Molecular phylogeny of Sterrhinae moths (Lepidoptera: Geometridae): towards a global classification

A multigene phylogenetic study was carried out to test current, mostly morphology-based hypotheses on Sterrhinae phylogeny with additional material included from further geographical areas and morphologically different lineages. A maximum likelihood analysis (11 molecular markers and 7665 bp) was conducted on 76 species and 41 genera using iq-tree software. The resulting phylogenetic hypothesis is well resolved and branches have high support values. Results generally agree with earlier hypotheses at tribal levels and support the hypothesis that Sterrhinae comprises two major lineages. Based on the molecular phylogeny and extensive morphological examination, nine tribes are considered valid and the following taxonomic changes are introduced to recognize monophyletic groups: Mecoceratini Guenée, 1858 (= Ametridini Prout, 1910) is transferred from Desmobathrinae to Sterrhinae, and it is considered valid at tribal level new classification ; Haemaleini Sihvonen & Brehm is described as a new tribe and deemed sister to Scopulini + Lissoblemmini; Lissoblemmini Sihvonen & Staude is described as a new tribe and sister to Scopulini; Lythriini Herbulot, 1962 is now a junior synonym of Rhodometrini Agenjo, 1952 syn.n. ; and Rhodostrophiini Prout, 1935 is now a junior synonym of Cyllopodini Kirby, 1892 syn.n. In addition, 48 taxa are transferred from other geometrid subfamilies to Sterrhinae, or within Sterrhinae from one tribe to another, or they are classified into a tribe for the first time, or a new genus classification is proposed. The results demonstrate the limited explanatory power of earlier classifications, particularly at the tribal level. This is probably a result of earlier classifications being based on superficial characters and biased towards the European and North American fauna. The species richness and distribution of Sterrhinae and its constituent tribes are reviewed, showing that the globally distributed Sterrhinae are most diverse in the Neotropics (31% of global fauna). They are species-rich in the Palaearctic (22%), Afrotropics (19%) and Indo-Malay (16%) regions, whereas they are almost absent in Oceania (1%). In terms of the described fauna, the most species-rich tribes are Scopulini (928 species), Sterrhini (876 species) and Cosymbiini (553 species), all of which have a cosmopolitan distribution. Mecoceratiini and Haemaleini are almost entirely Neotropical. Timandrini and Lissoblemmini, by contrast, are absent in the Neotropics. We present a revised classification of the global Sterrhinae fauna, which includes about 3000 putatively valid species, classified into nine tribes and 97 genera. Four genera are of uncertain position within Sterrhinae. Our results highlight the compelling need to include more genera from a global perspective in molecular phylogenetic studies, in order to create a stable global classification for this subfamily. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:zoobank.org :pub:A66F5DDD-06D6-4908-893E-E8B124BB99B1.     

Molecular phylogeny of swallowtail butterflies of the tribe Papilionini (Papilionidae, Lepidoptera).

Swallowtail butterflies of the tribe Papilionini number about 225 species and are currently used as model organisms in several research areas, including genetics, chemical ecology and phylogenetics of host plant utilization and mimicry, mechanisms of speciation, and conservation. We have inferred phylogenetic relationships for a sample of 18 species of the genus Papilio (sensu lato) and five outgroup taxa by sequencing two stretches of mitochondrial DNA that correspond to segments 12886-13370 and 12083-12545 of Drosophila melanogaster mitochondrial DNA and consist of sections of the genes for the large ribosomal RNA and subunit 1 of NADH-dehydrogenase. Our data support the monophyly of Papilio and, within it, of several traditionally recognized subgroups. Species belonging to groups that utilize primarily Rutaceae as larval foodplants form two clusters, corresponding to Old World and American taxa, respectively, while two previously recognized clades-of American and South Asian-Austronesian origin-whose members were known to feed mostly on Lauraceae and Magnoliaceae, are observed to form a clade. The sister group of Papilio is found to be the South Asian genus Meandrusa, which also happens to feed on Lauraceae. The latter plant family is therefore the probable larval host of the ancestor Papilio and the shift to Rutaceae (which four-fifths of extant Papilio species use as foodplants) is more likely to have occurred only after the initial diversification of the genus.     

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.     

Exploring phenotypic plasticity and biogeography in emerald moths: A phylogeny of the genus Nemoria (Lepidoptera: Geometridae)

The moth genus Nemoria (Lepidoptera: Geometridae) includes 134 described species whose larvae and adults display a considerable range of phenotypic plasticity in coloration and morphology. We reconstructed the phylogeny of 54 species of Nemoria and seven outgroups using characters from the mitochondrial genes, Cytochrome Oxidase I and II (COI and COII), and the nuclear gene, Elongation Factor-alpha (EF-1alpha). Maximum parsimony, maximum likelihood and Bayesian inference were used to infer the phylogeny. The 54 ingroup species represented 13 of the 15 recognized species groups of Nemoria [Ferguson, D.C., 1985. Fasc. 18.1, Geometroidea: Geometridae (in part). In: Dominick, R.B. (Ed.), The Moths of America North of Mexico, Fasc. 18.1. Wedge Entomological Research Foundation, Washington; Pitkin, L.M., 1993. Neotropical emerald moths of the genera Nemoria, Lissochlora and Chavarriella, with particular reference to the species of Costa Rica (Lepidoptera: Geometridae, Geometrinae). Bull. Br. Mus. Nat. Hist. 62, 39-159], and the seven outgroups came from four tribes of Geometrinae. These data support Nemoria as a monophyletic group and largely recover the species groupings proposed in previous taxonomic analyses using morphological characters. Phenotypic plasticity of larvae is not correlated with plasticity of adults among those species of Nemoria where life histories are known, and appears to be evolutionarily labile for both life history stages: Species exhibiting larval phenotypic plasticity, such as N. arizonaria and N. outina, are placed in several distinct clades, suggesting that this trait has evolved multiple times, and species displaying adult phenotypic plasticity are likewise distributed throughout the phylogeny. A comparative analysis of the biogeographic history of Nemoria supports a South American origin for the genus with multiple introductions into North America, and an application of published substitution rates to the phylogram provides an age estimate of 7.5 million years.     

Molecular phylogeny and evolutionary biology of Acrodipsas (Lepidoptera: Lycaenidae)

Most butterflies in the family Lycaenidae associate with ants but fewer than 3% are myrmecophagous. Larvae of the Australian endemic butterfly genus Acrodipsas parasitise the nests of ants from two subfamilies and, thus, constitute an interesting and uncommon myrmecophagous radiation within the Lepidoptera. Phylogenetic relationships among Acrodipsas species were inferred from fragments of mitochondrial cytochrome oxidase subunits I and II totalling 1155 bp using maximum parsimony and a neighbour joining method. Monophyly of the genus was confirmed, as was the sister genus status of Lucia. Acrodipsas myrmecophila was established as the plesiotypic Acrodipsas species, which together with Acrodipsas brisbanensis parasitises the ancestral dolichoderine host-ants. A speciation event associated with a radical host-ant shift to Myrmicinae occurred in an ancestor of Acrodipsas cuprea but subsequent speciation events may have been driven by climatic fluctuations during the Pleistocene. Modifications to leg morphology in several species were found to be a synapomorphic state, which arose subsequent to the host-shift to Myrmicinae. Minimal genetic variation detected in allopatric species exhibiting divergent morphology suggests that phenotypic variability has been driven by strong environmental selective pressures. As a result, morphological differences between closely related allopatric species have evolved faster than genetic differences, most notably between Acrodipsas arcana and Acrodipsas illidgei. In contrast, sympatric lineages of A. brisbanensis across similar habitats show considerable genetic differentiation, yet have remained phenotypically indistinguishable. Successful amplification of short overlapping fragments of DNA from museum specimens confirms their utility for phylogenetic analysis when the availability of fresh tissue is limited.     

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-1α (Ef-1α; 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 .).     

The genus Cleora Curtis (Lepidoptera: Geometridae) in Fiji

An account is given of the 10 species of the genus Cleora known to inhabit Fiji; two of these are described as new and two are transferred to Cleora from other genera. Species previously known from Fiji are re-described, and a key to the males is included.     

Molecular phylogeny of Eois (Lepidoptera,Geometridae): evolution of wing patterns and host plant use in a species‐rich group of Neotropical moths

Strutzenberger, P., Brehm, G., Bodner, F. & Fiedler K. (2010). Molecular phylogeny of Eois (Lepidoptera, Geometridae): evolution of wing patterns and host plant use in a species‐rich group of Neotropical moths. —Zoologica Scripta, 39, 603–620. Eois is a pantropical genus of Geometridae moths with currently 250 valid described species, the majority of which occur in the Neotropics. Eois is a prominent component of Andean moth communities locally accounting for up to ～10% of geometrid individuals. We address the evolution of wing patterns and host plant use in Neotropical Eois and provide a preliminary assessment on the monophyly and biogeographic history of the entire genus as well as affinities within the subfamily Larentiinae. We applied Bayesian, maximum likelihood and maximum parsimony methods of phylogenetic reconstruction to a 142 taxon dataset of partial COI (1220 bp) and Ef1α (1066 bp) sequences resulting in the largest taxon set of geometrid moths analyzed in a molecular phylogenetic study so far. Monophyly of Eois was always strongly supported. Ten monophyletic clades were found with good support, seven of which have characteristic wing pattern phenotypes. Only one wing pattern type occurs in two clades. Trophic associations with representatives of the family Piperaceae occur in all 8 (of 9) Neotropical clades for which host information is available. Apart from feeding on Piper, at least two Eois species in Ecuador feed on Peperomia, and one on Manekia (all Piperaceae); two further species live on Hedyosmum (Chloranthaceae). Species feeding on Peperomia, Manekia and Hedyosmum are usually nested in Piper‐associated clades. Single records of associations with Gesneriaceae and Monimiaceae are scattered in otherwise Piperaceae‐associated clades. These patterns suggest multiple parallel host shifts away from Piper as ancestral food plant. Old World Eois were recovered as monophylum and sister to Neotropical Eois. Within the subfamily Larentiinae the genus Eois has previously been placed close to the tribe Eupitheciini, but this was not supported in our phylogenetic analyses.     

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.     

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).     

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.     

Embryonic development in two species of Chesias (Lepidoptera: Geometridae)

The embryonic development of Chesias legatella Schiffermiiller and C. rufata Fabricius is described and compared. Eggs of C. rufata develop in approximately nine days at 20°C; those of C. legatella overwinter and hatch after five or six months. The morphology of the developing embryo is described in detail and is essentially similar in both species except during the formation and early development of the germ band. Morphogenesis is divided into a number of arbitrary stages and the relative duration of the different stages is compared in these and other species.     

Temperature-dependent oviposition model of Scopula subpunctaria (Lepidoptera: Geometridae)

Scopula subpunctaria, is one of important defoliating insect pests of tea plants in China. However, the effect of temperature on its longevity and fecundity had not been reported, and the temperature-dependent oviposition model is lacking. The reproductive characteristics of S. subpunctaria were investigated under eight constant temperatures (13, 16, 19, 22, 25, 28, 31, and 33℃), with a 14L: 10D photoperid and 65–85% RH. The results showed that temperature significantly influence oviposition period, longevity, and fecundity of S. subpunctaria. The mean longevity of female adults increased from 10.8 days (33℃) to 33.9 days (13℃). Male lived shorter than female did. The mean longevity of males was shortest at 33℃ (8.1 days) and longest at 23.8℃ (17.6 days). Total fecundity of S. subpunctaria females ranged from 15.4 eggs at 33℃ to 279.9 eggs at 22℃. The temperature-dependent oviposition model of S. subpunctaria was constructed based on four sub-models (developmental rate model, total fecundity model, oviposition rate model, and survival rate model). The results of this study would be helpful in forecasting and predicting population occurrence of S. subpunctaria in tea plantations.     

茶尺蠖和灰茶尺蠖内共生菌Wolbachia的分子检测及序列分析

【目的】对茶尺蠖Ectropis obliqua及其近缘种灰茶尺蠖E.grisescens体内共生菌Wolbachia进行分子鉴定,确定两者体内Wolbachia的感染率及其进化地位,为进一步探讨其对茶尺蠖和灰茶尺蠖的潜在影响提供科学依据。【方法】采用Wolbachia的16S r RNA、fts Z和wsp基因特异性引物,通过PCR扩增法检测了我国3个茶尺蠖地理种群(浙江杭州、余杭和江苏无锡)和3个灰茶尺蠖地理种群(浙江新昌、湖北浠水和江西南昌)中Wolbachia的感染情况,并进行测序和序列分析。【结果】茶尺蠖和灰茶尺蠖都感染了Wolbachia,灰茶尺蠖的Wolbachia感染率为100%,但茶尺蠖的Wolbachia感染率在22%~95%,且PCR产物电泳得到的条带微弱。wsp序列在茶尺蠖和灰茶尺蠖种间、种内无差异;但16S r RNA序列在茶尺蠖和灰茶尺蠖种间、种内差异为0.362%~0.727%之间;茶尺蠖样本未成功扩增出fts Z序列,灰茶尺蠖样本获得2条fts Z基因序列差异为1.647%。基于Wolbachia的16S r RNA和wsp基因构建的系统发育树表明,本研究中茶尺蠖和灰茶尺蠖种群所感染的Wolbachia全部属于B组的Pip亚组。【结论】茶尺蠖和灰茶尺蠖均被B组Pip亚组的Wolbachia感染,但感染率相差很大,这为研究Wolbachia对茶尺蠖和灰茶尺蠖生物学及生态学的影响奠定了基础。     

Molecular phylogenetic and morphological studies on the systematic position of Heracula discivitta reveal a new subfamily of Pseudobistonidae (Lepidoptera: Geometroidea)

Heracula discivitta Moore is an uncommon moth species currently recorded from India, Nepal and China. Although this species has traditionally been placed in Lymantriinae, its systematic position in Macroheterocera has been enigmatic due to its unique morphological features. Here we used molecular and morphological data to explore the systematic position of H. discivitta. Our molecular phylogenetic analyses indicate that this species is sister to Pseudobiston pinratanai Inoue, a member of a recently established monotypic family Pseudobistonidae. The examinations of morphological features further show that H. discivitta shares synapomorphies with Pseudobistonidae. Based on the analysis results, we propose a new subfamily of Pseudobistonidae (Heraculinae subfam.n. ) to accommodate H. discivitta. The resemblance of the habitus to that of the brahmaeid genus Calliprogonos Mell & Hering is discussed. This published work has been registered on ZooBank, http://zoobank.org/urn:lsid:urn:lsid:zoobank.org:pub:63D17850‐6D51‐4E03‐A5D6‐F9EF6E7AF402 .