Within these lineages, several monophyletic groups match, to some extent, species groups defined by previous authors, but certain groups proposed by those authors are polyphyletic or paraphyletic.     

Egg morphology as an indirect method to identify Anopheles benarrochi, Anopheles oswaldoi and Anopheles rangeli (Diptera: Culicidae)

In the Department of Putumayo in southern Colombia, malaria transmission has continued in the absence of the 4 traditional Latin American vector species--Anopheles darlingi, Anopheles nuneztovari, Anopheles albimanus or Anopheles trinkae. Human bait collections yielded Anopheles mosquitoes and a morphological variant of Anopheles benarrochi, the adult females of which can easily be misidentified as Anopheles oswaldoi. Species identification of females of Anopheles in the subgenus Nyssorhynchus is generally difficult due to overlapping morphological characters; therefore, progeny of field collected females were link-reared to assess species identity. Herein a robust method is presented to identify the species Anopheles benarrochi, Anopheles oswaldoi and Anopheles rangeli from southern Colombia, using the morphology of the eggs induced from wild-caught females. Eggs of A. rangeli and A. benarrochi were differentiated on the basis of the anterior crown. In A. rangeli, this feature is positioned apically with high walls. In A. benarrochi, anterior crown is positioned more ventrally with comparatively shorter walls. No crown is present in A. oswaldoi. These differences are clear with the aid of a dissecting microscope and make accurate species determination possible even in field conditions. Egg morphology is shown to be an accurate, albeit indirect, method for the taxonomic determination for the three southern Colombian species and may also be useful in other regions of Latin America where the morphological variant of A. benarrochi is sympatric with A. oswaldoi.     

Evolutionary diversification of spiny rats (genus Trinomys, Rodentia: Echimyidae) in the Atlantic Forest of Brazil

Phylogenetic and phylogeographic relationships within and among species of the Atlantic Forest spiny rat Trinomys (family Echimyidae) were examined using cytochrome b sequence data. Levels of sequence divergence among species of Trinomys are as high as those found among taxa of echimyids that are recognized as different genera. Trinomys contains three distinct monophyletic clades that show a striking concordance with vegetational distribution. Haplotypes of clade 1 are distributed along the coastal margins of southeastern Brazil, following the moist tropical forest. Members of clade 2 are found in the semi-deciduous tropical forest. T. albispinus represents clade 3 and is found in a more xeric vegetation. Estimates of divergence times separating the three clades are very deep and range from 1.6 to 7.4 millions of years, predating the climatic fluctuations of the Pleistocene. Therefore, the proposed Late Pleistocene refugia in the Atlantic Forest cannot account for the divergence of the clades of Trinomys , but most likely shaped the modern distribution of species. The current taxonomy of this group does not reflect the diversity and phylogenetic relationships of the named species. However, morphological characters are congruent with the phylogeny uncovered by the molecular data. An extensive taxonomic rearrangement is suggested, reflecting phylogenetic relationships of monophyletic entities within the genus Trinomys , degree of sequence differences, and morphological diagnosability.     

Phylogenetic relationships among bumble bees (Bombus Latr.): a reappraisal of morphological evidence

Abstract. Cladistic analyses of morphological characters of adult bumble bees show significantly higher congruence among characters of the male genitalia than among other characters of males and females. Phylogenetic interpretations of groups supported by the combined morphological evidence (even with minimal assumptions) include (1) that the earliest diverging species belong to the para-phyletic subgenus Mendacibombus , together with Bombus nevadensis and B.confusus ; (2) that, in contrast, species of the subgenus Bombus (sensu stricto) share a relatively recent common ancestor with B.pressus and B.rufipes ; and (3) that Psithyrus (the social parasites) is a monophyletic group, and the sister-group of B.persicus. For genus-group nomenclature a single genus Bombus (sensu lato) is recommended, to include Psithyrus as another subgenus. Names for sections and for groups of subgenera are synonymized with subgeneric names.     

A STUDY ON PHYLOGENETIC RELATIONSHIPS AMONG THE SUB-GENERA OF BUMBLE BEES (HYMENOPTERA:APIDAE)

Abstract Phylogenetic relationships among the 28 subgenera of bumble bees were studied by using morphological and behavioral characters. The results suggest that the traditional Bombus Latreille and Psithyrus Lepeletier are monophyletic groups. However, the position of Mendacibombus in cladograms is different from that of the traditional view. In addition, according to their geographical distribution and relative relationships Eurasian—Asian group and Eurasian—American group are recognized for Bombus Latreille.     

A revision of Benjaminia (Hymenoptera: Ichneumonidae, Campopleginae)

Abstract The campoplegine genus Benjuminia is herein revised, with ten species being described as new, in addition to the five previously known species. Phylogenetic relationships of Benjuminia to other genera are discussed, and a cladogram presented for the species. Two monophyletic species groups are recognized, one being confined to the Palearctic and the other to the Nearctic. Examination of larval morphology not only fails to reveal larval characters for distinguishing Benjuminia but leads to the conclusion that use of larval morphology to identify campoplegine species is a dubious practice.     

Phylogenetic relationships of Palaeacanthocephala (Acanthocephala) inferred from SSU and LSU rDNA gene sequences

The Palaeacanthocephala is traditionally represented by 2 orders, Echinorhynchida and Polymorphida, with 10 and 3 families, respectively. To test the monophyly of the class, these 2 orders, and certain families, phylogenies were inferred using nuclear small-subunit (SSU) and large-subunit (LSU) ribosomal DNA sequences obtained for 29 species representing 10 families, 2 other classes of acanthocephalans, and 3 rotifer outgroups. Phylogenetic relationships were inferred by analyzing combined SSU and LSU sequences using maximum parsimony (MP) and maximum likelihood (ML) methods. Parsimony and ML trees inferred from combined analysis of these rDNA data strongly supported monophyly of Palaeacanthocephala and provided good resolution among species. Neither Polymorphida nor Echinorhynchida was monophyletic. Gorgorhynchoides bullocki (Echinorhynchida) was nested within the 6 species representing Polymorphida, and this clade was nested within species representing Echinorhynchida. Three of 4 palaeacanthocephalan families that could be evaluated were not monophyletic, and this finding was strongly supported. These results indicate that the family level classification of palaeacanthocephalans, which is mainly based on combinations of shared characters (not shared derived characters), needs to be reevaluated with respect to comprehensively sampled phylogenetic hypotheses.     

Remotely-sensed land use patterns and the presence of Anopheles larvae (Diptera: Culicidae) in Sukabumi, West Java, Indonesia

Land use patterns and the occurrence of Anopheles species larvae were studied in Sukabumi District, West Java, Indonesia, from October 2004 to September 2005. Two land use maps derived using remote sensing were used. One map derived from Quickbird satellite images of 150 km2 of the Simpenan and Ciemas subdistricts (106 degrees 27' 53"-106 degrees 38' 38" E and 6 degrees 59' 59"-7 degrees 8' 46" S) in Sukabumi and one using ASTER images covering 4,000 km2 of Sukabumi District from 106 degrees 22' 15"-107 degrees 4' 1" E and 6 degrees 42' 50" - 7 degrees 26' 13" S. There was a total of 11 Anopheles spp. collected from 209 sampling locations in the area covered by the Quickbird image and a total of 15 Anopheles spp. collected from 1,600 sampling locations in the area covered by the ASTER map. For the area covered by the land use maps, ten species were found to have statistically positive relationships between land use class and species presence: Anopheles aconitus, An. annularis, An. barbirostris. An. flavirostris, An. insulaeflorum, An. kochi, An. maculatus, An. subpictus, An. sundaicus, and An. vagus. Quickbird and ASTER satellite images both produced land maps that were adequate for predicting species presence in an area. The land use classes associated with malaria vector breeding were rice paddy (An. aconitus, An. subpictus), plantation located near or adjacent to human settlements (An. maculatus), bush/shrub (An. aconitus, An. maculatus, An. sundaicus), bare land, and water body land use on the coast located < or = 250 m of the beach (An. sundaicus). Understanding the associations of habitat and species in one area, predictions of species presence or absence can be made prior to a ground survey allowing for accurate vector survey and control planning.     

Reconsideration of anopheline mosquito phylogeny (Diptera: Culicidae: Anophelinae) based on morphological data

The phylogeny of anopheline mosquitoes (Culicidae: Anophelinae) is re‐examined using morphological data derived from adults, fourth‐instar larvae and pupae. Based on the data set of Sallum et al. (2000), we add some previously missing data and simplify and recode characters to eliminate ambiguities and more accurately reflect homologies, with special emphasis on characters of the male genitalia that provide the main criteria for the subgeneric classification of genus Anopheles. The principal aim of the study is to assess objectively the phylogenetic relationships and classification of two taxa not included by Sallum et al. (2000): Anopheles corethroides, a representative of the Australasian Stigmaticus Group, and An. kyondawensis, an unusual Oriental species whose adult and pupal stages were only recently discovered. The revised data set consists of 167 characters for 66 species representing the three traditionally recognised genera of Anophelinae, the six traditionally accepted subgenera of genus Anopheles and all informal series and most species groups of subgenera Anopheles, Cellia and Nyssorhynchus. The data are analysed using equal weighting (EW) and implied weighting (IW). Analysis under EW generates a strict consensus tree with principal lineages consistent with those reported by Sallum et al. (2000). Analysis under IW supports the monophyly of Anophelinae, the basal position of Chagasia, the monophyly of subgenera Cellia, Kerteszia and Nyssorhynchus, and the sister relationship of Kerteszia + Nyssorhynchus, but otherwise yields relationships that differ significantly in one respect or another from those obtained in all previous analyses of both morphological and molecular data. Subgenus Anopheles is arrayed as a polyphyletic lineage basal to a monophyletic clade comprising the Neotropical Kerteszia + Nyssorhynchus and the Old World Cellia in a sister‐group relationship. Bironella, Lophopodomyia and Stethomyia are firmly nested within subgenus Anopheles, which would nevertheless still be paraphyletic if these taxa were subsumed within it. Anopheles kyondawensis is well supported as the sister group of Bironella + all other Anopheles. Bironella, Stethomyia, An. corethroides and several other Anopheles clades are each strongly supported in a pectinate series of relationships, terminating in the clade comprising subgenera Cellia, Kerteszia and Nyssorhynchus. These relationships and other aspects of the phylogeny are discussed in relation to the formal and informal classification of genus Anopheles.