Phylogeny and classification of tribe Aedini (Diptera: Culicidae)

The phylogeny and classification of tribe Aedini are delineated based on a cladistic analysis of 336 characters from eggs, fourth‐instar larvae, pupae, adult females and males, and immature stage habitat coded for 270 exemplar species, including an outgroup of four species from different non‐aedine genera. Analyses of the data set with all multistate characters treated as unordered under implied weights, implemented by TNT version 1.1, with values of the concavity constant K ranging from 7 to 12 each produced a single most parsimonious cladogram (MPC). The MPCs obtained with K values of 7–9 were identical, and that for K = 10 differed only in small changes in the relationships within one subclade. Because values of K < 7 and > 10 produced large changes in the relationships among the taxa, the stability of relationships exemplified by the MPC obtained from the K = 9 analysis is used to interpret the phylogeny and classification of Aedini. Clade support was assessed using parsimony jackknife and symmetric resampling. Overall, the results reinforce the patterns of relationships obtained previously despite differences in the taxa and characters included in the analyses. With two exceptions, all of the groups represented by two or more species were once again recovered as monophyletic taxa. Thus, the monophyly of the following genera and subgenera is corroborated: Aedes, Albuginosus, Armigeres (and its two subgenera), Ayurakitia, Bothaella, Bruceharrisonius, Christophersiomyia, Collessius (and its two subgenera), Dahliana, Danielsia, Dobrotworskyius, Downsiomyia, Edwardsaedes, Finlaya, Georgecraigius (and its two subgenera), Eretmapodites, Geoskusea, Gilesius, Haemagogus (and its two subgenera), Heizmannia (and subgenus Heizmannia), Hopkinsius (and its two subgenera), Howardina, Hulecoeteomyia, Jarnellius, Kenknightia, Lorrainea, Macleaya, Mucidus (and its two subgenera), Neomelaniconion, Ochlerotatus (subgenera Chrysoconops, Culicelsa, Gilesia, Pholeomyia, Protoculex, Rusticoidus and Pseudoskusea), Opifex, Paraedes, Patmarksia, Phagomyia, Pseudarmigeres, Rhinoskusea, Psorophora (and its three subgenera), Rampamyia, Scutomyia, Stegomyia, Tanakaius, Udaya, Vansomerenis, Verrallina (and subgenera Harbachius and Neomacleaya), Zavortinkius and Zeugnomyia. In addition, the monophyly of Tewarius, newly added to the data set, is confirmed. Heizmannia (Mattinglyia) and Verrallina (Verrallina) were found to be paraphyletic with respect to Heizmannia (Heizmannia) and Verrallina (Neomacleaya), respectively. The analyses were repeated with the 14 characters derived from length measurements treated as ordered. Although somewhat different patterns of relationships among the genera and subgenera were found, all were recovered as monophyletic taxa with the sole exception of Dendroskusea stat. nov. Fifteen additional genera, three of which are new, and 12 additional subgenera, 11 of which are new, are proposed for monophyletic clades, and a few lineages represented by a single species, based on tree topology, the principle of equivalent rank, branch support and the number and nature of the characters that support the branches. Acartomyia stat. nov. , Aedimorphus stat. nov. , Cancraedes stat. nov. , Cornetius stat. nov. , Geoskusea stat. nov. , Levua stat. nov. , Lewnielsenius stat. nov. , Rhinoskusea stat. nov. and Sallumia stat. nov., which were previously recognized as subgenera of various genera, are elevated to generic status. Catageiomyia stat. nov. and Polyleptiomyia stat. nov. are resurrected from synonymy with Aedimorphus, and Catatassomyia stat. nov. and Dendroskusea stat. nov. are resurrected from synonymy with Diceromyia. Bifidistylus gen. nov. (type species: Aedes lamborni Edwards) and Elpeytonius gen. nov. (type species: Ochlerotatus apicoannulatus Edwards) are described as new for species previously included in Aedes (Aedimorphus), and Petermattinglyius gen. nov. (type species: Aedes iyengari Edwards) and Pe. (Aglaonotus) subgen. nov. (type species: Aedes whartoni Mattingly) are described as new for species previously included in Aedes (Diceromyia). Four additional subgenera are recognized for species of Ochlerotatus, including Oc. (Culicada) stat. nov. (type species: Culex canadensis Theobald), Oc. (Juppius) subgen. nov. (type species: Grabhamia caballa Theobald), Oc. (Lepidokeneon) subgen. nov. (type species: Aedes spilotus Marks) and Oc. (Woodius) subgen. nov. (type species: Aedes intrudens Dyar), and seven are proposed for species of Stegomyia: St. (Actinothrix) subgen. nov. (type species: Stegomyia edwardsi Barraud), St. (Bohartius) subgen. nov. (type species: Aedes pandani Stone), St. (Heteraspidion) subgen. nov. (type species: Stegomyia annandalei Theobald), St. (Huangmyia) subgen. nov. (type species: Stegomyia mediopunctata Theobald), St. (Mukwaya) subgen. nov. (type species: Stegomyia simpsoni Theobald), St. (Xyele) subgen. nov. (type species: Stegomyia desmotes Giles) and St. (Zoromorphus) subgen. nov. (type species: Aedes futunae Belkin). Due to the unavailability of specimens for study, many species of Stegomyia are without subgeneric placement. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species.     

Phylogeny and classification of Ochlerotatus and allied taxa (Diptera: Culicidae: Aedini) based on morphological data from all life stages

The phylogenetic relationships and generic assignments of ‘Ochlerotatus’ and related taxa of uncertain taxonomic position in the classification of Aedini previously proposed by the authors in 2004 and 2006 are explored using 297 characters from eggs, fourth‐instar larvae, pupae, adults and immature habitat coded for 158 exemplar species. The ingroup comprises 54 species and the outgroup includes four non‐aedine species and 100 aedine species, 21 of which were previously classified as incertae sedis. Data are analysed in a total‐evidence approach using implied weighting. The analysis produced 158 most parsimonious cladograms. The strict consensus tree (SCT) corroborates the monophyly of the 30 generic‐level taxa recognized previously that are included in the analysis. Overall, the results show remarkable congruence with those obtained previously despite differences in the taxa and morphological characters analysed in this and the two previous studies. All species of Ochlerotatus s.s., subgenus ‘Ochlerotatus’sensu auctorum, Geoskusea, Levua, Pseudoskusea and Rhinoskusea included in the analysis fall within a single clade that is treated as genus Ochlerotatus; thus, the last four taxa are restored to their previous subgeneric rank within this genus. Nine additional subgenera, of which four are new, are proposed for monophyletic clades of Ochlerotatus species based on the strength of character support and application of the principle of equivalent rank. Acartomyia stat. nov. , Culicelsa stat. nov. , Gilesia stat. nov. , Protoculex stat. nov. and Chrysoconops stat. nov. are resurrected from synonymy with Ochlerotatus; and Empihals subgen. nov. (type species: Culex vigilax Skuse), Pholeomyia subgen. nov. (type species: Aedes calcariae Marks), Buvirilia subgen. nov. (type species: Aedes edgari Stone & Rosen) and Sallumia subgen. nov. (type species: Aedes hortator Dyar & Knab) are described as new. The sister group of Ochlerotatus includes a number of species that were previously regarded as incertae sedis in ‘Oc. (Finlaya)’ and ‘Oc. (Protomacleaya)’. Based on previous observations, refined relationships and new character support, three additional genera are recognized for species previously included in ‘Finlaya’, i.e. Danielsia stat. nov . (type species: Danielsia albotaeniata Leicester), Luius gen. nov. (type species: Aedes fengi Edwards) and Hopkinsius gen. nov. (type species: Aedes ingrami Edwards). Additionally, Alloeomyia subgen. nov. (type species: Culex pseudotaeniatus Giles) and Yamada subgen. nov. (type species: Aedes seoulensis Yamada) are introduced as subgenera of Collessius and Hopkinsius, respectively. As is usual with generic‐level groups of Aedini, the newly recognized genera and subgenera are polythetic taxa that are diagnosed by unique combinations of characters. The analysis corroborates the previous observation that ‘Oc. (Protomacleaya)’ is a polyphyletic assemblage of species. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 153 , 29–114.     

EOCENE RECORDS OF BEE FLIES (INSECTA, DIPTERA, BOMBYLIIDAE, COMPTOSIA): THEIR PALAEOBIOGEOGRAPHIC IMPLICATIONS AND REMARKS ON THE EVOLUTIONARY HISTORY OF BOMBYLIIDS

Abstract:  The first fossil records of Holarctic representatives of the genus Comptosia Macquart, 1840 from the middle Eocene Messel Pit, Germany, and the upper Eocene of Florissant, USA, are reported. The fossil from Messel, Comptosia pria sp. nov., is represented by a well-preserved wing, displaying characteristic wing venation, remains of the second wing and elements of the thorax. The Florissant fossil, C. miranda comb. nov., is preserved almost completely with both wings and most parts of the body visible. Extant members of the genus Comptosia are currently found only in Australia, and close relatives are known from southern South America. Previously this group of genera has been thought to be of Gondwanan origin, but these Eocene fossil representatives of Comptosia from the Holarctic region argue against this interpretation, and suggest that Comptosia and its relatives were more widespread in both Northern and Southern hemispheres during the Palaeogene. The fossil record of bee flies suggests a major radiation of bombyliids in the late Cretaceous or early Cenozoic. This assumption is supported by the comparatively young geological age of many bombyliid host taxa. A key factor driving the specialization of many bombyliid larvae on ground-dwelling hosts may have been avoidance of competition with tachinid flies and parasitic hymenopterans.     

Apioceridae (Insecta: Diptera): cladistic reappraisal and biogeography

Twenty members of the fly families Apioceridae and Mydidae are compared in terms of 77 adult characters. Cladistic analysis of 91 synapomorphies provides a well corroborated reconstruction of lineage relationships, and reveals that the Apioceridae is paraphyletic with respect to the Mydidae. In order to render the Apioceridae monophyletic, the genus Rhaphiomidas and the subfamily Megascelinae of the Apioceridae are transferred to the Mydidae. The subfamily Rhaphiomidinae is reinstated to accommodate Rhaphiomidas , comprising the most plesiomorphic mydids. The relationships of the remaining subfamilies of Mydidae are discussed in the context of these findings. The genus Apiocera is divided into four subgenera, the type subgenus Apiocera Westwood in Australia, Pyrocera subgen. nov. in North America, and the subgenera Ripidosyrma Hermann and Anypenus Philippi are applied to the South African and South American species, respectively. A key to the four subgenera of Apiocera is provided. The biogeographic relationships of the subgenera and genera of the Apioceridae and the Megascelinae are discussed. Although considered an example of a transantarctic or Gondwanan group, we argue that the distribution of the Apioceridae predates the Mesozoic supercontinent Gondwanaland and extends onto sections of Pangaea, and should be termed 'Pangaean'. The cladistic relationships between the genera of Apioceridae and Megascelinae are consistent with the geological vicariance of the fragments of Pangaea on which they now occur.     

Phylogeny and classification of Muscini (Diptera, Muscidae)

Worldwide in distribution, the tribe Muscini comprises 21 accepted genera and about 350 species. In the present study, a cladistic analysis based upon adult morphological characters is carried out in order to discuss the monophyly of the tribe and its genera, the intergeneric relationships and, in some cases, also the intrageneric relationships. As a result, Muscini is supported as a monophyletic tribe sister-group of Stomoxyini. Except for Morellia Robineau-Desvoidy, Curranosia Paterson, and Eudasyphora Townsend, all the remaining genera are monophyletic. The results are dubious for Polietes Rondani, which was then provisionally kept unchanged. Morellia was broadened to include the Neotropical endemic genera Parapyrellia Townsend, Trichomorellia Stein, and Xenomorellia Malloch. Therefore, a new classification is proposed for Morellia in which it is divided into four subgenera: Morellia s.s. , Parapyrellia , Trichomorellia , and Xenomorellia . Furthermore, the previously proposed subgenus Dasysterna Zimin is given new status as a genus; however, as it is preoccupied by Dasysterna Dejean, the new replacement name Ziminellia nom. nov. is proposed herewith. Eudasyphora was found to be a paraphyletic group relative to Dasyphora Robineau-Desvoidy; both genera are hence synonymized, and Dasyphora is classified in three subgenera: Dasyphora s.s. , Eudasyphora , and Rypellia Malloch. The analysis demonstrated that the traditional classification of Musca Linnaeus into subgenera is artificial and, moreover, that the use of characters from male genitalia could be strongly informative for classifying the genus in phylogeny-supported species groups. Finally, the new classification proposal for Muscini recognizes 18 genera and, furthermore, two undescribed genus-ranked taxa are indicated.  © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society , 2007, 149 , 493–532.     

Phylogeny and taxonomic revision of all genera of Conopidae (Diptera) based on morphological data

All global genera of the fly family Conopidae are revised here. A cladistic analysis of 117 morphological characters recorded from 154 species, including representatives of 59 genera and subgenera, recovers a phylogenetic hypothesis for the family. This hypothesis is used as the basis of a new classification for the family. Both Sicini and Zodionini are removed from Myopinae and elevated to subfamilial status. A new tribe, Thecophorini, is proposed within Myopinae to accommodate Thecophora, Scatoccemyia, and Pseudoconops. Two genera, Pseudomyopa and Parazodion, are removed from Dalmanniinae and placed in Myopinae and Zodioninae, respectively. Conopinae is divided into 11 tribes, seven of which are newly described (Asiconopini, Caenoconopini, Gyroconopini, Microconopini, Neoconopini, and Siniconopini). Some examined species are transferred to different or new genera and subgenera. A new genus, Schedophysoconops gen. nov. , and subgenus Asiconops (Aegloconops) subgen. nov. within Conopinae are described. A review of character evolution and phylogeography is included in light of the new classification. A catalogue of all genus‐group names is included with new emendations noted.     

Morphological classification of loaches (Nemacheilinae)

Comprehensive comparative morphological analysis of loaches from the subfamily Nemacheilinae is conducted. Forty significant phylogenetic characters are suggested, and phylogenetic relationships are reconstructed. The subfamily Nemacheilinae is subdivided into five tribes (Vaillantellini, Lefuini nov., Yunnanilini nov., Triplophysini nov., and Nemacheilini), and the relationships between these tribes can be expressed by the following formula: Vaillantellini (Lefuini (Yunnanilini (Triplophysini + Nemacheilini))). The classification of highland Asian loaches (Triplophysini) is specified, and the revised diagnoses of the genera from this tribe are given. Heterogeneity of the genus Triplophysa is demonstrated. This genus can be divided into several subgenera, and three new subgenera (Labiatophysa subgen. nov., Indotriplophysa subgen. nov., and Tarimichthys subgen. nov.) are described. A new species Hedinichthys grummorum sp. n is described from the Turpan Depression (northwest China).     

Notes on the Chenopodiaceae of Ethiopia, Somalia and Southern Arabia

Atriplex griffithii is divided into two subspecies, ssp. griffithii and ssp. stocksii , stat. nov. Atriplex farinosa is typified and divided into two subspecies, ssp. farinosa and ssp. keniensis , stat. nov. A new species of Atriplex, A. nogalensis is described from Somalia. The subtribe Sevadinae is revised and found to consist of the two monotypic genera Sevada and Fadenia and the genus Lagenantha with two species, L. cycloptera , comb. nov. (which includes the types of Gyroptera and Cycloptera) , and L. gillettii , comb. nov.     

Spiders of the genus Lachesana and a new storenoid genus from Israel (Araneae: Zodariidae)

Zodariid spiders of two genera in Israel are revised. The genus Lachesana is represented by two species. Only one male of these was ever depicted while the females of both and their hitherto unknown retreats were never described. Pax gen. nov. has been erected to accommodate five species, some of which have formerly been placed in Slorena, Selamia or Habronestes . Diagnostic characters of the West Mediterranean Selamia and the Australian Habronestes are newly illustrated. No drawings of Pax libani and P. islamita were ever published; the male of the latter is described here for the first time. Pax palmonii sp. nov. and P. engediensis sp. nov. are described and affinities of all taxa are discussed. Illustrations, records of distribution and keys to the species are provided.     

雪雀属系统发育关系的研究（雀形目：文鸟科)

雷雀属Montifringilla7种36个形态特征作为分类单元,进行支序分析。以麻雀属Passer、石雀属Petronia部分种类作为比较外群,讨论雪雀属的分类地位及其与近缘属麻雀属、石雀属系统发育关系;分析了雪雀属内种间的系统发育。支序分析的结果表明雪雀属为单系群,建议将雪雀属划分成两个亚属,讨论了雪雀属种类在进货上的位置,在3个近缘属的系统发育关系中,麻雀属类群较原始,雪雀属类群较进化,石雀属在进货中介于二之间。     

Phylogeny, classification, and biogeography of the cycloteline Therevinae (Insecta: Diptera: Thereavidae)

Twenty-one members of the Laurasian group of Therevinae (Diptera: Therevidae) are compared using 65 adult morphological characters. Cladistic analysis using parsimony on the 17 ingroup and 4 outgroup taxa provides a well-supported hypothesis of relationships among taxa within the Gyclotelini, tribe nov. The Cyclotelini is a monophyletic assemblage of mostly New World genera, including Anolinga , gen. nov. , Breviperna Irwin, Coleiana , gen. nov. , Crebraseta , gen. nov. , Cyclotelus Walker, Mesonana , gen.nov. , and Ozodiceromyia Bigot. In addition, three Old World genera, Ammothereva Lyneborg, Bugulaverpa , gen. nov. , and Procyclotelus Nagatomi & Lyneborg, are included in the tribe. These ten genera are divided into two monophyletic genus-groups, the Brevipema-group and the Cyclotelus-group. Keys are provided for the genera of Cyclotelini. The tribe, the two informal genus-groups, and all genera are diagnosed; five new genera and six new species are proposed. The biogeographical histories of the genera are discussed in terms of their cladistic relationships using methods of cladistic biogeography. Two major vicariant events account for the current distribution of the tribe. The first relates to the Beringian land bridge connecting western North America and eastern Asia. Second, New World cyclotelines were profoundly affected by the Early Eocene breakup of the archipelagic bridge between North and South America, and the distributions support the hypotheses favouring the continental origin of the Greater Antilles.     

A phylogenetic study of the Plantaginaceae

In a study based on morphological, embryological and chemical data of the Plantaginaceae, within the subclass Sympetalae or Asteridae, the superorder Lamianae is shown to be monophyletic. However, it was not possible to reconstruct the phylogeny within Lamianae or to find a sister-group for the monophyletic Plantaginaceae – Hydrostachyaceae for the latter is rejected. Three or rarely four genera have previously been recognized within Plantaginaceae, but in both cases Plantago appears as polyphyletic, which is considered unacceptable. Six clades are recognized as subgenera within Plantago: subgen. Plantago (c. 131 species), subgen. Coronopus (c. 11 species), subgen. Littorella (three species), subgen. Psyllium (c. 16 species), subgen. Bougueria (one species), and subgen. Albicans (51 species). Within P. subgen. Plantago, the paraphyletic sect. Plantago (c. 42 species) is found in all parts of the world except South America, New Zealand, Australia, and New Guinea, where sect. Oliganthos and sect. Mesembryniae vicariate. It is not possible to infer the phylogeny within sect. Plantago, for example between the endemic species from distant Pacific Islands; their common ancestor might be a species that once had a very wide distribution. Keys to genera, subgenera, sections and series are given. Only one genus, Plantago L., is recognized. The following proposals are made in the Appendix: P. ser. Oliganthos Rahn, ser. nov.; P. ser. Carpophorae (Rahn) Rahn, stat. nov. (= sect. C.); P. ser. Microcalyx (Pilg.) Rahn, stat. nov. (= sect. M.); P. unibrackteata Rahn, nom. nov. (=P. uniflora Hook.f. non L.); P. subgen. Littorella (P. J. Bergius) Rahn, stat. nov. (=Littorella P. J. Bergius); P. araucana Rahn, nom. nov. (=Littorella australis Griseb. non Plantago australis Lam.); P. americana (Fernald) Rahn, comb. nov.(=Littorella a.); P. subgen. Bougueria (Decne.) Rahn, stat. nov. (=Bougueria Decne.); P. nubicola (Decne.) Rahn, comb. nov. (=Bougueria n. Decne.); P. subgen. Albicans Rahn, subgen. nov. Ten lectotypes are selected.     

A taxonomic revision of Limacoccus Bondar with a cladistic analysis of its relationships with other scale insects (Hemiptera: Coccoidea)

The scale insect genus Limacoccus and its four palm pest species are revised and all instars with sexual dimorphism are described and illustrated. A key to Limacoccus species is provided. The unique life-history of this genus with construction of a protective tunnel by the second-instar female is described. A cladistic analysis of Limacoccus and eleven representative species of Diaspididae, Halimococcidae, Phoenicococcidae and Beesoniidae, was undertaken using fifty-four traditional and new characters from first and second instars and adult females. Character polarity was determined by outgroup comparison with Margarodidae, Pseudococcidae, Eriococcidae and Asterolecaniidae as outgroups. Nine most parsimonious trees were found. Analysis of the eleven terminal taxa of the strict consensus tree using Phoenicococcidae, Halimococcidae and Diaspididae as outgroups produced one parsimonious tree. As result of this analysis, Limacoccus is believed to be a monophyletic group of characterized by seven autapomorphies and is assigned to the family Beesoniidae fam.rev. The redefined family is divided into two tribes: Beesoniini stat.nov. with genera Beesonia and Mangalorea and Limacocciini stat.nov. with the genus Limacoccus. The Beesoniidae are here regarded as a sister group of Phoenicococcidae.     

Phylogeny and taxonomy of the Prenolepis genus‐group of ants (Hymenoptera: Formicidae)

Abstract. We investigated the phylogeny and taxonomy of the Prenolepis genus‐group, a clade of ants we define within the subfamily Formicinae comprising the genera Euprenolepis, Nylanderia, gen. rev. , Paraparatrechina, gen. rev. & stat. nov. , Paratrechina, Prenolepis and Pseudolasius. We inferred a phylogeny of the Prenolepis genus‐group using DNA sequence data from five genes (CAD, EF1αF1, EF1αF2, wingless and COI) sampled from 50 taxa. Based on the results of this phylogeny the taxonomy of the Prenolepis genus‐group was re‐examined. Paratrechina (broad sense) species segregated into three distinct, robust clades. Paratrechina longicornis represents a distinct lineage, a result consistent with morphological evidence; because this is the type species for the genus, Paratrechina is redefined as a monotypic genus. Two formerly synonymized subgenera, Nylanderia and Paraparatrechina, are raised to generic status in order to provide names for the other two clades. The majority of taxa formerly placed in Paratrechina, 133 species and subspecies, are transferred to Nylanderia, and 28 species and subspecies are transferred to Paraparatrechina. In addition, two species are transferred from Pseudolasius to Paraparatrechina and one species of Pseudolasius is transferred to Nylanderia. A morphological diagnosis for the worker caste of all six genera is provided, with a discussion of the morphological characters used to define each genus. Two genera, Prenolepis and Pseudolasius, were not recovered as monophyletic by the molecular data, and the implications of this result are discussed. A worker‐based key to the genera of the Prenolepis genus‐group is provided.     

Punctelia, a new lichen genus in the Parmeliaceae

The genus Punctelia Krog gen. nov., comprising subgen. Punctelia (the Parmelia borreri group) and subgen. Flavopunctelia Krog subgen. nov. (the Parmelia flaventior group), is here segregated from Parmelia s. str. The main differences between the two genera lie in the development of the pseudocyphellae, the chemical properties of the medulla, and the centres of distribution. Parmelia s. str. has its highest number of species in East Asia and the Australian region, Punctelia in the Americas and Africa.