Showing papers in "Systematic Entomology in 2003"

Phylogenetic analysis of higher‐level relationships of Odonata

Abstract: . This is the most comprehensive analysis of higher-level relationships in Odonata conducted thus far. The analysis was based on a detailed study of the skeletal morphology and wing venation of adults, complemented with a few larval characters, resulting in 122 phylogenetically informative characters. Eighty-five genera from forty-five currently recognized families and subfamilies were examined. In most cases, several species were chosen to serve as exemplars for a given genus. The seven fossil outgroup taxa included were exemplar genera from five successively more distant odonatoid orders and suborders: Tarsophlebiidae (the closest sister group of Odonata, previously placed as a family within ‘Anisozygoptera’), Archizygoptera, Protanisoptera, Protodonata and Geroptera. Parsimony analysis of the data, in which characters were treated both under equal weights and implied weighting, produced cladograms that were highly congruent, and in spite of considerable homoplasy in the odonate data, many groupings in the most parsimonious cladograms were well supported in all analyses, as indicated by Bremer support. The analyses supported the monophyly of both Anisoptera and Zygoptera, contrary to the well known hypothesis of zygopteran paraphyly. Within Zygoptera, two large sister clades were indicated, one comprised of the classical (Selysian) Calopterygoidea, except that Amphipterygidae, which have traditionally been placed as a calopterygoid family, nested within the other large zygopteran clade comprised of Fraser's ‘Lestinoidea’ plus ‘Coenagrionoidea’ (both of which were shown to be paraphyletic as currently defined). Philoganga alone appeared as the sister group to the rest of the Zygoptera in unweighted cladograms, whereas Philoganga + Diphlebia comprised the sister group to the remaining Zygoptera in all weighted cladograms. ‘Anisozygoptera’ was confirmed as a paraphyletic assemblage that forms a ‘grade’ towards the true Anisoptera, with Epiophlebia as the most basal taxon. Within Anisoptera, Petaluridae appeared as the sister group to other dragonflies.

Phylogeny of Syrphidae (Diptera) inferred from combined analysis of molecular and morphological characters

Abstract: Syrphidae (Diptera) commonly called hoverflies, includes more than 5000 species world-wide. The aim of this study was to address the systematic position of the disputed elements in the intrafamilial classification of Syrphidae, namely the monophyly of Eristalinae and the placement of Microdontini and Pipizini, as well as the position of particular genera (Nausigaster, Alipumilio, Spheginobaccha). Sequence data from nuclear 28S rRNA and mitochondrial COI genes in conjunction with larval and adult morphological characters of fifty-one syrphid taxa were analysed using optimization alignment to explore phylogenetic relationships among included taxa. A species of Platypezidae, Agathomyia unicolor, was used as outgroup, and also including one representative (Jassidophaga villosa) of the sister-group of Syrphidae, Pipunculidae. Sensitivity of the data was assessed under six different parameter values. A stability tree sum- marized the results. Microdontini, including Spheginobaccha, was placed basally, and Pipizini appeared as the sister-group to subfamily Syrphinae. The monophyly of subfamily Eristalinae was supported. The results support at least two independ- ent origins of entomophagy in syrphids, and frequent shifts between larval feeding habitats within the saprophagous eristalines.

Wing geometry in Triatoma infestans (Klug) and T. melanosoma Martinez, Olmedo & Carcavallo (Hemiptera: Reduviidae)

Abstract: . Geometric morphometrics is a novel approach to biological shape analysis, and its application to medical entomology is just beginning. Here, we use it to examine geographical and interspecific variation in the Triatoma infestans complex, vectors of Chagas disease in southern Latin America. Using six landmarks defining the membranous part of the hemelytra, we analysed the geographical patterns of variation in several populations of T. infestans, including the recently described ‘dark morph’ of T. infestans from hollow trees in the Bolivian Chaco. As a potential outgroup, T. melanosoma was added to the sample. A consistent geographical differentiation was observed, but no evidence was found in either sex to consider either the dark morph or T. melanosoma as a species distinct from T. infestans. Triatoma melanosoma is relegated to synonymy with T. infestans (syn.n.).

A new genus and species of bothriurid scorpion from the Brandberg Massif, Namibia, with a reanalysis of bothriurid phylogeny and a discussion of the phylogenetic position of Lisposoma Lawrence

Abstract: .Brandbergia haringtoni, a new genus and species of bothriurid scorpion, is described from the Brandberg Massif, northwestern Namibia. A cladistic analysis, based on seventy-four morphological characters scored for thirty-one exemplar species representing all genera of Bothriuridae, and one genus from each of the six remaining families of Scorpionoidea, was conducted to test the phylogenetic placement of the new genus and whether it affects the internal relationships of Bothriuridae. The available data demonstrate, under a range of weighting regimes, that the new genus is the most basal bothriurid, and confirm the scheme of relationships among the remaining bothriurid genera that was recovered in a previous analysis: (Brandbergia (Lisposoma (Thestylus (Phoniocercus (Tehuankea (Cercophonius + Urophonius) (Bothriurus + Brachistosternus +Orobothriurus + Centromachetes (Timogenes + Vachonia))))))). On the basis of this evidence, Lourenco's recent proposal of family Lisposomidae for Lisposoma is rejected and Lisposomidae is synonymized with Bothriuridae. The implications of the phylogeny for understanding the biogeography of Bothriuridae are discussed.

Phylogenetic relationships of Phyciodes butterfly species (Lepidoptera: Nymphalidae): complex mtDNA variation and species delimitations

Abstract: . Mitochondrial DNA variation was studied in the butterfly genus Phyciodes (Lepidoptera: Nymphalidae) by sequencing 1450 bp of the COI gene from 140 individuals of all eleven currently recognized species. The study focused on four species in particular that have been taxonomically difficult for the past century, P. tharos, P. cocyta, P. batesii and P. pulchella. A cladistic analysis of ninety-eight unique haplotypes showed that Phyciodes forms a monophyletic group with P. graphica as the most basal species. Of the three informal species groups described for Phyciodes, only one (the mylitta-group) is unambiguously monophyletic. Within the tharos-group, seven well supported clades were found that correspond to three taxa, P. tharos, P. pulchella and a grade consisting of P. cocyta and P. batesii haplotypes interdigitated with each other. None of the clades is formed exclusively by one species. The patterns of haplotype variation are the result of both retained ancient polymorphism and introgression. Introgression appears to be most common between P. cocyta and P. batesii; however, these two species occur sympatrically and are morphologically and ecologically distinct, suggesting that the level of current introgression does not seem to be enough to threaten their genetic integrity. The results indicate that mitochondrial DNA sequences must be used with great caution in delimiting species, especially when infraspecific samples are few, or introgression seems to be rampant.

Phylogeny of Habronattus jumping spiders (Araneae: Salticidae), with consideration of genital and courtship evolution

Abstract: . DNA sequences from the mitochondrial (including ND1, 16S) and nuclear (EF-1α) genomes of about ninety-four species were obtained to reconstruct phylogenetic relationships of Habronattus jumping spiders. Maximum parsimony trees were sought with both separate (mitochondrial, nuclear) and combined analyses; maximum likelihood trees were sought with both separate (ND1, 16S, EF-1α introns, EF-1α exons) and combined (mitochondrial, nuclear) analyses. All analyses agreed on some fundamental aspects of the tree, including the monophyly of the previously recognized agilis, amicus, dorotheae and americanus species groups. The deep phylogenetic structure is well resolved, placing the agilis, amicus, tranquillus and dorotheae groups basally. Several other previously unrecognized clades were well supported, including a newly formulated decorus group. The large group of species with modified male first and third legs was supported as monophyletic except for the surprising placement elsewhere of three species of the group. The phenotypic similarities between these three and the others are so detailed and precise that convergence in ornamentation can probably be ruled out. There are hints of phylogenetically distant genetic introgression involving the coecatus group. The combination Habronattus paratus is restored based on the species falling within Habronattus. Regarding patterns of character evolution, there was consistent support for the basal placement of several species groups with a long embolus, suggesting that there were more evolutionary reductions in embolus length than postulated in a previous morphological phylogeny. This is in accord with the expectation that there is a bias to an overly conservative interpretation of a character's evolution if it is interpreted on a phylogeny based in part on that same character. In contrast, the molecular phylogeny did not suggest any instances of the evolutionary transformation of one complex style of courtship into another, a possibility that could have been difficult to detect using the morphological phylogeny because of the same bias to conservativism.

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

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

Morphological and molecular characterization anopheles (anopheles) sacharovi favre, a primary vector of malaria in the middle east

Abstract: . Combined morphological and molecular studies were conducted on Anopheles (Anopheles) sacharovi Favre, an important malaria vector of the Palaearctic Maculipennis Complex. Specimens collected in Greece and Iran were identified on the basis of morphology and confirmed by correlation of their nuclear internal transcribed spacer 2 (ITS2) rDNA sequences with those available in GenBank. The progeny of females collected in Greece were used for detailed morphological study. The morphology of the adults, pupa, fourth-instar larva and egg are described and distinguished from those of An. maculipennis, the nominotypical member of the Maculipennis Complex. In addition to sequence data for the nuclear ITS2 region, partial sequence data are provided for the mitochondrial cytochrome oxidase I gene. The taxonomy, systematics, bionomics and distribution of the species are reviewed. This work provides the first fully integrated assessment of An. sacharovi as a basis for comparative studies of the Maculipennis Complex.

Cladistic analysis of the Neotropical butterfly genus Adelpha (Lepidoptera: Nymphalidae), with comments on the subtribal classification of Limenitidini

Abstract: . A two-stage cladistic analysis of 114 characters from adult and immature stage morphology provided phylogenetic hypotheses for the diverse Neotropical nymphalid butterfly genus Adelpha Hubner. Higher-level cladistic relationships were inferred for thirty Adelpha species and twenty other species of Limenitidini, confirming the monophyly of Adelpha as currently conceived and indicating several montane Asian species as potential sister taxa for the genus. Cladistic relationships between all eighty-five Adelpha species were then inferred using three outgroup combinations. Basal and terminal nodes were reasonably resolved and supported, but a low proportion of non-wing pattern characters resulted in weak resolution and support in the middle of the tree. The most basal members of Adelpha feed on the temperate or montane plant family Caprifoliaceae, suggesting that a switch from this family early in the evolutionary history was important in subsequent diversification into tropical lowland habitats. The cladograms confirm suspicions of earlier authors that dorsal mimetic wing patterns have convergently evolved a number of times in Adelpha. The subtribal classification of Limenitidini is discussed and both Lebadea (from Parthenina) and Neptina are transferred to Limenitidina, whereas Cymothoe, Bhagadatta and Pseudoneptis (all formerly Limenitidina) are regarded as incertae sedis.

Morphological and molecular characterization of Anopheles (Anopheles) maculipennis Meigen, type species of the genus and nominotypical member of the Maculipennis Complex

Abstract: . Adults and larvae of Anopheles maculipennis Meigen were collected in Greece in May and June 2001. Larvae and the progeny of wild-caught females were individually reared to obtain samples of all life stages for integrated morphological and molecular study. Specimens were identified on the basis of egg morphology and correlation of their ITS2 rDNA sequences with those in GenBank. The adult, pupal, larval (fourth-instar) and egg stages are described, and all stages except the egg are illustrated. Partial sequence data are provided for the mitochondrial cytochrome c oxidase I (COI) gene and complete sequences for the nuclear ITS2 region. Additionally, the bionomics and distribution of the species are reviewed, and its taxonomy and systematics are discussed. This study provides the first fully integrated morphological and molecular assessment of An. maculipennis, the nominotypical member of the Maculipennis Complex, and the type species of genus Anopheles, and serves as a foundation for further studies of the complex and subfamily Anophelinae.

Specificity of cicada calling songs in the genus Tibicina (Hemiptera: Cicadidae)

Abstract: Males of Tibicina cicada species produce a sustained and monotonous calling song by tymbal activity. This acoustic signal constitutes the first step in pair formation, attracting females at long range, and is involved in male-male interactions. The specificity of this signal was investigated for the first time for seven species and one subspecies of Tibicina occurring in France. This analysis was achieved by describing tymbal anatomy, tymbal mechanism and calling song structure. Male calling songs are emitted following the same general scheme: tymbals are activated alternately and the successive buckling of the sclerotized ribs that they bear produces a regular succession of groups of pulses. The struc- tural and mechanical properties shared by Tibicina species and subspecies lead to a considerable uniformity of the signal shape. Nevertheless, a principal component analysis applied to eight temporal and three frequency parameters revealed differ- ences between the signals of the species studied. In particular, calling songs differed in groups of pulse rate and/or in peak of the second frequency band (carrier frequency). These acoustic differences are probably linked to differences in the numbers of tymbal ribs and body size. Groups of pulse rate and/or peak of the second frequency band could encode specific information. However, Tibicina calling songs may not act as distinct specific-mate recognition systems and may not play a leading role in the mating isolation process; rather, they might merely belong to a complex set of specific spatial, ecological, ethological and morpho- logical characters that ensure syngamy.

Molecular systematics of Psocomorpha (Psocoptera)

Abstract: Previous classification of the insect order Psocoptera has relied on morphological characters Psocoptera are generallydivided into three suborders: Trogiomorpha, Troctomorpha, and Psocomorpha Traditional classification divides the Psocomorpha into four infraorders (Homilopsocidea, Caeciliusetae, Psocetae and Epipsocetae), but a recent morphological cladistic studyremoved Archipsocidae from Homilopsocidea and Hemipsocidae from Psocetae We investigated the phylogenetic relationships within the suborder Psocomorpha using DNA sequences from the nuclear 18S and mitochondrial 16S, 12S and cytochrome oxidase I genes Phylogenetic analyses of these gene sequences supported monophyly for Psocomorpha In addition, monophyly of the trad- itional subgroups Caeciliusetae and Psocetae was generallysupported Monophyly of Homilopsocidea was not supported, and Archipsocus is removed from this group Although the molecular phylogeny is generally consistent with recent cladistic studies of morphological characters, we found no evidence that Hemi- psocidae should be removed from Psocetae

Molecular phylogenetics of the Oriental members of the Myzomyia Series of Anopheles subgenus Cellia (Diptera: Culicidae) inferred from nuclear and mitochondrial DNA sequences

Abstract: . The phylogenetic relationships of fifteen Oriental and two Afrotropical taxa of the Myzomyia Series of Anopheles subgenus Cellia and two outgroup species, An. maculatus (Neocellia Series) and An. dirus A (Neomyzomyia Series), were inferred from nucleotide sequences of the entire 685 bp of the mitochondrial cytochrome oxidase subunit II locus (COII) and 372 bp of the third domain (D3) of the 28S rDNA locus, both separately and together. Alignment of the D3 sequences was achieved with the aid of secondary structure comparisons, and the pattern of nucleotide substitution was best explained by the GTR + I + G model for either separate or combined datasets. Maximum likelihood and maximum parsimony analyses robustly identified five monophylies: An. fluviatilis U and T; An. fluviatilis U and T + An. minimus A, C, E and #157 + An. leesoni; An. filipinae + An. mangyanus; An. filipinae + An. mangyanus + An. aconitus; and An. culicifacies A and B. The results confirm the specific status of An. flavirostris, the close relationship of An. leesoni with the Minimus Complex, and the exclusion of An. jeyporiensis, An. culicifacies s.l and An. funestus from the Minimus Group. All of the species classified as members of the Minimus Group on morphological grounds formed a single clade, which comprised two subgroups: the Minimus Subgroup, including An. minimus s.l., An. fluviatilis s.l., An. leesoni and An. flavirostris, and the Aconitus Subgroup, including An. filipinae, An. mangyanus, An. aconitus, An. pampanai and An. varuna. However, these clades are only weakly supported by the present dataset.

A lacewing with the wanderlust: the European song species ‘Maltese’, Chrysoperla agilis, sp.n., of the carnea group of Chrysoperla (Neuroptera: Chrysopidae)

Abstract: . A fifth European member of the Holarctic carnea sibling species group, Chrysoperla agilis, sp.n. is described. This species was initially recognized because of its unique song phenotype. Both sexes produce one type of song, characterized by a single very long volley averaging 4.5 s in duration and repeated every 9.5 s. The unit of exchange during heterosexual duets (the shortest repeated unit) is the single volley. The carrier frequency is upwardly, then downwardly, modulated during each volley. The maximum frequency is higher (around 110 Hz) than in other members of the carnea group. Each volley ends with an audible, percussive ‘rattle’. Song features vary only slightly over a large geographical range that extends from northern Iran westward to the Azores and northward to southern Switzerland. Discriminant function analysis confirms the uniqueness of the C. agilis song in comparison with nine other song species. Morphologically, the adult of C. agilis is very similar to C. carnea and C. pallida and is generally intermediate between them with respect to the size of the pretarsal basal dilation, the shape of the genital lip and chin, the colour and type of setae on the abdomen, and colour markings on the head and mouthparts. No single feature will distinguish C. agilis from either C. carnea or C. pallida, but a suite of physical attributes can be used with reasonable success. Larval morphology fails to distinguish clearly C. agilis from other song species. Ecologically, C. agilis shares an affinity for low vegetation with C. lucasina and C. carnea, and a preference for milder, Mediterranean climates with C. mediterranea. Adults undergo a colour change to pale yellowish brown during winter diapause, and eggs are laid singly rather than in groups. Based on its ecological characteristics, C. agilis may prove to be an effective agent of biological control in the agricultural fields of warmer, drier parts of southern Europe and the Middle East, where other members of the carnea group are less common or absent.

Review of competing hypotheses of phylogenetic relationships of Paussinae (Coleoptera: Carabidae) based on larval characters

Abstract: Larval structural characters are an important component in phylogenetic reconstruction of the subfamily Paussinae. Based on larval characters, two main cladistic hypotheses have been proposed concerning basal relationships within Paussinae: (1) Metriini as sister-group of Ozaenini + Paussini, with Ozaenini as a paraphyletic group and the genus Physea as sister-group of Paussini; and (2) Metriini as sister-group only of the monophyletic Ozaenini, with the Paussini as sister-group of Metriini + Ozaenini. We present here a review of these hypotheses and a new cladistic analysis based on larvae, emphasizing the phylogeny of Ozaenini. The analysis includes the following taxa: Metrius (Metriini), Itamus, Pachyteles, Sphaerostylus, Physea (Ozaenini), Platyrhopalopsis and Paussus (Paussini). For both Physea and Platyropalopsis larvae we provide new detailed descriptions, because previous descriptions are inadequate to allow an advanced phylogenetic discussion. A total of fifty-six characters are analysed. Most characters show alternative states organized into sequences of character states, each of which is logically derivable from its neighbour in the sequence, resulting in an almost completely pectinate cladogram. Metrius is the sister-group of all other Paussinae, and Paussini are a highly derived offshoot within the ozaenine grade. According to the present analysis, Physea, regarded traditionally as a typical ozaenine, represents the ozeanine sister to Paussini, showing that Ozaenini are paraphyletic. As our analysis is based on the few paussine genera sufficiently known at the larval stage, which represent a low number of the extant paussine genera, the results are merely illustrative of the evolution of the major clades within the subfamily. More detailed information about the phylogeny of ozaenines could arise from a more comprehensive analysis, mainly including adult morphological characters.

Evolution, mitochondrial DNA phylogeny and systematic position of the Macaronesian endemic Hydrotarsus Falkenstrom (Coleoptera: Dytiscidae)

Abstract: Hydrotarsus Falkenstrom (Coleoptera, Dytiscidae) is the only genus of aquatic beetle considered to be endemic to the Atlantic islands of Macaronesia Its three known species (H lundbladi Falkenstrom from Madeira, and H compunctus (Wollaston) and H pilosus Guignot from the Canary Islands) are revised, and their phylogenetic position studied based on sequences of the 16S rRNA and Cytochrome Oxidase I genes of mitochondrial DNA Our results clearly indicate that the species of Hydrotarsus fall deep within the genus Hydroporus Clairville, in a clade formed by (in addition to Hydrotarsus) species currently included in the H fuscipennis, marginatus, nigrita and tessellatus groups, and in consequence a new synonymy is proposed: Hydrotarsus Falkenstrom ¼ Hydroporus Clairville synn The origin of the species of 'Hydrotarsus', based on molecular clock estimations, is late Miocene, relatively recent in the history of the Atlantic islands They are therefore not palaeoendemics but highly derived, recently evolved elem- ents in the Macaronesian fauna The estimated ages of divergence among species are much more recent than the emergence of any of the islands on which they are found, suggesting rare long-range dispersal as the mechanism driving the diversi- fication of the lineage Morphological modifications of the species of 'Hydrotar- sus' are discussed, as well as those of other dytiscids from hygropetric habitats

Evolutionary trends and patterns in centipede segment number based on a cladistic analysis of Mecistocephalidae (Chilopoda: Geophilomorpha)

Abstract: . Evolutionary changes in segment number during the radiation of Mecistocephalidae, a group of geophilomorph centipedes with segment number usually invariant at the species level, were explored based on a cladistic analysis of forty-six mecistocephalid species, representative of the extant diversity in segment number. The data matrix included 118 morphological characters. Trends were recognized in the evolution of segment number and discussed in relation to the underlying ontogenetic mechanisms of segmentation. The basic trend was towards an increasingly higher number of leg-bearing segments, from (most probably) forty-one to sixty-five (101 in one exceptional case). Changes always involved even sets of segments. Additions of two, four or eight segments usually occurred, but a case of overall duplication of the whole number was also documented. Most changes occurred starting from values belonging to the arithmetical series forty-one, forty-five, forty-nine, whereas the intermediate values forty-three, forty-seven, fifty-one were often evolutionary dead-ends. This evidence suggests a multiplicative mechanism of segmentation involving one or more final run of duplication, as well as a precise control of the final number of segments which produces absolute number stability, except for a single, highly derived species with an exceptionally high number of segments. These ideas contribute to a more general model of arthropod segmentation recently developed by Minelli. A taxonomic revision of mecistocephalids is presented: three subfamilies are proposed (Arrupinae, Dicellophilinae and Mecistocephalinae) and Sundarrup is recognized as a junior synonym of Anarrup.

Phylogeny of the bee genus Agapostemon (Hymenoptera: Halictidae)

Abstract: . A phylogeny for genus Agapostemon (Hymenoptera: Halictidae) is reconstructed from morphological characters, using all species, with four representatives of Agapostemonini as outgroup taxa. Parsimony analysis using 150 unordered characters resulted in eighteen equally parsimonious trees, a strict consensus of which is highly resolved for Agapostemon. Based on this analysis, Agapostemonoides Roberts & Brooks is returned to generic status, and a new subgenus is described: Notagapostemon, which includes the rhopalocerus, heterurus, atrocaeruleus, erebus, kohliellus, nasutus and intermedius groups of Moure & Hurd. Species groups are proposed for Notagapostemon and revised in A. (Agapostemon). Agapostemon cockerelli Crawford is synonymized with A. obliquus (Provancher), A. epichryseus (Morelos) with A. leunculus Vachal, A. inca Roberts with A. heterurus Cockerell, A. hispaniolicus Roberts female with A. centratus (Vachal) and A. hispaniolicus Roberts male with A. alayoi Roberts. Optimization of known behaviours suggests communal ancestry for the genus and infers the prevalence of this behaviour in Agapostemon. Optimization of geographical range suggests a Central American or Caribbean origin of Agapostemon, with one invasion of North America and multiple invasions of South America and the Caribbean. One invasion of the Caribbean suggests arrival from North America via Florida.

Phylogeny of the death's head hawkmoths, Acherontia[Laspeyres], and related genera (Lepidoptera: Sphingidae: Sphinginae: Acherontiini)

Abstract: . Adult death's head hawkmoths (Acherontia species) have a unique feeding biology as cleptoparasites of honeybees, stealing honey from the combs, rather than imbibing nectar from flowers. The moths have a range of features, both morphological and behavioural, that enable them to successfully enter, feed and escape from the colonies. These adaptations may vary among the three Acherontia species and allow them each to target different species of honeybee. A cladistic analysis is presented of the hawkmoths of tribe Acherontiini. The study aims to resolve the relationships of the genera and species of Acherontiini, with a particular focus on the three species of Acherontia. The dataset comprises sixty-five characters derived from adult, larval and pupal morphology, and larva host-plant biology. These data are analysed using equal weighting and implied weighting. Acherontiini and each constituent genus are recovered as monophyletic. However, within Coelonia, there is ambiguity in that the sister-species relationships C. brevis+C. fulvinotata and C. fulvinotata + C. solani are equally parsimonious under both weighting schemes. Furthermore, under equal weighting Agrius is placed equally parsimoniously as the sister group of either Acherontia + Coelonia or Callosphingia. Under implied weighting, however, only the latter relationship is most parsimonious (fit). Within Acherontia, A. atropos and A. styx are always recovered as sister species to the exclusion of A. lachesis. The results of the phylogenetic analysis provide an objective basis for future studies of the unique cleptoparasitic association of these moths.

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

Abstract: . 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.

Ocoidae, a new family of asiloid flies (Diptera: Brachycera: Asiloidea), based on Ocoa chilensis gen. and sp.n. from Chile, South America

Abstract: . The monotypic new family Ocoidae is described to accommodate Ocoa gen.n., small, elongate, delicate, asiloid flies known from west-central Chile, South America. Both sexes of adult O. chilensis sp.n. are described and illustrated; immature stages are unknown. Diagnostic morphological features of adults include the antennal postpedicel comprising a single, undivided structure consisting of a bulbous base and a long, threadlike terminal element; vein M3 lacking, anal vein complete; acropod heterodactylus; spurs lacking on all tibiae; and anterior surface of hind coxa with strong, knoblike, bulbous projection; and abdominal tergite 2 lacking sensory patches of hairs. The epandrium of the male terminalia is divided along the midline into two sicklelike sections; gonostyli articulate in a horizontal plane. The female terminalia has well-developed acanthophorite spines; posterior margin of hypoproct with several ventrally projecting, elongate, needlelike setae; two large, poorly sclerotized spermathecae; and a spermathecal sac that is smaller than each spermatheca. The relationships of this enigmatic family are discussed in relation to recent findings on the phylogeny and classification of the Asiloidea. Ocoidae are similar to Therevidae and close relatives Scenopinidae and Apsilocephalidae, comprising the therevoid group of families; however, they lack defining synapomorphies of those families. Morphological evidence supports a sister-group relationship between Ocoidae and Scenopinidae. Molecular evidence from 28S rDNA provides further support for the monophyly of the therevoid group of families, and suggests that Ocoidae belongs within this clade, in agreement with the morphological data. The nucleotide data place Ocoidae as sister to the Scenopinidae + Therevidae, but the morphology-based hypothesis of relationships (Ocoidae + Scenopinidae) is only two additional steps (0.2%) on the optimal trees from the molecular data.

A revision of the Encarsia longifasciata species group (Hymenoptera: Aphelinidae)

Abstract: . The Encarsia longifasciata group is established and revised, and its relationship to other species groups of Encarsia is discussed with reference to biological, morphological and molecular evidence. Principal component analysis on all available specimens provided supporting evidence for the existence of five species, two of which are described herein as new: E. dewa Pedata & Polaszek, sp.n and E. prinslooi Pedata & Polaszek, sp.n. All species are either diagnosed or described, and illustrated, and information is given on their distribution and host range. Additional information on the biology and larval stages, and scanning electron micrographs of morphological details, are provided for E. arabica Hayat. A preliminary phylogenetic study using 28S D2 rDNA sequence data for E. arabica was inconclusive, but suggested possible affinities between the longifasciata and parvella species groups. An illustrated dichotomous key to all species of the E. longifasciata group is given.

Resolving relationships over a wide taxonomic range in Delphacidae (Homoptera) using the COI gene

Abstract: Using a combination of different methods to investigate the suitability of a fragment of the cytochrome c oxidase I gene (COI), we succeeded in partially resolving phylogenetic relationships in Delphacidae from the level of species to subfamily. Spectral analysis applied to the relatively noisy COI data proved to be especially useful. It clearly showed when phylogenetic signals were not completely randomized and it was very helpful for identifying problem areas in the dataset. Relationships among the four sampled subfamilies were completely resolved. In contrast to the tree based on morphological characters, we found evidence that Asiraca and Ugyops are sister groups (supporting monophyly of Asiracinae) and that Stenocraninae are the sister group of Kelisiinae. Contradictory signals were observed within Delphacini, but there are characters that support a close relationship between Conomelus and Megamelus. Other than this, the COI data gave support for the monophyly of Kelisiinae, Delphacinae, Chloriona and Javesella. Although third codon positions may appear to be saturated within the 'modern' Delphacidae (Delphacini), they still contain important phylogenetic signals at the deepest taxonomic level. The easiest explanation for this is the difference in amino acid usage between Asiracinae and non-Asiracinae. Overall, this fragment of the COI gene seems to be useful for a rather wide taxonomic range in Delphacidae, except maybe for resolving generic relationships in the large tribe Delphacini.

Systematics and cladistic analysis of Cerophytidae (Elateroidea: Coleoptera)

Abstract: . The elateroid family Cerophytidae, originally based on Cerophytum Latreille, is characterized and revised. Detailed morphological studies of adults and a cladistic analysis permitted the recognition of three well-defined lineages: one in the Holarctic Region, and the other two in the Neotropical Region, each one distributed from southern Mexico to South America. The resulting cladogram for these genera is ((Cerophytum) (Brachycerophytum, Phytocerum)). Cerophytum includes four species: C. elateroides (Latreille) (type species, southern and central Europe), C. convexicolle (LeConte) (western U.S.A.), C. japonicum Sasaji (Japan) and C. pulsator (Haldeman) (eastern U.S.A.). Brachycerophytum gen.n. contains two Neotropical species: B. fuscicorne (Bonvouloir) comb.n. (type species, Mexico to Colombia) and B. sinchona sp.n. (Peru and Bolivia). Phytocerum gen.n. includes fifteen species: P. golbachi sp.n. (type species, Argentina), P. alleni sp.n. (Costa Rica, Nicaragua), P. belloi sp.n. (Brazil), P. birai sp.n. (Brazil), P. boliviense (Golbach) comb.n. (Bolivia), P. burakowskii sp.n. (Trinidad), P. cayennense (Bonvouloir) comb.n. (Mexico, French Guyana, Brazil), P. distinguendum (Soares & Peracchi) comb.n. (Brazil, Paraguay and Argentina), P. serraticorne sp.n. (Guatemala), P. golbachi sp.n. (Argentina), P. ingens sp.n. (Brazil), P. inpa sp.n. (Brazil), P. minutum (Golbach) comb.n. (Argentina), P. simonkai sp.n. (Trinidad), P. trinidadense (Golbach) comb.n. (Trinidad) and P. zikani (Soares & Peracchi) comb.n. (Brazil). Diagnoses for the genera, redescriptions and descriptions for the species, as well as keys to genera and species, are provided. The larva of C. elateroides is redescribed.

Revision and phylogenetic analysis of the Central American endemic genus Phalangogonia Burmeister (Coleoptera: Scarabaeidae: Rutelinae: Anoplognathini)

Abstract: Phalangogonia Burmeister is revised and now includes eight species: P. dispar Ohaus, P. james- onae, sp.n., P. lacordairei Bates, P. obesa Burmeister, P. parilis Bates, P. punctata Franz, P. ratcliff ei, sp.n. and P. sperata Sharp. Phalangogonia debilidens Ohaus is placed in synonymy with P. sperata. Lectotypes are designated for the following nominal species: P. dispar Ohaus, P. lacordairei Bates, P. parilis Bates and P. championi Bates. Neotypes are designated for: P. obesa Burmeister, P. sperata Sharp, P. stipes Sharp and P. debilidens Ohaus. A cladistic analysis of the species of Phalangogonia was executed using thirty-two mor- phological characters of adults.

Assessment of monophyly of species-groups within Afrotropical Empidini (Diptera: Empididae: Empidinae), with a cladistic analysis of the Empis setitarsus-group

Abstract: . The Afrotropical Empidini are reviewed from a phylogenetic point of view. Seven monophyletic groups are recognized within the tribe, namely Empis (Coptophlebia) bivittata, E. (C ) chrysocera, E. (C ) hyalea, E. (C ) namwamba, E. (C ) poecilosoma, E. (Disneyempis) and E. setitarsus. Their monophyly is assessed by comparison with the remaining Empidini. The E. setitarsus-group is cladistically analysed to study the evolution of the length of vein M1 by optimization on the cladogram, and the status of subgenus Coptophlebia Bezzi is discussed. A key to the seven Afrotropical species-groups of Empidini is provided, genus Rhamphomyia is no longer recognized for the Afrotropical Region, and four new species belonging to the E. setitarsus-group, namely E. cinerarius sp.n., E. pseudosetitarsus sp.n., E. spinosa sp.n and E. spungaberaensis sp.n., are described and included in a key to all known species of this group.