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 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 biogeography of the oil-collecting bee subgenus Centris (Wagenknechtia) (Hymenoptera,Apidae)

Centris (Wagenknechtia) is the only subgenus of centridine bees that occurs almost exclusively in the Andean Region. This study investigates the monophyly of C. (Wagenknechtia) proposing a hypothesis for the relationships among its species using 42 morphological characters of adults. The analysis resulted in one most parsimonious tree corroborating the monophyly of this group of oil-collecting bees, with the relationships among its species as follows: ((C. escomeli + C. moldenkei) (C. muralis (C. rhodophthalma + C. vardyorum) (C. cineraria + C. orellanai))). Based on these results plus available distributional records, a cladistic biogeographic analysis was performed. The resulting hypothesis of the relationships between the endemism areas in the Andean Region is as follows: (((Atacama–Puna) (Desert–Atacama–Puna)) ((Pampean–Chaco–Puna) ((Monte–Coquimban) (Coquimban (Patagonian–Valdivian Forest–Coquimban–Maule–Monte–Santiago–Magellanic Forest))))).     

Phylogeny and classification of Aedini (Diptera: Culicidae), based on morphological characters of all life stages

Higher‐level relationships within Aedini, the largest tribe of Culicidae, are explored using morphological characters of eggs, fourth‐instar larvae, pupae, and adult females and males. In total, 172 characters were examined for 119 exemplar species representing the existing 12 genera and 56 subgenera recognized within the tribe. The data for immature and adult stages were analysed separately and in combination using equal (EW) and implied weighting (IW). Since the classification of Aedini is based mainly on adult morphology, we first tested whether adult data alone would support the existing classification. Overall, the results of these analyses did not reflect the generic classification of the tribe. The tribe as a whole was portrayed as a polyphyletic assemblage of Aedes and Ochlerotatus within which eight (EW) or seven (IW) other genera were embedded. Strict consensus trees (SCTs) derived from analyses of the immature stages data were almost completely unresolved. Combining the adult and immature stages data resulted in fewer most parsimonious cladograms (MPCs) and a more resolved SCT than was found when either of the two data subsets was analysed separately. However, the recovered relationships were still unsatisfactory. Except for the additional recovery of Armigeres as a monophyletic genus, the groups recovered in the EW analysis of the combined data were those found in the EW analysis of adult data. The IW analysis of the total data yielded eight MPCs consisting of three sets of two mutually exclusive topologies that occurred in all possible combinations. We carefully studied the different hypotheses of character transformation responsible for each of the alternative patterns of relationship but were unable to select one of the eight MPCs as a preferred cladogram. Overall, the relationships within the SCT of the eight MPCs were a significant improvement over those found by equal weighting. Aedini and all existing genera except Ochlerotatus and Aedes were recovered as monophyletic. Ochlerotatus formed a polyphyletic assemblage basal to Aedes. This group included Haemagogus and Psorophora, and also Opifex in a sister‐group relationship with Oc. (Not.) chathamicus. Aedes was polyphyletic relative to seven other genera, Armigeres, Ayurakitia, Eretmapodites, Heizmannia, Udaya, Verrallina and Zeugnomyia. With the exception of Ae. (Aedimorphus), Oc. (Finlaya), Oc. (Ochlerotatus) and Oc. (Protomacleaya), all subgenera with two or more species included in the analysis were recovered as monophyletic. Rather than leave the generic classification of Aedini in its current chaotic state, we decided a reasonable and conservative compromise classification would be to recognize as genera those groups that are ‘weighting independent’, i.e. those that are common to the results of both the EW and IW analyses of the total data. The SCT of these combined analyses resulted in a topology of 29 clades, each comprising between two and nine taxa, and 30 taxa (including Mansonia) in an unresolved basal polytomy. In addition to ten genera (Armigeres, Ayurakitia, Eretmapodites, Haemagogus, Heizmannia, Opifex, Psorophora, Udaya, Verrallina and Zeugnomyia), generic status is proposed for the following: (i) 32 existing subgenera of Aedes and Ochlerotatus, including nine monobasic subgenera within the basal polytomy, i.e. Ae. (Belkinius), Ae. (Fredwardsius), Ae. (Indusius), Ae. (Isoaedes), Ae. (Leptosomatomyia), Oc. (Abraedes), Oc. (Aztecaedes), Oc. (Gymnometopa) and Oc. (Kompia); (ii) three small subgenera within the basal polytomy that are undoubtedly monophyletic, i.e. Ae. (Huaedes), Ae. (Skusea) and Oc. (Levua), and (iii) another 20 subgenera that fall within the resolved part of the SCT, i.e. Ae. (Aedes), Ae. (Alanstonea), Ae. (Albuginosus), Ae. (Bothaella), Ae. (Christophersiomyia), Ae. (Diceromyia), Ae. (Edwardsaedes), Ae. (Lorrainea), Ae. (Neomelaniconion), Ae. (Paraedes), Ae. (Pseudarmigeres), Ae. (Scutomyia), Ae. (Stegomyia), Oc. (Geoskusea), Oc. (Halaedes), Oc. (Howardina), Oc. (Kenknightia), Oc. (Mucidus), Oc. (Rhinoskusea) and Oc. (Zavortinkius). A clade consisting of Oc. (Fin.) kochi, Oc. (Fin.) poicilius and relatives is raised to generic rank as Finlaya, and Downsiomyia Vargas is reinstated from synonymy with Finlaya as the generic name for the clade comprising Oc. (Fin.) leonis, Oc. (Fin.) niveus and their relatives. Three other species of Finlaya?Oc. (Fin.) chrysolineatus, Oc. (Fin.) geniculatus and Oc. (Fin.) macfarlanei? fall within the basal polytomy and are treated as Oc. (Finlaya) incertae sedis. Ochlerotatus (Ochlerotatus) is divided into three lineages, two of which, Oc. (Och.) atropalpus and Oc. (Och.) muelleri, are part of the basal polytomy. The remaining seven taxa of Oc. (Ochlerotatus) analysed, including the type species, form a reasonably well‐supported group that is regarded as Ochlerotatus s.s. Ochlerotatus (Rusticoidus) is retained as a subgenus within Ochlerotatus s.s. Ochlerotatus (Nothoskusea) is recognized as a subgenus of Opifex based on two unique features that support their sister‐group relationship. A new genus, Tanakaius gen. nov. , is proposed for Oc. (Fin.) togoi and the related species Oc. (Fin.) savoryi. The taxonomic status and generic placement of all currently valid species of Aedini are listed in an appendix. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 142 , 289?368.     

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.     

Phylogeny,natural groups and nemertean classification

Contemporary practice in the classification of nemerteans (phylum Nemertea) is critically discussed. It is argued that basing higher taxa on the existence of a unique combination of characters in a species (or genus) is unlikely to lead to monophyletic taxa, and that this approach should be abandoned in favour of a classification based on explicit hypotheses of phylogeny. These hypotheses should be based on all available characters and characters should not be excluded before the analysis. The classification should be based on a reconstruction of the phylogeny and reflect this phylogeny in an unambiquous way.     

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 of Palaearctic Syrphidae (Diptera): evidence from larval stages

We estimated the phylogeny of Palaearctic Syrphidae using 187 larval morphological characters obtained from about 65% of the fauna (85 supraspecific taxa represented by 118 species) and based the analysis at the generic, level. The root of the syrphid tree was established from an outgroup consisting of other Aschizan families: the Platypezidae, Phoridae and Pipunculidae, with the tree rooted on the Lonchopteridae. The Syrphidae was the most derived Aschizan family. The Pipunculidae was the sister group to the Syrphidae. Eumerus was basal within Syrphidae. A trend exists towards increasing complexity of integumental folds and grooves across the Aschiza. In movement, the integument collapses along the line of these grooves. Grooves are evidence of muscles forming functional groups. Elaboration of independent groups of muscles appears to underlie much of the evolution of larval form within Aschiza. The basal feeding modes of syrphid larvae are mycophagy and phytophagy. Above these feeding modes, all remaining syiphids fall into one of two lineages comprising entomophages and saprophages, each of which has a single origin within the apparently polyphyletic Volucella. Major morphological innovation is associated with shifts between feeding modes; within feeding modes, change is gradual and tends towards increasing complexity. Change is mostly in structures associated with the mouthparts, thorax, anal segment and locomotory organs. Generic diagnoses with biological and taxonomic notes and a key to genera using larval characters are provided.     

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.     

Phylogeny of the Cocculinoidea (Mollusca, Gastropoda)

Abstract. The superfamily Cocculinoidea is a group of marine, deep-water, limpet-like gastropods. Recent speculation surrounding their affinities has concentrated on their placement within the Gastropoda. However, phylogenetic relationships within the Cocculinoidea, especially the monophyly of families and genera within the group, remain poorly understood. Phylogenetic analysis of 31 morphological characters for 15 cocculinoidean taxa and 2 outgroups resulted in a single most parsimonious tree, length=70, CI=0.62, and RI=0.71. Monophyly of the Cocculinoidea, Cocculinidae, and the genera Cocculina and Coccopigya was supported; Paracocculina and Coccocrater were found to be paraphyletic. Character optimization demonstrates that many characters often cited as diagnostic of various taxa, are often homoplastic and/or synapomorphies at different hierarchical levels.     

Phylogenetic reassessment and biogeography of the Ectateus generic group (Coleoptera: Tenebrionidae: Platynotina)

Owing to the reinterpretation of its morphological synapomorphies, the taxonomic composition of the Ectateus generic group had been ambiguous. The present study scrutinized all existing taxonomic concepts of the group based on a cladistic analysis of the adult morphology of all of the Afrotropical platynotoid Platynotina genera. The phylogenetic relationships were reconstructed using parsimony and Bayesian inference. The results show that all previous taxonomic concepts of the Ectateus generic group concerned paraphyletic entities. The cladistic analysis revealed the following synapomorphies for the taxon: (1) presence of basal indentations of the pronotal disc, (2) ratio of prothorax width to its maximal height > 6.0, and (3) ratio of maximal height of the prothorax to total height < 0.3. Moreover, phylogenetic studies revealed the existence of the Upembarus generic group, a sister‐taxon group to the Ectateus generic group, within the Afrotropical platynotoid Platynotina. Autapomorphic and synapomorphic character mapping show that several taxonomic and nomenclatural changes are needed to consider the particular generic‐level entities traditionally assigned to Afrotropical platynotoid Platynotina as monophyletic lineages. The following taxonomic and nomenclatural adjustments are made in this paper: P teroselinus gen. nov. is erected to accommodate a single species that was previously assigned to Zidalus: Pteroselinus insularis comb. nov. Additionally, the following synonymies are proposed: Anchophthalmops (= Platykochius syn. nov. ), Angolositus (= Aberlencus syn. nov. , = Platymedvedevia syn. nov. ), Glyptopteryx (= Microselinus syn. nov. , = Quadrideres syn. nov. , = Synquadrideres syn. nov. ). In addition, Kochogaster is lowered in rank and is treated as one of the subgenera of Anchophthalmus. Moreover, Pseudoselinus is treated as a subgenus of Upembarus. An identification key to all Afrotropical platynotoid Platynotina genera and subgenera is presented. Zoogeographical analyses revealed the following dispersal barriers for the Ectateus generic group: (1) the Sahara (northern barrier); (2) the dry ecosystems of Botswana, Namibia, and South Africa (southern barrier); and (3) the Congolian rainforests (internal distributional gap). The ancestor of the taxon probably originated in East African ecoregions that predominantly contained wattletrees (acacias) and Commiphora Jacq. Moreover, past climate changes seem to have had a great impact on the observed generic distribution. © 2015 The Linnean Society of London     

Phylogeny,classification and biogeography of bombyliine bee flies (Diptera,Bombyliidae)

The Bombyliinae comprises over 1100 described species in 73 known genera distributed worldwide. It is one of the largest subfamilies of bee flies (Diptera: Bombyliidae). We present the first phylogenetic hypothesis for this subfamily, based on 157 adult morphological characters scored for 123 species representing 60 genera, including all the tribes of Bombyliinae, and the related subfamilies Lordotinae and Toxophorinae. Four most parsimonious trees were generated from our analysis under equal weighting schemes. The monophyly of Bombyliinae is supported, and Lordotinae is sister to the Bombyliinae. Within Bombyliinae, Conophorini is sister to the remaining tribes. Five previously recognized tribes are revised and four new tribes are erected. We placed almost all genera in our tribal classification, based on our phylogenetic results and available character evidence. The genus Parabombylius is proposed as a synonym of Bombylius. The Gondwanan origin for the major lineages of Bombyliinae is strongly indicated by our biogeographic analysis which reconstructs ancestral areas. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub: 1EC5C827‐34D5‐4A95‐BA78‐4ACF457F6D40.     

Phylogeny and biogeography of Crinum L. (Amaryllidaceae) inferred from nuclear and limited plastid non-coding DNA sequences

The genus Crinum L. is the only pantropical genus of the Amaryllidaceae. Phylogenetic and biogeographical analyses of nrDNA ITS and plastid trnL-F sequences for all continental groups of the genus Crinum and related African genera are presented, with the genus Amaryllis used as outgroup. ITS indicates that C. baumii is more closely related to Ammocharis and Cybistetes than to Crinum sensu stricto . Three clades are resolved in Crinum s.s. One unites a monophyletic American group with tropical and North African species. The second includes all southern African species and the Australian endemic C. flaccidum . The third includes monophyletic Madagascar, Australasian and Sino-Himalayan clades, with southern African species. The trnL-F phylogeny resolves an American and an Asian/Madagscar clade, and confirms the relationship of C. flaccidum with species endemic to southern Africa. The salverform, actinomorphic perianths of subg. Crinum appear to have evolved several times in the genus from ancestors with zygomorphic perianths (subg. Codonocrinum ), thus neither subgenus is monophyletic. Biogeographical analyses place the origin of Crinum in southern Africa, as the region is optimized at all ancestral nodes in the tree topology, and in basal interior nodes of all but one of the major clades. The genus underwent three major waves of radiation corresponding to the three main clades resolved in our trees. Two entries into Australia for the genus are indicated, as are separate Sino-Himalayan and Australasian dispersal events.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141 , 349–363.     

Phylogeny of Heteronychia: the largest lineage of Sarcophaga (Diptera: Sarcophagidae)

Sarcophaga Meigen is one of the megadiverse genera of true flies, with approximately 850 valid species worldwide. The genus is divided into about 160 subgenera, the validity of a vast majority of which has never been verified using cladistic methods. This paper deals with the mainly Palaearctic subgenus Heteronychia Brauer & Bergenstamm, which comprises 89 species and is thus the largest subunit of Sarcophaga. We performed a cladistic analysis of the group based exclusively on male morphological characters. Parsimony analyses were run on a matrix of 84 characters for 88 species. Species of the subgenera Discachaeta Enderlein and Notoecus Stein were also included in the matrix. A further analysis was carried out using a subset of characters from the terminalia alone (70 characters). The results show that the clade formed by Heteronychia, Discachaeta, and Notoecus is monophyletic, with Discachaeta emerging as polyphyletic whereas Sarcophaga (Notoecus) longestylata Strobl is nested within the Sarcophaga filia‐group. Character states supporting Heteronychia and the few well‐supported species‐groups are discussed in detail. The following synonymies are proposed: Discachaeta = Heteronychia ( syn. nov. ) and Notoecus = Heteronychia ( syn. nov. ). The paper also includes a historical background of the taxon in relation to the classification of the genus Sarcophaga over the past two centuries, as well as a terminological review of the male terminalia, particularly of the distiphallus. © 2013 The Linnean Society of London     

Systematics and biogeography of the New World species of Trichadenotecnum Enderlein (Insecta: Psocodea: 'Psocoptera': Psocidae)

The taxonomy of the New World species of the genus Trichadenotecnum is revised. A total of 44 species, including 29 new species, were treated. These species are classified into 12 monophyletic species groups, eight of them newly proposed here. Two species previously assigned to Trichadenotecnum , T. pichincha New & Thornton and T. sylvaticum Turner, are recognized as not belonging to this genus. Phylogenetic relationships among 16 previously and presently proposed species groups are estimated based on a data matrix of 58 morphological characters. Trees from these analyses support monophyly of Trichadenotecnum and the proposed species groups. The New World species were divided into three major clades. Based on the phylogenetic hypothesis and distributional pattern of the species groups, the biogeographical history of the New World Trichadenotecnum is discussed.  © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society , 2008, 153 , 651–723.     

Phylogeny of Oestridae (Insecta: Diptera)

The phylogeny of Oestridae was analysed at the generic level using 118 characters from all developmental stages and including morphology, ontogeny, physiology and behaviour. Four major clades were given subfamilial rank with the phylogenetic relationship (Cuterebrinae (Gasterophilinae (Hypodermatinae + Oestrinae))). The subdermal parasites of the African elephant, Neocuterebra squamosa Grünberg and Ruttenia loxodontis Rodhain, had their most probable affiliation subordinate to the clade of stomach parasites, although their exact position needs further investigation. Genus Ochotonia Grunin, which is known from a single third‐instar larva only, was the probable sister group of all other Hypodermatinae. Twenty‐five oestrid genera were recognised as valid and those containing more than one species were defined through lists of autapomorphies. Cuterebra Clark was proposed as a senior synonym of Alouattamyia Townsend, Andinocuterebra Guimarães, Pseudogametes Bischof and Rogenhofera Brauer. The clade of hypodermatine ungulate parasites (Hypoderma Latreille +Pallasiomyia Rubtzov +Pavlovskiata Grunin +Przhevalskiana Grunin +Strobiloestrus Brauer) remained largely unresolved, and genus Przhevalskiana emerged without defining characters.     

Phylogeny of Ptychopteroidea (Insecta: Diptera)

A phylogenetic tree is proposed for the superfamily Ptychopteroidea, reconstructed taking into account both extinct and extant taxa and based mainly on characters of wing venation.     

Phylogeny of the orb-web building spiders (Araneae, Orbiculariae: Deinopoidea, Araneoidea)

This phylogenetic analysis of 31 exemplar taxa treats the 12 families of Araneoidea (Anapidae, Araneidae, Cyatholipidae, Linyphiidae, Mysmenidae, Nesticidae, Pimoidae, Symphytognathidae, Synotaxidae, Tetragnathidae, Theridiidae, and Theridiosomatidae). The data set comprises 93 characters: 23 from male genitalia, 3 from female genitalia, 18 from cephalothorax morphology, 6 from abdomen morphology, 14 from limb morphology, 15 from the spinnerets, and 14 from web architecture and other behaviour. Criteria for tree choice were minimum length parsimony and parsimony under implied weights. The outgroup for Araneoidea is Deinopoidea (Deinopidae and Uloboridae). The preferred shortest tree specifies the relationships ((Uloboridae, Deinopidae) (Araneidae (Tetragnathidae ((Theridiosomatidae (Mysmenidae (Symphytognathidae, Anapidae))) ((Linyphiidae, Pimoidae) ((Theridiidae, Nesticidae) (Cyatholipidae, Synotaxidae))))))). The monophyly of Tetragnathidae (including metines and nephilines), the symphytognathoids, theridiid-nesticid lineage, and Synotaxidae are confirmed. Cyatholipidae are sister to Synotaxidae, not closely related to either the Araneidae or Linyphiidae, as previously suggested. Four new clades are proposed: the cyatholipoids (Cyatholipidae plus Synotaxidae), the 'spineless femur clade' (theridioid lineage plus cyatholipoids), the 'araneoid sheet web builders' (linyphioids plus the spineless femur clade), and the 'reduced piriform clade' (symphytognathoids plus araneoid sheet web builders). The results imply a coherent scenario for web evolution in which the monophyletic orb gives rise to the monophyletic araneoid sheet, which in turn gives rise to the gumfoot web of the theridiid-nesticid lineage. While the spinning complement of single pairs of glands does not change much over the evolution of the group, multiple sets of glands are dramatically reduced in number, implying that derived araneoids are incapable of spinning many silk fibers at the same time.     

Phylogeny, biogeography, and host-plant association in the subfamily Apaturinae (Insecta: Lepidoptera: Nymphalidae) inferred from eight nuclear and seven mitochondrial genes

The subfamily Apaturinae consists of 20 genera and shows disjunct distributions and unique host-plant associations. Most genera of this subfamily are distributed in Eurasia South-East Asia and Africa, whereas the genera Doxocopa and Asterocampa are distributed mainly in South America and North America, respectively. Although the Apaturinae larvae mainly feed on the Cannabaceae, those of the genus Apatura are associated with Salix and Populus (Salicaceae), which are distantly related to the Cannabaceae. Here, we infer the phylogeny of Apaturinae and reconstruct the history of host shifting and of colonization in the New World. We analyzed 9761 bp of nuclear and mitochondrial DNA sequence data, including the genes encoding EF1a, Wg, ArgK, CAD, GAPDH, IDH, MDH, RpS5, COI, COII, ATPase8, ATPase6, COIII, ND3, and ND5 for 12 apaturine genera. We also inferred the phylogeny with six additional genera using mitochondrial sequence data alone. Within the Apaturinae, two major clades are recovered in all the datasets. These clades separate the New World genera, Doxocopa and Asterocampa, indicating that dispersal to the New World occurred at least twice. According to our divergence time estimates, these genera originated during the Early Oligocene to the Early Miocene, implying that they migrated across the Bering Land Bridge rather than the Atlantic Land Bridge. The temporal estimates also show that host shifting to Salix or Populus in Apatura occurred more than 15 million years after the divergence of their host plants. Our phylogenetic results are inconsistent with the previously accepted apaturine genus groups and indicate that their higher classification should be reconsidered.