Phylogeny of New Zealand hepialid moths (Lepidoptera: Hepialidae) inferred from a cladistic analysis of morphological data

The phylogeny of the New Zealand hepialid moths was estimated from a cladistic analysis of sixty‐three morphological characters, from all life cycle stages. One hundred and sixteen maximum parsimony trees were produced. The phylogenetic reconstruction indicated that the currently recognized generic concepts, and the four informal lineages hypothesized in a previous morphological taxonomic revision, were monophyletic. The relationships of species within genus Wiseana were not fully resolved. Analysis of a data set of thirty‐nine adult male characters from the New Zealand taxa and the Australian genera Jeana, Oxycanus and Trictena supported the monophyly of the New Zealand ‘Oxycanus’ s.s lineage.     

A cladistic analysis of the nomadine bees (Hymenoptera: Apoidea)

Abstract. This study compares the results of Rozen's cladistic analysis of the larvae of fifteen genera of cleptoparasitic bees in the subfamily Nomadinae with an independent data set of adult characters for the same genera. Adult characters exhibited considerably higher levels of homoplasy and poorer resolution of cladistic relationships, with multiple equally parsimonious cladograms. However, comparison of a Nelson consensus tree based on adult characters with the cladogram based on larval characters reveals three components consistently supported in both analyses (the tribes Epeolini and Ammobatini, and Neopasites + Neolarra) , one component supported only by adult characters (Isepeolus + Protepeolus) , and one terminal component supported only by larval characters (Nomada + Ammobatini), as well as several more inclusive groupings based on larval characters that are difficult to compare with the adult consensus tree because it shows so much less resolution. When adult and larval characters are combined in a single data matrix, the resulting cladogram closely resembles the cladogram based on larval characters alone, although levels of homoplasy are considerably higher than in the larval analysis.
A preliminary analysis of adult characters for thirty-four genera in the Nomadinae also exhibited high levels of homoplasy and very large numbers of equally parsimonious cladograms. Nevertheless, certain consistent monophyletic groupings, most notably the Epeolini and Ammobatini, were also supported in this analysis. The one currently recognized tribe whose monophyly has received no support from any analysis is the Nomadini.
The relevance of these phylogenetic hypotheses to our understanding of host associations and variable features of egg morphology and oviposition behaviour in nomadine bees is briefly discussed.     

Molecular phylogeny of the brachyuran crab superfamily Majoidea indicates close congruence with trees based on larval morphology

In this study, we constructed the first molecular phylogeny of the diverse crab superfamily Majoidea (Decapoda: Pleocyemata: Brachyura), using three loci (16S, COI, and 28S) from 37 majoid species. We used this molecular phylogeny to evaluate evidence for phylogenetic hypotheses based on larval and adult morphology. Our study supports several relationships predicted from larval morphology. These include a monophyletic Oregoniidae family branching close to the base of the tree; a close phylogenetic association among the Epialtidae, Pisidae, Tychidae, and Mithracidae families; and some support for the monophyly of the Inachidae and Majidae families. However, not all majoid families were monophyletic in our molecular tree, providing weaker support for phylogenetic hypotheses inferred strictly from adult morphology (i.e., monophyly of individual families). This suggests the adult morphological characters traditionally used to classify majoids into different families may be subject to convergence. Furthermore, trees constructed with data from any single locus were more poorly resolved than trees constructed from the combined dataset, suggesting that utilization of multiple loci are necessary to reconstruct relationships in this group.     

Systematic analysis of the Diplostomidae and Strigeidae (Trematoda)

A systematic analysis of the genera in the Diplostomidae and Strigeidae was made using the Proterodiplostomidae as the outgroup. The Proterodiplostomidae was the family with the greatest preponderance of primitive characters and its monophyly was supported by the unique paraprostate gland. However, no character state supported the monophyly of the Diplostomidae sensu Dubois, 1970. That paraphyletic taxon was composed of 3 monophyletic groups: the pseudosuckerless Neodiplostomidae n. fam. had the most primitive character states of the 3 and its monophyly was based on characters in the neascus/neodiplostomulum metacercariae; the Bolbophoridae n. fam., with pseudosuckers, had its monophyly supported by characters present in the newly named prodiplostomulum metacercaria; and the emended Diplostomidae, also with pseudosuckers, had the most derived states and its monophyly was supported by characters present in the diplostomulum. The presence of pseudosuckers united the Bolbophoridae n. fam., the emended Diplostomidae, and the Strigeidae as a monophyletic assemblage. The Strigeidae had the most derived characters of these 3 taxa and its monophyly was supported by characters in the tetracotyle and the cup-shaped forebody and bilobed tribocytic organ of the adult. In general, the adult stages of these strigeoid families showed very conserved morphology and it was the metacercariae that possessed the innovations. The conserved adult morphology was typical of what one might expect if the intramolluscan stages were analyzed. Thus, the data were concordant with the view that the mollusc and vertebrate definitive host were the original hosts to the Digenea and that the second intermediate host and metacercaria were more recently intercalated. More specifically, the phylogeny of these groups suggested that ancestral bisegmented strigeoids originally infected reptiles, they subsequently radiated into birds with which they coevolved extensively, and on 5 separate occasions they radiated into mammals. The radiations into mammals were, by all available evidence, preceded by second intermediate host shifts from fish to amphibians.     

Phylogeny of the Neotropical moth tribe Josiini (Notodontidae: Dioptinae): a hidden case of Miillerian mimicry

The Neotropical moth tribe Josiini (Notodontidae: Dioptinae) contains over 100 described species in 11 genera. All are diurnal, with brightly-coloured, presumably aposematic wing patterns. Larval hostplants are exclusively in the genus Passiflora (Passifloraceae) except for two new records, reported here, from Tumera (Turneraceae). A comparative morphological study of 26 representative josiine species yielded 86 characters from adults, larvae and pupae, all of which are figured and discussed. Phylogenetic analysis of these data produced a single most-parsimonious cladogram. According to the phylogenetic results: (1) monophyly of the Josiini is strongly supported; (2) the currently accepted generic classification is in disarray; (3) morphological character variation is extensive, and adult traits reflect phylogeny more effectively than do those of immature stages; (4) wing pattern types have undergone convergent evolution. A rare phenotype, longitudinal wing stripes, appears in two widely divergent clades, suggesting the evolution of Miillerian mimicry within the Josiini.     

Phylogeny of the tachyporine group subfamilies and 'basal' lineages of the Aleocharinae (Coleoptera: Staphylinidae) based on larval and adult characteristics

Abstract. This paper reports the conclusions of studies into the phylogeny of tachyporine group subfamilies and the ‘basal’ lineages of the subfamily Aleocharinae (Coleoptera: Staphylinidae) based on both larval and adult morphological data (133 adult characters, twenty-seven larval characters). Representatives of forty species of the tachyporine group were used in the analysis, including representatives of the Aleocharinae, Trichophyinae, Habrocerinae, Phloeocharinae, Olisthaerinae, and Tachyporinae. The Aleocharinae included representatives of the tribes Gymnusini, Deinopsini, Mesoporini, the ‘subfamily’ Trichopseniinae, and representatives of nine major tribes in the ‘higher’ Aleocharinae (Athetini, Hoplandriini, Falagriini, Lomechisini, Oxypodini, Aleocharini, Myllaenini, Homalotini, and Hypocyphtini). Analyses were performed first with adult characters alone and then with both larval and adult characters in a simultaneous analysis. The analysis based on adult characters produced eighty-five equally parsimonious trees (length = 499, consistency index = 42; retention index = 69). In the consensus tree, the Tachyporinae are not monophyletic, and the sister-group relationship between the Trichophyinae + Habrocerinae and the Aleocharinae is not resolved. The Aleocharinae are monophyletic, but, among the ‘basal’ Aleocharinae, the relationships of Gymnusini + Deinopsini, the Mesoporini, and the Trichopseniinae are unresolved. The combined adult and larval data, using Tachinus as the outgroup, produced six equally parsimonious trees (tree length = 588; consistency index = 43; retention index = 69). The strict consensus tree of the combined larval and adult data supports the following conclusions: (1) larval characters substantially stabilize the tree; (2) the subfamily Tachyporinae is not supported to be monophyletic; (3) the subfamilies Trichophyinae and Habrocerinae are sister groups, and together they are sister to the Aleocharinae; (4) the ‘basal’ Aleocharinae are not a monophyletic group, but the ‘higher’ Aleocharinae are monophyletic; (5) the sister group of the remaining Aleocharinae is a lineage made up of genera currently in the tribes Gymnusini and Deinopsini; (6) within the Gymnusini–Deinopsini lineage, the monophyly of the Gymnusini is weakly supported, but the monophyly of the Deinopsini is strongly supported; (7) the subfamily Trichopseniinae is strongly supported to be a member of the ‘basal’ Aleocharinae; (8) the Myllaenini are resolved well within the ‘higher’ Aleocharinae; (9) strong support for the monophyly of some tribes of ‘higher’ Aleocharinae suggests that morphological characters provide substantial phylogenetic signal for analysis of higher-level phylogeny of the Aleocharinae in spite of the preliminary nature of the analysis at this taxonomic level.     

Molecular phylogeny of the moonseed family (Menispermaceae): implications for morphological diversification

We used the chloroplast gene ndhF to reconstruct the phylogeny of the moonseed family (Menispermaceae), a morphologically diverse and poorly known cosmopolitan family of dioecious, primarily climbing plants. This study includes a worldwide sample of DNA sequences for 88 species representing 49 of the 70 genera of all eight traditionally recognized tribes. Phylogenetic relationships were estimated, and the Shimodaira-Hasegawa test was used to compare the likelihood of alternative phylogenetic hypotheses and to evaluate the monophyly of tribes currently in use. The monospecific Indo-Malesian Tinomiscium is sister to the remaining members of the family, within which are two major clades. Within these two clades, well-supported clades correspond to four of the eight traditionally recognized tribes, while others, such as Menispermeae, are polyphyletic. Mapping of major morphological characters on the phylogeny indicates that the crescent-shaped seed is derived from a straight seed, the tree habit has arisen multiple times, endosperm has been lost many times, but unicarpellate flowers evolved only once. Morphological synapomorphies for Menispermaceae include the presence of a condyle, a large embryo, and druplets. The phylogeny provides for the first time a detailed molecular-based assessment of relationships in Menispermaceae and clarifies our understanding of morphological diversification within the family.     

Phylogeny of the Nymphalidae (Lepidoptera)

A generic-level phylogeny for the butterfly family Nymphalidae was produced by cladistic analysis of 234 characters from all life stages. The 95 species in the matrix (selected from the 213 studied) represent all important recognized lineages within this family. The analysis showed the taxa grouping into six main lineages. The basal branch is the Libytheinae, with the Danainae and Ithomiinae on the next branch. The remaining lineages are grouped into two main branches: the Heliconiinae-Nymphalinae, primarily flower-visitors (but including the fruit-attracted Coeini); and the Limenitidinae (sensu strictu), Biblidinae, and the satyroid lineage (Apaturinae, Charaxinae, Biinae, Calinaginae, Morphinae, Brassolinae, and Satyrinae), primarily fruit-attracted. Data partitions showed that the two data sets (immatures and adults) are very different, and a partitioned Bremer support analysis showed that the adult characters are the main source of conflict in the nodes of the combined analysis tree. This phylogeny includes the widest taxon coverage of any morphological study on Nymphalid butterflies to date, and supports the monophyly and relationships of most presently recognized subgroups, providing strong evidence for the presently accepted phylogenetic scheme.     

Evaluating the monophyly and phylogenetic relationships of Brassolini genera (Lepidoptera, Nymphalidae)

Abstract.  This study uses 80 morphological characters and cladistic analysis to evaluate the monophyly and phylogenetic relationships of 18 genera that constitute the butterfly tribe Brassolini. Most characters derive from genitalia, confirming previous generic definitions based mainly on wing characters, and showing that 16 of 18 genera are monophyletic. Mimoblepia Casagrande, syn.n. was subsumed within Opoptera Aurivillius to address the paraphyly of the latter, but resolution of the status of Aponarope Casagrande requires further study. The results suggest that the Brassolini includes six suprageneric groups/clades. Although this study verifies some genus-level relationships put forward over 100 years ago, some new hypotheses of relationships are proposed. Tracing larval host plant use onto the Brassolini phylogeny indicates that species in this tribe retain the use of Arecaceae and Poaceae from their 'satyroid' ancestors.     

A phylogenetic study on genera of Cidariini from the Holarctic and the Indo-Australian areas (Lepidoptera: Geometridae: Larentiinae)

A cladistic analysis of forty-one species, belonging to ten genera, of the Cidariini sensu Herbulot from the Holarctic and the Indo-Australian areas, was performed using seventy-seven characters including larval and pupal data. Eight most parsimonious cladograms were found (length 398, CI 0.30, RI 0.70). The monophyly of the Cidariini is demonstrated, using selected species of Xanthorhoini sensu Herbulot as the outgroup. The relationships among the genera are as follows: ( Ecliptopera ( Eulithis ( Cidaria (( Plemyria ( Chloroclysta , Dysstroma ))(( Thera , Pennithera ) ( Heterothera ))))). This result suggests some taxonomic changes: Dysstroma Hübner, stat. rev. and Chloroclysta Hübner stat. rev. are sister taxa; Heterothera Inoue stat. rev. includes Viidaleppia Inoue, syn.n., Heterothera firmata (Hübner) comb.n. is transferred from Pennithera Viidalepp to Heterothera sensu lato . The results of confirmation and incongruence tests suggest that the characters from adult and immature stages exhibit the same evolutionary pattern. Thus the phylogeny derived from the combined data matrix does not give a misleading conclusion, even though there are many missing states in the larval data set.     

A phylogeny of extant penguins (Aves: Sphenisciformes) combining morphology and mitochondrial sequences

The phylogenetic relationships among the penguins have received little attention, despite their well‐known anatomy and the conspicuous nature of the group. Previous attempts have included datasets limited to few, mostly osteological characters, and one study was based on integumentary and breeding characters. We developed a morphological matrix comprising 159 morphological characters of osteology (70 characters), myology (15), digestive tract (1), integument (66), and breeding (7 characters), scored in 18 extant forms (all currently recognized species plus one distinct subspecies). A gaviiform was placed at the root, and 11 species of representative procellariiform groups completed the outgroup. A heuristic parsimony analysis under equal weights was performed. We also compiled DNA sequences available in GenBank for the mitochondrial genes 12S rDNA and cytochrome b. We included the two data partitions in a combined analysis under direct optimization. Both analyses recovered the monophyly of Sphenisciformes and all the traditional polytypic genera. Morphological characters performed optimally at the ordinal and generic nodes, also providing resolution and varying degrees of support at supra‐ and intrageneric nodes. The comparison of molecular and morphological results indicated that the most significant problem in the phylogeny of extant penguins is rooting the ingroup. The mutual interaction of molecular and morphological data decreases the ambiguity regarding the placement of the root, and provides a resolved, relatively well‐supported phylogeny of extant penguins. Biogeographical patterns based on breeding ranges and derived from the combined analysis show that the major intercontinental vicariance events detected are consistent with cold marine current patterns of the Southern Hemisphere. © The Willi Hennig Society 2005.     

Phylogenetic analysis of the family Ariidae (Ostariophysi: Siluriformes), with a hypothesis on the monophyly and relationships of the genera

Ariid monophyly and intrafamilial relationships are investigated based on cladistic analysis of 230 morphological characters. Terminal taxa examined include whenever possible type‐species, or the most morphologically similar species to the type‐species of the nominal genera, and the largest possible number of species, including cleared and stained specimens, available in zoological collections. Previous hypotheses about monophyly of the Ariidae are strongly corroborated by new synapomorphies discovered in the present study. The subfamily Galeichthyinae and the remaining ariids are strongly supported by new morphological characters. The monotypic subfamily Bagreinae is recognized as the sister group to all nongaleichthyin ariids, supported by a large series of exclusive synapomorphies. A new concept of Ariinae is presented: the subfamily is found to be unequivocally monophyletic and includes all ariid genera, except Galeichthys and Bagre. New data supporting the monophyly of the genera included in the Ariinae are introduced and previous hypotheses of monophyly, species composition, morphological definition, and relationships are reviewed and discussed.     

Phylogenetic reassessment of the five forewing radial-veined tribes of Riodininae (Lepidoptera: Riodinidae)

The relationships among the genera and tribal groupings of Riodininae with five forewing radial veins, and between these and tribes with four forewing radial veins, were examined using a phylogenetic analysis. Using the type species from all sixteen genera in the tribal groupings Eurybiini, Mesosemiini and incertae sedis (a presumed paraphyletic group of loosely related genera), and representatives from the four forewing radial‐veined riodinine tribes, thirty‐five new and traditional characters were coded from adult ecology, wing venation and pattern, the adult head and body, male and female genitalia, and early stage ecology and morphology. The majority of characters are illustrated. Phylogenetic analysis of these data produced five equally most parsimonious cladograms using equal weights and after successive weighting. The strict consensus of these confirms the monophyly of Eurybiini and Mesosemiini as currently conceived, but also indicates several higher‐level relationships not previously hypothesized. Mesosemiini is here more broadly defined to also include the entire incertae sedis section, and the tribe is divided into Mesosemiina, for the previously delimited Mesosemiini plus Eunogyra and Teratophthalma, and Napaeina, subtr.n. for the incertae sedis section minus these two genera. The following hypothesis of relationships is tentatively proposed for the basal clades of Riodininae: Mesosemiini + (Eurybiini + remainder of Riodininae). These new hypotheses, and the characters supporting them, are discussed and compared with those previously proposed.     

Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters

In a previous study of the phylogeny of basal Hymenoptera, Vilhelmsen (2001; Zool. J. Linn. Soc . 131 : 393–442) compiled an extensive morphological data matrix for a phylogenetic analysis of basal Hymenoptera, comprising 38 hymenopteran genera. In this study, his characters are revised. This results in a cladogram whose relationships largely agree with those proposed by Vilhelmsen, except that the relationships at the base of the Hymenoptera are unresolved. The revised data matrix is expanded by 17 sawfly and three apocritan taxa. Moreover, 112 new morphological characters from different parts of the larval and adult morphology are also added to the data matrix, including 82 from a recent study of the terminal abdominal segments of male Hymenoptera. The addition of the new characters leads to Xyelidae, again, being the sister-group of all other Hymenoptera. The relationships among the sawfly families as proposed by Vilhelmsen are confirmed, except that the relationships among Syntexis , Siricidae and Xiphydriidae + Vespina are unresolved and that the monophyly of Apocrita is not convincingly supported. A separate analysis is performed which includes all extant genera of Xyelidae. The internal phylogeny of Xyelidae is determined as (( Macroxyela Megaxyela ) Xyelecia ( Xyela Pleroneura )).  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 209–243.     

The osteology and phylogeny of the Hawaiian finch radiation (Fringillidae: Drepanidini), including extinct taxa

The monophyly and phylogeny of the adaptive radiation of Hawaiian finches (Fringillidae: Drepanidini; honeycreepers, auct.) were studied using parsimony analysis of comparative osteology, combined with Templeton (Wilcoxon signed‐ranks) tests of alternative phylogenetic hypotheses. Eighty‐four osteological characters were scored in 59 terminal taxa of drepanidines, including 24 fossil forms, and in 30 outgroup species. The optimal phylogenetic trees show considerable agreement, and some conflict, with independently derived ideas about drepanidine evolution. The monophyly of a large Hawaiian radiation was upheld, although one fossil taxon from Maui fell outside the drepanidine clade. The finch‐billed species were placed as basal drepanidine taxa, and continental cardueline finches (Carduelini) were identified as the radiation's closest outgroups. The study found anatomical as well as phylogenetic evidence that the radiation had a finch‐billed ancestor. The optimal trees identify the red‐and‐black plumage group as a clade, and suggest that the tubular tongue evolved only once in the radiation. Because comparative osteology provides too few characters to strongly support all the nodes of the tree, it was helpful to evaluate statistical support for alternative hypotheses about drepanidine relationships using the Templeton test. Among the alternatives that received significant statistical support are a relationship of the drepanidines with cardueline finches rather than with the Neotropical honeycreepers (Thraupini), classification of the controversial genera Paroreomyza and Melamprosops as drepanidines, and a secondary loss of the tubular tongue in Loxops mana. The hypothesis of monophyly for all the Hawaiian taxa in the study was not rejected statistically. The study provides a framework for incorporating morphological and palaeontological information in evolutionary studies of the Drepanidini. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141 , 207–255.     

Monophyly of Euaesthetinae (Coleoptera: Staphylinidae): phylogenetic evidence from adults and larvae,review of austral genera,and new larval descriptions

Abstract We develop a morphological dataset for the rove beetle subfamily Euaesthetinae comprising 167 morphological characters (135 adult and 32 larval) scored from 30 terminal taxa including 25 ingroup terminals (from subfamilies Euaesthetinae and Steninae) and five outgroups. Four maximum parsimony analyses using different sets of terminals and character sets were run to test the monophyly of (1) Euaesthetinae, (2) Steninae, (3) Euaesthetinae + Steninae, (4) euaesthetine tribes Austroesthetini, Alzadaesthetini, Euaesthetini, Fenderiini and Stenaesthetini, and (5) the ten currently known austral endemic genera together. Analyses of adult and larval character sets separately and in combination recovered the monophyly of Euaesthetinae, Steninae, and both subfamilies together, with strong support. Analysis of 13 ingroup terminals for which complete data were available suggests that monophyly of Euaesthetinae is supported by 19 synapomorphies (13 adult, six larval), of Steninae by 23 synapomorphies (14 adult, nine larval), and of both subfamilies together by 24 synapomorphies (21 adult, three larval). Within Euaesthetinae, only the tribe Stenaesthetini was recovered as monophyletic based on adult characters, and in no analyses were the ten austral endemic genera recovered as a monophyletic group. Phylogenetic relationships among euaesthetine genera were weakly supported, although analyses including adult characters supported monophyly of Octavius and Protopristus separately, and of Octavius + Protopristus, Austroesthetus + Chilioesthetus and Edaphus + Euaesthetus. Steninae may include a third genus comprising two undescribed species probably possessing a ‘stick–capture’ method of prey capture, similar to that in Stenus. These two species formed a strongly supported clade recovered as the sister group of Stenus based on adult characters. Diagnoses and a key to adults are provided for the 15 euaesthetine genera currently known from the austral region (Australia, New Zealand, South Africa and southern South America). Euaesthetine larvae previously were known only for Euaesthetus, and we describe the larvae of nine more genera and provide the first larval identification key for genera of Euaesthetinae.     

Phylogeny and classification of the Culicidae (Diptera)

The generic relationships and higher classification of the family Culicidae are examined on the basis of a phylogenetic analysis. New and traditional morphological characters studied and compared throughout the Culicidae resulted in the acquisition of character data relative to the majority of species within each genus. Polymorphisms and morphological observations are discussed and additional information and illustrations are provided for the majority of characters and their character states. The analysis of seventy-three adult, pupal and fourth-instar larval characters coded for the thirty-eight currently recognized genera of mosquitoes resulted in relationships and groupings which differ significantly from traditional hypotheses. The analysis supports the monophyly of the subfamily Anophelinae and the tribes Culicini and Sabethini. The Anophelinae form the most basal clade of the family. The results indicate that Aedini is a paraphyletic assemblage with respect to the Mansoniini, each of which is monophyletic in itself. The Aedini + Mansoniini form a sister group to the Culicini + Sabethini, with the Aedini and the Culicini placed in ancestral relationships to the Mansoniini and the Sabethini, respectively. Based on the topography of generic relationships among more 'generalized' mosquitoes, the boundaries and relationships of the tribes Aedeomyiini, Uranotaeniini, Ficalbiini, Hodgesiini, Orthopodomyiini and Culisetini appear to be problematic. Relationships between genera of the tribe Aedini are generally poorly resolved due to a significant amount of polymorphism, especially within the genus Aedes as currently defined. There is no support for separate subfamily recognition for the genus Toxorhynchites , which is downgraded to tribal status as a result of the analysis. The results are discussed in relation to previous hypotheses based on subjective inference and cladistic analyses.     

Phylogenetic relationships of Central European wolf spiders (Araneae: lycosidae) inferred from 12S ribosomal DNA sequences

We have analyzed a sequence dataset of a portion of mitochondrial 12S rRNA gene of the ribosomal small subunit for 27 species of the family Lycosidae (wolf spiders) from Central Europe, belonging to six genera (Alopecosa, Arctosa, Pardosa, Pirata, Trochosa, and Xerolycosa) and four subfamilies (Evippinae, Lycosinae, Pardosinae and Venoniinae). Phylogenetic analyses were performed in two steps and corroborate the monophyly of all the genera analyzed with strong bootstrap support. In the first step focusing on the most ancestral splits the genus Pirata consistently emerged as the most ancestral branch, followed by the two genera Arctosa and Xerolycosa, with conflicting branching order, however. The second step of analysis placed Xerolycosa more ancestral than Arctosa. Arctosa appeared as sister group to the genera Alopecosa, Trochosa, and Pardosa. The palearctic genus Xerolycosa was not yet included in previous studies derived from morphological characters, but its placement based on mtDNA sequences is in good agreement to that according to current diagnostic morphological features. Further, the single representative of the genus Arctosa examined in our study was placed at a more ancestral position than in a previous investigation based on phenotypic characters. The superimposition of the currently used diagnostic phenotypic characters on the DNA-based phylogeny shows that both character sets are widely congruent; only 3 out of 16 phenotypic characters were resolved as homoplasious, suggesting their parallel evolution and/or reversal. Among the three different styles of predation found in the Lycosids, tube builders appear to be the most ancestral from which burrow dwellers descended and from which two groups of vagrant hunters evolved in parallel.     

Afrochlus Freeman: an African gondwanan midge and the phylogeny of the Podonominae (Diptera: Chironomidae)

The immature stages and male adult of Afrochlus harrisoni Freeman are described for the first time, from the type locality, a granite outcrop in Zimbabwe. A phylogenetic position within the subfamily Podonominae as sister to South African-western/central Australian genus Archaeochlus Brundin, suggested in 1966 by Brundin from the adult female alone, is confirmed by cladistic analysis of all life history stages combined. Within the Podonominae, the tribes Boreochlini and Podonomini erected by Brundin are monophyletic only if Trichotanypus Kieffer is transferred from the former to the latter. Little or none of the structure provided by the combined evidence from all stages is retrieved by cladistic analyses of characters derived solely from the adult stage. The implications of erection of monotypic genera for adult-only taxa are discussed.     

Testing the monophyly of the New Zealand and Australian endemic family Conoesucidae Ross based on combined molecular and morphological data (Insecta: Trichoptera: Sericostomatoidea)

Conoesucidae (Trichoptera, Insecta) are restricted to SE Australia, Tasmania and New Zealand. The family includes 42 described species in 12 genera, and each genus is endemic to either New Zealand or Australia. Although monophyly has been previously assumed, no morphological characters have been proposed to represent synapomorphies for the group. We collected molecular data from two mitochondrial genes (16S and cytochrome oxidase I), one nuclear gene (elongation factor 1-α) (2237–2277 bp in total), and 12 morphological characters to produce the first phylogeny of the family. We combined the molecular and morphological characters and performed both a maximum parsimony analysis and a Bayesian analysis to test the monophyly of the family, and to hypothesize the phylogeny among its genera. The parsimony analysis revealed a single most parsimonious tree with Conoesucidae being a monophyletic taxon and sistergroup to the Calocidae. The Bayesian inference produced a distribution of trees, the consensus of which is supported with posterior probabilities of 100% for 15 out of 22 possible ingroup clades including the most basal branch of the family, indicating strong support for a monophyletic Conoesucidae. The most parsimonious tree and the tree from the Bayesian analysis were identical except that the ingroup genus Pycnocentria changed position by jumping to a neighbouring clade. Based on the assumption that the ancestral conoesucid species was present on both New Zealand and Australia, a biogeographical analysis using the dispersal-vicariance criteria demonstrated that one or two (depending on which of the two phylogenetic reconstructions were applied) sympatric speciation events took place on New Zealand prior to a single, late dispersal from New Zealand to Australia.