Musculature of the male terminalia in geometrid moths of the subfamily Larentiinae (Lepidoptera,Geometridae)

The morphology of the male terminalia of 16 species belonging to 12 tribes of Larentiinae was studied; for 12 species and 4 tribes it was done for the first time. No clear diagnosis of Larentiinae based on the features of the male genital musculature can be given since the apomorphic position of M1, M4, and M7 muscles may originate independently in different tribes. The primitive Larentiinae retain the pattern of genital musculature plesiomorphic for Geometridae. Muscles M11 (Melanthiini: Anticollix sparsata) and M22 (Asthenini: Euchoeca nebulata and Hydrelia flammeolaria) are described for the first time for the family Geometridae.     

Revision of Scotocyma Turner (Lepidoptera: Geometridae: Larentiinae)

Abstract  The Australasian genus Scotocyma Turner is revised, containing the species S. albinotata (Walker), S. legalis (Warren), S. asiatica Holloway, S. scotopepla Prout, stat. n., S. manusensis Prout, stat. n., S. mimula (Warren), stat. n. and S. miscix Prout. Scotocyma euryochra Turner, syn. n., S. idioschema Turner, syn. n., S. ischnophrica Turner, syn. n. and S. transfixa Turner, syn. n. are regarded as synonyms of S. albinotata . Four species are described as new: S. samoensis , sp. n., S. sumatrensis , sp. n., S. rutilimixta , sp. n. and S. longiuncus , sp. n. Lectotypes are designated for S. scotopepla , S. manusensis and S. miscix . All species are illustrated, and keys to species and distribution maps are provided. A phylogenetic analysis was performed to test the monophyly of the genus and to examine distribution patterns of the species. A biogeographical discussion is included. The tribal position of the genus is clarified and relationships to closely related genera are discussed.     

Systematics of the genus Cidaria Treitschke (Lepidoptera, Geometridae, Larentiinae)

The genus Cidaria Treitschke is revised. Eight species of the genus which occur widely in the Palaearctic and northern India, are recognized, of which one, Cidaria luteata sp. nov. , is described as new while two subspecies of C. fulvata (Forster) are elevated to species, C. nugata Felder stat. rev., and C. distinctata Staudinger stat. nov. C. ochripennis Prout is proposed as a junior synonym of C. ochreata Staudinger. C. deletaria Hampson is excluded from the genus. All species of Cidaria and their genitalia are described and illustrated. The cladistic analysis of these eight species of Cidaria is carried out using two data matrices, the first comprising morphological characters alone and the second morphological and distributional characters. The most parsimonious cladogram of Cidaria is selected, and its monophyly defined. The character analysis shows that some distribution characters contribute to resolving the in group node. The choice of multiple trees is discussed.     

韩国波尺蛾亚科三种幼虫记述（鳞翅目：尺蛾科）

本文详细地记述了波尺蛾亚科Eupithecia abietaria debrunneata Staudinger, E. spadix Inoue 及Telenomeuta punctimarginaria (Leech) 幼虫的形态特征.     

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

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.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 473−530.     

Systematic position of Lythriini revised: transferred from Larentiinae to Sterrhinae (Lepidoptera, Geometridae)

The tribe Lythriini is a small group of diurnally active geometrid moths consisting of a single Palaearctic genus Lythria with five species. The systematic placement of Lythriini has remained controversial: though traditionally it has been placed into the subfamily Larentiinae, a number of morphological characters link this tribe with the subfamily Sterrhinae. A molecular phylogenetic study was conducted to verify the systematic position of Lythriini, using sequences of both mitochondrial and nuclear genes: elongation factor 1α ( EF-1α ), wingless ( wgl ), 28S rRNA expansion segment D2 ( 28S D2 ), cytochrome oxidase subunit 1 ( COI ) and NADH dehydrogenase subunit 1 ( ND1 ) (a total of 3784 bp). Phylogenetic analysis reliably demonstrated that Lythriini belong to the subfamily Sterrhinae. Therefore, we propose to remove tribe Lythriini from Larentiinae and unite it with Sterrhinae. Moreover, our analysis supports the monophyly of both Sterrhinae and Larentiinae. However, although both morphological data and interspecific genetic distances insinuated that Lythria cruentaria and L. sanguinaria are sister species, the latter formed a clade of sister taxa together with L. purpuraria .     

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

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

Revision of the genus Helastia sensu stricto with description of a new genus (Lepidoptera: Geometridae: Larentiinae)

Abstract

Helastia Guenée, 1868 is redefined and redescribed. New Zealand species previously placed in that genus but not congeneric with the type species are reassigned to either the available genera Epyaxa Meyrick, 1883, Asaphodes Meyrick, 1885 and Xanthorhoe Hübner, [1825] or placed in a newly described genus, Gingidiobora. Six Australian species placed in Xanthorhoe are shown to be congeneric with three New Zealand species, previously placed in Helastia and here transferred to Epyaxa.

Eight new species are described in Helastia: Helastia alba n. sp.; H. angusta n. sp.; H. christinae n. sp.; H. cryptica n. sp.; H. mutabilis n. sp.; H. ohauensis n. sp.; H. salmoni n. sp.; H. scissa n. sp. The following new combinations and synonymies are proposed: Asaphodes chlorocapna (Meyrick, 1925) n. comb.; A. citroena (Clark, 1934) n. comb.; A. glaciata (Hudson, 1925) n. comb.; A. ida (Clark, 1926) n. comb; Epyaxa agelasta (Turner, 1904) n. comb.; E. centroneura (Meyrick, 1890) n. comb.;

E. epia (Turner, 1922) n. comb.; E. hyperythra (Lower, 1892) n. comb.; E. lucidata (Walker, 1862) n. comb.; E. sodaliata (Walker, 1862) n. comb.; E. subidaria (Guenée, 1857) n. comb.; E. venipunctata (Walker, 1863) n. comb.; Gingidiobora nebulosa (Philpott, 1917) n. comb.; G. subobscurata (Walker, 1862) n. comb.; Helastia clandestina (Philpott, 1921) n. comb.; H. corcularia (Guenée, 1868) n. comb. (= Larentia infantaria Guenée, 1868 n. syn.); H. expolita (Philpott, 1917) n. comb.; H. siris (Hawthorne, 1897) n. comb.; H. triphragma (Meyrick, 1883) n. comb.     

Phylogeny and tribal classification of Sterrhinae with emphasis on delimiting Scopulini (Lepidoptera: Geometridae)

Abstract. The phylogenetic relationships of tribes of the geometrid subfamily Sterrhinae (Lepidoptera) were studied, with special emphasis on finding delimiting characters for the tribe Scopulini. Two cladistic analyses were conducted for fifty‐nine species representing all previously recognized Sterrhinae tribes and covering the geographical range of the subfamily. In the first analysis, twelve putative synapomorphies of Scopulini, taken from the literature, were coded for actual specimens in order to test their ability to support the monophyly of the group. The resulting strict consensus cladogram was totally unresolved. In the second analysis, the twelve characters were combined with additional information from the morphology and ecology of adults and immature stages. Analysis of these ninety‐six characters resulted in a well‐resolved cladogram. The tribes were found to be monophyletic, except Cosymbiini and Rhodostrophiini. There are two main lineages within Sterrhinae: Cosymbiini + Rhodometrini + Timandrini and Rhodostrophiini + Cyllopodini + Sterrhini + Scopulini. Aletini and Problepsini lay within the concept of Scopulini. The association of the included Larentiinae taxa with the Cosymbiini + Rhodometrini + Timandrini lineage questions the monophyly of Sterrhinae. A majority of the recovered synapomorphic characters had been recognized previously, but several new phylogenetically informative characters were found, especially from the thorax. No unique characters diagnosing the tribe Scopulini were found, but many homoplastic synapomorphic features were found which diagnose parts of it. All recognized Sterrhinae genera are assigned tentatively to tribes and problematic cases are discussed.     

Phylogeny of Cephalobina (Nematoda): molecular evidence for recurrent evolution of probolae and incongruence with traditional classifications

Nematodes of the suborder Cephalobina include an ecologically and morphologically diverse array of species that range from soil-dwelling microbivores to parasites of vertebrates and invertebrates. Despite a long history of study, certain of these microbivores (Cephaloboidea) present some of the most intractable problems in nematode systematics; the lack of an evolutionary framework for these taxa has prevented the identification of natural groups and inhibited understanding of soil biodiversity and nematode ecology. Phylogenetic analyses of ribosomal (LSU) sequence data from 53 taxa revealed strong support for monophyly of taxa representing the Cephaloboidea, but do not support the monophyly of most genera within this superfamily. Historically these genera have primarily been recognized based on variation in labial morphology, but molecular phylogenies show the same general labial (probolae) morphotype often results from recurrent similarity, a result consistent with the phenotypic plasticity of probolae previously observed for some species in ecological time. Phylogenetic analyses of LSU rDNA also recovered strong support for some other groups of cephalobs, including taxa representing most (but not all) Panagrolaimoidea. In addition to revealing homoplasy of probolae, molecular trees also imply other unexpected patterns of character evolution or polarity, including recurrent similarity of offset spermatheca presence, and representation of complex probolae as the ancestral condition within Cephaloboidea. For Cephalobidae, molecular trees do not support traditional genera as natural groups, but it remains untested if deconstructing probolae morphotypes or other structural features into finer component characters may reveal homologies that help delimit evolutionary lineages.     

Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae)

Background

The moth family Geometridae (inchworms or loopers), with approximately 23 000 described species, is the second most diverse family of the Lepidoptera. Apart from a few recent attempts based on morphology and molecular studies, the phylogeny of these moths has remained largely uninvestigated.

Methodology/Principal Findings

We performed a rigorous and extensive molecular analysis of eight genes to examine the geometrid affinities in a global context, including a search for its potential sister-taxa. Our maximum likelihood analyses included 164 taxa distributed worldwide, of which 150 belong to the Geometridae. The selected taxa represent all previously recognized subfamilies and nearly 90% of recognized tribes, and originate from all over world. We found the Geometridae to be monophyletic with the Sematuridae+Epicopeiidae clade potentially being its sister-taxon. We found all previously recognized subfamilies to be monophyletic, with a few taxa misplaced, except the Oenochrominae+Desmobathrinae complex that is a polyphyletic assemblage of taxa and the Orthostixinae, which was positioned within the Ennominae. The Sterrhinae and Larentiinae were found to be sister to the remaining taxa, followed by Archiearinae, the polyphyletic assemblage of Oenochrominae+Desmobathrinae moths, Geometrinae and Ennominae.

Conclusions/Significance

Our study provides the first comprehensive phylogeny of the Geometridae in a global context. Our results generally agree with the other, more restricted studies, suggesting that the general phylogenetic patterns of the Geometridae are now well-established. Generally the subfamilies, many tribes, and assemblages of tribes were well supported but their interrelationships were often weakly supported by our data. The Eumeleini were particularly difficult to place in the current system, and several tribes were found to be para- or polyphyletic.     

Phylogeny of the Nymphalidae (Lepidoptera)

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

Phylogeny of feather mite subfamily Avenzoariinae (Acari: Analgoidea: Avenzoariidae) inferred from combined analyses of molecular and morphological data.

Phylogenetic relationships among feather mites of the subfamily Avenzoariinae (Acari: Analgoidea: Avenzoariidae) were reconstructed by parsimony analysis of a combined data matrix. We analyzed 41 morphological characters and 246 molecular characters from a fragment of the 16S rDNA. Morphological trees were well supported at deep branches (genera and above), but showed much less support and resolution within genera. Molecular analyses produced trees with better resolution and support on terminal branches and worse support on basal branches. I(MF) index for the combined matrix pointed to the significant congruence of both data subsets with the whole of the data. The topology of the combined tree was close to the morphological tree in the deep branches and had well-resolved terminal branches as in the molecular tree. This suggests a considerable level of complimentarity between the two data sets. An analysis of association patterns of the mites and their hosts was conducted based on the results of the combined analyses for the Avenzoariinae and a phylogeny of their charadriiform hosts (compiled from various bird phylogeny hypotheses). The trees could be reconciled by the invoking of 12-13 cospeciation events, 6-7 duplications, 2 host shifts, and 26-29 sorting events. This suggests a high degree of cospeciation.     

Additions to the geometrid fauna (Lepidoptera: Geometridae) of Chile

Three species of geometrid moths are reported from the extreme North of Chile. All three are new for the Chilean fauna: Scopula umbilicata (Fabricius, 1794) (Sterrhinae), Cataspilates grisescens Warren, 1897, and Pero obtusaria Prout, 1928 (Ennominae).     

Two new species of the Hagnagora anicata complex (Geometridae, Larentiinae) from Costa Rica

Examination of the lectotype (here designated) reveals that Hagnagora anicata (Felder & Rogenhofer) does not occur in Costa Rica. Instead two new species are described, Hagnagora eliannesp. n. and Hagnagora unniasp. n., and their distribution is discussed. The previous treatment of Hagnagora anicata as a single widespread species ranging from Jamaica and Mexico to Bolivia needs to be critically evaluated.     

Maintenance of the phenology of the winter moth (Lepidoptera: Geometridae)

Adult winter moths ( Operophtera brumata (L.)) are active in late autumn or early winter. The eggs overwinter in the canopy of trees and hatch simultaneously with the bursting of host tree buds. Many young larvae disperse on the wind on silk strands. Larvae are polyphagus and feed until late spring when they pupate in soil or leaf litter. The duration of the egg and pupal stages is genetically determined and varies with latitude. The egg stage is long in the north and short in the south, while the pupal stage is short in the north and long in the south.
The literature on the ecology and physiology of winter moth is reviewed. The factors maintaining the unusual phenology are discussed. It is concluded that the larval stage is early because mature leaves of many host trees are unsuitable as food, because parasitism against later larvae is more intense, and because summer temperatures may be injurious to larvae. The adult period is late in the year so that the final stages of pupal development occur in cool conditions and so that adults emerge after most insect predators have ceased activity. Throughout most of the range retarding the adult emergence period would cause activity to be impeded by severe winter weather; in the south this is not so and it is suggested that eggs must be on the trees for a minimum period to ensure synchronization of egg hatch with bud burst. The protracted adult emergence period may be an adaptation reducing predation by birds.