# references

title:
Description and phylogeny of Staminodeus, a new genus of Derelomini (Coleoptera: Curculionidae) associated with Cyclanthaceae

citation:
Franz, N.M. 2001. Description and phylogeny of Staminodeus, a new genus of Derelomini (Coleoptera: Curculionidae) associated with Cyclanthaceae. Coleopterists' Bulletin 55: 397-418.


title:
The Indo-Australian Agaoninae (pollinators of figs)

citation:
Wiebes, J.T. 1994. The Indo-Australian Agaoninae (pollinators of figs). Koninklijke Nederlandse Akademie van Wetenschappen, Amsterdam.


title:
Deductions from the present day host plants of aphids and related insects

journal:
Insect/Plant Relationships: Symposia of the Royal Entomological Society of London

citation:
Eastop, V.F. 1971. Deductions from the present day host plants of aphids and related insects. Pp. 157- in: van Emden, H.F. [ed.], Insect/Plant Relationships: Symposia of the Royal Entomological Society of London. Blackwell Scientific Publications, Oxford.


author:
Zhang, A.-B.; Kong, X.-B.; Li, D.-M.; Liu, Y.Q.

title:
DNA fingerprinting evidence for the phylogenetic relationship of eight species and suspecies of Dendrolimus (Lepidoptera: Lasiocampidae) in China

citation:
Zhang, A.-B., Kong, X.-B., Li, D.-M. and Liu, Y.Q. 2004. DNA fingerprinting evidence for the phylogenetic relationship of eight species and suspecies of Dendrolimus (Lepidoptera: Lasiocampidae) in China. Acta Entomologica Sinica 47: 236-242.


title:
Speciation of the Enchenopa binotata complex (insecta: Homoptera: Membracidae)

citation:
Wood, T.K. 1993. Speciation of the Enchenopa binotata complex (insecta: Homoptera: Membracidae). Pp. 300-317 in: Lees, D.R. and Edwards, D. [eds.], Evolutionary Patterns and Processes. Academic Press, London.


author:
Kergoat, G.J.; Alvarez, N.; Hossaert-McKey, M.; Faure, N.; Silvain, J.-F.

title:
Parallels in the evolution of the two largest New and Old World seed-beetle genera (Coleoptera, Bruchidae)

citation:
Kergoat, G.J., Alvarez, N., Hossaert-McKey, M., Faure, N. and Silvain, J.-F. 2005. Parallels in the evolution of the two largest New and Old World seed-beetle genera (Coleoptera, Bruchidae). Molecular Ecology : (in press).


title:
The Asphondylia (Cecidomyiidae: Diptera) of creosote bush (Larrea tridentata) in North America

citation:
Gagné, R.J. and Waring, G.L. 1990. The Asphondylia (Cecidomyiidae: Diptera) of creosote bush (Larrea tridentata) in North America. Proceedings of the Entomological Society of Washington 92: 649-671.


title:
Phylogenetic relationships, ecology, and ecological genetics of cecidogenous Tephritidae

journal:
Biology, Ecology, and Evolution of Gall-inducing Arthropods

citation:
Korneyev, V., Zwölfer, H. and Seitz, A. 2005. Phylogenetic relationships, ecology, and ecological genetics of cecidogenous Tephritidae. Biology, Ecology, and Evolution of Gall-inducing Arthropods 1: 321-371.


title:
Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae)

abstract:
The phylogenetic relationships of the genera in the geometrid tribe Scopulini (Lepidoptera: Sterrhinae) were examined using 141 characters of adult morphology and ecology. The study material included 92 species, representing all previously recognized genera and covering the morphological variation and full geographical range of the tribe. The cladistic analysis resulted in 20 equally parsimonious trees and a strict consensus cladogram based on these was well resolved. A majority of the recovered synapomorphic characters have been used previously in the taxonomy of the tribe. However, many novel characters were found in the sclerotized structures of the thorax. Many previously recognized genera were found to be nonmonophyletic and based on the present revised, synapomorphy-based classification, the number of recovered genera is reduced considerably. Twenty new generic synonyms and 90 new or revived species combinations are proposed. Seven genera are considered valid, with the large genus Scopula Schrank including over 85% of all species in the tribe. The taxonomic history of the tribe is reviewed and the problems of earlier classifications are discussed. A key to the genera is presented, although an informal diagnosis is preferred. All recognized genera are illustrated and a revised world checklist of the Scopulini is presented.

citation:
Pasi, S. 2005. Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae). Zoological Journal of the Linnean Society 143: 473-530.


title:
Phylogenetic analysis of Cyanophrys Clench, a synopsis of its species, and the potentially threatened C. bertha (Jones) (Lycaenidae: Theclinae: Eumaeini)

abstract:
A species level phylogenetic analysis of Cyanophrys Clench was performed using 14 characters of the frons, wing venation, wing shape, androconia, and genitalia. There were 15 most parsimonious cladograms, and a successive weighting iteration resulted in three of these 15. Cyanophrys is characterized in this paper by two hypothesized synapomorphies (an expanded hindwing anal lobe that is tan-brown and a pair of dorsal brush organs), and adults can be readily identified in the field. Cyanophrys has been divided into Plesiocyanophrys K. Johnson, Eisele and MacPherson, Apophrys K. Johnson and Le Crom, Antephrys K. Johnson, Eisele and MacPherson, Mesocyanophrys K. Johnson, and Cyanophrys (as delimited in Johnson and Le Crom 1997a), but only the monotypic Antephrys is monophyletic on the most parsimonious cladograms. A synopsis of Cyanophrys species includes notes on their distribution, habitat, identification, nomenclature, larval food plants, and male behavior. Cyanophrys bertha, which occurs in moist evergreen and seasonally deciduous forests in the mountains of southern Brazil from 800 to 1,400 m elevation, has been proposed for threatened status. It is "cladistically distinct" (sensu Vane-Wright et al. 1991) in that its sister is a lineage of five montane species or seven primarily lowland species.

citation:
Robbins, R.K. and Duarte, M. 2005. Phylogenetic analysis of Cyanophrys Clench, a synopsis of its species, and the potentially threatened C. bertha (Jones) (Lycaenidae: Theclinae: Eumaeini). Proceedings of the Entomological Society of Washington 107: 398-416.


author:
Nice, C.C.; Anthony, N.; Gelembiuk, G.; Raterman, D.; Ffrench-Constant, R.

title:
The history and geography of diversification within the butterfly genus Lycaeides in North America

abstract:
The Lycaeides butterfly species complex in North America consists of two nominal, morphologically defined species. These butterflies are ecologically diverse and appear to be distributed as a geographically complex mosaic of locally differentiated populations that may be undergoing adaptive radiation. We asked whether patterns of molecular genetic variation within the species complex are congruent with currently recognized morphological species and whether the distribution of molecular variation is consistent with the hypothesis that Pleistocene climate changes contributed to the process of differentiation within the genus. Variation in the form of the genitalia from 726 males from 59 populations clearly distinguishes both species with only six populations containing morphologically intermediate or ambiguous individuals. However, partitioning of molecular variance in a 236 bp section of the mitochondrial AT-rich region from 628 individuals (57 populations) surveyed using single strand conformation polymorphism analysis (SSCP) indicates that only 26% of the total genetic variation is distributed along nominal species boundaries as defined by morphology. Instead, three phylogeographical groups were detected, represented by three major haplotype clades, which account for 90% of the total genetic variance. Pleistocene glaciations appear to have fostered divergence during glacial maxima, while postglacial range expansions created opportunities for gene exchange and reticulation along suture zones between geographical groups. Data presented here allow us to make inferences about the history of the species complex. However, evidence of ancestral polymorphism and reticulation limit our ability to define species boundaries based on mitochondrial DNA sequence variation.

citation:
Nice, C.C., Anthony, N., Gelembiuk, G., Raterman, D. and Ffrench-Constant, R. 2005. The history and geography of diversification within the butterfly genus Lycaeides in North America. Molecular Ecology 14: 1741-1754.


title:
A systematic revision of the New World species of Trypeta Meigen (Diptera: Tephritidae)

abstract:
The tephritid genus Trypeta is revised for the New World and a key, descriptions, illustrations, and phylogeny provided for the eighteen species recognized: Trypeta bifasciata, sp.n., californiensis, sp.n., chiapasensis, sp.n., concolor (Wulp), costaricana, sp.n., denticulata, sp.n., flaveola Coquillett, flavifasciata, sp.n., footei, sp.n., fractura (Coquillett), inclinata, sp.n., maculata, sp.n., maculosa (Coquillett), melanoura, sp.n., reducta, sp.n., rufata (Wulp), striata (Wulp), and wulpi, sp.n. Lectotypes are designated for Spilographa fractura Coquillett and Spilographa maculosa Coquillett. Trypeta (Acidia) tortile Coquillett, Acidia sigma Phillips, and Trypeta angustigena Foote are synonymized with Trypeta flaveola Coquillett. Based upon many specimens, we redefine Trypeta flaveola as a widely distributed and morphologically highly variable species. New host plant records are provided for five species, and host data and other information on the biology and immature stages of all of the New World Trypeta species are summarized and discussed. Our phylogenetic analysis suggests that the majority of New World Trypeta species form a monophyletic group that diversified in the southwestern U.S.A. and Mesoamerica.

citation:
Han, H.-Y. and Norrbom, A.L. 2005. A systematic revision of the New World species of Trypeta Meigen (Diptera: Tephritidae). Systematic Entomology 30: 208-347.


author:
Brändle, M.; Knoll, S.; Eber, S.; Stadler, J.; Brandl, R.

title:
Flies on thistles: support for synchronous speciation?

abstract:
Synchronous speciation of hosts and herbivorous insects predicts a congruent topology of host and insect phylogenies and similar evolutionary ages of host and insect taxa. To test these predictions for the specialized herbivorous fly genus Urophora (Diptera: Tephritidae), we used three different approaches. (i) We generated a phylogenetic tree of 11 European Urophora species from allozyme data and constructed a phylogeny of their hosts from published sources. Superimposing the Urophora tree on the host-plant tree we found no evidence for general congruence. (ii) We correlated genetic distances (Nei distances) of the host plants vs. the genetic distances of associated Urophora species. Overall, the relationship was not positive. Nevertheless, for some pairs of Urophora species and host plants genetic distances were in the same order of magnitude. (iii) We collected allozyme data for pairs of thistle taxa and pairs of herbivores on thistles together with independent time estimates. With these data we calibrated a molecular clock. There was a non-linear relationship between phylogenetic age and genetic distance, rendering the dating of deep events in thistle-insect evolution difficult. Nevertheless the derived molecular clock showed that the split of insect taxa lagged behind the split of hosts.

citation:
Brändle, M., Knoll, S., Eber, S., Stadler, J. and Brandl, R. 2005. Flies on thistles: support for synchronous speciation? Biological Journal of the Linnean Society 84: 775-783.


title:
Reconstruction of phylogeny in leaf-rolling weevils (Coleoptera, Rhynchitidae, Attelabidae) using the SYNAP method. Report 2.

abstract:
Phylogenetic relations between tribes of the family Attelabidae were reconstructed. The independence of the subfamily Apoderinae is confirmed. The most important in the cladogram obtained is a serial branching of tribes from the main stem that shows their isolation from each other and a consecutive purchase of apomorphies.;

citation:
Legalov, A.A. 2005. Reconstruction of phylogeny in leaf-rolling weevils (Coleoptera, Rhynchitidae, Attelabidae) using the SYNAP method. Report 2.. Zoologicheskii Zhurnal 84: 190-194.


author:
Johnson, D.G.; Carstens, B.C.; Sheppard, W.S.; Zack, R.S.

title:
Phylogeny of leafhopper subgenus Errhomus (Erronus) (Hemiptera: Cicadellidae) based on mitochondrial DNA sequences

abstract:
The Nearctic leafhopper genus Errhomus Oman (Hemiptera: Cicadellidae) exhibits high host-plant selectivity and female brachyptery, which may limit gene flow. Two subgenera, Errhomus (Errhomus) and Errhomus (Erronus), are recognized based on morphological and behavioral differences. We used rnitochondrial DNA sequencing to elucidate the phylogenetic relationships among members of the subgenus Errhomus (Erronus). Portions of the NADH dehydrogenase subunit 1 (ND1) and the leucine (CUN) transferRNA (tRNA-Leu) genes were sequenced and were analyzed with parsimony and maximum likelihood. Our results generally verified the findings of Hamilton and, Zack (1999); however, both optimality criteria revealed a paraphyletic Errhomus (Erronus). Errhomus (Erronus) calvus does not seem to be a member of the subgenus Errhomus (Erronus) but seems to be more closely aligned with the subgenus Errhomus (Errhomus). We tested two a priori null hypotheses: 1) that the phylogeny estimated from the molecular data matched a previous phylogeny based on morphology and biogeography from Hamilton and Zack (1999) and 2) that diagnostic characters of the male genitalia (serrated aedeagal processes and triangular aedeagal shaft tip) were synapomorphies that defined major clades. We rejected these hypotheses with both Bayesian tests and parametric bootstraps.

citation:
Johnson, D.G., Carstens, B.C., Sheppard, W.S. and Zack, R.S. 2005. Phylogeny of leafhopper subgenus Errhomus (Erronus) (Hemiptera: Cicadellidae) based on mitochondrial DNA sequences. Annals of the Entomological Society of America 98: 165-172.


title:
Molecular systematics of nuclear gene period in genus Anastrepha (Tephritidae)

abstract:
The nuclear gene period is used to reconstruct the phylogeny of Anastrepha, a genus of tephritid fruit flies. Specifically, a region of period termed C3C5 that is located in the dCLK: CYC inhibition domain (CCID) is analyzed. This is the first study using this C-terminal region of the period gene for phylogenetics. Variation in the gene is informative for this genus, and period trees are compared with phylogenies based on initochondrial and morphological data sets. In general, branches that are supported in the other data sets are recovered in the period tree; moreover, trees built using the period gene support branching patterns previously unresolved with 16S rRNA gene data. Important relationships recovered in the period gene tree include a paraphyletic placement of Anastrepha with respect to Toxotrypana, support for the separation of Anastrepha species groups into two evolutionary sections (with the exception of the pseudoparallela species group), and evidence that several species groups are not monophyletic.

citation:
Barr, N.B., Cui, L. and McPheron, B.A. 2005. Molecular systematics of nuclear gene period in genus Anastrepha (Tephritidae). Annals of the Entomological Society of America 98: 173-180.


title:
Revision of the fruit fly genus Capparimyia (Diptera, Tephritidae)

abstract:
Capparimyia Bezzi is revised and eight species recognized, of which four are new: C. aenigma sp. nov., C. maerua sp. nov., C. mirabilis sp. nov., and C. spatulata sp. nov. A lectotype is designated for C. bipustulata (Bezzi, 1923), C. savastani (Martelli) and C. melanaspis (Bezzi). Capparimyia aristata nom. nov. is proposed as a replacement name for Pardalaspis (now Capparimyia) bipustulata Bezzi, 1924 (preoccupied Bezzi, 1923). All species are described or re-described, with illustrations of mesonotal and wing patterns and male and female terminalia. A key to species is provided. Phylogenetic relationships within the genus are analysed cladistically and considered vis-a-vis the relationships between the host plants. Distribution patterns are discussed.

citation:
DeMeyer, M. and Freidberg, A. 2005. Revision of the fruit fly genus Capparimyia (Diptera, Tephritidae). Zoologica Scripta 34: 279-303.


title:
Colour pattern evolution in butterflies: a phylogenetic analysis of structural ultraviolet and melanic markings in North American sulphurs

abstract:
The males of many butterflies are more brightly coloured than conspecific females and this sexual dichromatism often extends into ultraviolet (UV) wavelengths. Sexual selection, acting either in the context of intra- or intersexual interactions. is thought responsible for the relative brightness of the male visage: however, this hypothesis has been difficult to test experimentally. In this study, we undertake an indirect evaluation of this idea by analysing the phylogenetic distribution of two elements of male colour pattern - UV reflectance and melanic markings - in North American sulphur butterflies. Only the UV element, which derives from nanometre-scale surface structures and is extremely bright and spectrally pure, is expected to be of sexually selected origin. We therefore predicted, based upon prior theoretical treatments. that UV colour pattern should exhibit a polyphyletic distribution typical of a sexually selected trait, whereas melanic patterns should not. We charted the distribution of each colour element across the dorsal hind- and fore-wing surfaces of 26 species (spanning four genera), and summarized the resulting intraspecific variation in colour pattern using principal components analysis. Analysis of among- versus within-sister species group variance confirmed, as predicted, a largely polyphyletic distribution for UV but not melanic patterns. We also found that variation in fore-wing UV patterning could be considered orthogonal of hind-wing variation. These findings shed interesting light on the evolution of structural coloration in this group and add strength to the idea that brilliant male coloration is principally of sexually selected origin.

citation:
Kemp, D.J., Rutowski, R.L. and Mendoza, M. 2005. Colour pattern evolution in butterflies: a phylogenetic analysis of structural ultraviolet and melanic markings in North American sulphurs. Evolutionary Ecology Research 7: 133-141.


author:
Hundsdoerfer, A.K.; Kitching, I.J.; Wink, M.

title:
A molecular phylogeny of the hawkmoth genus Hyles (Lepidoptera: Sphingidae, Macroglossinae)

abstract:
The hawkmoth genus Hyles is one of 15 genera in the subtribe Choerocampina of the subfamily Macroglossinae. Due to a remarkable uniformity, morphological characters usually used to identify and classify Lepidoptera at the species level cannot be used in this genus. Instead, we used DNA sequences comprising about 2300 bp derived from the mitochondrial genes COX 1, COX 11, and tRNA-leucine to elucidate the phylogeny of Hyles. The results corroborate the monophyly of Hyles but conflict with previous internal classifications of the genus based on morphology. Hyles seems to have evolved in the Neotropics during the Oligocene/Eocene epochs and the molecular data (which evolved clock-like) confirm the hypothesis that it is a very young genus that radiated on a global scale rather quickly. We hypothesize its sister group to be one of the genera Deilephila, Theretra or Xylophanes. The Nearctic may have been colonized rapidly by Hyles once the land bridge formed during the Pliocene, since within this same Epoch, the invasion of the Palaearctic appears to have proceeded from the East, via the Bering route. The colonization of Australia appears to have occurred rather early in Hyles radiation, although the route is not clear. We propose that the radiation of the Hyles euphorbiae-complex s. str. (HEC) occurred as recently as the Pliocene/Pleistocene boundary and that its roots can still be reconstructed in Asia. Hyles dahlii is closely related to the HEC, but a sister group relationship to the HEC s. str. cannot be corroborated unequivocally. HEC population ranges appear to have tracked climate oscillations during the Pleistocene Ice Ages, resulting in hybridization around the Mediterranean Sea as they repeatedly intermingled. Comparison of the phylogeny with food plant affiliations leads us to hypothesize that Euphorbia monophagy evolved at least two times independently within Hyles.

citation:
Hundsdoerfer, A.K., Kitching, I.J. and Wink, M. 2005. A molecular phylogeny of the hawkmoth genus Hyles (Lepidoptera: Sphingidae, Macroglossinae). Molecular Phylogenetics and Evolution 35: 442-458.


title:
Gelechioidea (Insecta: Lepidoptera) systematics: A reexamination using combined morphology and mitochondrial DNA data

abstract:
Phylogenies of Gelechioidea (Insecta: Lepidoptera) historically have been in disagreement and deWnitions vary at the family and subfamily levels. Addition of new taxa or new characters drastically changes relationships indicating that current phylogenetic schemes require more investigation. This study is the Wrst phylogenetic analysis of Gelechioidea to include molecular data. Here we present a combined analysis using Maximum Parsimony to investigate sister-group relationships within Gelechioidea. The addition of Cytochrome oxidase I and II to revised published morphological matrices gives 453 parsimony informative characters for the 42 taxa for which we have sequence data. The combined analysis resulted in two trees with mostly novel sister-group relationships. These results challenge current concepts of Gelechioidea, suggesting that traditional morphological characters that have united taxa may not be homologous structures and are in need of further investigation. A combination of morphological data with new molecular; data will be the most robust method of study for Gelechioidea phylogenetics.

citation:
Bucheli, S.R. and Wenzel, J. 2005. Gelechioidea (Insecta: Lepidoptera) systematics: A reexamination using combined morphology and mitochondrial DNA data. Molecular Phylogenetics and Evolution 35: 380-394.


author:
Yen, S.H.; Robinson, G.S.; Quicke, D.L.J.

title:
The phylogenetic relationships of Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae)

abstract:
The chalcosiine zygaenid moths constitute one of the most striking groups within the lower-ditrysian Lepidoptera, with highly diverse mimetic patterns, chemical defence systems, scent organs, copulatory mechanisms, hostplant utilization and diapause biology, plus a very disjunctive biogeographical pattern. In this paper we focus on the genus-level phylogenetics of this subfamily A cladistic study was performed using 414 morphological and biochemical characters obtained from 411 species belonging to 186 species-groups of 73 genera plus 21 outgroups. Phylogenetic analysis using maximum parsimony leads to the following conclusions: (1) neither the current concept of Zygaenidae nor that of Chalcosiinae is monophyletic; (2) the previously proposed sister-group relationship of Zygaeninae + Chalcosiinae is rejected in favour of the relationship (Zygaeninae + ((Callizygaeninae + Cleoda) + (Heteropan + Chalcosiinae))); (3) except for the monobasic Aglaopini, none of the tribes sensu Alberti (1954) is monophyletic; (4) chalcosiine synapomorphies include structures of the chemical defence system, scent organs of adults and of the apodemal system of the male genitalia. A paired metathoracic androconial organ and a series of abdominal tergal corematal organs have been discovered, both being new to Lepidoptera. Due to highly homoplastic patterns in copulatory structures and wings that demonstrate significant sexual dimorphism, polymorphism and mimicry, 17 of the 69 'true' chalcosiine genera (c. 25%) are shown to be either paraphyletic or polyphyletic. The present classification is therefore very misleading. Reductions of various parts of the male genitalia in some groups are accompanied by morphological and functional replacement involving the 8th abdominal segment. A prominent but convergent lock and key mechanism is revealed.

citation:
Yen, S.H., Robinson, G.S. and Quicke, D.L.J. 2005. The phylogenetic relationships of Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae). Zoological Journal of the Linnean Society 143: 161-341.


author:
Cesari, M.; Marescalchi, O.; Francardi, V.; Mantovani, B.

title:
Taxonomy and phylogeny of European Monochamus species: first molecular and karyological data.

abstract:
The worldwide distributed genus Monochamus Megerle, 1821 (Coleoptera Cerambicydae) comprises beetles that may become pests of economic importance in conifer stands in the Nearctic and Palearctic Regions. Besides direct damage due to the larval tunnelling habits, they have also been recognized as main vectors of the phytoparasitic nematode Bursaphelenchus xylophilus (Steiner & Buhrer, 1934) (Nematoda Aphelenchoididae). We analysed the complete mitochondrial cytochrome oxidase I gene and a fragment of the small subunit RNA gene sequences (1536 base pairs) in the five European species. These are: Monochamus galloprovincialis (Olivier, 1795), morphologically distinguished in two subspecies M. galloprovincialis galloprovincialis (Olivier, 1795) and M. galloprovincialis pistor (Germar, 1818); Monochamus sutor (Linneus 1758); Monochamus saltuarius (Gebler 1830); Monochamus sartor (Fabricius, 1787) and Monochamus urussovi (Fischer, 1806). For appropriate comparisons, also the Asiatic Monochamus alternatus Hope, 1842 and a Japanese M. saltuarius sample have been analysed. Both genes show an absolute identity between the two subspecies of M. galloprovincialis and a strong affinity between M. sartor and M. urussovi: the morphological subdivisions of the former taxon in two subspecies and of the latter in two entities of specific level are therefore not supported genetically. On the other hand, the Italian and the Japanese samples of M. saltuarius always cluster together in all trees, and for the remaining taxa, no doubt about their rank of specific differentiation emerges from present analyses. From a phyletic point of view, tree topology indicates the Japanese M. alternatus as the most differentiated taxon and the Euroasiatic M. saltuarius as basal to all other strictly European entities. Chromosome analyses show that the diploid autosomal complement ranges from 18 in M. saltuarius to 20 in M. galloprovincialis, and 22 in M. sartor, but a XX-Xyp sex determining system is shared by all analysed taxa. The M. saltuarius karyotype appears as the most primitive from which the others may be derived through Robertsonian fissions. Karyological data therefore agree with molecular analyses in indicating a basal position of Euroasiatic M. saltuarius with respect to the group of European Monochamus taxa; among these, M. galloprovincialis and M. sartor represent two clearly diverging evolutionary units. Furthermore, karyotype analyses substantiate molecular conclusions about the identity between M. galloprovincialis galloprovincialis and M. galloprovincialis pistor.

citation:
Cesari, M., Marescalchi, O., Francardi, V. and Mantovani, B. 2005. Taxonomy and phylogeny of European Monochamus species: first molecular and karyological data.. Journal of Zoological Systematics and Evolutionary Research 43: 1-7.


title:
The impact of redefined species limits in Palame (Coleoptera: Cerambycidae: Lamiinae: Acanthocinini) on assessments of host, seasonal, and stratum  specificity

abstract:
Assumptions about the host and stratum specificity of tropical insects are routinely incorporated into estimates of global species richness, but few empirical studies reliably assess tropical insect specificity. In French Guiana, cerambycid beetles in the genus Palame reproduce exclusively in trees belonging to the Brazil nut family (Lecythidaceae). During a year-long rearing project, Palame anceps (Bates) emerged exclusively from a single host species, but P. crassimana Bates and P. mimetica Monné appeared to make seasonal changes in host affiliation. In the case of P. crassimana, dry season specimens could be distinguished from rainy season specimens by subtle differences in pubescence, but it was difficult to find diagnostic morphological characters supporting their delimitation as separate species. In order to detect genetic differentiation in Palame, 1049 bp of mitochondrial DNA (COI gene) were sequenced. The three nominal species of Palame yielded six haplotype groups, with sequence divergences; among groups ranging from 8.27 to 17.56%. Species with apparent seasonal changes in host use included multiple distinct haplotype groups that I interpret to represent multiple species-level taxa. Insects considered generalized owing to temporal or geographical changes in host association may, with improved resolution in species limits, prove to be assemblages including relatively specialized cryptic species.

citation:
Berkov, A. 2002. The impact of redefined species limits in Palame (Coleoptera: Cerambycidae: Lamiinae: Acanthocinini) on assessments of host, seasonal, and stratum  specificity. Biological Journal of the Linnean Society 76: 195-209.


title:
Le genre Bruchidius (Coleoptera: Bruchidae) est-il monophylétique? Apports des méthodes de parcimonie, maximum de vraisemblance et inférence bayésienne

citation:
Kergoat, G.J. and Silvain, J.-F. 2004. Le genre Bruchidius (Coleoptera: Bruchidae) est-il monophylétique? Apports des méthodes de parcimonie, maximum de vraisemblance et inférence bayésienne. Biosystema 22: 113-125.


author:
Weingartner, E.; Wahlberg, N.; Nylin, S.

title:
Dynamics of host plant use and species diversity in Polygonia butterflies (Nymphalidae)

abstract:
The ability of insects to utilize different host plants has been suggested to be a dynamic and transient phase. During or after this phase, species can shift to novel host plants or respecialize on ancestral ones. Expanding the range of host plants might also be a factor leading to higher levels of net speciation rates. In this paper, we have studied the possible importance of host plant range for diversification in the genus Polygonia (Nymphalidae, Nymphalini). We have compared species richness between sistergroups in order to find out if there are any differences in number of species between clades including species that utilize only the ancestral host plants ('urticalean rosids') and their sisterclades with a broader (or in some cases potentially broader) host plant repertoire. Four comparisons could be made, and although these are not all phylogenetically or statistically independent, all showed clades including butterfly species using other or additional host plants than the urticalean rosids to be more species-rich than their sisterclade restricted to the ancestral host plants. These results are consistent with the theory that expansions in host plant range are involved in the process of diversification in butterflies and other phytophagous insects, in line with the general theory that plasticity may drive speciation.

citation:
Weingartner, E., Wahlberg, N. and Nylin, S. 2005. Dynamics of host plant use and species diversity in Polygonia butterflies (Nymphalidae). Journal of Evolutionary Biology : (in press?).


title:
Mutualists with attitude: coevolving fig wasps and figs

abstract:
The intimate mutualism between fig wasps and figs has long captivated biologists, and new phylogenies are now uncovering its evolutionary history. Fig-pollinating wasps evolved just once, but fig parasitism has evolved repeatedly and convergently. Figs and their pollinators appear to have co-speciated considerably, but not invariably, because the famous one-to-one rule of specificity is often broken. Some key traits of figs and pollinators; show impressive correlated evolution, but the resolution of conflicts that threaten the stability of the mutualism remains controversial.

citation:
Cook, J.M. and Rasplus, J.-Y. 2003. Mutualists with attitude: coevolving fig wasps and figs. Trends in Ecology and Evolution 18: 241-248.


title:
The adaptive significance of insect gall morphology

abstract:
Insect galls are dramatic examples of extended phenotypes: although composed of host plant tissues, their development is largely controlled by insect genes. Adaptive explanations for gall traits should thus be expressed in terms of impacts on insect fitness, but the extent to which interspecific variation in gall structure is adaptive, and the possible selective pressures driving diversification in gall form remain controversial. In colonial aphids and thrips, gall structures probably diversified in response to selection for enhancement of the surface area available for feeding. In other taxa, such as gall wasps and gall midges, diversity is expressed predominantly in non-nutritive tissues, particularly those on the gall surface. All natural enemies attack the occupants of closed galls by penetrating gall tissue, and modifications that reduce enemy attack rates should thus be favoured. Recent studies of intraspecific variation in gall form strongly support a defensive role for several traits, but, to date, there is little empirical support for enemies as a cause of interspecific variation in gall form. Selection imposed by enemies nevertheless remains the most probable adaptive explanation for the; evolution of diversity. We suggest that this hypothesis has yet to be tested explicitly, and discuss the requirements for an appropriate cross-species analysis.

citation:
Stone, G.N. and Schonrogge, K. 2003. The adaptive significance of insect gall morphology. Trends in Ecology and Evolution 18: 512-522.


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

abstract:
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)).

citation:
Schulmeister, S. 2003. Review of morphological evidence on the phylogeny of basal Hymenoptera (Insecta), with a discussion of the ordering of characters. Biological Journal of the Linnean Society 79: 209-243.


author:
Martin, J.-F.; Gilles, A.; Lortscher, M.; Descimon, H.

title:
Phylogenetics and differentiation among the western taxa of the Erebia tyndarus group (Lepidoptera: Nymphalidae)

abstract:
The phylogenetic relationships among six members of the 'tyndarus' group in the Erebia genus of Satyrid butterflies, i.e. E. tyndarus, E. cassioides, E. nivalis, E. calcaria, E. hispania and E. ottomana were analysed using data from 19 presumptive enzyme loci, as well as 440 and 429 bp, respectively, from the mitochondrial large ribosomal subunit (16S) and subunit 1 from the NADH dehydrogenase (ND1) genes. The two types of molecular data (allozymes and mtDNA) yielded largely congruent tree topologies. The two basal, independent lineages formed by E. ottomana and E. hispania are contrasted by a group of genetically closely related taxa, suggestive of a recent radiation associated with allopatric speciation, and competitive exclusion. The time of divergence for the radiation event is similar for both allozymes and mtDNA with an estimation of 440000 years ago. The lineages involved in this radiation do not comply with all the criteria necessary to assign to each of them full species rank, but they can no more be included in one single species unit. Such situations involving more than two allo- or parapatric lineages may explain why polytomies are so often met in  phylogenetic reconstructions, after the lineages have reached full species rank.

citation:
Martin, J.-F., Gilles, A., Lortscher, M. and Descimon, H. 2002. Phylogenetics and differentiation among the western taxa of the Erebia tyndarus group (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society 75: 319-332.


author:
Farrell, B. D.; Sequeira, A.S.; O'Meara, B.C.; Normark, B.B. ; Chung,J.H.; Jordal, B.H.

title:
The evolution of agriculture in beetles (Curculionidae: Scolytinae and Platypodinae)

abstract:
Beetles in the weevil subfamilies Scolytinae and Platypodinae are unusual in that they burrow as adults inside trees for feeding and oviposition. Some of these beetles are known as ambrosia beetles for their obligate mutualisms with asexual fungi-known as ambrosia fungi-that are derived from plant pathogens in the ascomycete group known as the ophiostomatoid fungi. Other beetles in these subfamilies are known as bark beetles and are associated with free-living, pathogenic ophiostomatoid fungi that facilitate beetle attack of phloem of trees with resin defenses. Using DNA sequences from six genes, including both copies of the nuclear gene encoding enolase, we performed a molecular phylogenetic study of bark and ambrosia beetles across these two subfamilies to establish the rate and direction of changes in life histories and their consequences for diversification. The ambrosia beetle habits have evolved repeatedly and are unreversed. The subfamily Platypodinae is derived from within the Scolytinae, near the tribe Scolytini. Comparison of the molecular branch lengths of ambrosia beetles and ambrosia fungi reveals a strong correlation, which a fungal molecular clock suggests spans 60 to 21 million years. Bark beetles have shifted from ancestral association with conifers to angiosperms and back again several times. Each shift to angiosperms is associated with elevated diversity, whereas the reverse shifts to conifers are associated with lowered diversity. The unusual habit of adult burrowing likely facilitated the diversification of these beetle-fungus associations, enabling them to use the biomass-rich resource that trees represent and set the stage for at least one origin of eusociality.

citation:
Farrell, B. D., Sequeira, A.S., O'Meara, B.C., Normark, B.B. , Chung,J.H. and Jordal, B.H. 2001. The evolution of agriculture in beetles (Curculionidae: Scolytinae and Platypodinae). Evolution 55: 2011-2027.


title:
Mitochondrial DNA variation and identification of bark weevils in the Pissodes strobi species group in western Canada (Coleoptera: Curculionidae)

citation:
Langor,D.W. and Sperling, F.A.H. 1995. Mitochondrial DNA variation and identification of bark weevils in the Pissodes strobi species group in western Canada (Coleoptera: Curculionidae). Canadian Entomologist 127: 895-911.


author:
Sperling, F.A.H.; Landry, J.-F.; Hickey, D.A.

title:
DNA-based identification of introduced ermine moth species in North America (Lepidoptera: Yponomeutidae)

citation:
Sperling, F.A.H., Landry, J.-F. and Hickey, D.A. 1995. DNA-based identification of introduced ermine moth species in North America (Lepidoptera: Yponomeutidae). Annals of the Entomological Society of America 88: 155-162.


title:
Molecular phylogenetics of the genus Ceratitis (Diptera: Tephritidae)

abstract:
The Afrotropical fruit fly genus Ceratitis MacLeay is an economically important group that comprises over 89 species, subdivided into; six subgenera. Cladistic analyses of morphological and host use characters have produced several phylogenetic hypotheses for the genus.; Only monophyly of the subgenera Pardalaspis and Ceratitis (sensu stricto) and polyphyly of the subgenus Ceratalaspis are common to all; of these phylogenies. In this study, the hypotheses developed from morphological and host use characters are tested using gene trees produced; from DNA sequence data of two mitochondrial genes (cytochrome oxidase I and NADH-dehydrogenase subunit 6) and a nuclear; gene (period). Comparison of gene trees indicates the following relationships: the subgenus Pardalaspis is monophyletic, subsection A of; the subgenus Pterandrus is monophyletic, the subgenus Pterandrus may be either paraphyletic or polyphyletic, the subgenus Ceratalaspis; is polyphyletic, and the subgenus Ceratitis s. s. might not be monophyletic. In addition, the genera Ceratitis and Trirhithrum do not form; reciprocally monophyletic clades in the gene trees. Although the data statistically reject monophyly for Trirhithrum under the Shimodaira-; Hasegawa test, they do not reject monophyly of Ceratitis.

citation:
Barr, N.B. and McPheron, B.A. 2006. Molecular phylogenetics of the genus Ceratitis (Diptera: Tephritidae).  38: 216-230.


author:
von Dohlen, C.D.; Rowe, C.A.; Heie, O.E.

title:
A test of morphological hypotheses for tribal and subtribal relationships of Aphidinae (Insecta: Hemiptera: Aphididae) using DNA sequences

abstract:
Aphidinae is the most diverse major lineage of aphids (Aphididae). Aphidinae currently dominate the temperate, northern-hemisphere fauna, but only since the late Tertiary, and few species are native to the southern hemisphere. The success of Aphidinae may be linked to the evolution of an unusual life cycle, host alternation. The classification and phylogeny of Aphidinae have been controversial; schemes based on morphology have been confounded by widespread homoplasy. Here we present the first phylogenetic study of higher-level Aphidinae relationships based on molecular data (elongation factor-1, leucine tRNA, and cytochrome oxidase II sequences). Analyses supported the monophyly of Aphidini and its subtribes, Aphidina and Ropalosiphina, but revealed novel relationships concerning Pterocommatini and Macrosiphini, with the former nested within the latter tribe as the sister to Cavariella. Several relationships within Pterocommatini+Macrosiphini corresponded better with host-plant affliations than with aphid classiWcation. Overall, relationships found here challenge several traditional views of Aphidinae evolution: they suggest more than one origin of host alternation in the family, and they question the assumption that Aphidinae originated in the northern hemisphere.

citation:
von Dohlen, C.D., Rowe, C.A. and Heie, O.E. 2006. A test of morphological hypotheses for tribal and subtribal relationships of Aphidinae (Insecta: Hemiptera: Aphididae) using DNA sequences. Molecular Phylogenetics and Evolution 38: 316-329.


author:
Lopez-Vaamonde, C.; Wikstrom, N.; Labandeira, C.; Godfray, H.C.J.; Goodman, S.J.; Cook, J.M.

title:
Fossil-calibrated molecular phylogenies reveal that leaf-mining moths radiated millions of years after their host plants

abstract:
Coevolution has been hypothesized as the main driving force for the remarkable diversity of insect-plant associations. Dating of insect and plant; phylogenies allows us to test coevolutionary hypotheses and distinguish between the contemporaneous radiation of interacting lineages vs. insect 'host tracking' of previously diversified plants. Here, we used nuclear DNA to reconstruct a molecular phylogeny for 100 species of Phyllonorycter leaf-mining moths and 36 outgroup taxa. Ages for nodes in the moth phylogeny were estimated using a combination of a penalized likelihood method and a Bayesian approach, which takes into account phylogenetic uncertainty. To convert the relative ages of the moths into dates, we used an absolute calibration point from the fossil record. The age estimates of (a selection of) moth clades were then compared with fossil-based age estimates of their host plants. Our results show that the principal radiation of Phyllonorycter leafmining moths occurred well after the main radiation of their host plants and may represent the dominant associational mode in the fossil record.

citation:
Lopez-Vaamonde, C., Wikstrom, N., Labandeira, C., Godfray, H.C.J., Goodman, S.J. and Cook, J.M. 2006. Fossil-calibrated molecular phylogenies reveal that leaf-mining moths radiated millions of years after their host plants. Journal of Evolutionary Biology : (in press).


author:
Jousselin, E.; Van Noort, S.; Rasplus, J.-Y.; Greeff, J.M.

title:
Patterns of diversification of Afrotropical Otiteselline fig wasps: phylogenetic study reveals a double radiation across host figs and conservatism of host association

abstract:
We studied the phylogenetic relationships of Otiteselline fig wasps associated with Ficus in the Afrotropical region using rDNA sequences. African fig species usually host two species of Otiteselline fig wasps. Phylogenetic analyses reveal that this pattern of association results from the radiation of two clades of wasps superimposed on the fig system. Within each clade, wasp species generally cluster according to their host classification. The phylogenies of the two clades are also generally more congruent than expected by chance. Together these results suggest that Otiteselline wasp speciation is largely constrained by the diversification of their hosts. Finally, we show a difference in ovipositor length between the two Otiteselline species coexisting in the same Ficus species, which probably corresponds to ecological differences. The diversification of ecological niches within the fig is probably, with cospeciation, one of the key factors explaining the diversification and maintenance of species of parasites of the fig/pollinator system.

citation:
Jousselin, E., Van Noort, S., Rasplus, J.-Y. and Greeff, J.M. 2006. Patterns of diversification of Afrotropical Otiteselline fig wasps: phylogenetic study reveals a double radiation across host figs and conservatism of host association. Journal of Evolutionary Biology 19: 253-266.


author:
Jiang, Z.-F.; Huang, D.-W.; Zhu, C.-D.; Zhen, W.-Q.

title:
New insights into the phylogeny of fig pollinators; using Bayesian analyses

abstract:
The interaction between figs and fig pollinators is one of the most species-specific mutualisms. Recently, phylogenies of both partners; based on molecular data provided insights into a wide spectrum of co-evolutionary questions. However, for the phylogeny of fig pollinators,; there are some discrepancies between different studies and left some relationships unresolved, especially for deep nodes. The phylogenetic; uncertainties of pollinators prohibit our further understanding of the history of the mutualism. Here, we present phylogenetic; analyses of a larger COI sequence dataset that includes previously published datasets and our sequences from 20 species using Bayesian; method and maximum parsimony. The analyses using different methods share similar topologies. Bayesian analyses provide high level of; confidence for most internal nodes in terms of posterior probability. This study also clarifies some discrepancies between previous studies.; After rooting with Tetrapus, other pollinators split into two clades. Wiebesia and Blastophaga are at basal positions in respective clade.; Ceratosolen is not monophyletic because Kradibia and Liporrhopalum fall inside this group. Three subgenera of Ceratosolen: subgen.; Ceratosolen, subgen. Rothropus, and subgen. Strepitus are not supported. Therefore, Ceratosolen is suggested to be re-divided into three; groups. Urostigma pollinators (including Dolichoris and Blastophaga psenes) are clustered together. The monophylies of Wiebesia, Blastophaga,; Dolichoris are not supported in this analysis. This study also provides a new framework for re-evaluating character evolution; and re-inspecting the deffnition of some genera.

citation:
Jiang, Z.-F., Huang, D.-W., Zhu, C.-D. and Zhen, W.-Q. 2006. New insights into the phylogeny of fig pollinators; using Bayesian analyses. Molecular Phylogenetics and Evolution 38: 306-315.


abstract:
The phylogenetic relationships of the genera in the geometrid tribe Scopulini (Lepidoptera: Sterrhinae) were examined using 141 characters of adult morphology and ecology. The study material included 92 species, representing all previously recognized genera and covering the morphological variation and full geographical range of the tribe. The cladistic analysis resulted in 20 equally parsimonious trees and a strict consensus cladogram based on these was well resolved. A majority of the recovered synapomorphic characters have been used previously in the taxonomy of the tribe. However, many novel characters were found in the sclerotized structures of the thorax. Many previously recognized genera were found to be nonmonophyletic and based on the present revised, synapomorphy-based classification, the number of recovered genera is reduced considerably. Twenty new generic synonyms and 90 new or revived species combinations are proposed. Seven genera are considered valid, with the large genus Scopula Schrank including over 85% of all species in the tribe. The taxonomic history of the tribe is reviewed and the problems of earlier classifications; are discussed. A key to the genera is presented, although an informal diagnosis is preferred. All recognized genera are illustrated and a revised world checklist of the Scopulini is presented.

citation:
Sihvonen, P. 2005. Phylogeny and classification of the Scopulini moths (Lepidoptera: Geometridae, Sterrhinae). Zoological Journal of the Linnean Society 143: 473-530.


title:
A New Species of Iraella Nieves-Aldrey (Hymenoptera: Cynipidae), from Northeastern Spain, Inducing Galls on Papaver (Papaveraceae)

abstract:
A new species of Iraella Nieves-Aldrey, Iraella hispanica n. sp. (Hymenoptera: Cynipidae),; is described from northeastern Spain. The new species induces plurilocular galls on terminal; flower buds of the common poppies Papaver rhoeas L. and Papaver dubium L. (Papaveraceae). The; terminal-instar larva also is described. I. hispanica represents the second known species of the genus; Iraella and is related to Iraella luteipes (Thomson), a species inducing cryptic galls in stems of Papaver; somniferum L. Characters separating adults, larvae, and galls of both species are given as well as life; cycle data. Comments on an associated parasitic species of Figitidae, Parnips sp. cf nigripes (Barbotin),; are provided. The phylogenetic relationship of the new species with a clade of cynipids associated to; Papaver plant species is discussed.

citation:
Nieves-Aldrey, J.L. 2005. A New Species of Iraella Nieves-Aldrey (Hymenoptera: Cynipidae), from Northeastern Spain, Inducing Galls on Papaver (Papaveraceae). Annals of the Entomological Society of America 98: 853-860.


title:
A classification of Danaus butterflies (Lepidoptera: Nymphalidae) based upon data from morphology and DNA

abstract:
Classification of the cosmopolitan butterfly genus Danaus (Nymphalidae: Danainae) is revised at subgeneric, specificand subspecific levels, combining for the first time mitochondrial and nuclear DNA sequence information with morphologicaldata. Tree topologies based on the nuclear genome (allozymes, pheromone components, the morphology ofall life history stages and nuclear DNA sequences), on the one hand, and mitochondrial DNA, on the other, are incongruentand challenge the current taxonomy of the genus. Although earlier classifications, based on adult morphology alone, are, in general, well supported by an analysis of total evidence, the mitochondrial phylogeny shows that the species D. chrysippus and its subgenus; Anosia are deeply paraphyletic. Subspecies dorippus of D. chrysippus is the basal clade of the genus and is reinstated as the species D. dorippus. The former species D. plexaure is demoted to a subspecies of D. eresimus . The specific status of D. erippus , as distinct from D. plexippus, is tentatively supported. On the strength of the new data, division of the monophyletic genus Danaus s.l. into three subgenera Danaus s.s., Salatura; and Anosia is unsustainable and is abandoned. Of the 15 terminal clades (taxa) of Danaus s.l. included in the study, 11 are species that broadly conform to the biological species concept. (The West Indian species D. cleophile, missing from our analysis, is the twelfth species). The remaining terminal clades are subspecies of D. chrysippus comb. nov. and D. dorippus stat. rev. Two sympatric Neotropical species, D. eresimus and D. gilippus, are morphologically distinct and sexually isolated but have nearly identical mitochondrial genomes. In contrast, two partially sympatric Palaeotropical species, D. chrysippus and D. dorippus, are cryptic species that share structural morphology and hybridize but have highly differentiated mitochondrial genomes. D. dorippus is polymorphic for two anciently diverged haplotypes and its history has possibly involved recombinational speciation and/or hybridism.

citation:
Smith, D.A.S., Lushai, G. and Allen, J.A. 2005. A classification of Danaus butterflies (Lepidoptera:; Nymphalidae) based upon data from morphology and DNA. Zoological Journal of the Linnean Society 14: 191-212.


title:
Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae)

abstract:
We infer for the first time the phylogenetic relationships of genera and tribes in the ecologically and evolutionarily well-studied subfamily Nymphalinae using DNA sequence data from three genes: 1450 bp of cytochrome oxidase subunit I (COI) (in the mitochondrial genome), 1077 bp of elongation factor 1-alpha (EF1-a) and 400-403 bp of wingless (both in the nuclear genome). We explore the influence of each gene region on the support given to each node of the most parsimonious tree derived from a combined analysis of all three genes using Partitioned Bremer Support. We also explore the influence of assuming equal weights for all characters in the combined analysis by investigating the stability of clades to different transition/transversion weighting schemes. We find many strongly supported and stable clades in the Nymphalinae. We are also able to identify 'rogue' taxa whose positions are weakly supported (the different gene regions are in conflict with each other) and unstable. Our main conclusions are: (1) the tribe Coeini as currently constituted is untenable, and Smyrna, Colobura and Tigridia are part of Nymphalini; (2) 'Kallimini' is paraphyletic with regard to Melitaeini and should be split into three tribes: Kallimini s.s., Junoniini and Victorinini; (3) Junoniini, Victorinini, Melitaeini and the newly circumscribed Nymphalini are strongly supported monophyletic  groups, and (4) Precis and Junonia are not synonymous or even sister groups. The species Junonia coenia, a model; system in developmental biology, clearly belongs in the genus Junonia. A dispersal-vicariance analysis suggests that dispersal has had a major effect on the distributions of extant species, and three biotic regions are identified as being centres of diversification of three major clades: the Palaearctic for the Nymphalis-group, the Afrotropics for Junoniini and the Nearctic for Melitaeini.

citation:
Wahlberg, N., Brower, A.V.Z. and Nylen, S. 2005. Phylogenetic relationships and historical biogeography of tribes and genera in the subfamily Nymphalinae (Lepidoptera: Nymphalidae). Biological Journal of the Linnean Society 86: 227-251.


author:
Auger-Rozenberg, M.-A.; Kerdelhué, C.; Magnoux, E.; Turgeon, J.; Rasplus, J.-Y.; Roques, A.

title:
Molecular phylogeny and evolution of host-plant use in conifer seed chalcids in the genus Megastigmus (Hymenoptera: Torymidae)

abstract:
Phylogenetic relationships amongst Megastigmus species (Chalcidoidea: Torymidae) associated with conifer seeds were inferred from DNA sequence data. Twenty-nine species of seed chalcids were analysed using two different genes, cytochrome b (mitochondrial DNA) and the D2 domain of the 28S ribosomal DNA. Maximum-parsimony and maximum-likelihood analyses showed that taxa formed two monophyletic groups, one clade comprising all species associated with Cupressaceae and Taxodiaceae hosts with the exception of Chamaecyparis, and the other clade composed of species associated with Pinaceae. Species infesting Cupressaceae and Taxodiaceae seemed to be specialized to particular host genera or even to be species specific, which was consistent with a taxonomic radiation following initial host adaptation. By contrast, Megastigmus species associated with; Pinaceae appeared capable of shifting onto different congeneric species or even onto a new host genus, with their evolution apparently less constrained by plant association. We hypothesized that the Megastigmus group associated with Pinaceae may have a much higher invasive potential than that related to Cupressaceae. The study also confirmed the presence of invasive Nearctic species in the Palaearctic, and demonstrated the existence of a cryptic species complex.

citation:
Auger-Rozenberg, M.-A., Kerdelhue, C., Magnoux, E., Turgeon, J., Rasplus, J.-Y. and Roques, A. 2006. Molecular phylogeny and evolution of host-plant use in conifer seed chalcids in the genus Megastigmus (Hymenoptera: Torymidae). Systematic Entomology 31: 47-64.


author:
Yen, S.-H.; Robinson, G.S.; Quicke, D.L.J.

title:
Phylogeny, systematics and evolution of mimetic wing patterns of Eterusia mothsa (Lepidoptera, Zygaenidae, Chalcosiinae)

citation:
Yen, S.-H., Robinson, G.S. and Quicke, D.L.J. 2005. Phylogeny, systematics and evolution of mimetic wing patterns of Eterusia mothsa (Lepidoptera, Zygaenidae, Chalcosiinae). Systematic Entomology 30: 358-397.


title:
Phylogenetic analysis of the Abana group of genera (Hemiptera: Cicadellidae: Cicadellinae: Proconiini)

citation:
Ceotto, P.C. and Mejdalani, G. 2005. Phylogenetic analysis of the Abana group of genera (Hemiptera: Cicadellidae: Cicadellinae: Proconiini). Systematic Entomology 30: 480-496.


title:
Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy

abstract:
The hemipteran infraorder Cicadomorpha comprises the superfamilies Cicadoidea (cicadas), Cercopoidea (spittlebugs or froghoppers) and  Membracoidea (leafhoppers and treehoppers). Earlier attempts to determine relationships among these three monophyletic lineages using either morphological or molecular data suffered from insufficient sampling (taxonomic and data) and problematic tree rooting, leading to discordant results. Presented here are phylogenetic reconstructions within Cicadomorpha based on DNA nucleotide sequence data from multiple genetic markers (18S rDNA, 28S rDNA, and histone 3) sequenced from representative taxa of Cicadidae, Tettigarctidae, Cercopidae, Aphrophoridae, Clastopteridae, Machaerotidae, Epipygidae, Cicadellidae, Membracidae, Myerslopiidae and Aetalionidae. To test the robustness of the phylogenetic signal, these sequence data were analysed separately and in combination under various alignment parameters using both manual alignment (of both attenuated and full sequences) and alignment via CLUSTAL X. The results demonstrate clearly that, despite the alignment method used, basing a phylogeny on a single gene region is often misleading. Analyses of the combination of datasets support the major relationships within Cicadomorpha as (Membracoidea (Cicadoidea, Cercopoidea)). Internal relationships recovered within each superfamily shows evidence for: (1) the placement of Myerslopiidae as the sister group of the remaining Membracoidea; (2) the paraphyly of Cicadellidae; (3) the sister-group relationship between Machaerotidae and Clastopteridae; (4) the monophyly of Cercopidae; (5) the diversification of Epipygidae from within the possibly paraphyletic Aphrophoridae.

citation:
Cryan, J.R. 2005. Molecular phylogeny of Cicadomorpha (Insecta: Hemiptera: Cicadoidea, Cercopoidea and Membracoidea): adding evidence to the controversy.  30: 563-574.


title:
Cladistic analysis of moon moths using morphology, molecules, and behaviour: Actias Leach, 1815; Argema Wallengren, 1858; Graellsia Grote, 1896 (Lepidoptera: Saturniidae)

abstract:
A phylogenetic analysis of 16 moon moth species was conducted using morphology, molecules and behavior.  A data matrix of morphology and behaviour together was comprised of 93 characters from the larvae, pupae, cocoons, and adults in all ingroup species and two outroups.  Molecular data included 2662 nucleotides from elongation factor 1-alpha (EF 1-a) and dopa decarboxylase (DDC) protein coding nuclear genes of six ingroups and the two outgroups.  Both data matrices were analyzed separately, compared, and combined.  The total evidence analysis includeing all characters reveals the following generic relationships: (outgourps (Argema(Graellsia+Actias))). Character evolution indicates that the short hindwing tail evolved once and lengthened multiple times in different lineages of moon moths. Results support retaining Graellsia as a genus separate from Actias.

citation:
Ylla, J., Peigler, R.S. and Kawahara, A.Y. 2005. Cladistic analysis of moon moths using morphology, molecules, and behaviour: Actias Leach, 1815; Argema Wallengren, 1858; Graellsia Grote, 1896 (Lepidoptera: Saturniidae). SHILAP Revista de Lepidopterologia 33: 299-317.


title:
Biochemical genetic differentiation among seed chalcid species of genus Megastigmus (Hymenoptera: Torymidae)

citation:
Roux, G. and Roques, A. 1996. Biochemical genetic differentiation among seed chalcid species of genus Megastigmus (Hymenoptera: Torymidae). Experientia 52: 522-530.


author:
Kerdelhué, C.; Le Clainche, I.; Rasplus, J.-Y.

title:
Molecular phylogeny of the Ceratosolen species pollinating Ficus of the subgenus Sycomorus sensu stricto: biogeographical history and origins of the species-specificity breakdown cases

citation:
Kerdelhue', C., Le Clainche, I. and Rasplus, J.-Y. 1999. Molecular phylogeny of the Ceratosolen species pollinating Ficus of the subgenus Sycomorus sensu stricto: biogeographical history and origins of the species-specificity breakdown cases. Molecular Phylogenetics and Evolution 11: 401-414.


author:
Rasplus, J.-Y.; Kerdelhué, C.; Le Clainche, I.; Mondor, G.

title:
Molecular phylogeny of fig wasps: Agaonidae are not monophyletic

journal:
Comptes Rendus de l'Académie des Sciences Série III - Sciences de la Vie-Life Science

citation:
Rasplus, J.-Y., Kerdelhué, C., Le Clainche, I. and Mondor, G. 1998. Molecular phylogeny of fig wasps: Agaonidae are not monophyletic. Comptes Rendus de l'Académie des Sciences Série III - Sciences de la Vie-Life Science 321: 517-527.


title:
Phylogenetic relationships and character evolution among selected species of Bactrocera (Diptera: Tephritidae) based on multiple mitochondrial genes

citation:
Smith, P.T., Kambhampati, S. and Armstrong, K.A. 2005. Phylogenetic relationships and character evolution among selected species of Bactrocera (Diptera: Tephritidae) based on multiple mitochondrial genes. Insect Systematics and Evolution 36: 343-360.


title:
Phylogenetic relationships within the fruit fly genus Ceratitis MacLeay (Diptera: Tephritidae), derived from morphological and host plant evidence

citation:
De Meyer, M. 2005. Phylogenetic relationships within the fruit fly genus Ceratitis MacLeay (Diptera: Tephritidae), derived from morphological and host plant evidence. Insect Systematics and Evolution 36: 459-480.


author:
Brown, K.S.; Trigo, J.R.; Francini, A.B.B.; Motta, P.C.

title:
Aposematic insects on toxic host plants: coevolution, colonization, and chemical emancipation

editor:
Price, P.W.; Lewinsohn, T.M.; Fernandes, G.W.; Ben

citation:
Brown, K.S., Trigo, J.R., Francini, A.B.B. and Motta, P.C. 1991. Aposematic insects on toxic host plants: coevolution, colonization, and chemical emancipation. Plant-Animal Interactions: Evolutionary Ecology in Tropical and Temperate Regions : .

taxkey:
Ithomiinae; Heliconiinae; Troidini; Papilionidae; Nymphalidae; Papilionoidea


title:
Origin and radiation of the Auchenorrhyncha

series:
Proceedings, Thomas Say Publications in Entomology

citation:
Blocker, H.D. 1996. Origin and radiation of the Auchenorrhyncha. Studies on Hemipteran Phylogeny : 46-64.


title:
Adjustment of gene flow at the population, species, and ecosystem level: thistles and their herbivores

journal:
Flux Control in Biological Systems: From Enzymes to Populations and Ecosystems

citation:
Jensen, U. and Zwölfer, H. 1994. Adjustment of gene flow at the population, species, and ecosystem level: thistles and their herbivores. Pp. 447-467 in: Schulze, E.-D. [ed.], Flux Control in Biological Systems: From Enzymes to Populations and Ecosystems. Academic Press, San Diego.


title:
Sequential evolution of Euphilotes (Lycaenidae: Scolitantidini) on their plant host Eriogonum (Polygonaceae: Eriogonoideae)

abstract:
The relationship between the butterfly genus Euphilotes and their host plant genus Eriogonum in western North America is suggested to be one of sequential evolution rather than coevolution.  Eriogonum, a genus of nearly 250 species, probably had a Miocene origin, but has had its modern distribution significantly influenced by recent Pleistocene glatiaion.  The evolution of Euphilotes, as a distinct genus of four sibling species, apparetnly postdates the establishment and recetn proliferaqtion of Eriogonum.  Successful speciation in Euphilotes has been accomplished mainly through modifications in genitalia of those butterflies using a single species of Eriogonum.  The subsequent proliferation of Euphilotes subspecies has been the result of host switching coupled with geographic isolation onto individual species of Eriogonum acting as restricted biogeographic islands.  In the first instance, direct evolutionary competition for a limited resource (one species of Eriogonum) leads to partitioning of that resource by the butterflies whose entire life cycle is associated with that plant species.  In the second instance, host switching and isolation have permitted establishment of minor subspecies without significant interaction with other subspecies of the same species.  In instances where interspecific subspecies competition exists, resource partitioning, coupled with more pronounced genetic isolation, seem to have occurred resulting in more readily distinct subspecies.  We speculate that the success of subspeciation in Euphilotes is dependent upon the numberic size and geographic extent of the host species.  Euphilotes subspecies on plants of restricted distributions are themselves seemingly limited in their evolutionary potential as the most dynamic evolution of Euphilotes subspecies is that associated with widespread and variable Eriogonum species.  In all instances, the tempo and mode of evolution in Euphilotes appears to be sequential as it follows and is seemingly dependent upon what has already occurred in Eriogonum.

citation:
Shields, O. and Reveal, J.L. 1988. Sequential evolution of Euphilotes (Lycaenidae: Scolitantidini) on their plant host Eriogonum (Polygonaceae: Eriogonoideae). Biological Journal of the Linnean Society 33: 51-93.


author:
Als, T.D.; Vila, R.; Kandul, N.P.; Nash, D.R.; Yen, S.-H.; Hsu, Y.-F.; Mignault, A.A.; Boomsma, J.J.

title:
The evolution of alternative parasitic life histories in large blue butterflies

abstract:
Large blue (Maculinea) butterflies are highly endangered throughout the Palaearctic region, and have been the focus of intense conservation research.  In addition, their extraordinary parasitic lifestyles make them ideal for studies of life history evolution. Early instars consume flower buds of specific host plants, but later instars live in ant nests where they either devour the brood (predators), or are fed mouth-to-mouth by the adult ants (cuckoos).  Here we present the phylogeny for the group, which shows that it is a monophyletic clade nested within Phengaris, a rare Oriental genus whose species have similar life histories. Cuckoo species are likely to have evolved from predatory ancestors.  As early as five million years ago, two Maculinea clades diverged, leading to the different parasititc strategies seen in the genus today.  Contrary to current belief, the two recognized cuckoo species show little genetic divergence and are probably a single ecologically differentiated species.  On the other hand, some of the predatory morphospecies exhibit considerable genetic divergence and may contain cryptic species. These findings have important implications for conservation and reintroduction efforts.

citation:
Als, T.D., Vila, R., Kandul, N.P., Nash, D.R., Yen, S.-H., Hsu, Y.-F., Mignault, A.A., Boomsma, J.J. and Pierce, N. 2004. The evolution of alternative parasitic life histories in large blue butterflies. Nature 432: 386-390.


author:
Nyman, T.; Zinovjev, A.G.; Vikberg, V.; Farrell, B.D.

title:
Molecular phylogeny of the sawfly subfamily Nematinae (Hymenoptera: Tenthredinidae)

abstract:
Nematinae is one of the largest subfamilies in the sawfly family Tenthredinidae, but internal relationships are unknown in the absence of any formal phylogenetic analysis. To understand the internal phylogeny of Nematinae, we sequenced a portion of the mitochondrial cytochrome oxidase I gene and the nuclear elongation factor-1a gene from thirteen outgroup taxa and sixty-eight nematine species, the ingroup taxa of which represent all major genera and subgenera within the subfamily. Maximum parsimony and Bayesian phylogenetic analyses of the DNA sequence data show that: (1) Nematinae are monophyletic in a broad sense which includes Hoplocampa, Susana and the tribe Cladiini, which have been classified often into separate subfamilies; together with Craterocercus, these taxa form a paraphyletic basal grade with respect to the remaining Nematinae, but among- group relationships within the grade remain weakly resolved; (2) the remainder of the ingroup, Nematinae s. str., is monophyletic in all combined-data analyses; (3) within Nematinae s. str., the 'Higher' Nematinae is divided into three groups, Mesoneura and the large tribes Nematini and Pristiphorini; (4) although the traditional classifications at the tribal level are largely upheld, some of the largest tribes and genera are obviously para- or polyphyletic; (5) according to rate-smoothed phylogenies dated with two fossil calibration points, Nematinae originated 50-120 million years ago. In addition, the results from all Bayesian analyses provide strong and consistent support for the monophyly of Tenthredinidae, which has been difficult to demonstrate in  previous parsimony analyses of morphological and molecular data.


author:
Ronsted, N.; Weiblen, G.D.; Cook, J.M.; Salamin, N.; Machado, C.A.; Savolainen, V.

title:
60 million years of co-divergence in the fig-wasp symbiosis

abstract:
Figs (Ficus; ca 750 species) and fig wasps (Agaoninae) are obligate mutualists: all figs are pollinated by agaonines that feed exclusively on figs. This extraordinary symbiosis is the most extreme example of specialization in a plant-pollinator interaction and has fuelled much speculation about co-divergence. The hypothesis that pollinator specialization led to the parallel diversification of fig and pollinator lineages (co-divergence) has so far not been tested due to the lack of robust and comprehensive phylogenetic hypotheses for both partners. We produced and combined the most comprehensive molecular phylogenetic trees to date with fossil data to generate independent age estimates for fig and pollinator lineages, using both non-parametric rate smoothing and penalized likelihood dating methods. Molecular dating of ten pairs of interacting lineages provides an unparalleled example of plant-insect co-divergence over a geological time frame spanning at least 60 million years.

citation:
Ronsted, N., Weiblen, G.D., Cook, J.M., Salamin, N., Machado, C.A. and Savolainen, V. 2005. 60 million years of co-divergence in the fig-wasp symbiosis. Proceedings of the Royal Society B 272: 2593-2599.


title:
Montane speciatiion patterns in Ithomiola butterflies (Lepidoptera: Riodinidae): are they consistently moving up in the world?

abstract:
Tropical lowland areas have often been seen as the centres of terrestrial species proliferation, but recent evidence suggests that young species may be more frequent in montane areas. Several montane speciation modes have been proposed, but their relative frequencies and predominant evolutionary sequence remain unclear because so few biogeographic and phylogenetic studies have tested such questions. I use morphological data to generate a phylogenetic hypothesis for all 11 species of the riodinid butterfly genus Ithomiola (Riodininae: Mesosemiini: Napaeina). These species are shown here to be all strictly geographically and elevationally allo- or parapatrically distributed with respect to their closest relatives in lowland and montane regions throughout the Neotropics. The overwhelming pattern in Ithomiola is of repeated upward parapatric speciation across an elevational gradient, and the genus appears to provide the clearest example to date of vertical montane speciation. All of the young derived species are montane and all of the old basal species are confined to the lowlands, supporting the hypothesis of montane regions largely as 'species pumps' and lowland regions as 'museums'. Possible reasons for the post-speciation maintenance of parapatric ranges in Ithomiola are discussed.

citation:
Hall, J.P.W. 2005. Montane speciatiion patterns in Ithomiola butterflies (Lepidoptera: Riodinidae): are they consistently moving up in the world?. Proceedings of the Royal Society B 272: 2457-2466.


title:
Phylogenetic framework for Dioryctria (Lepidoptera: Pyralidae: Phycitinae) based on combined analysis of mitochondrial DNA and morphology

abstract:
Coneworms of the genus Dioryctria Zeller are important lepidopterous pests of conifer cones throughout the Holarctic region. Seventy-nine Dioryctria species are currently recognized and arranged into 11 species groups, but a globally unified classification of these species groups has not been attained. We surveyed 14 Dioryctria species belonging to 7 species groups recognized as being taxonomically problematic. Mitochondrial DNA sequences and morphological characters were used to resolve relationships among and within species groups and species. Sequences were obtained for 2.3 kb of the mitochondrial COI+COII genes and related to 52 morphological characters...

citation:
Du, Y., Roe, A. D. and Sperling, F. A. H. 2005. Phylogenetic framework for Dioryctria (Lepidoptera: Pyralidae: Phycitinae) based on combined analysis of mitochondrial DNA and morphology. Canadian Entomolgist 137: 685-711.


title:
Biology, ecology, and evolution of gall-inducing sawflies (Hymenoptera: Tenthredinidae and Xyelidae)

editor:
Raman, A.; Schaefer, C. W.; Withers, T. M.

citation:
Roininen, H., Nyman, T. and Zinovjev, A. 2005. Biology, ecology, and evolution of gall-inducing sawflies (Hymenoptera: Tenthredinidae and Xyelidae). Pp. 467-494 in: Raman, A., Schaefer, C. W. and Withers, T. M. [eds.], Biology, Ecology, and Evolution of Gall-inducing Arthropods. Science Publishers, Enfield, NH.


title:
Biology, ecology, and evolution of gall-inducing Cynipidae

citation:
Csóka, G., Stone, G. and Melika, G. 2005. Biology, ecology, and evolution of gall-inducing Cynipidae. Pp. 573-642 in: Raman, A., Schaefer, C. W. and Withers, T. M. [eds.], Biology, Ecology, and Evolution of Gall-inducing Arthropods. Science Publishers, Enfield, NH.


abstract:
Miners are concealed feeding phytophagous insect larvae which live on particular plant tissue, but without an ability to modify plant growth. In this chapter special attention has been paid to morphological adaptations of dipteran larvae to the leaf-mining feeding habit. Due to morphological constraints, the larvae of phytophagous Diptera are prone to endophytic feeding behavior. This is especially true for the subgroup Cyclorrhapha. Morphological patterns of phytophagy in dipteran larvae in general, and leaf miners in particular have been outlined.;   The phylogeny of the predominantly leaf-mining dipteran family Agromyzidae has been used to test recent theories on the adaptive significance of the leaf-mining habit. The agromyzid species are specialized to various plant parts, and several species also have the ability to induce galls. Character optimization on the phylogenetc tree revealed that leaf-mining is ancestral in Agromyzidae. Such other internal feeding habits as stem or seed boring emerged repeatedly in various agromyzid clades. These results are consistent with ecological theories. However, observations of larval morphology and phylogeny also yield different levels of specialization within the leaf-mining habit, which clearly shows that leaf mining is a viable lifestyle within extant phytophagous communities.;   Gall inducers among Agromyzidae have been examined to obtain evidence for leaf mining being ancestral to gall induction. Although probable transformations from leaf miners to gall inducers can be assumed, gall inducers are species poor within developent except for plants that respond extraordinarily actively. Possible explanations and selective advantages of nongalling over galling species are suggested.

citation:
Dempewolf, M. 2005. Dipteran Leaf Miners. Pp. 407-429 in: Raman, A., Schaefer, C. W. and Withers, T. M. [eds.], Biology, Ecology, and Evolution of Gall-inducing Arthropods. Science Publishers, Enfield, NH.


title:
Towards a phylogenetic system of derelomine flower weevils (Coleoptera: Curculionidae)

abstract:
A revised phylogenetic classification for derelomine flower weevils (Coleoptera: Curculionidae: Curculioninae: Derelomini Lacordaire) is proposed, based on a cladistic analysis of 115 outgroup and ingroup taxa and 155 primarily morphological characters. The single most-parsimonious cladogram (length = 271, consistency index = 65, retention index = 95) indicates that several genera must be excluded from the tribe, as they lack certain modifications of the mouthparts and a primary reproductive association with the inflorescences of palms. These include Araucarietus Kuschel, Eisingius Kuschel, and Planus Kuschel, new placements (now all Trypetidini); Euryscapoides Wibmer & O'Brien, new placement (Curculioninae incertae sedis); Neopsilorhinus Bovie, new placement (Erirhinidae: Erirhinini); and Pedetinus Faust, new placement (Eugnomini). Five subtribes are recognized within Derelomini. The oldest African and South American Derelomina have a carinate rostrum and lamellate dorsal scales. Grasidius Champion (previously Erirhinini) and Terires Champion (Storeini) are placed therein, new placements.Neoderelomus Hoffmann is nested within Derelomus Schoenherr, syn.n., and its only species is therefore renamed as D. piriformis (Hoffmann), comb.n. The predominantly Asian Acalyptina are reduced from tribal status, stat.n., to now be part of Derelomini, new placement. They are defined (inter alia) by truncate elytra. They also contain Eudela Pascoe and Eudelodes Zimmermann, new placements (previously Curculioninae incertae sedis). The New World Notolomina, subtr.n., are distinguished by bifurcate maxillary lacinial teeth and other mouthpart characters. Andranthobius mariahelenae (Bondar) (formerly Derelomus), comb.n., is added to this entity. Phyllotrogina, subtr.n., are a very diverse and mostly Neotropical lineage. They have two-segmented maxillary palps, a densely pubescent prosternum, and long macrosetae along the basal fifth of the posterior wing margin. Within the subtribe there are independent transitions to host plant associations with various dicots (Phyllotrox Schoenherr), Cyclanthaceae (e.g. Perelleschus Wibmer & O'Brien and Systenotelus Anderson & Gómez), and Araceae (Cyclanthura Franz). Their life history traits show related changes, including losses of the ability to pollinate, and more detrimental (herbivorous, seed-predating) larval developments (e.g. Cotithene Voss). Phyllotrox tatianae (Bondar) (formerly Derelomus), comb.n., is assigned to Phyllotrogina. Hypoleschus Fall is nested within Phyllotrox, syn.n., and thereby the latter now contains P. atratus (Fall), comb.n. The Central American palm-associated Androtrox Franz, gen.n., is proposed to accommodate A. megalops (Champion) (formerly Phyllotrox), comb.n., a species with large contiguous eyes and a set of mouthpart attributes convergently present in Andranthobius Kuschel (Notolomina) and Phyllotrox. The Neotropical Staminodeina, subtr.n., with a labial prementum with two triangular projections and a large anal lobe on the wing, probably represent a young clade of weevils specialized to oviposit into the ephemeral staminodes of Cyclanthaceae inflorescences. The apparent phylogenetic sequence of the subtribes is (Derelomina ((Acalyptina, Staminodeina) (Notolomina, Phyllotrogina))), with a Juano- rhinini-Trypetidini clade as the most immediate outgroup, followed by various curculionine lineages. The revised system thus improves the taxonomy of derelomine weevils, and clarifies our understanding of their roles in the evolution of several tropical plant lineages - palms and cyclanths in particular.

citation:
Franz, N.M. 2006. Towards a phylogenetic system of derelomine flower weevils (Coleoptera: Curculionidae). Systematic Entomology 31: 220-287.


title:
The green-veined white (Pieris napi L.), its Pierine relatives, and the systematics dilemmas of divergent character sets (Lepidoptera, Pieridae)

abstract:
The butterfly Pieris napi (L.) and relatives exemplify recently evolving taxa, exhibiting variation that makes their evolutionary dynamics interesting, but their systematics difficult. Wing-pattern characters commonly used to distinguish these Holarctic insects display both genetic polymorphism and environmentally-cued polyphenism. Often, these causes of variation are confounded, impairing the characters' phylogenetic usefulness. DNA sequences of four mitochondrial genes offer an independent view of pierine phylogeny. Sampling diverse relatives within family Pieridae assists resolution of the P. napi complex, suggests previous underestimation of clade diversity in subfamily Pierinae, and shows that other genera near Pieris also display confusions of wing-pattern-based phylogenetic inference. The European P. napi is sister to all North American taxa, and is well diverged from them all in sequences. The North American taxa comprise a northern subclade including Pieris oleracea, and questionably distinct Pieris ; 'angelika', and a southern subclade including distinct Pieris virginiensis, Pieris marginalis, and Pieris macdunnoughii, and other regional entities yet to be clarified. Weak bootstrap support for some nodes in this group arises from a closeness of sequence identity rather than character conflict; more sequence data and denser geographical sampling may resolve these nodes more clearly. Evidence of reproductive isolation, from other experimental hybridization studies, agrees with the DNA results where these conflict with other divergent character sets. The system offers much promise for a deeper understanding of character evolution in relation to phyletic differentiation.

citation:
Chew, F.S. and Watt, W.B. 2006. The green-veined white (Pieris napi L.), its Pierine relatives, and the systematics dilemmas of divergent character sets (Lepidoptera, Pieridae). Biological Journal of the Linnean Society 88: 413-435.


title:
A preliminary cladistic analysis of Nipponoserica, with implications for phylogenetic relationships among sericine chafers (Coleoptera: Scarabaeidae:; Sericini)

abstract:
A cladistic analysis of the species of Nipponoserica Nomura, 1972, was executed using 36 morphological charactersofadults.Fromthegenerated phylogenetic tree, the monophyly of the genus is supported, when Pseudosericania Kobayashi, 1980 is included. Consequently, Pseudosericania Kobayashi, 1980 is considered a junior subjective synonym of Nipponoserica Nomura, 1972. Interestingly, the analysis indicates the monophyly of the East Asian Nipponoserica and the North American Stilbolemma. A new genus, Xenoserica gen. nov., is established for the basal monophyletic clade found for the Himalayan species so far assigned to Nipponoserica including the first brachypterous species of Sericini from the Himalaya. Two new; species are described: Xenoserica brachyptera sp. nov. from Nepal, and Nipponoserica dahongshanica sp. nov. from Hubei (China).

citation:
Ahrens, D. 2005. A preliminary cladistic analysis of Nipponoserica, with implications for phylogenetic relationships among sericine chafers (Coleoptera: Scarabaeidae: Sericini). Systematics and Biodiversity 3: 265-279.


author:
Brower, A.V.Z.; Freitas, A.V.L.; Lee, M.-M.; Silva-Brandao, K.L.

title:
Phylogenetic relationships among the Ithomiini; (Lepidoptera: Nymphalidae) inferred from one; mitochondrial and two nuclear gene regions

abstract:
A phylogenetic hypothesis for the tribe Ithomiini (Lepidoptera: Nymphalidae: Danainae) is presented, based on sequences of the mitochondrial; cytochrome oxidase subunits I and II (COI-COII) region and regions of the nuclear genes wingless and Elongation factor 1-alpha. Branch support for each clade is assessed, and a partition congruence index is used to explore conflict among gene regions. The monophyly of the clade is strongly supported, as are many of the traditionally recognized subtribes and genera. The data imply paraphyly of some genera and tribes, but largely support recent classifications and phylogenetic hypotheses based on morphological characters.

citation:
Brower, A.V.Z., Freitas, A.V.L., Lee, M.-M. and Silva-Brandao, K.L. 2006. Phylogenetic relationships among the Ithomiini (Lepidoptera: Nymphalidae) inferred from one mitochondrial and two nuclear gene regions. Systematic Entomology 31: 288-301.


author:
Bugrov, A.; Novikova, O.; Mayorov, V.; Adkison, L.; Blinov, A.

title:
Molecular phylogeny of Palaearctic genera of; Gomphocerinae grasshoppers (Orthoptera, Acrididae)

abstract:
Molecular phylogenetic methods were used to examine morphologically based hypotheses concerning the taxonomic structure and relationships of the; grasshopper subfamily Gomphocerinae. Two mitochondrial gene (cytochrome b and cytochrome oxidase subunit I) sequences were determined for twenty-five species representing eleven Palaearctic genera. The studied Gomphocerinae species constituted a monophyletic group; furthermore, the earlier division of Gomphocerinae into tribes was supported, with each tribe monophyletic. There was no support for various systems uniting Stenobothrini and Gomphocerini into one tribe. Two separate clusters were discerned in Gomphocerini and two tribes were distinguished - Gomphocerini (genera Aeropus, Stauroderus, Chorthippus); and Stenobothrini (genera Omocestus, Stenobothrus).

citation:
Bugrov, A., Novikova, O., Mayorov, V., Adkison, L. and Blinov, A. 2006. Molecular phylogeny of Palaearctic genera of; Gomphocerinae grasshoppers (Orthoptera, Acrididae). Systematic Entomology 31: 362-368.


title:
Systematics and evolution of Iolania (Hemiptera:; Fulgoromorpha: Cixiidae) from Hawai'i

abstract:
Within the planthopper taxon Cixiidae, which is distributed worldwide, only two lineages have colonized the Hawaiian Islands: Iolania Kirkaldy, 1902, and Oliarus Staøª l, 1862, and subsequently given rise to endemic species. Neither radiation has been studied in depth hitherto. Here the degree of speciation within Iolania is assessed and a taxonomic revision including a key to the species based on the male genitalic characters is provided. Six endemic species are recognized: I. perkinsi Kirkaldy, I. koolauensis Giffard, I. oahuensis Giffard, I. lanaiensis Giffard, I. mauiensis Giffard and I. kraussohana sp.n. A lectotype is designated for the type-species I. perkinsi Kirkaldy, and I. perkinsi notata Kirkaldy is interpreted as an invalid name. Morphological arguments for the monophyly of Iolania and phylogenetic relationships among the species are discussed. A plausible scenario for the sequence of speciation events and history of colonization within the Hawaiian Islands is attempted. Combined information from taxon- and area-cladograms suggests progressive inter-island dispersal from older to younger islands in the Hawaiian chain as the major pattern of colonization; and speciation.

citation:
Hoch, H. 2006. Systematics and evolution of Iolania (Hemiptera: Fulgoromorpha: Cixiidae) from Hawai'i. Systematic Entomology 31: 302-320.


title:
A molecular phylogenetic study of armoured scale; insects (Hemiptera: Diaspididae)

abstract:
Armoured scale insects are economically important parasites of woody plants and grasses. They are promising subjects for the evolutionary study of physiology (no complete gut), genetics (chimerism, paternal genome elimination, frequent parthenogenesis) and coevolution (with host plants, parasitoids, Septobasidium fungi, endosymbiotic bacteria). Little phylogenetic work has been accomplished with armoured scales, and uncertainty surrounds their classification. Here, we report the phylogenetic results of Bayesian and parsimony analyses of 705 base pairs of Elongation Factor 1a and 660 base pairs of 28S from eighty-nine species of armoured scale insects, representing forty-seven genera and five tribes in the subfamilies Diaspidinae and Aspidiotinae, together with two outgroups. 28S was aligned based on a secondary structural model. Our results broadly corroborate the major features of the existing classification, although we do not find perfect monophyly of any of the traditionally recognized subfamilies or tribes. The subfamily Aspidiotinae is paraphyletic with respect to the subfamily Diaspidinae. Diaspidinae consists of two main clades that only roughly correspond to the tribes Lepidosaphidini and Diaspidini. Diaspidini is nearly monophyletic, except that it includes a single aspidiotine species. Other members of the tribe Aspidiotini form a clade, except that the clade includes a single species of Leucaspidini and excludes Maskellia and Pseudaonidia. Our results weakly support the hypothesis that the most recent common ancestor of the Diaspididae had adult females that were permanently enclosed within the derm of the second instar (the pupillarial habit) and had diploid adult males that eliminated their paternal genomes during spermatogenesis (late paternal genome elimination).

citation:
Morse, G.E. and Normark, B.B. 2006. A molecular phylogenetic study of armoured scale insects (Hemiptera: Diaspididae). Systematic Entomology 31: 338-349.


author:
Niehuis, O.; Yen, S.-H.; Naumann, C.M.; Misof, B.

title:
Higher phylogeny of zygaenid moths (Insecta: Lepidoptera) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence

abstract:
Zygaenid moths are capable of releasing hydrogen cyanide in their defense by enzymatic break-down of cyanoglucosides, but only larvae of chalcosiine and zygaenine moths store cyanogenic compounds in cuticular cavities and thus are able to discharge defense droplets, which effectively deter potential predators. A previously proposed phylogeny of Zygaenidae hypothesized a sister group relationship of chalcosiine and zygaenine moths because of their similar larval defense system. Not all chalcosiine taxa possess cuticular cavities, however, and a comparable defense mechanism has been reported in larvae of the zygaenoid family Heterogynidae. Considering sequence data of seven molecular loci, the present study estimates the posterior probability of phylogenetic hypotheses explaining the occurrence of larval cuticular cavities. The molecular data confirm the previous exclusion of Himantopteridae from Zygaenidae and suggest their close affinity to Somabrachyidae. The sequence data also corroborate the recently proposed exclusion of the Phaudinae from the Zygaenidae, because this subfamily is recovered in a reasonably well supported species cluster consisting of members of the families Lacturidae, Limacodidae, Himantopteridae, and Somabrachyidae. We consequently agree to raise Phaudinae to family rank. Within Zygaenidae, the subfamilies Callizygaeninae, Chalcosiinae, and Procridinae most likely constitute a monophyletic group, which is sister to the Zygaeninae. Our results imply that cuticular cavities were probably present in the larvae of the most recent common ancestor of Zygaenidae. Heterogynidae cannot be confirmed as sister taxon to this family, but appear at the very first split of the Zygaenoidea, although with poor support. The specific pattern of taxa in the molecular phylogeny showing larval cuticular cavities opens the possibility that these structures could have been already present in the most recent common ancestor of the Zygaenoidea.

citation:
Niehuis, O., Yen, S.-H., Naumann, C.M. and Misof, B. 2006. Higher phylogeny of zygaenid moths (Insecta: Lepidoptera) inferred from nuclear and mitochondrial sequence data and the evolution of larval cuticular cavities for chemical defence. Molecular Phylogenetics and Evolution 39: 812-829.


title:
A phylogenetic reassessment of Hyalenna Forbes and Dircenna Doubleday, with a revision of Hyalenna; (Lepidoptera: Nymphalidae: Ithomiinae)

abstract:
A morphological study of adults and immature stages and a cladistic analysis were conducted to clarify the relationships amongst members of the sister nymphalid butterfly genera Dircenna Doubleday and Hyalenna Forbes. Two species formerly included in Dircenna, Ithomia paradoxa Staudinger and Dircenna hugia Schaus, clustered with Ithomia perasippa, the type species of Hyalenna. Therefore, the first two species are transferred from Dircenna to Hyalenna (comb.n.). Hyalenna and Dircenna as now conceived are both monophyletic and well supported by morphological apomorphies and branch support measures. The taxonomy and classification of Hyalenna are reviewed and seven species and twenty-four taxa are recognized, including one new species, Hyalenna buckleyi sp.n., and eleven new subspecies: Hyalenna paradoxa incachaca ssp.n., H. perasippa valencia ssp.n., H. perasippa ortygiosa ssp.n., H. perasippa solitaria ssp.n., H. buckleyi pomacocha ssp.n., H. alidella exsulans ssp.n., H. alidella cinereola ssp.n., H. alidella vesca ssp.n., H. sulmona balsamica ssp.n., H. sulmona hyalina ssp.n. and H. sulmona tersa ssp.n. A key for the identification of all; taxa, and illustrations of male and female genitalia for all species and adult specimens of all taxa (where known), are presented. The natural history of each species is summarized and the immature stages of H. paradoxa and H. sulmona are described for the first time.

citation:
Willmott, K. R. and Lamas, G. 2006. A phylogenetic reassessment of Hyalenna Forbes and Dircenna Doubleday, with a revision of Hyalenna (Lepidoptera: Nymphalidae: Ithomiinae). Systematic Entomology 31: 419-468.


title:
Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae: Limenitis)

abstract:
The evolution of wing pattern diversity in butterXies has emerged as a model system for understanding the origins and maintenance of adaptive phenotypic novelty. Admiral butterXies (genus Limenitis) are an attractive system for studying wing pattern diversity because mimicry is common among the North American species and hybrid zones occur wherever mimetic and non-mimetic wing pattern races meet. However, the utility of this system has been limited because the evolutionary relationships among these butterXies remain unclear. Here I present a robust species-level phylogeny of Limenitis based on 1911 bp of two mitochondrial genes (COI and COII) and 904 bp of EF1- for all Wve of the Nearctic species/wing pattern races, the majority of the Palearctic species, and three outgroup genera; Athyma, Moduza (Limenitidini), and Neptis (Limenitidinae: Neptini). Maximum-likelihood and Bayesian analyses indicate that the North American species are a well-supported, monophyletic lineage that is most closely related to the widespread, Palearctic, Poplar admiral (L. populi). Within North America, the Viceroy (L. archippus) is the basal lineage while the relationships among the remaining species are not well resolved. A combined maximum-likelihood analysis, however, indicates that the two western North America species (L. lorquini and L. weidemeyerii) are sister taxa and closely related to the wing pattern subspecies of the polytypic Limenitis arthemis species complex.; These results are consistent with (1) an ancestral host-shift to Salicaceae by the common ancestor of the Poplar admiral and the Nearctic admiral lineage, (2) a single colonization of the Nearctic, and (3) a subsequent radiation of the North American forms leading to at least three independent origins of mimicry.

citation:
Mullen, S.P. 2006. Wing pattern evolution and the origins of mimicry among North American admiral butterflies (Nymphalidae: Limenitis). Molecular Phylogenetics and Evolution 39: 747-758.


title:
Linking patterns and processes of species diversification in the cone flies Strobilomyia; (Diptera: Anthomyiidae)

abstract:
Phytophagous insects provide useful models for the study of ecological speciation. Much attention has been paid to host shifts, whereas situations where closely related lineages of insects use the same plant during different time periods have been relatively neglected in previous studies of insect diversification. Flies of the genus Strobilomyia are major pests of conifers in Eurasia and North America. They are specialized feeders in cones and seeds of Abies (fir), Larix (larch) and Picea (spruce). This close association is accompanied by a large number of sympatric Strobilomyia species coexisting within each tree genus. We constructed a molecular phylogeny with a 1320 base-pair fragment of mitochondrial DNA that demonstrated contrasting patterns of speciation in larch cone flies, as opposed to spruce and fir cone flies; this despite their comparable geographic distributions and similar resource quality of the host. Species diversity is the highest on larch, and speciation is primarily driven by within-host phenological shifts, followed by allopatric speciation during geographical expansion. By contrast, fewer species exploit spruce and fir, and within-host phenological shifts did not occur. This study illustrates within-host adaptive radiation through phenological shifts, a neglected mode of sympatric speciation.

citation:
Sachet, J.-M., Roques, A. and Després, L. 2006. Linking patterns and processes of species diversification in the cone flies Strobilomyia (Diptera: Anthomyiidae). Molecular Phylogenetics and Evolution: in press.


title:
Tropical forests are both evolutionary cradles and museums of leaf beetle diversity

abstract:
The high extant species diversity of tropical lineages of organisms is usually portrayed as a relatively recent and rapid development or as a consequence of the gradual accumulation or preservation of species over time. These explanations have led to alternative views of tropical forests as evolutionary ''cradles'' or ''museums'' of diversity, depending on the organisms under study. However, biogeographic and fossil evidence implies that the evolutionary histories of diversification among tropical organisms may be expected to exhibit characteristics of both cradle and museum models.; This possibility has not been explored in detail for any group of terrestrial tropical organisms. From an extensively sampled molecular phylogeny of herbivorous Neotropical leaf beetles in the genus Cephaloleia, we present evidence for (i) comparatively ancient Paleocene-Eocene adaptive radiation associated with global warming and Cenozoic maximum global temperatures, (ii) moderately ancient lineage-specific diversification coincident with; the Oligocene adaptive radiation of Cephaloleia host plants in the genus Heliconia, and (iii) relatively recent Miocene-Pliocene diversification; coincident with the collision of the Panama arc with South America and subsequent bridging of the Isthmus of Panama.These results demonstrate that, for Cephaloleia and perhaps other lineages of organisms, tropical forests are at the same time both evolutionary cradles and museums of diversity.

citation:
McKenna, D.D. and Farrell, B.D. 2006. Tropical forests are both evolutionary cradles and museums of leaf beetle diversity. Proceedings of the National Academy of Sciences, USA 103: 10947-10951.


author:
Nyman, T.; Farrell, B.D.; Zinovjev, A.G.; Vikberg, V.

title:
Larval habits, host-plant associations, and speciation in nematine sawflies (Hymenoptera: Tenthredinidae)

citation:
Nyman, T., Farrell, B.D., Zinovjev, A.G. and Vikberg, V. 2006. Larval habits, host-plant associations, and speciation in nematine sawflies (Hymenoptera: Tenthredinidae). Evolution : (in press).


author:
Shapiro, L.H.; Strazanac, J.S.; Roderick, G.K.

title:
Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago

abstract:
The extant endemic katydids (Orthoptera: Tettigoniidae) of the Hawaiian Archipelago include one to three species per high island and a single species on Nihoa, all currently placed in the genus Banza. These acoustic insects provide an excellent opportunity for investigating the evolution of reproductive isolation and speciation, but such studies require an understanding of phylogenetic relationships within the group. We use maximum parsimony, likelihood-based Bayesian inference, and maximum likelihood to infer phylogenetic relationships among these taxa, based on ~2 kb of; mitochondrial cytochrome oxidase I and cytochrome-b. Our results strongly support two distinct high island clades: one clade ("Clade I") composed of species from Kauai, Oahu, Molokai, and Lanai and another clade ("Clade II") composed of species from Maui and Hawaii (B. unica, from Oahu, may be basal to both of these clades, but its placement is not well resolved). Within these clades, some inferred relationships are strongly supported, such as the sister status of B. kauaiensis (Kauai) and B. parvula (Oahu) within Clade I, but other relationships remain more ambiguous, such as the relative position of B. brunnea (Maui) within Clade II. Although a detailed reconstruction of the historical biogeography of the Hawaiian katydids is difficult, we; use our genetic data combined with the known geological history of the Hawaiian Islands to set limits on plausible historical scenarios for diversification of this group. Beyond these historical biogeographic inferences, our results indicate possible cryptic speciation on both Oahu and Hawaii, as well as what may be unusually high average rates of nucleotide substitution. The present work sets the stage for future genetic and experimental investigations of this group.

citation:
Shapiro, L.H., Strazanac, J.S. and Roderick, G.K. 2006. Molecular phylogeny of Banza (Orthoptera: Tettigoniidae), the endemic katydids of the Hawaiian Archipelago. Molecular Phylogenetics and Evolution : in press.


title:
Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing the re-interpretation of morphology and a revised generic classification

abstract:
The scale insect tribe Iceryini (Coccoidea: Monophlebidae) is a group of relatively large and polyphagous insects found worldwide. Currently, the tribe contains about 80 named species placed in seven genera, which are diagnosed largely on features associated with egg protection. We reconstruct the phylogeny of the Iceryini on the basis of nucleotide sequence data from nuclear ribosomal (18S and D2, D3 and D10 regions of 28S) and protein-coding (histone H3) gene regions of 40 iceryine species representing six of the seven genera and seven outgroup taxa, mostly from two other tribes of Monophlebidae. Bayesian and maximum parsimony analyses recover a monophyletic tribe and clades that correspond more to geography than to the existing morphology-based classification. Gueriniella Fernald is sister to the rest of the Iceryini and the genera Crypticerya Cockerell, Icerya Signoret and Steatococcus Ferris are not monophyletic. Our data imply that the distinctive iceryine reproductive strategies, such as protecting eggs in a waxy ovisac or inside a marsupium, are poor indicators of relationships. On the basis of molecular relationships and the re-examination of morphological characters, we recognize only five genera of Iceryini - Crypticerya, Echinicerya Morrison, Gigantococcus Pesson & Bielenin, Gueriniella and Icerya - and substantially revise the generic concepts of Crypticerya, Gigantococcus and Icerya. We provide a key to the genera based on adult females. We redescribe and illustrate the adult female and first-instar nymph of the type species Crypticerya rosae (Riley & Howard), Echinicerya anomala Morrison, Gigantococcus maximus (Newstead) (adult female only), Gueriniella serratulae (Fabricius) and Icerya seychellarum (Westwood). We recognize Auloicerya Morrison as a junior synonym (syn.n.) of Icerya, and transfer the two Auloicerya species to Icerya as I. acaciae (Morrison & Morrison) comb.n. and I. australis Maskell comb.rev. We recognize Steatococcus and Proticerya Cockerell as junior synonyms (syn.n.) of Crypticerya. From Steatococcus, we transfer five species to Crypticerya [C. mexicana Cockerell & Parrott comb.rev., C. morrilli (Cockerell) comb.n., C. tabernicola (Ferris) comb.n., C. townsendi Cockerell comb.rev., C. tuberculata (Morrison) comb.n.], four species to Gigantococcus [Gi. euphorbiae (Brain) comb.n., Gi. gowdeyi (Newstead) comb.n., Gi. madagascariensis (Mamet) comb.n., Gi. theobromae (Newstead) comb.n.] and three species to Icerya [I. assamensis (Rao) comb.n., I nudata Maskell comb.rev., I. samaraia (Morrison) comb.n.]. From Icerya, we transfer 14 species to Crypticerya [C. brasiliensis (Hempel) comb.n., C. colimensis (Cockerell) comb.n., C. flava (Hempel) comb.n., C. flocculosa (Hempel) comb.n., C. genistae (Hempel) comb.n., C. littoralis (Cockerell) comb.n., C. luederwaldti (Hempel) comb.n., C. minima (Morrison) comb.n., C. montserratensis (Riley & Howard) comb.n., C. palmeri (Riley & Howard) comb.n., C. rileyi (Cockerell) comb.n., C. similis (Morrison) comb.n., C. subandina (Leonardi) comb.n., C. zeteki (Cockerell) comb.n.] and nine species to Gigantococcus [Gi. alboluteus (Cockerell) comb.n., Gi. bimaculatus (De Lotto) comb.n., Gi. brachystegiae (Hall) comb.n., Gi. longisetosus (Newstead) comb.n., Gi. nigroareolatus (Newstead) comb.n., Gi. pattersoni (Newstead) comb.n., Gi. schoutedeni (Vayssière) comb.n., Gi. splendidus (Lindinger) comb.n., Gi. sulfureus (Lindinger) comb.n.]. From Crypticerya, we transfer seven species to Icerya [I. clauseni (Rao) comb.n., I. jacobsoni Green comb.rev., I. jaihind (Rao) comb.n., I. kumari (Rao) comb.n., I. mangiferae (Tang & Hao) comb.n., I. natalensis (Douglas) comb.rev., I. nuda Green comb.rev.] and five species to Gigantococcus [Gi. bicolor (Newstead) comb.n., Gi. cajani (Newstead) comb.n., Gi. caudatus (Newstead) comb.n., Gi. ewarti (Newstead) comb.n., Gi. rodriguesi (Castel-Branco) comb.n.]. Both I. hyperici (Froggatt) and Palaeococcus dymocki (Froggatt) are syn.n. of I. nudata (all previously placed in Steatococcus). We recognize I. maynei Vayssière as a syn.n. of Gi. nigroareolatus, I. tremae Vayssière as a syn.n. of Gi. schoutedeni and I. townsendi plucheae Cockerell as a syn.n. of C. townsendi. We revalidate the species name I. crocea Green stat.reval. In addition, we transfer I. taunayi Hempel to Laurencella Foldi (Monophlebidae: Llaveiini) as L. taunayi (Hempel) comb.n. Four species, Coccus hirticornis Boyer de Fonscolombe, I. chilensis Hempel, I. insulans Hempel and I. paulista Hempel, are considered incertae sedis. We designate lectotypes for C. rosae, E. anomala and I. candida (a junior synonym of I. seychellarum). Following this revision, we recognize 74 species of Iceryini, distributed as follows: 22 in Crypticerya, one in Echinicerya, 19 in Gigantococcus, two in Gueriniella and 30 in Icerya.

citation:
Unruh, C.M 2008. Molecular data reveal convergent reproductive strategies in iceryine scale insects (Hemiptera: Coccoidea: Monophlebidae), allowing the re-interpretation of morphology and a revised generic classification. Systematic Entomology 33: 8-50.


author:
Hardy, N.B.; Gullan, P.J.; Hodgson, C.J.

title:
A subfamily-level classification of mealybugs (Hemiptera: Pseudococcidae) based on integrated molecular and morphological data

abstract:
The mealybugs (Hemiptera: Coccoidea: Pseudococcidae) are a speciose and ubiquitous group of sap-sucking plant parasites, many of which are very serious agricultural pests. There has been much work on the alpha-level taxonomy, amounting to the description of more than 2000 species, but suprageneric relationships remain poorly known. Downie & Gullan reviewed proposed schemes for a mealybug subfamily-level classification and used DNA sequence data from three nuclear genes to infer the mealybug phylogeny. They recognized three subfamilies: the Rhizoecinae, the Phenacoccinae, and the Pseudococcinae; and within the Pseudococcinae, recognized the tribes Pseudococcini, Trabutinini, and Planococcini. Excepting the Trabutinini, none of these groupings was well-supported. We improve our estimation of the mealybug phylogeny by: (i) increasing the taxon sampling by 50%; (ii) adding a morphological character matrix; and (iii) performing mixed model Bayesian and maximum likelihood inference procedures. We recover two primary clades within the Pseudococcidae, to which we apply the subfamily names Phenacoccinae and Pseudococcinae. Within the Phenacoccinae, we recover support for the monophyly of the hypogaeic mealybugs (Rhizoecini), and within the Pseudococcinae, we find support for the tribes Pseudococcini, Trabutinini, and Planococcini. Our results suggest a clear sequence of (i) ß-Proteobacteria primary endosymbionts infecting the mycetome of the most recent common ancestor of the Pseudococcinae, followed by (ii) ?-Proteobacteria secondary endosymbionts infecting the primary ß-Proteobacteria endosymbionts. For each subfamily, we provide a morphological diagnosis and a comprehensive list of included genera. We resurrect the genus Ceroputo Sulc, currently under synonymy with the genus Puto Signoret (Putoidae), and place it in the mealybug subfamily Phenacoccinae. Greenoripersia kaiseri Bodenheimer is transferred from the Pseudococcidae to the Eriococcidae.

citation:
Hardy, N.B., Gullan, P.J. and Hodgson, C.J. 2008. A subfamily-level classification of mealybugs (Hemiptera: Pseudococcidae) based on integrated molecular and morphological data. Systematic Entomology 33: 51-71.


author:
Regier, J.C.; Cook, C.P.; Mitter, C.; Hussey, A.

title:
A phylogenetic study of the 'bombycoid complex' (Lepidoptera) using five protein-coding nuclear genes, with comments on the problem of macrolepidopteran phylogeny

abstract:
This study had two aims. First, we tested the monophyly of and relationships within the 'bombycoid complex', an assembly of approximately 5300 species postulated by Minet to represent 12 families in three superfamilies, by sequencing five protein-coding nuclear gene regions (CAD, DDC, enolase, period, wingless; approximately 6750 bp total) in 66 representatives of most of the subfamilies and tribes. Second, we sought initial evidence on the utility of these genes for estimating relationships among Macrolepidoptera more broadly (11 superfamilies total), by adding representatives of eight families from four other superfamilies, and by assessing the phylogenetic information content of the individual genes and partitions thereof. Analysis of the combined data by likelihood and parsimony upholds monophyly for the bombycoid complex and for Bombycoidea sensu stricto (includes Anthelidae, see below), but with weak bootstrap support. Minet's assignment of Phiditiinae to Bombycoidea rather than to Noctuoidea is strongly upheld, but Anthelidae, placed in Lasiocampoidea by Minet, group securely within Bombycoidea sensu stricto. Within the latter, the basal split segregates a strongly supported 'BALE' group [Apatelodinae + (Eupterotidae + (Brahmaeidae + Lemoniidae))]. The remaining families form a consistently but weakly supported clade, within which the basal split segregates the very strongly supported 'CAPOPEM' group [Carthaeidae, Anthelidae, Phiditiinae, (Prismostictini + (Endromidae + (Oberthueriini + Mirinidae)))]. The remaining bombycoids are grouped, very weakly, as Sphingidae + (Bombycinae + Saturniidae). All multiply-sampled families are strongly recovered, in both outgroups and ingroups, except that Bombycidae sensu Minet are rendered decisively polyphyletic. All genes make important contributions to the combined data results, and there is little strong conflict among genes or between synonymous and nonsynonymous change, although two instances of inter-gene conflict were notable, one in Lasiocampidae and one in Mimallonidae. Overall, about 75% of nodes are strongly supported (i.e. bootstrap value =80%). Superfamilies are recovered, but not always strongly, whereas relationships among superfamilies are recovered only weakly and inconsistently; even within the reasonably well-sampled Bombycoidea sensu stricto, a (to us) surprising number of interfamily relationships remain uncertain. Thus, it seems clear that substantially more genes, plus additional taxon sampling in most superfamilies, will be required to resolve macrolepidopteran phylogeny.

citation:
Regier, J.C., Cook, C.P., Mitter, C. and Hussey, A. 2008. A phylogenetic study of the 'bombycoid complex' (Lepidoptera) using five protein-coding nuclear genes, with comments on the problem of macrolepidopteran phylogeny. Systematic Entomology 33: 175-189.


title:
Molecular phylogeny of Anthelidae and other bombycoid taxa (Lepidoptera: Bombycoidea)

abstract:
Based on DNA sequences of the fusion protein carbamoylphosphate synthetase/aspartate transcarbamylase/dihydroorotase (CAD; 680 bp) and elongation factor-1a (Ef-1a; 1240 bp); the first molecular phylogeny of the moth family Anthelidae and its placement within the Bombycoidea sensu Brock (1971) (= bombycoid complex sensu Minet, 1994) is proposed. The results strongly support the monophyly of the family Anthelidae and its subfamilies Munychryiinae and Anthelinae, but demonstrate the vast polyphyly of its main genus Anthela Walker, 1855. The proposed phylogeny suggests that grass feeding, as apparent from some pest records, probably is an ancestral trait within the subfamily Anthelinae. Evolutionary relationships of the family Anthelidae and of most parts of the Bombycoidea remain obscure. However, the results contradict many of the widely accepted phylogenetic hypotheses within the Bombycoidea proposed by Minet (1994: Entomologica scandinavica, 25, 63-88). The Brahmaeidae are paraphyletic relative to the Lemoniidae (syn.nov.), and the current concept of Bombycidae is polyphyletic, with the bombycid subfamily Apatelodinae being part of a monophylum comprising Brahmaeidae / Lemoniidae, Eupterotidae and Apatelodidae (stat.rev.).

citation:
Zwick, A. 2008. Molecular phylogeny of Anthelidae and other bombycoid taxa (Lepidoptera: Bombycoidea). Systematic Entomology 33: 190-209.


author:
Acs, Z.; Melika, G.; Pénzes, Z.; Pujade-Villar, J.; Stone, G.N.

title:
The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini)

abstract:
Oak gallwasps (Hymenoptera; Cynipidae, tribe Cynipini) are cyclically parthenogenetic insects that induce galls on specific plant hosts in the family Fagaceae. Understanding the processes underlying the evolution of specific oak associations requires knowledge of the phylogenetic relationships among oak gallwasp genera. Although three major lineages of oak gallwasps have been identified, the status and relationships of several species-poor but biologically significant genera remain unresolved. Two such genera are Chilaspis and Dryocosmus, whose western palaearctic species all gall oaks in the section Cerris. Dryocosmus is particularly significant biologically because it includes: (a) the only palaearctic gallwasp to gall chestnuts, Castanea, and (b) nearctic species. The oak section Cerris is wholly absent from the nearctic, and the relationship between palaearctic and nearctic Dryocosmus is significant for patterns of host plant evolution in the tribe as a whole. We examined the relationships between Chilaspis, Dryocosmus and other oak cynipid genera using cladograms from sequence data for two mitochondrial loci (cytochrome c oxidase subunit I and cytochrome b) and two nuclear loci (the 28S ribosomal gene regions D2 and D3-5). Our analyses support the following conclusions: (1) palaearctic Chilaspis and Dryocosmus species form an intermingled monophyletic group. (2) We propose that Chilaspis Mayr, 1881 is a syn.n. of Dryocosmus Giraud, 1859 and propose the name D. mayri as a comb.rev. for the species previously named C. mayri, and D. nitidus and D. israeli as comb.n. of C. nitida and C. israeli, respectively. (3) We reassess the utility of morphological characters previously regarded as diagnostic for these genera. (4) Two species previously known only from a single generation represent two halves of a single species lifecycle. Dryocosmus nervosus is here designated a syn.n. of D. cerriphilus. (5) The nearctic species D. favus lies outside the palaearctic Chilaspis/Dryocosmus clade, and Dryocosmus as currently recognized is not a monophyletic group. (6) Dryocosmus/Chilaspis is closely related to the other oak gallwasp taxa (Aphelonyx, Plagiotrochus, Pseudoneuroterus, Trichagalma, and some Neuroterus species) galling section Cerris oaks. This implies an early branching evolution of this oak association within this group, and supports previous work showing the rarity of oak gallwasp host shifts.; ;

citation:
Acs, Z., Melika, G., Pénzes, Z., Pujade-Villar, J. and Stone, G.N. 2007. The phylogenetic relationships between Dryocosmus, Chilaspis and allied genera of oak gallwasps (Hymenoptera, Cynipidae: Cynipini). Systematic Entomology 32: 70-80.


title:
Assessing the phylogenetic usefulness of a previously neglected morphological structure through elliptic Fourier analyses: a case study in Bruchus seed-beetles (Coleoptera: Chrysomelidae: Bruchinae)

abstract:
We address the systematics of Bruchus seed-beetles through the use of a geometric morphometric outline approach, namely elliptic Fourier analysis. We found that a previously neglected genitalic structure, the ventral plate, provides new evidence in the discussion of taxonomic issues raised by recent molecular studies. Three methods of hierarchical clustering allow investigation of the phylogenetic relationships of the key species that cause the paraphyly of two species groups in recent molecular studies. The resulting reconstructions reveal the phylogenetic usefulness of the structure of interest in recovering consistent relationships of Bruchus. Our analyses support the monophyly of the species group whose paraphyletic status was weakly supported by statistical tests in molecular analyses. Our results agree with those molecular and morphological studies that indicate, with relatively strong support, the paraphyletic status of the other species group. We highlight the need to reappraise the use of neglected or presumably uninformative (in traditional morphometrics) morphological characters with geometric morphometrics methods. In addition, we assess the utility of the combination of morphometric descriptors with other sources of phylogenetic information by analysing together an extant molecular dataset and matrix representations based on the results of the elliptic Fourier analyses (to our knowledge our study is the first to investigate such a combination of datasets within a Bayesian framework). Combining morphometric descriptors with other information can improve phylogenetic reconstructions, as suggested by the results of the corresponding analyses we performed using a published molecular dataset.

citation:
Kergoat, G.J. and Alvarez, N. 2008. Assessing the phylogenetic usefulness of a previously neglected morphological structure through elliptic Fourier analyses: a case study in Bruchus seed-beetles (Coleoptera: Chrysomelidae: Bruchinae). Systematic Entomology : (in press).


title:
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.; ;

citation:
Penz, C.M. 2007. Evaluating the monophyly and phylogenetic relationships of Brassolini genera (Lepidoptera, Nymphalidae). Systematic Entomology 32: 668-689.


title:
Systematic status of the genus Formosaphis Takahashi and the evolution of galls based on the molecular phylogeny of Pemphigini (Hemiptera: Aphididae: Eriosomatinae)

abstract:
Aphids of the tribe Pemphigini (Hemiptera: Aphididae: Eriosomatinae) can induce pseudo-galls or galls on their primary host plants. Those belonging to subtribe Prociphilina often produce pseudo-galls, and those of the other subtribe, Pemphigina, can form not only pseudo-galls but also true galls. Pseudo-galls are all formed on the leaves, whereas true galls, although all similar in shape, can be located on the joint of the leaf blade and the petiole, the middle of the petiole and the branches of the host plant. From a first phylogenetic analysis of Pemphigini based on nuclear elongation factor-1a (EF-1a) sequences, no support was found for the monophyly of Pemphigini, but subtribe Pemphigina was monophyletic with robust support. Formosaphis obviously clusters with Pemphigina which has Populus as a primary host. Formosaphis belongs to Pemphigini and its unknown primary host is probably Populus. The evolution of galls in Pemphigina is discussed based on the phylogenetic tree drawn from EF-1a sequences and mitochondrial cytochrome oxidase subunit I (COI) sequences. The results suggest that, in this subtribe, the closed gall is preceded by an open pseudo-gall, and the galls move their locations from the joint of the leaf blade and the petiole to the middle of the petiole and, eventually, to the branch of the host plant. Such an evolutionary tendency may provide aphids with more protection and nutrition.

citation:
Zhang, H. and Qiao, G. 2007. Systematic status of the genus Formosaphis Takahashi and the evolution of galls based on the molecular phylogeny of Pemphigini (Hemiptera: Aphididae: Eriosomatinae). Systematic Entomology 32: 690-699.


author:
Frick, Z.; Wahlberg, N.; Pech, P.; Zrzavy, J.

title:
Phylogeny and classification of the Phengaris-Maculinea clade (Lepidoptera: Lycaenidae): total evidence and phylogenetic species concepts

abstract:
Total evidence analysis, based on a combination of morphological and ecological characters with two mitochondrial sequences (cytochrome c oxidase subunits I and II) and one nuclear (elongation factor-1a) sequence, provides a new phylogeny of the uniquely obligate ant parasitic Phengaris-Maculinea butterflies. The clade, including all species of Maculinea and Phengaris, is very stable and well supported. However, various analyses suggest that either Phengaris or Maculinea is not monophyletic with regard to the other, which necessitates generic reclassification of the clade. Application of the diagnostic and monophyletic 'phylogenetic' species concepts leads to species-level reclassification, including ten species (P. alcon comb.n. including 'P. rebeli', P. daitozana, P. albida, P. atroguttata, P. kurentzovi comb.n., P. nausithous comb.n., P. teleius comb.n., P. arion comb.n., P. arionides comb.n., P. takamukui comb.n.) and one unresolved metaspecies ('P. cyanecula' comb.n.) in four monophyletic species groups. The existence of further or additional cryptic species is possible within P. nausithous and P. teleius. Maculinea Van Eecke, 1915 syn.n. is considered a junior synonym of Phengaris Doherty, 1891.

citation:
Frick, Z., Wahlberg, N., Pech, P. and Zrzavy, J. 2007. Phylogeny and classification of the Phengaris-Maculinea clade (Lepidoptera: Lycaenidae): total evidence and phylogenetic species concepts. Systematic Entomology 32: 558-567.


author:
Soulier-Perkins, A.; Ouvrard, D.; Attié, M.; Bourgoin, T.

title:
Evolutionary patterns in biogeography and host plant association: 'taxonomic conservatism' in Lophopidae (Hemiptera, Fulgoromorpha)

abstract:
The association between the Lophopidae (Hemiptera, Fulgoromorpha) and their host plants was studied within a phylogenetic framework. Host plant use was optimized on Lophopidae phylogeny and the most parsimonious hypothesis is presented. This hypothesis describes the evolution of host plant use by the Lophopidae, and postulates the ancestral plant family used. This scenario is discussed within the biogeographical evolutionary context of the fulgoromorphan families, and is corroborated by information from both insect and host plant fossils. The association of the Lophopidae and their host plants is made by comparing the angiosperms and Lophopidae phylogenies, demonstrating at this level of comparison that the insects show 'taxonomic conservatism' for their host plants.

citation:
Soulier-Perkins, A., Ouvrard, D., Attié, M. and Bourgoin, T. 2007. Evolutionary patterns in biogeography and host plant association: 'taxonomic conservatism' in Lophopidae (Hemiptera, Fulgoromorpha). Systematic Entomology 32: 305-311.


author:
Velázquez de Castro, A.J.; Alonso-Zarazaga, M.A.; Outerelo, R.

title:
Systematics of Sitonini (Coleoptera: Curculionidae: Entiminae), with a hypothesis on the evolution of feeding habits

abstract:
A new classification of Sitonini Gistel into ten genera is proposed, derived from a phylogenetic analysis based on morphology. Two subgenera of Sitona, Charagmus and Coelositona, are promoted to genus level. Sitona is restricted to the former nominotypical subgenus and redefined using apomorphies, and several species previously included in it are transferred to the genus Coelositona González. The species composition of the genus Charagmus Schönherr is unchanged. Andrion gen.n. is described (type species: Curculio regensteinensis Herbst, 1794). Catachaenus Schönherr, 1840 is synonymized with Eugnathus Schönherr, 1834 (syn.n.). Homalorhinus lutosus Hochhuth, 1847 is removed from Sitonini. The relationships represented by the new classification correspond with the evolution of Fabaceae (Leguminosae), mostly with the group of 'temperate herbaceous tribes' or Hologalegina. This group is divided into the sister clades, Loteae s.l. and IRLC (inverted repeat-lacking clade) - a clade well defined by the loss of a structural mutation in the genome of the chloroplast. Andrion does not feed on Hologalegina. Charagmus and Coelositona feed on species of Loteae s.l., whereas members of the genus Sitona feed on species in IRLC. This ability to feed on IRLC plants seems to have been a key trait that enabled the radiation of Sitona into more than 100 species. A key to the genera of Sitonini and illustrations of important morphological features are provided. A new structure is described from the internal sac, the 'hamuli'. The variability of hamuli between the species of Sitonini is an important new tool in the taxonomy of this tribe. New combinations: Eugnathus circulus (Eydoux & Souleyet, 1839), E. mangarinicus (Voss, 1925), E. scintillans (Pascoe, 1874), E. sulcifrons (Heller, 1934), Charagmus gressorius (Fabricius, 1792), Ch. intermedius Küster, 1847, Ch. griseus (Fabricius, 1775), Ch. cachectus Gyllenhal, 1834, Ch. stierlini Reitter, 1903, Ch. variegatus (Fåhraeus, 1840), Coelositona ribesi (González, 1971), C. cambricus (Stephens, 1831), C. cinerascens (Fåhraeus, 1840), C. limosus (Rossi, 1792), C. ocellatus (Küster, 1849), C. latipennis (Gyllenhal, 1834), C. puberulus (Reitter, 1903), C. villosus (Allard, 1869), Andrion regensteinense (Herbst, 1794) and Anemeroides lutosus (Hochhuth, 1847).

citation:
Velázquez de Castro, A.J., Alonso-Zarazaga, M.A. and Outerelo, R. 2007. Systematics of Sitonini (Coleoptera: Curculionidae: Entiminae), with a hypothesis on the evolution of feeding habits. Systematic Entomology 32: 312-331.


title:
Systematics, biogeography and diversification of the Indo-Australian genus Delias Hübner (Lepidoptera: Pieridae): phylogenetic evidence supports an 'out-of-Australia' origin

abstract:
Two alternative hypotheses for the origin of butterflies in the Australian Region, that elements dispersed relatively recently from the Oriental Region into Australia (northern dispersal hypothesis) or descended from ancient stocks in Gondwana (southern vicariance hypothesis), were tested using methods of cladistic vicariance biogeography for the Delias group, a diverse and widespread clade in the Indo-Australian Region. A phylogenetic hypothesis of the twenty-four species-groups recognized currently in Delias and its sister genus Leuciacria is inferred from molecular characters generated from the nuclear gene elongation factor-1 alpha (EF-1a) and the mitochondrial genes cytochrome oxidase subunits I and II (COI/COII) and NADH dehydrogenase 5 (ND5). Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined dataset (3888 bp, 1014 parsimony informative characters) confirmed the monophyly of Delias and recovered eight major lineages within the genus, informally designated the singhapura, belladonna, hyparete, chrysomelaena, eichhorni, cuningputi, belisama and nigrina clades. Species-group relationships within these clades are, in general, concordant with current systematic arrangements based on morphology. The major discrepancies concern the placement of the aganippe, belisama and chrysomelaena groups, as well as several species-groups endemic to mainland New Guinea. Two species (D. harpalyce (Donovan), D. messalina Arora) of uncertain group status are currently misplaced based on strong evidence of paraphyly, and are accordingly transferred to the nigrina and kummeri groups, respectively. Based on this phylogeny, a revised systematic classification is presented at the species-group level. An historical biogeographical analysis of the Delias group revealed that the most parsimonious reconstruction is an origin in the Australian Region, with at least seven dispersal events across Wallacea to the Oriental Region. The eight major clades of Delias appear to have diverged rapidly following complete separation of the Australian plate from Gondwana and its collision with the Asian plate in the late Oligocene. Further diversification and dispersal of Delias in the Miocene-Pliocene are associated with major geological and climatic changes that occurred in Australia-New Guinea during the late Tertiary. The 'out-of-Australia' hypothesis for the Delias group supports an origin of the Aporiina in southern Gondwana (southern vicariance hypothesis), which proposes that the ancestor of Delias + Leuciacria differentiated vicariantly on the Australian plate.;

citation:
Braby, M.F. and Pierce, N.E. 2007. Systematics, biogeography and diversification of the Indo-Australian genus Delias Hübner (Lepidoptera: Pieridae): phylogenetic evidence supports an 'out-of-Australia' origin. Systematic Entomology 32: 2-25.


title:
Out-of-Africa origin and dispersal-mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae)

abstract:
The relative importance of dispersal and vicariance in the diversification of taxa has been much debated. Within butterflies, a few studies published so far have demonstrated vicariant patterns at the global level. We studied the historical biogeography of the genus Junonia (Nymphalidae: Nymphalinae) at the intercontinental level based on a molecular phylogeny. The genus is distributed over all major biogeographical regions of the world except the Palaearctic. We found dispersal to be the dominant process in the diversification of the genus. The genus originated and started diversifying in Africa about 20 Ma and soon after dispersed into Asia possibly through the Arabian Peninsula. From Asia, there were dispersals into Africa and Australasia, all around 5 Ma. The origin of the New World species is ambiguous; the ancestral may have dispersed from Asia via the Beringian Strait or from Africa over the Atlantic, about 3 Ma. We found no evidence for vicariance at the intercontinental scale. We argue that dispersal is as important as vicariance, if not more, in the global diversification of butterflies.; ;

citation:
Kodandaramaiah, U. and Wahlberg, N. 2007. Out-of-Africa origin and dispersal-mediated diversification of the butterfly genus Junonia (Nymphalidae: Nymphalinae). Journal of Evolutionary Biology 20: 2181-2191.


title:
The taxonomy, biogeography and host plant relationships of jumping plant-lice (Hemiptera: Psyllidae) associated with creosote bushes (Larrea spp., Zygophyllaceae)

abstract:
The Neotropical genus Panisopelma (Psyllidae: Aphalaroidinae) is revised and its internal phylogeny analysed. The constituent species, including five new ones, are described and illustrated. Keys are provided for the adults and the last instar larvae. Eight species are associated with creosote bushes (Larrea, Zygophyllaceae): five with L. nitida and three with L. divaricata. There is evidence that another three species, the larvae of which are unknown, also develop on L. divaricata. Seven species are restricted to Argentina, one to Bolivia and three to Chile. The cladistic analysis based on male, female and larval morphological characters yielded a single most-parsimonious tree. The species associated with L. nitida form a monophyletic clade, those on L. divaricata, by contrast, are paraphyletic. One clade with three species is restricted to Argentina, but three clades each contain a species from Argentina and Chile. Although a close association exists between Panisopelma and Larrea, there is no evidence for cospeciation, but rather an initial shift from an unknown host to L. divaricata and a second shift from L. divaricata to L. nitida. In three species pairs of Panisopelma, the distribution patterns suggest geographical vicariance between Argentina and Chile.

citation:
Burckhardt, D. and Ouvrard, D. 2007. The taxonomy, biogeography and host plant relationships of jumping plant-lice (Hemiptera: Psyllidae) associated with creosote bushes (Larrea spp., Zygophyllaceae). Systematic Entomology 32: 136-155.


title:
Speciation in Pararge (Satyrinae: Nymphalidae) butterflies - North Africa is the source of ancestral populations of all Pararge species

abstract:
The genus Pararge comprises three species: P. aegeria, distributed in Europe and North Africa; P. xiphia, endemic to Madeira; and P. xiphioides, endemic to the Canary Islands. Two subspecies are recognized in P. aegeria, P. a. tircis and P. a. aegeria, distributed in northern and southern Europe, respectively. In the 1970s, P. aegeria appeared on Madeira. However, despite the status of P. aegeria as a model species in ecological studies, the evolutionary history of Pararge remains unknown. We studied the phylogenetic relationships of the three Pararge species, using the mitochondrial gene cytochrome oxidase subunit I and the nuclear gene wingless to infer modes and times of speciation. On the basis of our analyses, Pararge forms a strongly supported monophyletic group, with the DNA haplotypes of the three species also forming well-supported monophyletic groups. We found that P. xiphia diverged first from the common ancestor a maximum of five million years ago, with P. xiphioides and P. aegeria being sister species that diverged a maximum of three million years ago. The two subspecies, P. a. tircis and P. a. aegeria, were not distinguishable on the basis of DNA haplotypes; instead, our data clearly distinguished between European specimens and those from North Africa. Madeiran P. aegeria has North African haplotypes and thus originated from there rather than from Europe. We hypothesize that the Mediterranean Sea forms a strong barrier to dispersal for Pararge butterflies, and has done so for approximately the past one million years.

citation:
Weingartner, E., Wahlberg, N. and Nylin, S. 2006. Speciation in Pararge (Satyrinae: Nymphalidae) butterflies - North Africa is the source of ancestral populations of all Pararge species. Systematic Entomology 31: 621-632.


author:
Sultana, F.; Hu, Y.-G.; Toda, M.J.; Takenaka, K.; Yafuso, M.

title:
Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants

abstract:
Colocasiomyia, a moderate-sized genus in the subfamily Drosophilinae, comprises seventy (twenty-six described and forty-four undescribed) species. Several Colocasiomyia species have evolved intimate mutualisms with specific host plants, especially of the family Araceae: the flies depend throughout the entire life cycle, oviposition, larval growth, pupation, and adult feeding and mating, on inflorescences of their host plants, and in turn act as species-specific pollinators for their host plants. To understand the evolution of this mutualism between Colocasiomyia flies and their host plants, the phylogenetic relationships of this genus and some possibly related taxa are inferred from a cladistic analysis based on sixty-two characters of adult morphology. We conclude that Colocasiomyia is polyphyletic, with the C. arenga species group clearly separate. Colocasiomyia without the arenga group (Colocasiomyia proper) is sister to all other studied drosophilines, whereas the arenga group is relatively derived within the Drosophilinae. Within Colocasiomyia proper, four clades are recognized, three of which correspond to previously proposed species groups: the cristata, toshiokai and baechlii groups. The other clade, C. sp.1 aff. nepalensis + C. sp.2 aff. nepalensis, is defined as a new species group. Relationships amongst the four clades and three independent species (C. micheliae, C. gigantea and C. sp.K1) remain almost unresolved, except for a sister group relationship between the toshiokai and baechlii groups. The classification of species groups in Colocasiomyia is revised by erecting two new species groups (crassipes and zeylanica groups) in addition to the three known (baechlii, cristata and toshiokai) groups. Revision of the arenga group, which should be removed from Colocasiomyia, is left for future studies. The evolution of host plant selection in Colocasiomyia is discussed by mapping host plant taxa (families, subfamilies and tribes) on the phylogenetic tree deduced from the cladistic analysis. Cohabitation in the same host inflorescence by a pair of species with microallopatric niche separation on the spadix is hypothesized to have evolved independently at least more than twice in Colocasiomyia.; ;

citation:
Sultana, F., Hu, Y.-G., Toda, M.J., Takenaka, K. and Yafuso, M. 2006. Phylogeny and classification of Colocasiomyia (Diptera, Drosophilidae), and its evolution of pollination mutualism with aroid plants. Systematic Entomology 31: 684-702.


title:
Molecular phylogeny and historical biogeography of the Holarctic wetland leaf beetle of the genus Plateumaris

abstract:
Leaf beetles of the genus Plateumaris inhabit wetlands across the temperate zone of the Holarctic region. To explore the phylogeographic relationships among North American, East Asian, and European members of this genus and the origin of the species endemic to Japan, we studied the molecular phylogeny of 20 of the 27 species in this genus using partial sequences of mitochondrial cytochrome oxidase subunit I (COI) and the 16S and nuclear 28S rRNA genes. The molecular phylogeny revealed that three species endemic to Europe are monophyletic and sister to the remaining 11 North American and six Asian species. Within the latter clade, North American and Asian species did not show reciprocal monophyly. Dispersal-vicariance analysis and divergence time estimation revealed that the European and North America-Asian lineages diverged during the Eocene. Moreover, subsequent differentiation occurred repeatedly between North American and Asian species, which was facilitated by three dispersal events from North America to Asia and one in the opposite direction during the late Eocene through the late Miocene. Two Japanese endemics originated from different divergence events; one differentiated from the mainland lineage after differentiation from the North American lineage, whereas the other showed a deep coalescence from the North American lineage with no present-day sister species on the East Asian mainland. This study of extant insects provides molecular phylogenetic evidence for ancient vicariance between Europe and East Asia-North America, and for more recent (but pre-Pleistocene) faunal exchanges between East Asia and North America.

citation:
Sota, T., Bocak, L. and Hyashi, M. 2008. Molecular phylogeny and historical biogeography of the Holarctic wetland leaf beetle of the genus Plateumaris. Molecular Phylogenetics and Evolution 46: 183-192.


title:
Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae)

abstract:
The historical biogeography of insects in South America is largely unknown, as dated phylogenies have not been available for most groups. We have studied the phylogenetic relationships and historical biogeography of a subtribe of butterflies, Phyciodina in the family Nymphalidae, based on one mitochondrial gene (COI) and two nuclear gene regions (EF-1a and wingless). The subtribe comprises 89 species mainly found in tropical South America, with a few species in North America and the Greater Antilles. We find that the enigmatic genus Antillea is sister to the rest of Phyciodina, and suggest that it should be included in the subtribe. Several genera are found to be polyphyletic or nested within another genus, and are proposed to be synonymised. These are Dagon, Castilia, Telenassa and Janatella, which we propose should be synonymised with Eresia. Brazilian "Ortilia" form an independent lineage and require a new genus name. The diversification of Phyciodina has probably taken place over the past about 34 MYA. The ancestral phyciodine colonised South America from North America through a possible landspan that connected the Greater Antilles to South America about 34 MYA. A vicariance event left the ancestral Antillea on the Greater Antilles, while the ancestral 0e on South America colonised the Guyanan Shield and soon after the Brazilian Shield. We hypothesise that the Brazilian Shield was an important area for the diversification of Phyciodina. From there, the ancestor of Anthanassa, Eresia and Tegosa colonised NW South America, where especially Eresia diversified in concert with the rising of the Andes beginning about 20 MYA. Central America was colonised from NW South America about 15 MYA by the ancestors of Anthanassa and Phyciodes. Our study is the first to use a dated phylogeny to study the historical biogeography of a group of South American species of butterflies. ;

citation:
Wahlberg, N. and Freitas, A.V.L. 2007. Colonization of and radiation in South America by butterflies in the subtribe Phyciodina (Lepidoptera: Nymphalidae). Molecular Phylogenetics and Evolution 44: 1257-1272.


title:
Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes

abstract:
Geometridae is one of the most diverse families within the Lepidoptera, comprising nine subfamilies. Winter moths, which have a unique life history, are found in three subfamilies. To examine the phylogeny of the Geometridae at the subfamily level and determine the evolutionary history of winter moths, we constructed phylogenetic trees for all nine geometrid subfamilies using two mitochondrial and two nuclear gene sequences. Specimens of all subfamilies were sampled from Japan. Simultaneous analyses of the combined data from all genes revealed that the Geometridae comprised two major clades: one with subfamilies Larentiinae and Sterrhinae, and the other with the remaining seven subfamilies. The second clade included the largest subfamily, Ennominae, and the subfamily Archiearinae, which is traditionally considered to be an ancestral lineage of the Geometridae. The Larentiinae + Sterrhinae clade contained one winter moth lineage, and the second major clade consisted of three winter moth lineages, including Alsophilinae, which contains winter moths exclusively. Using a Bayesian inference of divergence times, we estimated that geometrids began to diverge 54 Mya (62-48 Mya), whereas winter moth lineages differentiated from non-winter moth lineages 34-12 Mya, during the global cooling events in the Oligocene and the early Miocene. The adaptation to cool climates may have been a preadaptation that facilitated the winter moth life cycle. ;

citation:
Yamamoto, S. and Sota, T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44: 711-723.


author:
Havill, N.P.; Foottit, R.G.; von Dohlen, C.D.

title:
Evolution of host specialization in the Adelgidae (Insecta: Hemiptera) inferred from molecular phylogenetics

abstract:
The Adelgidae form a small group of insects in the Aphidoidea. They are cyclically parthenogenetic with host alternating, multiple-generation complex life cycles and are restricted to certain host genera in the Pinaceae. Species that host alternate always have Picea as the primary host where sexual reproduction and gall formation occur, and another genus in the Pinaceae as the secondary host where a series of parthenogenetic generations are produced. Other species that do not host alternate complete their entire life cycle on one host and only reproduce parthenogenetically. We studied relationships within Adelgidae using DNA sequences from the mitochondrial COI, COII, and cytb genes, and the nuclear EF1a gene. Analysis of the combined data resulted in a well-resolved phylogeny in which the major adelgid clades correspond neatly to their association with secondary host genera. Specialization on each secondary host genus occurred only once and was followed by diversification on the host genus. Molecular dating of divergence times in the Adelgidae suggest that diversification among host genera occurred in the Late Cretaceous and Early Tertiary when the Pinaceae genera were diverging. It is not clear, however, whether the Adelgidae and Pinaceae co-diversified because the relationships among the Pinaceae genera are not fully resolved. We discuss implications for adelgid taxonomy, life cycle evolution, and evolution of the interaction between adelgids and their host plants. ;

citation:
Havill, N.P., Foottit, R.G. and von Dohlen, C.D. 2007. Evolution of host specialization in the Adelgidae (Insecta: Hemiptera) inferred from molecular phylogenetics. Molecular Phylogenetics and Evolution 44: 357-370.


author:
McLeish, M.J.; Crespi, B.J.; Chapman, T.W.; Schwarz, M.P.

title:
Parallel diversification of Australian gall-thrips on Acacia

abstract:
The diversification of gall-inducing Australian Kladothrips (Insecta: Thysanoptera) on Acacia has produced a pair of sister-clades, each of which includes a suite of lineages that utilize virtually the same set of 15 closely related host plant species. This pattern of parallel insect-host plant radiation may be driven by cospeciation, host-shifting to the same set of host plants, or some combination of these processes. We used molecular-phylogenetic data on the two gall-thrips clades to analyze the degree of concordance between their phylogenies, which is indicative of parallel divergence. Analyses of phylogenetic concordance indicate statistically-significant similarity between the two clades. Their topologies also fit with a hypothesis of some degree of host-plant tracking. Based on phylogenetic and taxonomic information regarding the phylogeny of the Acacia host plants in each clade, one or more species has apparently shifted to more-divergent Acacia host-plant species, and in each case these shifts have resulted in notable divergence in aspects of the phenotype including morphology, life history and behaviour. Our analyses indicate that gall-thrips on Australian Acacia have undergone parallel diversification as a result of some combination of cospeciation, highly restricted host-plant shifting, or both processes, but that the evolution of novel phenotypic diversity in this group is a function of relatively few shifts to divergent host plants. This combination of ecologically restricted and divergent radiation may represent a microcosm for the macroevolution of host plant relationships and phenotypic diversity among other phytophagous insects. ;

citation:
McLeish, M.J., Crespi, B.J., Chapman, T.W. and Schwarz, M.P. 2007. Parallel diversification of Australian gall-thrips on Acacia. Molecular Phylogenetics and Evolution 43: 714-725.


author:
Kergoat, G.J.; Silvain, J.-F.; Delobel, A.; Tuda, M.; Antone, K.-W.

title:
Defining the limits of taxonomic conservatism in host-plant use for phytophagous insects: Molecular systematics and evolution of host-plant associations in the seed-beetle genus Bruchus Linnaeus (Coleoptera: Chrysomelidae: Bruchinae)

abstract:
In this study, we have investigated the limits of taxonomic conservatism in host-plant use in the seed-beetle genus Bruchus. To reconstruct the insect phylogeny, parsimony and multiple partitioned Bayesian inference analyses were conducted on a combined data set of four genes. Permutation tests and both global and local maximum-likelihood optimizations of host preferences at distinct taxonomic levels revealed that host-fidelity is still discernible beyond the host-plant tribe level, suggesting the existence of more important than previously thought evolutionary constraints, which are further discussed in details. Our tree topologies are also mostly consistent with extant taxonomic groups. Through the analysis of this empirical data set we also provide meaningful insights on two methodological issues. First, Bayesian inference analyses suggest that partitioning by using codon positions greatly increase the accuracy of phylogenetical reconstructions. Regarding reconstruction of ancestral character states through maximum likelihood, the present study also highlights the usefulness of local optimizations. The issue of over-parameterization is also addressed, as the optimizations with the most parameter-rich models have returned the most counterintuitive results.

citation:
Kergoat, G.J., Silvain, J.-F., Delobel, A., Tuda, M. and Antone, K.-W. 2007. Defining the limits of taxonomic conservatism in host-plant use for phytophagous insects: Molecular systematics and evolution of host-plant associations in the seed-beetle genus Bruchus Linnaeus (Coleoptera: Chrysomelidae: Bruchinae). Molecular Phylogenetics and Evolution 43: 251-269.


author:
Coeur d'acier, A.; Jousselin, E.; Martin, J.-F.; Rasplus, J.-Y.

title:
Phylogeny of the Genus Aphis Linnaeus, 1758 (Homoptera: Aphididae) inferred from mitochondrial DNA sequences

abstract:
Aphis is the largest aphid genus in the world and contains several of the most injurious aphid pests. It is also the most reluctant aphid genus to any comprehensive taxonomic treatment: while most species are easily classified into "species groups" that form well defined entities, numerous species within these groups are difficult to tell apart morphologically and identification keys remain ambiguous and mostly rely on host plant affiliation. In this paper, we used partial sequences of COI/COII and CytB genes to reconstruct the first phylogeny of Aphis and discuss the present systematics. The monophyly of the subgenus Bursaphis and of the tree major species groups, Black aphid, Black backed aphid and frangulae-like species was recovered by all phylogenetic analyses. However our data suggested that the nominal subgenus was not monophyletic. Relationships between major species groups were often ambiguous but "Black" and "Black backed" species groups appeared as sister clades. The most striking result of this study was that our molecular data met the same limits as the morphological characters used in classifications: mitochondrial DNA did not allow the differentiation of species that are difficult to identify. Further, interspecies relationships within groups of species for which taxonomic treatment is difficult stayed unresolved. This suggests that species delineation in the genus Aphis is often ambiguous and that diversification might have been a rapid process.

citation:
Coeur d'acier, A., Jousselin, E., Martin, J.-F. and Rasplus, J.-Y. 2007. Phylogeny of the Genus Aphis Linnaeus, 1758 (Homoptera: Aphididae) inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution 42: 598-611.


author:
Scheffer, S.J.; Winkler, I.S.; Wiegmann, B.W.

title:
Phylogenetic relationships within the leaf-mining flies (Diptera: Agromyzidae) inferred from sequence data from multiple genes

abstract:
The leaf-mining flies (Diptera: Agromyzidae) are a diverse group whose larvae feed internally in leaves, stems, flowers, seeds, and roots of a wide variety of plant hosts. The systematics of agromyzids has remained poorly known due to their small size and morphological homogeneity. We investigated the phylogenetic relationships among genera within the Agromyzidae using parsimony and Bayesian analyses of 2965 bp of DNA sequence data from the mitochondrial COI gene, the nuclear ribosomal 28S gene, and the single copy nuclear CAD gene. We included 86 species in 21 genera, including all but a few small genera, and spanning the diversity within the family. The results from parsimony and Bayesian analyses were largely similar, with major groupings of genera in common. Specifically, both analyses recovered a monophyletic Phytomyzinae and a monophyletic Agromyzinae. Within the subfamilies, genera found to be monophyletic given our sampling include Agromyza, Amauromyza, Calycomyza, Cerodontha, Liriomyza, Melanagromyza, Metopomyza, Nemorimyza, Phytobia, and Pseudonapomyza. Several genera were found to be polyphyletic or paraphyletic including Aulagromyza, Chromatomyia, Phytoliriomyza, Phytomyza, and Ophiomyia. We evaluate our findings and discuss host-use evolution in light of current agromyzid taxonomy and two recent hypotheses of relationships based on morphological data.

citation:
Scheffer, S.J., Winkler, I.S. and Wiegmann, B.W. 2007. Phylogenetic relationships within the leaf-mining flies (Diptera: Agromyzidae) inferred from sequence data from multiple genes. Molecular Phylogenetics and Evolution 42: 756-775.


title:
Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea)

abstract:
The planthopper superfamily Fulgoroidea (Insecta: Hemiptera) comprises approximately 20 described insect families, depending on which classification is followed. Multiple competing hypotheses of fulgoroid phylogeny have been published, based on either morphological character coding or DNA sequence data; however, those hypotheses disagree in several key aspects regarding the evolution of planthoppers. The current paper seeks to test these hypotheses, including the Asche (Asche, M. 1987. Preliminary thoughts on the phylogeny of Fulgoromorpha (Homoptera Auchenorrhyncha). In: Proceedings of the 6th Auchenorrhyncha Meeting, Turin, Italy, 7-11 September, 1987, pp. 47-53.) hypothesis of a trend in ovipositor structure, which may be correlated with planthopper feeding ecology. Presented here are phylogenetic reconstructions of Fulgoroidea based on analysis of DNA nucleotide sequence data from four loci (18S rDNA, 28S rDNA, Histone 3, and Wingless) sequenced from 83 exemplar taxa representing 18 planthopper families and outgroups. Data sets were analyzed separately and in various combinations under the maximum parsimony criterion, and the total combined dataset was analyzed via both maximum parsimony and partitioned Bayesian criteria; results of the combined analyses were concordant across reconstruction paradigms. Relationships recovered suggest several major planthopper lineages, including: (1) Delphacidae + Cixiidae; (2) Kinnaridae + Meenoplidae; (3) Fulgoridae + Dictyopharidae; (4) Lophopidae + Eurybrachidae (possibly + Flatidae); (5) Ricaniidae + Caliscelidae (possibly + Tropiduchidae). Results also suggest the placement of Achilixiidae outside of Cixiidae and of Tettigometridae as one of the more recently diversified lineages within Fulgoroidea. The resulting phylogeny supports Asche's (1987) hypothesis of a functional trend in ovipositor structure across families. ;

citation:
Urban, J.M. and Cryan, J.R. 2007. Evolution of the planthoppers (Insecta: Hemiptera: Fulgoroidea). Molecular Phylogenetics and Evolution 42: 556-572.


author:
Nazari, V.; Zakharov, E.V.; Sperling, F.A.H.

title:
Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera, Papilionidae) based on morphology and seven genes

abstract:
We tested the taxonomic utility of morphology and seven mitochondrial or nuclear genes in a phylogenetic reconstruction of swallowtail butterflies in the subfamily Parnassiinae. Our data included 236 morphological characters and DNA sequences for seven genes that are commonly used to infer lepidopteran relationships (COI + COII, ND5, ND1, 16S, EF-1a, and wg; total 5775 bp). Nuclear genes performed best for inferring phylogenies, particularly at higher taxonomic levels, while there was substantial variation in performance among mitochondrial genes. Multiple analyses of molecular data (MP, ML and Bayesian) consistently produced a tree topology different from that obtained by morphology alone. Based on molecular evidence, sister-group relationships were confirmed between the genera Hypermnestra and Parnassius, as well as between Archon and Luehdorfia, while the monophyly of the subfamily was weakly supported. We recognize three tribes within Parnassiinae, with Archon and Luehdorfia forming the tribe Luehdorfiini Tutt, 1896 [stat. rev.]. Three fossil taxa were incorporated into a molecular clock analysis with biogeographic time constraints. Based on dispersal-vicariance (DIVA) analysis, the most recent common ancestor of Parnassiinae occurred in the Iranian Plateau and Central Asia to China. Early diversification of Parnassiinae took place at the same time that India collided into Eurasia, 65-42 million years ago. ;

citation:
Nazari, V., Zakharov, E.V. and Sperling, F.A.H. 2007. Phylogeny, historical biogeography, and taxonomic ranking of Parnassiinae (Lepidoptera, Papilionidae) based on morphology and seven genes. Molecular Phylogenetics and Evolution 42: 131-156.


author:
Arunkumar, K.P.; Metta, M.; Nagaraju, J.

title:
Molecular phylogeny of silkmoths reveals the origin of domesticated silkmoth, Bombyx mori from Chinese Bombyx mandarina and paternal inheritance of Antheraea proylei mitochondrial DNA

abstract:
Molecular phylogeny of some of the economically important silkmoths was derived using three mitochondrial genes, 12S rRNA, 16S rRNA, and COI, and the control region (CR). Maximum likelihood (ML) analyses showed two distinct clades, one consisting of moths from Bombycidae family and the other from Saturniidae family. The mitochondrial CR showed length polymorphisms with indels. The ML analyses for complete mitochondrial genome sequences of Bombyx mori (strains Aojuku, C108, Backokjam, and Xiafang), Japanese and Chinese strains of B. mandarina (Japanese mandarina and Chinese mandarina) and, Antheraea pernyi revealed two distinct clades, one comprising of B. mori strains and the other with B. mandarina, and A. pernyi forming an outgroup. Pairwise distances revealed that all of the strains of B. mori studied are closer to Chinese than to Japanese mandarina. Phylogenetic analyses based on whole mitochondrial genome sequences, the finding of a tandem triplication of a 126 bp repeat element only in Japanese mandarina, and chromosome number variation in B. mandarina suggest that B. mori must have shared its recent common ancestor with Chinese mandarina. Another wild species of the Bombycidae family, Theophila religiosa, whose phylogenetic status was not clear, clustered together with the other bombycid moths in the study. Analysis of the interspecific hybrid, A. proylei gave evidence for paternal inheritance of mitochondrial DNA.

citation:
Arunkumar, K.P., Metta, M. and Nagaraju, J. 2006. Molecular phylogeny of silkmoths reveals the origin of domesticated silkmoth, Bombyx mori from Chinese Bombyx mandarina and paternal inheritance of Antheraea proylei mitochondrial DNA. Molecular Phylogenetics and Evolution 40: 419-427.


author:
Peña, C.; Wahlberg, N.; Weingartner, E.; Kodandaramaiah, U.; Nylin, S.; Freitas, A.V.L.; Brower, A.V

title:
Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data

abstract:
We have inferred the first empirically supported hypothesis of relationships for the cosmopolitan butterfly subfamily Satyrinae. We used 3090 base pairs of DNA from the mitochondrial gene COI and the nuclear genes EF-1a and wingless for 165 Satyrinae taxa representing 4 tribes and 15 subtribes, and 26 outgroups, in order to test the monophyly of the subfamily and elucidate phylogenetic relationships of its major lineages. In a combined analysis, the three gene regions supported an almost fully resolved topology, which recovered Satyrinae as polyphyletic, and revealed that the current classification of suprageneric taxa within the subfamily is comprised almost completely of unnatural assemblages. The most noteworthy findings are that Manataria is closely related to Melanitini; Palaeonympha belongs to Euptychiina; Oressinoma, Orsotriaena and Coenonympha group with the Hypocystina; Miller's (1968). Parargina is polyphyletic and its components group with multiple distantly related lineages; and the subtribes Elymniina and Zetherina fall outside the Satyrinae. The three gene regions used in a combined analysis prove to be very effective in resolving relationships of Satyrinae at the subtribal and tribal levels. Further sampling of the taxa closely related to Satyrinae, as well as more extensive sampling of genera within the tribes and subtribes for this group will be critical to test the monophyly of the subfamily and establish a stronger basis for future biogeographical and evolutionary studies. ;

citation:
Peña, C., Wahlberg, N., Weingartner, E., Kodandaramaiah, U., Nylin, S., Freitas, A.V.L. and Brower, A.V.Z. 2006. Higher level phylogeny of Satyrinae butterflies (Lepidoptera: Nymphalidae) based on DNA sequence data. Molecular Phylogenetics and Evolution 40: 29-49.


title:
Adaptive radiation of gall-inducing insects within a single host-plant species

abstract:
Speciation of plant-feeding insects is typically associated with host-plant shifts, with subsequent divergent selection and adaptation to the ecological conditions associated with the new plant. However, a few insect groups have apparently undergone speciation while remaining on the same host-plant species, and such radiations may provide novel insights into the causes of adaptive radiation. We used mitochondrial and nuclear DNA to infer a phylogeny for 14 species of gall-inducing Asphondylia flies (Diptera: Cecidomyiidae) found on Larrea tridentata (creosote bush), which have been considered to be monophyletic based on morphological evidence. Our phylogenetic analyses provide strong support for extensive within-host plant speciation in this group, and it demonstrates that diversification has involved numerous shifts between different plant organs (leaves, buds, flowers, and stems) of the same host-plant species. Within-plant speciation of Asphondylia is thus apparently facilitated by the opportunity to partition the plant ecologically. One clade exhibits temporal isolation among species, which may have facilitated divergence via allochronic shifts. Using a novel method based on Bayesian reconstruction, we show that the rate of change in an ecomorphological trait, ovipositor length, was significantly higher along branches with inferred shifts between host-plant organs than along branches without such shifts. This finding suggests that Larrea gall midges exhibit close morphological adaptation to specific host-plant parts, which may mediate ecological transitions via disruptive selection.

citation:
Joy, J.B. and Crespi, B.J. 2007. Adaptive radiation of gall-inducing insects within a single host-plant species. Evolution 61: 784-795.


author:
Kandul, N.P.; Lukhtanov, V.A.; Pierce, N.E.

title:
Karyotypic diversity and speciation in Agrodiaetus butterflies

abstract:
That chromosomal rearrangements may play an important role in maintaining postzygotic isolation between well-established species is part of the standard theory of speciation. However, little evidence exists on the role of karyotypic change in speciation itself-in the establishment of reproductive barriers between previously interbreeding populations. The large genus Agrodiaetus (Lepidoptera: Lycaenidae) provides a model system to study this question. Agrodiaetus butterflies exhibit unusual interspecific diversity in chromosome number, from n = 10 to n = 134; in contrast, the majority of lycaenid butterflies have n = 23/24. We analyzed the evolution of karyotypic diversity by mapping chromosome numbers on a thoroughly sampled mitochondrial phylogeny of the genus. Karyotypic differences accumulate gradually between allopatric sister taxa, but more rapidly between sympatric sister taxa. Overall, sympatric sister taxa have a higher average karyotypic diversity than allopatric sister taxa. Differential fusion of diverged populations may account for this pattern because the degree of karyotypic difference acquired between allopatric populations may determine whether they will persist as nascent biological species in secondary sympatry. This study therefore finds evidence of a direct role for chromosomal rearrangements in the final stages of animal speciation. Rapid karyotypic diversification is likely to have contributed to the explosive speciation rate observed in Agrodiaetus, 1.6 species per million years.

citation:
Kandul, N.P., Lukhtanov, V.A. and Pierce, N.E. 2007. Karyotypic diversity and speciation in Agrodiaetus butterflies. Evolution 61: 546-559.


title:
Evolution of larval host plant associations and adaptive radiation in pierid butterflies

abstract:
Butterflies in the family Pieridae (Lepidoptera: Papilionoidea) feed as larvae on; plants belonging primarily to three distantly related angiosperm orders:; Fabales (legumes and allied plants), Brassicales (crucifers and related plants; containing mustard oil glucosides), and Santalales ('mistletoes'). However,; some utilize plants from 13 other families in a further eight orders. We; investigated the evolutionary history of host plant use of the Pieridae in the; context of a recent phylogenetic hypothesis of the family, using simple; character optimization. Although there is a close association between host; plant and butterfly higher classification, we find no evidence for cospeciation; but a pattern of repeated colonization and specialization. The ancestral host of; the family appears to be Fabaceae or Fabales, with multiple independent shifts; to other orders, including three to Santalales. The shift to Brassicales, which; contain secondary compounds (glucosinolates), promoted diversification and; adaptive radiation within the subfamily Pierinae. Subsequent shifts from; crucifers to mistletoes (aerial-stem hemiparasites) facilitated further diversification,; and more recent shifts from mistletoes to mistletoe host trees led to; exploitation of novel host plants outside the conventional three orders.; Possible mechanisms underlying these host shifts are briefly discussed. In the; Pierinae, a striking association between host plant, larval and adult behaviour,; adult phenotype, and mimicry calls for further research into possible; relationships between host specialization, plant chemistry and butterfly; palatability.

citation:
Braby, M.F. and Trueman, J.W.H. 2006. Evolution of larval host plant associations and adaptive radiation in pierid butterflies. Journal of Evoutionary Biology 19: 1677-1690.


title:
Evolution of larval food plant associations in Delias Hübner butterflies (Lepidoptera: Pieridae)

abstract:
A review of larval food plants of the genus Delias is presented. Larvae specialize primarily on aerial-stem; and root hemiparasites (¡°mistletoes¡±) in the order Santalales. Although butterfly food plant associations; have been recorded for only a small proportion of the genus (28 species or ¡«11%, representing 12/24; species-groups), available data suggest that the family Loranthaceae is used most frequently (77%), followed; by the Santalaceae sensu stricto (14%) and Viscaceae (8%). With the possible exception of Euphorbiaceae; (1%), almost all non-mistletoe records are considered to be erroneous and, in most cases, probably represent; the mistletoe host tree on which the larvae sometimes pupate. Of the eight major clades recognized in; Delias, food plants have been recorded for six of these, although the majority of records (89%) are for; three clades (hyparete, belladonna, nigrina). Optimization of the larval food plant data in the context; of recent phylogenetic hypotheses for both butterflies and plants revealed little evidence of cospeciation at; the higher systematic levels. The most parsimonious reconstruction was an origin of larval feeding on; Loranthaceae, followed by at least six independent colonizations to Santalaceae + Viscaceae. In contrast to; related pierids in the Aporiina associated with mistletoes in which further shifts from aerial-stem mistletoes; to distantly related plants (e.g. host trees parasitized by mistletoes) have facilitated differentiation at the; generic level, there is no firm evidence to indicate that such secondary, monomorphic shifts have evolved; in Delias. However, larvae of D. henningia (pasithoe group of belladonna clade) from Palawan and Luzon,; the Philippines, appear to be polymorphic, feeding on both Loranthaceae and Euphorbiaceae.

citation:
Braby, M.F. 2006. Evolution of larval food plant associations in Delias Hübner butterflies (Lepidoptera: Pieridae). Entomological Science 9: 383-398.


title:
Phylogeny and historical biogeography of the subtribe Aporiina (Lepidoptera: Pieridae): implications for the origin of Australian butterflies

abstract:
The Australian fauna is composed of several major biogeographical elements reflecting different spatial and temporal; histories. Two groups of particular interest are the Gondwanan Element, reflecting an ancient origin in Gondwana; or southern Gondwana (southern vicariance hypothesis), and the Asian Element, reflecting a more recent; origin in Asia, Eurasia or Laurasia (northern dispersal hypothesis). Theories regarding the origin and evolution of; butterflies (Hesperioidea, Papilionoidea) in Australia are controversial, with no clear consensus. Here, we investigate; the phylogenetic and historical biogeographical relationships of the subtribe Aporiina, a widespread taxon with disjunct; distributions in each of the major zoogeographical regions. Attention is paid to origins of the subtribe in the; Australian Region for which several conflicting hypotheses have been proposed for the Old World genus; Delias Hubner. Our phylogenetic reconstruction was based on analysis of fragments of two nuclear genes (elongation factor-1a, wingless) and one mitochondrial gene (cytochrome oxidase subunit I) for 30 taxa. Phylogenetic analyses based on maximum parsimony, maximum likelihood and Bayesian inference of the combined data set (2729 bp; 917 parsimony; informative characters) recovered six major lineages within the monophyletic Aporiina, with the following topology:; (Cepora+Prioneris+(Mylothris+(Aporia+Delias group+Catastictagroup))). Given a probable age of origin of the; stem-group near the Cretaceous/Tertiary boundary (69.54 Mya), followed by diversification of the crown-group in; the early to mid Tertiary (57.45 Mya), we show that an origin of the Aporiina in either southern Gondwana or Laurasia; is equally parsimonious, and that dispersal has played a major role in shaping the underlying phylogenetic pattern.; We tentatively conclude that an origin in southern Gondwanan is more likely; however, neither hypothesis; satisfactorily explains the present-day distribution, and additional lower-level phylogenies are needed to determine; the directionality of dispersal events of several taxa and to reject one hypothesis over the other. Dispersal is inferred; to have occurred primarily during cooler periods when land bridges or stepping-stones were available between many; of the zoogeographical regions.

citation:
Braby, M.F., Pierce, N.e. and Vila, R. 2007. Phylogeny and historical biogeography of the subtribe Aporiina (Lepidoptera: Pieridae): implications for the origin of Australian butterflies. Biological Journal of the Linnean Society 90: 413-440.


author:
Harris, K.M.; Sato, S.; Uechi, N.; Yukawa, J.

title:
Redefinition of Oligotrophus (Diptera: Cecidomyiidae) based on morphological and molecular attributes of species from galls on Juniperus (Cupressaceae) in Britain and Japan

abstract:
Morphological and molecular attributes of four species of Cecidomyiidae that induce galls on junipers,; Juniperus spp., in Britain and Japan were used to redefine the genus Oligotrophus Latreille, which now; includes Arceuthomyia Kieffer and Schmidtiella Rübsaamen (new synonyms). Illustrated morphological; descriptions are provided for the four species Oligotrophus juniperinus (Linnaeus), Oligotrophus panteli; Kieffer, Oligotrophus gemmarum (Rübsaamen) (new combination) and Oligotrophus nezu Kikuti; (= Arceuthomyia nakaharai Inouye (new synonymy)) and phylogenies based on partial sequencing of the; mitochondrial 12S rRNA gene are reported.

citation:
Harris, K.M., Sato, S., Uechi, N. and Yukawa, J. 2006. Redefinition of Oligotrophus (Diptera: Cecidomyiidae) based on morphological and molecular attributes of species from galls on Juniperus (Cupressaceae) in Britain and Japan. Entomological Science  9: 411-421.


abstract:
The various mutualistic and antagonistic symbioses between fig trees (Ficus : Moraceae) and chalcid wasps comprise; a community in microcosm. Phylogenetic estimates of figs and fig wasps show general topological correspondence,; making the microcosm a model system for cophylogeny. Incongruence between phylogenies from; associated organisms can be reconciled through a combination of evolutionary events. Cophylogeny mapping; reconciles phylogenies by embedding an associate tree into a host tree, finding the optimal combinations of events; capable of explaining incongruence and evaluating the level of codivergence. This review addresses the results of; cophylogeny analysis concerning Ficus and discusses the plausibility of different evolutionary events. Five different; associations encompassing fig-pollinator, fig-parasite and pollinator-parasitoid interactions are reconciled. The; method improves on previous comparisons by employing ' jungles ' to provide an exhaustive and quantitative; analysis of cophylogeny. A jungle is a mechanism for inferring host switches and obtaining all potentially optimal; solutions to the reconciliation problem. The results support the consensus that figs codiverge significantly; with pollinators but not non-pollinators. However, pollinators still appear to have switched between hosts in; contradiction to the traditional model of faithful codivergence. This emphasises the growing realisation that; evolutionary transitions in the microcosm are more flexible than previously thought and host specificity is; necessary but not sufficient for codivergence. The importance of sampling strategy is emphasised by the influence; of taxon set on the fig-pollinator and fig-parasite jungles. Spurious significant results for fig-parasite and; fig-parasitoid jungles indicate that the choice of congruence measure influences significance; the total number of; events required to reconcile two trees (' total cost ') is not a good measure of congruence when switches cannot be; realistically weighted.

citation:
Jackson, A.P. 2004. Cophylogeny of the Ficus microcosm. Biol. Rev. 79: 751-768.


title:
Assessment of the diversity and species specificity of the mutualistic association between Epicephala moths and Glochidion trees

abstract:
The obligate mutualisms between flowering plants and their seed-parasitic pollinators constitute fascinating examples of interspecific mutualisms, which are often characterized by high levels of species diversity and reciprocal species specificity. The diversification in these mutualisms has been thought to occur through simultaneous speciation of the partners, mediated by tight reciprocal adaptation; however, recent studies cast doubt over this general view. In this study, we examine the diversity and species specificity of Epicephala moths (Gracillariidae) that pollinate Glochidion trees (Phyllanthaceae), using analysis of mitochondrial and nuclear gene sequences. Phylogenetic analysis of Epicephala moths associated with five Glochidion species in Japan and Taiwan reveal six genetically isolated species that are also distinguishable by male genital morphology: (i) two species specific to single host species (G. acuminatum and G. zeylanicum, respectively); (ii) two species that coexist on G. lanceolatum; and (iii) two species that share two, closely-related parapatric hosts (G. obovatum and G. rubrum). Statistical analysis shows that the two species associated with G. lanceolatum are not sister species, indicating the colonization of novel Glochidion host in at least one lineage. Behavioural observations suggest that all six species possess the actively-pollinating habit, thus none of the studied species has become a nonmutualistic 'cheater' that exploits the benefit resulting from pollination by other species. Our results parallel recent findings in ecologically similar associations, namely the fig-fig wasp and yucca-yucca moth mutualisms, and contribute to a more general understanding of the factors that determine ecological and evolutionary outcomes in these mutualisms.

citation:
Kawakita, A. and Kato, M. 2006. Assessment of the diversity and species specificity of the mutualistic association between Epicephala moths and Glochidion trees. Molecular Ecology 15: 3567-3581.


author:
Kutty, S.N.; Bernasconi, M.V.; Sifner, F.; Meier, R.

title:
Sensitivity analysis, molecular systematics and natural history evolution of Scathophagidae (Diptera: Cyclorrhapha: Calyptratae)

abstract:
The 60 000 described species of Cyclorrhapha are characterized by an unusual diversity in larval life-history traits, which range from saprophagy over phytophagy to parasitism and predation. However, the direction of evolutionary change between the different modes remains unclear. Here, we use the Scathophagidae (Diptera) for reconstructing the direction of change in this relatively small family ( 250 spp.) whose larval habits mirror the diversity in natural history found in Cyclorrhapha. We subjected a mlecular data set for 63 species (22 genera) and DNA sequences from seven genes (12S, 16S, Cytb, COI, 28S, Ef1-alfa, Pol II) to an extensive sensitivity analysis and compare the performance of three different alignment strategies (manual, Clustal, POY). We find that the default Clustal alignment performs worst as judged by character incongruence, topological congruence and branch support. For this alignment, scoring indels as a fifth character state worsens character incongruence and topological congruence. However, manual alignment and direct optimization perform similarly well and yield near-identical trees, although branch support is lower for the direct-optimization trees. All three alignment techniques favor the upweighting of transversion. We furthermore confirm the independence of the concepts ''node support'' and ''node stability'' by documenting several cases of poorly supported nodes being very stable and cases of well supported nodes being unstable. We confirm the monophyly of the Scathophagidae, its two constituent; subfamilies, and most genera. We demonstrate that phytophagy in the form of leaf mining is the ancestral larval feeding habit for Scathophagidae. From phytophagy, two shifts to saprophagy and one shift to predation has occurred while a second origin of predation is from a saprophagous ancestor.

citation:
Kutty, S.N., Bernasconi, M.V., Sifner, F. and Meier, R. 2007. Sensitivity analysis, molecular systematics and natural history evolution of Scathophagidae (Diptera: Cyclorrhapha: Calyptratae). Cladistics 23 23: 64-83.


title:
Phylogenetic analysis of Zygaenoidea small-subunit rRNA structural variation implies initial oligophagy on cyanogenic host plants in larvae of the moth genus; Zygaena (Insecta: Lepidoptera)

journal:
Zoological Journal of the Linnean Societ

citation:
Niehuis, O. and Naumann, C.M. 2006. Phylogenetic analysis of Zygaenoidea small-subunit rRNA structural variation implies initial oligophagy on cyanogenic host plants in larvae of the moth genus; Zygaena (Insecta: Lepidoptera). Zoological Journal of the Linnean Societ 147: 367-381.


title:
The willow bud galler Euura mucronata Hartig; (Hymenoptera: Tenthredinidae): one polyphage or many monophages?

abstract:
The nematine sawfly Euura mucronata Hartig (Hymenoptera: Tenthredinidae) induces galls in the buds of over 30 willow species across the Holarctic region. This extensive host range is surprising, since the other Euura gallers are mostly monophagous; thus, the feeding habit of E. mucronata would represent a switch from monophagy to extreme polyphagy. Previous morphological studies have divided E. mucronata into separate species, but the feeding ranges of these species are unknown, and it is even doubtful whether multiple species really exist. To study whether or not E. mucronata consists of cryptic host-associated sibling species, an allozyme study was conducted using gallers collected from six willow species occurring in northern Fennoscandia. Electrophoretic data from seven variable enzyme loci show that: (1) 'E. mucronata' probably comprises at least three species with restricted host ranges, but the species may not be completely reproductively isolated from each other; (2) the pattern of host use is not explained by the phylogeny of willows; (3) the pattern of host use is not concordant with the overall chemical similarity of the hosts; and (4) simple allopatric speciation does not appear to explain the host associations. Consequently, it is possible that reasons such as differences in host phenology, habitat, or; morphology, are responsible for the limits in host use in the group.

citation:
Nyman, T. 2002. The willow bud galler Euura mucronata Hartig; (Hymenoptera: Tenthredinidae): one polyphage or many monophages? Heredity 88: 288-295.


author:
Regier, J. C.; Paukstadt, U.; Paukstadt, L.H.; Mitter, C.; Peigler, R.S.

title:
Phylogenetics of eggshell morphogenesis in Antheraea (Lepidoptera: Saturniidae): unique origin and repeated reduction of the aeropyle crown

citation:
Regier, J. C., Paukstadt, U., Paukstadt, L.H., Mitter, C. and Peigler, R.S.  2005. Phylogenetics of eggshell morphogenesis in Antheraea (Lepidoptera: Saturniidae): unique origin and repeated reduction of the aeropyle crown. Systematic Biology  54: 254-267.


title:
Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

abstract:
Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing; and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising; Yramea and Boloria s.l., Argynnina comprising Prokuekenthaliella, Issoria, Brenthis, and all the 'large fritillary' species joined in the genus Argynnis s.l. and a new subtribe Euptoietina comprising only the genus Euptoieta. The classical genus; Issoria s.l. is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and; hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella. Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas, Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.

citation:
Simonsen, T.J. 2006. Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini). Biological Journal of the Linnean Society 89: 627-673.


title:
Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonization and diversification

abstract:
We present a higher-level phylogenetic hypothesis for the diverse neotropical butterfly subfamily Ithomiinae, inferred from one of the largest non-molecular Lepidoptera data sets to date, including 106 species (105 ingroup) and 353 characters (306 informative) from adult and immature stage morphology and ecology. Initial analyses resulted in 1716 most parsimonious trees, which were reduced to a single tree after successive approximations character weighting. The inferred phylogeny was broadly consistent with other past and current work. Although some deeper relationships are uncertain, tribal-level clades were generally strongly supported, with two changes required to existing classification. The tribe Melinaeini is polyphyletic and Athesis + Patricia require a new tribe. Methona should be removed from Mechanitini into the restored tribe Methonini. Dircennini was paraphyletic in analyses of all data but monophyletic based on adult morphology alone, and its status remains to be confirmed. Hypothyris, Episcada, Godyris, Hypoleria and Greta are paraphyletic. A simulation analysis showed that relatively basal branches tended to have higher partitioned Bremer support for immature stage characters. Larval hostplant records were optimized on to a reduced, generic-level phylogeny and indicate that ithomiines moved from Apocynaceae to Solanaceae twice, or that Tithoreini re-colonized Apocynaceae after a basal shift to Solanaceae. Ithomiine clades have specialized on particular plant clades suggesting repeated colonization of novel hostplant niches consistent with adaptive radiation. The shift to Solanum, comprising 70% of neotropical Solanaceae, occurs at the base of a clade containing 89% of all ithomiines, and is interpreted as the major event in the evolution of ithomiine larval hostplant relationships.

citation:
Willmott, K.R. and Freitas, A.V.L. 2006. Higher-level phylogeny of the Ithomiinae (Lepidoptera: Nymphalidae): classification, patterns of larval hostplant colonization and diversification. Cladistics 22: 297-368.


author:
Beltrán, M.; Jiggins, C.D.; Brower, A.V.Z.; Bermingham, E.; Mallet, J.

title:
Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA; sequence data

abstract:
Phylogenetic information is useful in understanding the evolutionary history of adaptive traits. Here, we present; a well-resolved phylogenetic hypothesis for Heliconius butterflies and related genera. We use this tree to; investigate the evolution of three traits, pollen feeding, pupal-mating behaviour and larval gregariousness.; Phylogenetic relationships among 60 Heliconiina species (86% of the subtribe) were inferred from partial DNA; sequences of the mitochondrial genes cytochrome oxidase I, cytochrome oxidase II and 16S rRNA, and fragments; of the nuclear genes elongation factor-1a, apterous, decapentaplegic and wingless (3834 bp in total). The results; corroborate previous hypotheses based on sequence data in showing that Heliconius is paraphyletic, with Laparus; doris and Neruda falling within the genus, demonstrating a single origin for pollen feeding but with a loss of the; trait in Neruda. However, different genes are not congruent in their placement of Neruda; therefore, monophyly; of the pollen feeding species cannot be ruled out. There is also a highly supported monophyletic 'pupal-mating; clade' suggesting that pupal mating behaviour evolved only once in the Heliconiina. Additionally, we observed at; least three independent origins for larval gregariousness from a solitary ancestor, showing that gregarious larval; behaviour arose after warning coloration.

citation:
Beltrán, M., Jiggins, C.D., Brower, A.V.Z., Bermingham, E. and Mallet, J. 2007. Do pollen feeding, pupal-mating and larval gregariousness have a single origin in Heliconius butterflies? Inferences from multilocus DNA; sequence data. Biological Journal of the Linnean Society 92: 221-239.


author:
Snäll, N.; Tammaru, T.; Wahlberg, N.; Viidalepp, J.; Ruohomäki, K.; Savontaus, M.-L.; Huoponen, K.

title:
Phylogenetic relationships of the tribe Operophterini (Lepidoptera, Geometridae): a case study of the; evolution of female flightlessness

abstract:
A molecular phylogenetic analysis was conducted in order to reconstruct the evolution of female flightlessness in; the geometrid tribe Operophterini (Lepidoptera, Geometridae, Larentiinae). DNA variation in four nuclear gene; regions, segments D1 and D2 of 28S rRNA, elongation factor 1a, and wingless, was examined from 22 species; representing seven tribes of Larentiinae and six outgroup species. Direct optimization was used to infer a; phylogenetic hypothesis from the combined sequence data set. The results obtained confirmed that Operophterini; (including Malacodea) is a monophyletic group, and Perizomini is its sister group. Within Operophterini, the genus; Malacodea is the sister group to the genera Operophtera and Epirrita, which form a monophyletic group. This; relationship is also supported by morphological data. The results suggest that female flightlessness has evolved; independently twice: first in the lineage of Malacodea and, for the second time, in the lineage of Operophtera after; its separation from the lineage of Epirrita. An alternative reconstruction (i.e. recovery of flight ability in an; ancestor of Epirrita) appears unlikely for various reasons. The similarities shared by Epirrita with a basal; representative of Perizomini, Perizoma didymatum, allow the proposal of a sequence of evolutionary events that; has led to flightlessness. It is likely that the transition to female flightlessness in the two lineages of Operophterini; occurred after the colonization of stable forest habitats, followed by the evolution of a specific set of permissive; traits, including larval polyphagy, limited importance of adult feeding, and adult flight during the cold months of; the season.

citation:
Snäll, N., Tammaru, T., Wahlberg, N., Viidalepp, J., Ruohomäki, K., Savontaus, M.-L. and Huoponen, K. 2007. Phylogenetic relationships of the tribe Operophterini (Lepidoptera, Geometridae): a case study of the; evolution of female flightlessness.  92: 241-252.


title:
Genetic diversity, host-specificity and unusual; phylogeography of a cryptic, host-associated species complex of gall-inducing scale insects

abstract:
1. Australia has a unique and speciose gall-inducing scale insect fauna that; is primarily associated with Myrtaceae. Much of the diversity is currently undescribed; or uncharacterised.; 2. This study concerns Apiomorpha munita (Hemiptera), a scale insect that induces; characteristic four-horned galls on eucalypts of subgenus Symphyomyrtus and exhibits; extraordinary karyotypic diversity (2n = 6 - 2n > 100). The three described subspecies; of A. munita are each confined to hosts in different sections of Eucalyptus . Previous; chromosomal data, however, cast doubt on the validity of the groupings, as two of the; subspecies share multiple, different karyotypes (2n = 6, 20, 22, and 24).; 3. Allozyme data were used to examine species delimitation, chromosome evolution,; host associations and population structure in A. munita .; 4. A cryptic-species radiation was revealed, with at least five taxa each restricted to a; discrete set of host eucalypt species. This is consistent with host-associated speciation.; 5. Karyotypic variation within A. munita partially fits the five distinct genetic groups,; but there are additional chromosomal changes that are not accompanied by detected; genetic differentiation.; 6. The population structure of taxa within the A. munita species complex suggests; that there are high levels of inbreeding, as would be expected for scale insects in which; adult females are sessile. Some genotypes, however, are found over great distances; (up to 1100 km). This is an unusual population structure because it combines low; mobility and local differentiation with occasional long-distance dispersal, probably; mediated by wind-dispersal of first-instar nymphs.

citation:
Cook, L.G. and Rowell, D.M. 2007. Genetic diversity, host-specificity and unusual; phylogeography of a cryptic, host-associated species complex of gall-inducing scale insects. Ecological Entomology 32: 506-515.


author:
Niehuis, O.; Hofman, A.; Naumann, C.M.; Misof, B.

title:
Evolutionary history of the burnet moth genus Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae) inferred from nuclear and mitochondrial sequence data: phylogeny, host-plant association, wing pattern evolution and historical biogeography

abstract:
Burnet moths of the genus Zygaena are a striking group of primarily diurnal Lepidoptera displaying an exceptional; phenotypic plasticity. Previous attempts to elucidate the phylogenetic history of the group had been confounded by; a perplexing pattern of characters or insufficient taxon sampling. In the present study, we infer a phylogeny of the; genus Zygaena by analysing 5.4 kb of their nuclear and mitochondrial DNA. Eighty-four of the 98 currently; recognized species in this genus are considered, including representatives of all described species groups. RNA; coding sequences are aligned with reference to zygaenoid moth specific secondary structure models of corresponding; molecules. We conduct phylogenetic analyses within a Bayesian framework applying partition specific substitution; parameters; covariation of paired sites in RNA gene sequences is accommodated by using doublet; substitution models. The molecular data reveal that a considerable number of currently recognized species groups; in Zygaena are not monophyletic. The traditional subgeneric classification proves to be artificial as well; Agrumenia; and Zygaena (sensu stricto) are polyphyletic. Only the subgenus Mesembrynus can be confirmed as a monophyletic; species cluster. Optimization of larval host-plant associations and forewing patterns on sampled trees of the; Bayesian analyses suggest convergent evolution of similar wing pattern types in distantly related species clusters; and a shift from cyanogenic to acyanogenic host-plants. The phylogenetic results challenge the classic assumption; that early species diversification in Zygaena took place in the Irano-Turkestanian region. Rather, the molecular; data point to the western Mediterranean area as the geographical origin of the group and imply a subsequent; colonization of the Middle East and Central Asia. We discuss the apparently convergent evolution of similar wing; patterns in context with the chemical defence system of burnet moths and suggest a species group concept for the; genus Zygaena that accounts for the recent findings.

citation:
Niehuis, O., Hofman, A., Naumann, C.M. and Misof, B. 2007. Evolutionary history of the burnet moth genus Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae) inferred from nuclear and mitochondrial sequence data: phylogeny, host-plant association, wing pattern evolution and historical biogeography. Biological Journal of the Linnean Society 92: 501-520.


title:
Convergent evolution of chemical defence in; Galerucine larvae

abstract:
Selection pressure by natural enemies on phytophagous insect larvae is intense and has frequently triggered the; evolution of chemical defence as an effective counterstrategy. In the chrysomelid subfamily Galerucinae, glandular; structures and defensive fluids have been described for the tribe Sermylini Wilcox, 1965. Previous morphological; and ultrastructural studies raised doubts that these defensive devices in Sermylini can be traced back to a common; origin. The taxonomy of the Galerucinae cannot clear these doubts because the phylogeny of this taxon is a matter; of current debate. We therefore investigated the phylogeny of the Galerucinae based on approximately 1740 bp of; the mitochondrial 12S and 16S rRNA and the nuclear elongation factor 1 alpha genes. Our data support the; hypothesized close relationship between the subfamilies Galerucinae and Alticinae, yet, by contrast to other recent; analyses, the two groups are mostly resolved as monophyletic sister groups or, in some analyses, with the; Galerucinae nested paraphyletically within the Alticinae. Within the subfamily Galerucinae, only the tribe; Galerucini formed a monophyletic taxon, except for one species, Cerochroa brachialis Stal, 1858. In none of our; analyses were the Sermylini recovered as a monophyletic tribe. However, our data support monophyly of each of; the three groups within the Sermylini that have morphologically distinguishable larval defensive openings. We; conclude that the defensive structures in larvae of Sermylini have no common origin, but evolved independently.; Our data suggest that the tremendous selection pressure by natural enemies led to the recurrent evolution of; similar chemical defensive devices in Sermylini larvae.

citation:
Bünnige, M., Hilker, M. and Dobler, S. 2008. Convergent evolution of chemical defence in; Galerucine larvae. Biological Journal of the Linnean Society 93: 165-175.


title:
he molecular phylogeny of the Miarus campanulae (Coleoptera: Curculionidae) species group inferred from CO1 and ITS2 sequences

abstract:
Miarus is a Holarctic weevil genus with morphologically very similar species, all breeding on Campanula plants or their close relatives. Two European members of this genus, Miarus campanulae (L.), the type species, and Miarus graminis (Bohemann) have recently been split into several new species on the basis of slight external variations. The separation of these new forms has proved impossible and new data was needed. Molecular data were gathered from specimens from a number of locations in Finland, Estonia, Denmark and Sweden. The regions sequenced were mitochondrial CO1 and nuclear ITS2. Both combined and separate datasets were analyzed using the optimization alignment program POY, with parsimony as the optimality criterion. The recently separated Miarus species was found to be indistinguishable from the traditionally recognized form on the basis of this sequence data. On the other hand, the traditionally recognized species were characterized by numerous synapomorphies. Our data suggest that recent studies have underestimated the morphological variation in this genus. We propose that this may also be true for many taxonomically problematic beetle complexes in well-studied European regions. The idea that molecular evidence will inevitably reveal unnoticed cryptic variation may only apply to poorly known regions. Miarus fennicus Kangas, 1978 is placed as a junior synonym of Miarus campanulae (Linnaeus, 1767) syn. nov.

citation:
Vahtera, V. and Muona, J. 2006. he molecular phylogeny of the Miarus campanulae (Coleoptera: Curculionidae) species group inferred from CO1 and ITS2 sequences. Cladistics 22: 222-229.


abstract:
The butterfly Pieris napi (L.) and relatives exemplify recently evolving taxa, exhibiting variation that makes their evolutionary dynamics interesting, but their systematics difficult. Wing-pattern characters commonly used to distinguish these Holarctic insects display both genetic polymorphism and environmentally-cued polyphenism. Often, these causes of variation are confounded, impairing the characters' phylogenetic usefulness. DNA sequences of four mitochondrial genes offer an independent view of pierine phylogeny. Sampling diverse relatives within family Pieridae assists resolution of the P. napi complex, suggests previous underestimation of clade diversity in subfamily Pierinae, and shows that other genera near Pieris also display confusions of wing-pattern-based phylogenetic inference. The European P. napi is sister to all North American taxa, and is well diverged from them all in sequences. The North American taxa comprise a northern subclade including Pieris oleracea, and questionably distinct Pieris 'angelika', and a southern subclade including distinct Pieris virginiensis, Pieris marginalis, and Pieris macdunnoughii, and other regional entities yet to be clarified. Weak bootstrap support for some nodes in this group arises from a closeness of sequence identity rather than character conflict; more sequence data and denser geographical sampling may resolve these nodes more clearly. Evidence of reproductive isolation, from other experimental hybridization studies, agrees with the DNA results where these conflict with other divergent character sets. The system offers much promise for a deeper understanding of character evolution in relation to phyletic differentiation.

citation:
Chew, F.S. and Watt, W.B 2006. The green-veined white (Pieris napi L.), its Pierine relatives, and the systematics dilemmas of divergent character sets (Lepidoptera, Pieridae). Biological Journal of the Linnean Society 88: 413-435.


author:
Gómez-Zurita, J.; Funk, D.J.; Vogler, A.P.

title:
The evolution of unisexuality in Calligrapha leaf beetles: molecular and ecological insights on multiple origins via interspecific hybridization

abstract:
Interspecific hybridization is a well-established cause of unisexual origins in vertebrates. This mechanism is also suspected in other apomictic taxa, but compelling evidence is rare. Here, we evaluate this mechanism and other hypotheses for the evolutionary origins of unisexuality through an investigation of Calligrapha leaf beetles. This group provides an intriguing subject for studies of unisexual evolution because it presents a rare insect example of multiple apomictic thelytokous species within a primarily bisexual genus. To investigate unisexual evolution, this study conducts the first molecular systematic analysis of Calligrapha. This involved the collection and analysis of about 3000 bp of DNA sequences-representing RNA and protein-coding loci from mitochondrial and nuclear genomes-from 54 specimens of 25 Calligrapha species, including four unisexual tetraploid taxa. Phylogenetic and molecular clock analyses indicated independent and single evolutionary origins of each of these unisexual species during the Pleistocene. Significant phylogenetic incongruence was detected between mitochondrial and nuclear datasets and found to be especially associated with the asexual taxa. This pattern is expected when unisexual lineages arise via interspecific hybridization and thus represent genetic mosaics that possess certain nuclear alleles from the paternal species lineage and mitochondrial DNA (mtDNA) alleles from the maternal parent. Analyzing the mtDNA and nuclear relatedness of unisexuals with corresponding haplotypes of bisexual Calligrapha species allowed the putative identification of these maternal and paternal species lineages for each unisexual species. Strong phenotypic similarities between unisexual taxa and their paternal parent species supported a model that involves both backcrosses of interspecific hybrids with a paternal parent and unreduced gametes. This model accounts for the origins of apomixis, polyploidy, and an overrepresentation of paternal nuclear alleles (and associated phenotypes) in unisexuals. This model is also consistent with the tetraploid karyotypes of unisexual Calligrapha, in which three sets of chromosomes (of presumed paternal ancestry) are quite morphologically homogeneous compared to the fourth. Especially intriguing was a consistent association of unisexual species with the host plant of the paternal parent but never with the maternal host. The statistical implausibility of these patterns occurring by chance further supports our inference of parental species. Moreover, it points to a potentially critical role for host-association in the formation and preservation of unisexual lineages. These findings suggest that ecological factors are critical for the diversification of unisexual as well as bisexual taxa and thus point out new research directions in the area of ecological speciation.

citation:
Gómez-Zurita, J., Funk, D.J. and Vogler, A.P. 2006. The evolution of unisexuality in Calligrapha leaf beetles: molecular and ecological insights on multiple origins via interspecific hybridization. Evolution 60: 328-347.


