Phylogeny and taxonomy of thegenera of south-western North American Euctenizinae trapdoor spidersand their relatives (Araneae: Mygalomorphae, Cyrtaucheniidae)

The primary focus of this paper is to evaluate the monophylyand intergeneric relationships of the cyrtaucheniid subfamily Euctenizinaeand to a lesser degree the monophyly of Cyrtaucheniidae. Using 71morphological characters scored for 29 mygalomorph taxa our cladisticanalysis shows that Cyrtaucheniidae is likely paraphyletic withrespect to the Domiothelina, the clade that comprises the Migidae,Actinopodidae, Ctenizidae, and Idiopidae. Together, the Domiothelinaand Cyrtaucheniidae have been treated as the Rastelloidina clade.A strict interpretation of rastelloid classification based on ourcladogram would require the establishment of four additional spiderfamilies. However, we choose to use informal names for these cladesso that these taxa can be validated by subsequent studies of mygalomorphphylogeny before formal names are introduced. The phylogenetic analysisalso serves as a vehicle for examining the patterns of homoplasyobserved in mygalomorphs. The secondary focus of this paper is ataxonomic revision of Euctenizinae genera from the south-westernUnited States that includes a key to its genera. The cyrtaucheniid genera Enrico and Astrosoga areconsidered junior synonyms of Eucteniza. Actinoxia and Nemesoides arejunior ­synonyms of Aptostichus . At present theNorth American Euctenizinae comprises these seven nominal genera: Eucteniza, Neoapachella gen.nov. ( Neoapachella rothi sp. nov. ), Myrmekiaphila,Entychides, Promyrmekiaphila, Apomastus gen. nov. ( Apomastusschlingeri sp. nov. ), and Aptostichus .  © 2002The Linnean Society of London . Zoological Journal of the LinneanSociety, 2002, 136 , 487−534     

Molecular phylogenetics of the spider infraorder Mygalomorphae using nuclear rRNA genes (18S and 28S): conflict and agreement with the current system of classification

Mygalomorph spiders, which include the tarantulas, trapdoor spiders, and their kin, represent one of three main spider lineages. Mygalomorphs are currently classified into 15 families, comprising roughly 2500 species and 300 genera. The few published phylogenies of mygalomorph relationships are based exclusively on morphological data and reveal areas of both conflict and congruence, suggesting the need for additional phylogenetic research utilizing new character systems. As part of a larger combined evidence study of global mygalomorph relationships, we have gathered approximately 3.7 kb of rRNA data (18S and 28S) for a sample of 80 genera, representing all 15 mygalomorph families. Taxon sampling was particularly intensive across families that are questionable in composition-Cyrtaucheniidae and Nemesiidae. The following primary results are supported by both Bayesian and parsimony analyses of combined matrices representing multiple 28S alignments: (1) the Atypoidea, a clade that includes the families Atypidae, Antrodiaetidae, and Mecicobothriidae, is recovered as a basal lineage sister to all other mygalomorphs, (2) diplurids and hexathelids form a paraphyletic grade at the base of the non-atypoid clade, but neither family is monophyletic in any of our analyses, (3) a clade consisting of all sampled nemesiids, Microstigmata and the cyrtaucheniid genera Kiama, Acontius, and Fufius is consistently recovered, (4) other sampled cyrtaucheniids are fragmented across three separate clades, including a monophyletic North American Euctenizinae and a South African clade, (5) of the Domiothelina, only idiopids are consistently recovered as monophyletic; ctenizids are polyphyletic and migids are only weakly supported. The Domiothelina is not monophyletic. The molecular results we present are consistent with more recent hypotheses of mygalomorph relationship; however, additional work remains before mygalomorph classification can be formally reassessed with confidence-increased taxonomic sampling and the inclusion of additional character systems (more genes and morphology) are required.     

Chasing ghosts: the phylogeny of Amaurobioidinae ghost spiders (Araneae,Anyphaenidae)

The family Anyphaenidae, also known as ghost spiders, includes a diverse array of nocturnal cursorial spiders that actively hunt on vegetation. The family is mostly distributed in the Americas and has been traditionally divided into three subfamilies. The mostly tropical and North American Anyphaeninae and the Amaurobioidinae, primarily distributed in southern South America, hold the bulk of the diversity, while the Malenellininae includes a single Chilean species. Here, we use a combined morphological and molecular approach to infer the relationships of the subfamily Amaurobioidinae and examine the delimitation of contentious genera. The morphological characters include both genitalic and somatic morphology, whereas molecular data include four markers, two mitochondrial (COI, 16S) and two nuclear (28S, H3). All our analyses agree on the monophyly of Amaurobioidinae, Amaurobioidini, Gayennini, the genera Negayan, Amaurobioides, Josa, Araiya, Arachosia and Monapia, as well as the paraphyly of Anyphaeninae. The total evidence analysis supports the novel placement of Josa as the sister group of both tribes Amaurobioidini and Gayennini, most of the previously known intergeneric relationships within Gayennini, and a clade of Amaurobioidini with a projecting ocular area, including Aysenoides, Axyracrus, Amaurobioides and Aysenia. The sequence data solve the puzzling placement of Philisca puconensis, here transferred to Tomopisthes, and Tasata chiloensis, transferred to Oxysoma. The advantages of the total evidence phylogenetic approach and the evolution of the male copulatory organ are discussed.     

Phylogeny of the plant bug subfamily Mirinae (Hemiptera: Heteroptera: Cimicomorpha: Miridae) based on total evidence analysis

The first comprehensive phylogenetic analyses of the most diverse subfamily of plant bugs, Mirinae, is presented in this study, for 110 representative taxa based on total evidence analysis. A total of 85 morphological characters and 3898 bp of mitochondrial (16S, COI) and nuclear (18S, 28S) sequences were analysed for each partitioned and combined dataset based on parsimony, maximum likelihood and Bayesian inference. Major results obtained in this study include monophyly of the tribe Mecistoscelini. The largest tribe, Mirini, was recovered as polyphyletic, and Stenodemini was recovered as paraphyletic. The clade of Stenodemini + Mecistoscelini is the sister group of the remaining Mirinae. The monophyly of two complexes composed of superficially similar genera were tested; the Lygus complex was recovered as nonmonophyletic, and the Adelphocoris–Creontiades–Megacoelum complex was confirmed to be monophyletic. The generic relationships of the main clades within each tribe based on the phylogeny, as well as their supported morphological characters, are discussed.     

Molecular systematics of Eumolpinae and the relationships with Spilopyrinae (Coleoptera, Chrysomelidae)

The 3400 species of Eumolpinae constitute one of the largest subfamilies of leaf beetles (Chrysomelidae). Their systematics is still largely based on late 19th century monographs and remains highly unsatisfactory. Only recently, some plesiomorphic lineages have been split out as separate subfamilies, including the southern hemisphere Spilopyrinae and the ambiguously placed Synetinae. Here we provide insight into the internal systematics of the Eumolpinae based on molecular phylogenetic analyses of three ribosomal genes, including partial mitochondrial 16S and nuclear 28S and complete nuclear 18S rRNA gene sequences. Sixteen morphological characters considered important in the higher-level systematics of Eumolpinae were also included in a combined analysis with the molecular characters. All phylogenetic analyses were performed using parsimony by optimizing length variation directly on the tree, as implemented in the POY software. The data support the monophyly of the Spilopyrinae outside the clade including all sampled Eumolpinae, corroborating their treatment as a separate subfamily within the Chrysomelidae. The systematic placement of the Synetinae remains ambiguous but consistent with considering it a different subfamily as well, since the phylogenetic analyses using all the available evidence show the representative sequence of the subfamily also unrelated to the Eumolpinae. The Megascelini, traditionally considered a separate subfamily, falls within the Eumolpinae. Several recognized taxonomic groupings within Eumolpinae, including the tribes Adoxini or Typophorini, are not confirmed by molecular data; others like Eumolpini seem well supported. Among the morphological characters analyzed, the presence of a characteristic groove on the pygidium (a synapomorphy of the Eumolpini) and the shape of tarsal claws (simple, appendiculate or bifid) stand out as potentially useful characters for taxonomic classification in the Eumolpinae.     

Inferring species trees from gene trees in a radiation of California trapdoor spiders (Araneae, Antrodiaetidae, Aliatypus)

Background

The California Floristic Province is a biodiversity hotspot, reflecting a complex geologic history, strong selective gradients, and a heterogeneous landscape. These factors have led to high endemic diversity across many lifeforms within this region, including the richest diversity of mygalomorph spiders (tarantulas, trapdoor spiders, and kin) in North America. The trapdoor spider genus Aliatypus encompasses twelve described species, eleven of which are endemic to California. Several Aliatypus species show disjunct distributional patterns in California (some are found on both sides of the vast Central Valley), and the genus as a whole occupies an impressive variety of habitats.

Methodology/Principal Findings

We collected specimens from 89 populations representing all described species. DNA sequence data were collected from seven gene regions, including two newly developed for spider systematics. Bayesian inference (in individual gene tree and species tree approaches) recovered a general “3 clade” structure for the genus (A. gulosus, californicus group, erebus group), with three other phylogenetically isolated species differing slightly in position across different phylogenetic analyses. Because of extremely high intraspecific divergences in mitochondrial COI sequences, the relatively slowly evolving 28S rRNA gene was found to be more useful than mitochondrial data for identification of morphologically indistinguishable immatures. For multiple species spanning the Central Valley, explicit hypothesis testing suggests a lack of monophyly for regional populations (e.g., western Coast Range populations). Phylogenetic evidence clearly shows that syntopy is restricted to distant phylogenetic relatives, consistent with ecological niche conservatism.

Conclusions/Significance

This study provides fundamental insight into a radiation of trapdoor spiders found in the biodiversity hotspot of California. Species relationships are clarified and undescribed lineages are discovered, with more geographic sampling likely to lead to additional species diversity. These dispersal-limited taxa provide novel insight into the biogeography and Earth history processes of California.     

Assessing the taxonomic resolution of southern African trapdoor spiders (Araneae: Ctenizidae; Cyrtaucheniidae; Idiopidae) and implications for their conservation

Taxonomic classifications simultaneously represent hypotheses of taxon identity and relationships to taxonomists, and real, unchanging entities to users of taxonomic information. Taxonomic changes, while representing scientific progress, can be a source of frustration for users. A method for assessing confidence in the taxonomy of a group of organisms would assist users of the taxonomy. A method is presented for determining the degree of development of a taxonomy, a concept termed ‘taxonomic resolution’. The method was applied to six groups of southern African mygalomorph trapdoor spiders, namely Stasimopus Simon 1892 (Ctenizidae Thorell 1877), Ancylotrypa Simon 1889 (Cyrtaucheniidae Simon 1889), four genera of Idiopidae Simon 1889 assessed as a single group, Galeosoma Purcell 1903, the families Migidae Simon 1889 and Microstigmatidae Roewer 1942, and the burrowing scorpion genus Opistophthalmus C. L. Koch 1837 (Scorpionidae Latreille 1802). The method was based on the assumption that species delimitation in a group of organisms, the taxonomy of which is based on morphological characters, depends on whether the sample of material examined is adequate for assessing variation in those characters. Five assessment criteria were identified and scored for a group of species using the taxonomic literature. Estimates of the number of species remaining to be discovered and described in each group were also included in the assessment. The results obtained for the trapdoor spiders ranged from 15 to 29%, indicating a potentially significant degree of uncertainty in the taxonomy. Results for Migidae and Microstigmatidae were 51 and 78% respectively, whereas the result for Opistophthalmus was 93%. The applied value of a measure of taxonomic resolution, the limitations of the method, and a strategy for developing a more generally applicable method are discussed.     

Molecular phylogeny and biogeography of an ancient Holarctic lineage of mygalomorph spiders (Araneae: Antrodiaetidae: Antrodiaetus)

The mygalomorph spider genera Antrodiaetus and Atypoides (Antrodiaetidae) belong to an ancient lineage that has persisted since at least the Cretaceous. These spiders display a classic disjunct Holarctic distribution with species in the eastern Palaearctic plus the western and eastern Nearctic. Prior phylogenetic analyses of this group have been proposed on the basis of morphology, but lack strong support and independent corroboration. Here we present the first phylogenetic analysis of species-level relationships based on molecular data obtained from the mitochondrial (cytochrome c oxidase subunit I) and nuclear (18S and 28S rRNA) genomes. Analyses corroborate earlier findings that Atypoides forms a paraphyletic grade with respect to Antrodiaetus, and consequently, that genus is formally synonymized under Antrodiaetus. In addition, our results support the relatively early divergence of Antrodiaetus roretzi. Antrodiaetus pacificus is "paraphyletic" with respect to the A. lincolnianus group and is likely an assemblage of numerous species. The final topology based on a combined molecular dataset, in conjunction with two different molecular dating techniques (penalized likelihood plus a Bayesian approach) and ancestral distribution reconstructions, was used to infer the historical biogeography of these spiders. Trans-Beringian and trans-Atlantic routes appear to account for the present-day distribution of Antrodiaetus in Japan and North America. Future studies on Antrodiaetus phylogeny will be used to address questions regarding morphological stasis and the evolution of quantitative morphological characters.     

Insights into the phylogenetic relationships within Cixiidae (Hemiptera: Fulgoromorpha): cladistic analysis of a morphological dataset

Abstract.  According to the most recent classifications proposed, the planthopper family Cixiidae comprises three subfamilies, namely Borystheninae, Bothriocerinae and Cixiinae, the latter with 16 tribes. Here we examine morphological characters to present the first phylogenetic reconstructions within Cixiidae derived from a cladistic analysis. We scored 85 characters of the head, thorax, and male and female genitalia for 50 taxa representative of all cixiid subfamilies and tribes and for six outgroup taxa. Analyses were based on maximum parsimony – using both equally weighted and successive weighting procedures – and Bayesian inferences. The monophyly of most currently accepted tribes and subfamilies was investigated through Templeton statistical tests of alternative phylogenetic hypotheses. The cladistic analyses recover the monophyly of Cixiidae, the subfamily Bothriocerinae, and the tribes Pentastirini, Mnemosynini, and Eucarpiini. Successive weighting and Bayesian inference recover the monophyly of the tribe Gelastocephalini, but only Bayesian inference supports the monophyly of Semoniini. The relationships recovered support the groups [Stenophlepsini (Borystheninae + Bothriocerinae)] arising from the tribe Oecleini, and [Andini + Brixiidini + Brixiini (polyphyletic) + Bennini]. Templeton tests reject the alternative hypothesis of a monophyletic condition for the tribe Pintaliini as presently defined.     

Correlation with a limited set of behavioral niches explains the convergence of somatic morphology in mygalomorph spiders

Understanding the drivers of morphological convergence requires investigation into its relationship with behavior and niche space, and such investigations in turn provide insights into evolutionary dynamics, functional morphology, and life history. Mygalomorph spiders (trapdoor spiders and their kin) have long been associated with high levels of morphological homoplasy, and many convergent features can be intuitively associated with different behavioral niches. Using genus-level phylogenies based on recent genomic studies and a newly assembled matrix of discrete behavioral and somatic morphological characters, we reconstruct the evolution of burrowing behavior in the Mygalomorphae, compare the influence of behavior and evolutionary history on somatic morphology, and test hypotheses of correlated evolution between specific morphological features and behavior. Our results reveal the simplicity of the mygalomorph adaptive landscape, with opportunistic, web-building taxa at one end, and burrowing/nesting taxa with structurally modified burrow entrances (e.g., a trapdoor) at the other. Shifts in behavioral niche, in both directions, are common across the evolutionary history of the Mygalomorphae, and several major clades include taxa inhabiting both behavioral extremes. Somatic morphology is heavily influenced by behavior, with taxa inhabiting the same behavioral niche often more similar morphologically than more closely related but behaviorally divergent taxa, and we were able to identify a suite of 11 somatic features that show significant correlation with particular behaviors. We discuss these findings in light of the function of particular morphological features, niche dynamics within the Mygalomorphae, and constraints on the mygalomorph adaptive landscape relative to other spiders.     

Phylogeny of pholcid spiders (Araneae: Pholcidae): combined analysis using morphology and molecules

The spider family Pholcidae comprises a large number of mainly tropical, web-weaving spiders, and is among the most diverse and dominant spider groups in the world. The phylogeny of this family has so far been investigated exclusively using morphological data. Here, we present the first molecular data for the family analyzed in a phylogenetic context. Four different gene regions (12S rRNA, 16S rRNA, cytochrome c oxidase subunit I, 28S rRNA) and 45 morphological characters were scored for 31 pholcid and three outgroup taxa. The data were analyzed both for individual genes, combined molecular data, and molecular plus morphological data, using parsimony, maximum likelihood, and Bayesian methods. Some of the phylogenetic hypotheses obtained previously using morphology alone were also supported by our results, like the monophyly of pholcines and of the New World clade. On the other hand, some of the previous hypotheses could be discarded with some confidence (monophyly of holocnemines, the position of Priscula), and still others need further investigation (the position of holocnemines, ninetines, and Metagonia). The data obtained provide an excellent basis for future investigations of phylogenetic patterns both within the family and among spider families.     

On the systematics of the fungus gnat subfamily Mycetophilinae (Diptera): a combined morphological and molecular approach

The phylogenetic relationships within the fungus gnat subfamily Mycetophilinae (Diptera) are addressed using a combined morphological and molecular approach. Twenty-four species, representing nine genera of the tribe Mycetophilini and 15 genera of the tribe Exechiini, were included in the study. Analyses include nucleotide sequences of mitochondrial (cytochrome oxidase I and 16S), and nuclear (18S and 28S rDNA) genes, in addition to 65 morphological characters. A combined parsimony analysis, including all characters, supports the monophyly of the subfamily Mycetophilinae and two of its tribes, Exechiini and Mycetophilini. There is also statistical support for a Mycetophila- group and a Phronia- group within the tribe Mycetophilini. The Phronia- group includes the genera Phronia , Macrobrachius and Trichonta . The Mycetophila- group includes the genera Mycetophila , Epicypta , Platurocypta , Sceptonia and Zygomyia . A Bayesian analysis based on the nucleotide sequences alone also support these clades within Mycetophilini except for the position of Dynatosoma which is recovered as the sister taxon to the Phronia- group. A somewhat different pattern, however, is observed for the tribe Exechiini – neither molecular data nor the combined data set support unambiguously any intergeneric relationships within Exechiini.     

Dracula ant phylogeny as inferred by nuclear 28S rDNA sequences and implications for ant systematics (Hymenoptera: Formicidae: Amblyoponinae)

Ants are one of the most ecologically and numerically dominant families of organisms in almost every terrestrial habitat throughout the world, though they include only about 1% of all described insect species. The development of eusociality is thought to have been a driving force in the striking diversification and dominance of this group, yet we know little about the evolution of the major lineages of ants and have been unable to clearly determine their primitive characteristics. Ants within the subfamily Amblyoponinae are specialized arthropod predators, possess many anatomically and behaviorally primitive characters and have been proposed as a possible basal lineage within the ants. We investigate the phylogenetic relationships among the members of the subfamily, using nuclear 28S rDNA sequence data. Outgroups for the analysis include members of the poneromorph and leptanillomorph (Apomyrma, Leptanilla) ant subfamilies, as well as three wasp families. Parsimony, maximum likelihood, and Bayesian analyses provide strong support for the monophyly of a clade containing the two genera Apomyrma+Mystrium (100% bpp; 97% ML bs; and 97% MP bs), and moderate support for the monophyly of the Amblyoponinae as long as Apomyrma (Apomyrminae) is included (87% bpp; 57% ML bs; and 76% MP bs). Analyses did not recover evidence of monophyly of the Amblyopone genus, while the monophyly of the other genera in the subfamily is supported. Based on these results we provide a morphological diagnosis of the Amblyoponinae that includes Apomyrma. Among the outgroup taxa, Typhlomyrmex grouped consistently with Ectatomma, supporting the recent placement of Typhlomyrmex in the Ectatomminae. The results of this present study place the included ant subfamilies into roughly two clades with the basal placement of Leptanilla unclear. One clade contains all the Amblyoponinae (including Apomyrma), Ponerinae, and Proceratiinae (Poneroid clade). The other clade contains members from subfamilies Cerapachyinae, Dolichoderinae, Ectatomminae, Formicinae, Myrmeciinae, and Myrmicinae (Formicoid clade).     

Anatomy and phylogenetic analysis of the neotropical callichthyid catfishes (Ostariophysi, Siluriformes)

Based mainly on morphological characters, the phylogenetic relationships among genera and some species groups of the neotropical family Callichthyidae were examined. A study of the osteology of a generalized callichthyid, Callichthys callichthys (Linnaeus), with detailed comparisons among representatives of the remaining genera in the family, is presented and used as a basis for the phylogenetic analysis. A single most parsimonious tree supported the monophyly of the family Callichthyidae based on 28 derived features and the division of the family in the subfamilies Corydoradinae and Callichthyinae. In the subfamily Corydoradinae, the genus Aspidoras is the sister-group of the clade formed by Corydoras plus Brochis. Five derived features support the monophyly of this clade and four support the monophyly of Brochis. No characters, however, were found to support the genus Corydoras. In the subfamily Callichthyinae, Dianema and Hopbstemum are sister-taxa. Megalechis represents the sister-group of Dianema plus Hoplosternum and Lepthoplosternum represents the sister-group to Megalechis plus Dianema plus Hopbstemum. Finally, Callichthys is considered the least derived member of the subfamily, and is hypothesized as the sister-group of the remaining species. A key to all callichthyid genera is provided.     

Phylogenetic relationships and taxonomic ranking of pipizine flower flies (Diptera: Syrphidae) with implications for the evolution of aphidophagy

The taxonomic rank and phylogenetic relationships of the pipizine flower flies (Diptera: Syrphidae: Pipizini) were estimated based on DNA sequence data from three gene regions (COI, 28S and 18S) and 111 adult morphological characters. Pipizini has been treated as a member of the subfamily Eristalinae based on diagnostic adult morphological characteristics, while the larval feeding mode and morphology is shared with members of the subfamily Syrphinae. We analysed each dataset, both separately and combined, in a total evidence approach under maximum parsimony and maximum likelihood. To evaluate the influence of different alignment strategies of rDNA 28S and 18S genes on the resulting topologies, we compared the topologies inferred from a multiple alignment using fast Fourier transform (MAFFT) program with those topologies resulting from aligning the secondary structure of these rDNA genes. Total evidence analyses resolved pipizines as a sister group of the subfamily Syrphinae. Although the structural alignment and the MAFFT alignment differed in the inferred relationships of some clades and taxa, there was congruence in the placement of pipizines. The homogeneous morphology of the Pipizini clade in combination with their unique combination of characters among the Syrphidae suggest a change of rank to subfamily. Thus, we propose to divide Syrphidae into four subfamilies, including the subfamily Pipizinae stat. rev.     

Phylogenetic relationships of the spider family Tetragnathidae (Araneae, Araneoidea) based on morphological and DNA sequence data

The monophyly of Tetragnathidae including the species composition of the family (e.g., Are Nephila and their relatives part of this lineage?), the phylogenetic relationships of its various lineages, and the exact placement of Tetragnathidae within Araneoidea have been three recalcitrant problems in spider systematics. Most studies on tetragnathid phylogeny have focused on morphological and behavioral data, but little molecular work has been published to date. To address these issues we combine previous morphological and behavioral data with novel molecular data including nuclear ribosomal RNA genes 18S and 28S, mitochondrial ribosomal RNA genes 12S and 16S and protein‐coding genes from the mitochondrion [cytochrome c oxidase subunit I (COI)] and from the nucleus (histone H3), totaling ca. 6.3 kb of sequence data per taxon. These data were analyzed using direct optimization and static homology using both parsimony and Bayesian methods. Our results indicate monophyly of Tetragnathidae, Tetragnathinae, Leucauginae, the “Nanometa clade” and the subfamily Metainae, which, with the exception of the later subfamily, received high nodal support. Morphological synapomorphies that support these clades are also discussed. The position of tetragnathids with respect to the rest of the araneoid spiders remains largely unresolved but tetragnathids and nephilids were never recovered as sister taxa. The combined dataset suggests that Nephilidae is sister to Araneidae; furthermore, the sister group of Nephila is the clade composed by Herennia plus Nephilengys and this pattern has clear implications for understanding the comparative biology of the group. Tetragnathidae is most likely sister to some members of the “reduced piriform clade” and nephilids constitute the most‐basal lineage of araneids.     

Phylogeny of Anophelinae (Diptera: Culicidae) based on nuclear ribosomal and mitochondrial DNA sequences

Abstract Phylogenetic relationships among thirty-two species of mosquitoes in subfamily Anophelinae are inferred from portions of the mitochondrial genes COI and COII, the nuclear 18S small subunit rRNA gene and the expansion D2 region of the nuclear large subunit 28S rRNA gene. Sequences were obtained from the genera Anopheles , Bironella and Chagasia . Representatives of all six subgenera of Anopheles were included: Anopheles , Cellia , Kerteszia , Lophopodomyia , Nyssorhynchus and Stethomyia. Using parsimony and maximum likelihood methods, various combinations of these DNA sequence data were analysed separately: 18S, 28S, combined 18S and 28S, combined COI and COII, and combined 18S, 28S, COI and COII ('total evidence'). The combined rDNA data contain strong phylogenetic signal, moderately to strongly supporting most clades in MP and ML analyses; however, the mtDNA data (analysed as either nucleotide or amino acid sequences) contain little phylogenetic signal, except for relationships of very recently derived groups of species and, at the deepest level, for the monophyly of Anophelinae. The paraphyly of Anopheles relative to Bironella is confirmed by most analyses and statistical tests. Support for the monophyly of subgenera Anopheles , Cellia , Kerteszia and Nyssorhynchus is indicated by most analyses. Subgenus Lophopodomyia is reconstructed as the sister to Bironella , nested within a clade also containing Nyssorhynchus and Kerteszia . The most basal relationships within genus Anopheles are not well resolved by any of the data partitions, although the results of statistical analyses of the rDNA data (S-H-tests, likelihood ratio tests for monophyly and Bayesian MCMC analyses) suggest that the clade consisting of Bironella , Lophopodomyia , Nyssorhynchus and Kerteszia is the sister to the clade containing Cellia and Anopheles .     

基于形态学证据的长足虻科系统发育研究(双翅目,短角亚目)

试图根据成虫形态学证据探讨长足虻科各亚科之间的系统关系,同时检验各个亚科的单系性.在比较形态学研究基础上,同时参考前人有关长足虻科高阶元分类的研究结果,筛选出42个来自头部、胸部(包括足和翅)、腹部、雌性和雄性外生殖器在亚科水平的分类特征,为了考察亚科的单系性,也包括亚科的自有衍征;运用支序分类的方法,首次分析并讨论了世界长足虻科17个亚科之间的系统发育关系.结果表明,长足虻科是一个严格的单系群,其支持的共同衍征为体色金绿,亚前缘脉端部与第1径脉中部愈合,前缘脉接近肩横脉处有1个缺刻,第2基室与盘室愈合,臀室短小、终止于径脉分叉点之前,雄性外生殖器明显向下或向前弯折,生殖背板具生殖孔,下生殖板与第9背板愈合.金长足虻亚科Sciapodinae腋瓣发达,中脉分叉,为最基部的支系,是最原始的亚科;而长足虻科的其他亚科构成一单系群,其共同衍征为腋瓣不明显,中脉不分叉.斜脉长足虻亚科Plagioneurinae也比较原始,是靠基部的支系,支持其单系性的特征为腹部第7～8节膜质化,生殖孔基位.异长足虻亚科Diaphorinae和锥长足虻亚科Rhaphiinae以及斯长足虻亚科Stolidosomatinae和合长足虻亚科Sympycninae分别构成姊妹群关系,斯长足虻亚科Stolidosomatinae的两个属Pseudosympycnus和Stolidosoma系统地位还有待进一步研究.此外,巴长足虻亚科Babindellinae、聚脉长足虻亚科Medeterinae和寇长足虻亚科Kowmunginae构成单系群,其共同衍征为臀脉短或不明显,无后顶鬃.研究所用标本大部分保存在中国农业大学昆虫标本馆,包括与美国史密森研究院和澳大利亚博物馆交换而来的标本,部分标本保存在比利时皇家科学院.     

Phylogenetics, biogeography and classification of, and character evolution in, gamebirds (Aves: Galliformes): effects of character exclusion, data partitioning and missing data

The phylogenetic relationships, biogeography and classification of, and morpho‐behavioral (M/B) evolution in, gamebirds (Aves: Galliformes) are investigated. In‐group taxa (rooted on representatives of the Anseriformes) include 158 species representing all suprageneric galliform taxa and 65 genera. The characters include 102 M/B attributes and 4452 nucleic acid base pairs from mitochondrial cytochrome b (CYT B), NADH dehydrogenase subunit 2 (ND2), 12S ribosomal DNA (12S) and control region (CR), and nuclear ovomucoid intron G (OVO‐G). Analysis of the combined character data set yielded a single, completely resolved cladogram that had the highest levels of jackknife support, which suggests a need for a revised classification for the phasianine galliforms. Adding 102 M/B characters to the combined CYT B and ND2 partitions (2184 characters) decisively overturns the topology suggested by analysis of the two mtDNA partitions alone, refuting the view that M/B characters should be excluded from phylogenetic analyses because of their relatively small number and putative character state ambiguity. Exclusion of the OVO‐G partition (with > 70% missing data) from the combined data set had no effect on cladistic structure, but slightly lowered jackknife support at several nodes. Exclusion of third positions of codons in an analysis of a CYT B + ND2 partition resulted in a massive loss of resolution and support, and even failed to recover the monophyly of the Galliformes with jackknife support. A combined analysis of putatively less informative, “non‐coding” characters (CYT B/ND2 third position sites + CR +12S + OVO‐G sequences) yielded a highly resolved consensus cladogram congruent with the combined‐evidence cladogram. Traditionally recognized suprageneric galliform taxa emerging in the combined cladogram are: the families Megapodiidae (megapodes), Cracidae (cracids), Numididae (guineafowls), Odontophoridae (New World quails) and Phasianidae (pheasants, pavonines, partridges, quails, francolins, spurfowls and grouse) and the subfamilies Cracinae (curassows, chachalacas and the horned guan), Penelopinae (remaining guans), Pavoninae sensu lato (peafowls, peacock pheasants and argus pheasants), Tetraoninae (grouse) and Phasianinae (pheasants minus Gallus). The monophyly of some traditional groupings (e.g., the perdicinae: partridges/quails/francolins) is rejected decisively, contrasted by the emergence of other unexpected groupings. The most remarkable phylogenetic results are the placement of endemic African galliforms as sisters to geographically far‐distant taxa in Asia and the Americas. Biogeographically, the combined‐data cladogram supports the hypothesis that basal lineages of galliforms diverged prior to the Cretaceous/Tertiary (K‐T) Event and that the subsequent cladogenesis was influenced by the break‐up of Gondwana. The evolution of gamebirds in Africa, Asia and the Americas has a far more complicated historical biogeography than suggested to date. With regard to character evolution: spurs appear to have evolved at least twice within the Galliformes; a relatively large number of tail feathers (≥ 14) at least three times; polygyny at least twice; and sexual dimorphism many times. © The Willi Hennig Society 2006.