Forewing Colour Research Articles

The First Representative of the Roachoid Family Spiloblattinidae (Insecta, Dictyoptera) from the Late Pennsylvanian of the Iberian Peninsula.

Simple SummaryThe Palaeozoic–Early Mesozoic roachoid family Spiloblattinidae are valuable for understanding the stratigraphy of continental strata, thanks to their diversity of the forewing venation patterns of colouration. Here, the first Iberian representative of these roachoids is described as a new ‘form’ closely related to the Gzhelian–early-middle Asselian ‘zone species’ and ‘forms’. It supports the latest Gzhelian age of the concerned outcrop, obtained through stratigraphy and floral composition. It confirms the value of these insects for stratigraphic purposes.Sysciophlebia ‘sp. form Villablino’, the first Iberian representative of the Palaeozoic–Early Mesozoic family Spiloblattinidae, is described and illustrated. Its forewing colour pattern is strongly similar to those of the Gzhelian–early-middle Asselian species Sysciophlebia euglyptica, Sysciophlebia ilfeldensis, Sysciophlebia rubida, and ‘Sysciophlebia sp. form KBQ’, supporting the currently proposed Gzhelian age for its type locality. It supports the use of the representatives of the Spiloblattinidae for stratigraphic purposes. The diagnoses and limits of the families Subioblattidae, Phyloblattidae, Compsoblattidae, Spiloblattinidae, and of the spiloblattinid genera are discussed.

A new species of the genus Coptotriche (Lepidoptera: Tischeriidae) from Okinawa Island, Japan

A new Symplocos trumpet leafminer, Coptotriche yanbarensis Kim & Yagi sp. nov., feeding on S. stellaris and S. okinawensis (Symplocaceae) is described from Okinawa Island, Japan. The new species is distinguishable from the Japanese Symplocos feeding described species, C . symplocosella Kobayashi & Hirowatari, 2016 by the forewing coloration, the shape of genitalia, and also DNA barcodes.

Phenotypic variability in the Old World bollworm,Helicoverpa armigera(Hübner) populations in India

AbstractHelicoverpa armigera(Hübner) is one of the most prominent polyphagous species of the Heliothinae pest complex that inflicts severe damage to a wide range of crops in India. Knowledge regarding the population structure of the pest species, whether morphological or genetic, is considered as an essential tool in making effective management decisions. Thus here, we performed the phenotypic characterization ofH. armigerapopulations collected from varied geographic locations across India. Studied populations differed significantly for several external morphometric traits studied at larval, pupal and adult stages. Significant differences were also observed with respect to the intensity of black pigmentation on larval body as well as adult eye and forewing colour patterns. Besides external phenotypic traits, the length of genital organs like aedeagus and valva in males, and bursa copulatrix and bursa seminalis in females also differed significantly amongst populations. The dendrogram based on selected traits showed clear cut differentiation of studied populations into two major groups, one including all the South Zone populations and the other having populations from North and Central Zones. Differences based on phenotyping in the present study indicate the possibility of the existence of different subspecies within the Indian populations ofH. armigera.

Four new species and one new subspecies of the Ghoria albocinerea Moore, 1878 species-group from northern Indochina and China (Lepidoptera, Erebidae, Arctiinae, Lithosiini).

The Ghoria albocinerea Moore, 1878 species-group is established for a complex of species having the characteristic silver whitish forewing coloration and pattern presented mostly by a broad longitudinal shade along the anal wing margin. Four new species and one new subspecies of the species-group are described: Ghoria nanlingica dytiko ssp. n. (China: Sichuan), Ghoria dammay sp. n. (North Vietnam), Ghoria longivesica sp. n. (China: Yunnan and North Thailand), Ghoria mubupa sp. n. (China: Yunnan) and Ghoria synnefo sp. n. (North Vietnam). Adults, male and female genitalia of the new taxa and the related Ghoria albocinerea (the central and eastern Himalayas), Ghoria nanlingica Dubatolov, Kishida Wang, 2012 (China: Guangdong) and Ghoria sinotibetica Dubatolov, Kishida Wang, 2012 (China: Sichuan) are illustrated.

Whole-chromosome hitchhiking driven by a male-killing endosymbiont.

Neo-sex chromosomes are found in many taxa, but the forces driving their emergence and spread are poorly understood. The female-specific neo-W chromosome of the African monarch (or queen) butterfly Danaus chrysippus presents an intriguing case study because it is restricted to a single 'contact zone' population, involves a putative colour patterning supergene, and co-occurs with infection by the male-killing endosymbiont Spiroplasma. We investigated the origin and evolution of this system using whole genome sequencing. We first identify the 'BC supergene', a broad region of suppressed recombination across nearly half a chromosome, which links two colour patterning loci. Association analysis suggests that the genes yellow and arrow in this region control the forewing colour pattern differences between D. chrysippus subspecies. We then show that the same chromosome has recently formed a neo-W that has spread through the contact zone within approximately 2,200 years. We also assembled the genome of the male-killing Spiroplasma, and find that it shows perfect genealogical congruence with the neo-W, suggesting that the neo-W has hitchhiked to high frequency as the male-killer has spread through the population. The complete absence of female crossing-over in the Lepidoptera causes whole-chromosome hitchhiking of a single neo-W haplotype, carrying a single allele of the BC supergene and dragging multiple non-synonymous mutations to high frequency. This has created a population of infected females that all carry the same recessive colour patterning allele, making the phenotypes of each successive generation highly dependent on uninfected male immigrants. Our findings show how hitchhiking can occur between the physically unlinked genomes of host and endosymbiont, with dramatic consequences.

TWO NEW DARK COLORED SPECIES OF THE GENUS BARSINE WALKER, 1854 FROM CHINA AND INDOCHINA (LEPIDOPTERA, EREBIDAE, ARCTIINAE, LITHOSIINI)

In the present paper, two new species, Barsine persephone Volynkin & Černý, sp. nov. (N Thailand and N Vietnam), and Barsine asotoida Volynkin & Černý, sp. nov. (SW China and N Myanmar) are described. A new combination is established: Barsine specialis (Fang, 1991), comb. nov. Both new species belong to the Barsine prominens (Moore, 1878) species-group, and, as well as B. specialis have unusual for the genus dark brown forewing coloration. Adult, male and female genitalia are illustrated.

Aeolothrips neyrizi, a new species of the genus Aeolothrips (Thysanoptera: Aeolothripidae) from southern Iran

Aeolothrips neyrizi n. sp. is described and illustrated from Fars province, southern Iran, on the basis of two females collected from Alhagi persarum (Fabaceae). The fore wing colour is characteristic of the species. The new species is closely related to A. afghanus, a species that has been described from Afghanistan and also reported from Iran.

New species and stages description of Ulmeritoides Traver, 1959 (Ephemeroptera: Leptophlebiidae) from Roraima State, Northern Brazil

A new species of Ulmeritoides Traver, 1959 is described based on all life stages from Roraima State, Northern Brazil. U. amajari sp. nov. has a forewing coloration and penis shape similar to that of the widespread species U. uruguayensis (Traver) but it differs from the latter by the shape of the penis lobe apex in the male imago and by the lines of pectinate setae on the ventral surface of tibia III, in the nymph. The descriptions of nymph and female imago of U. passorum Gama-Neto & Hamada, 2014 are also presented.

Diversity of tortricid moths in apple orchards: evidence for a cryptic species of Grapholita (Lepidoptera: Tortricidae) from China.

Understanding herbivore diversity both at the species and genetic levels is a key to effective pest management. We examined moth samples from multiple locations from a major apple growing region in China. For specimen collection, we used a pheromone trap designed to attract Grapholita molesta (Busck) (Lepidoptera: Tortricidae). Surprisingly, we found a second species captured at high proportions. Its external morphology (e.g., male genitalia and forewing coloration) was the same as for Grapholita funebrana Treitschke (Lepidoptera: Tortricidae) specimens from Europe. However, the barcode sequence of the mitochondrial gene cytochrome oxidase I (COI) diverged markedly between specimens from China and Europe, and the genetic distance value between the specimens from the two regions as estimated using the Juke-Cantor (JC) model amounted to 0.067. These morphological and molecular findings together point to a cryptic species in G. funebrana from China. Further molecular analyses based on COI and COII genes revealed its extremely high genetic diversity, indicating that the origin of this species includes the sampling region. Moreover, molecular data suggest that this species passed through a recent population expansion. This is the first report on a cryptic species in G. funebrana, as well as the first report on its genetic diversity.

Occurrence and Ecological Characteristics of Chiasmia cinerearia (Bremer et Grey)(Lepidoptera: Geometridae) Attacking Sophora japonica

Chiasmia cinerearia (Bremer et Grey) belonging to the subfamily Ennominae(Lepidoptera: Geometridae) is a new forestry pest attacking to Sophora japonica in Gyeonggi province. Chiasmia cinerearia oviposited about 250~300 eggs per female on leaves. Larva had five instar stages. Adults emerged twice a year in early May and late July. The period of one generation was about 48 days. Body length of larvae was 25~30 mm and body color was light green with dark yellow lateral lines in abdomen. The wingspan of adults was 30 to 35 mm in length. The color of forewings was greyish brown with quadrilateral patterns on outer lines.

Two New Yellow-Banded Sister Species ofSyntomaulaMeyrick (Lepidoptera: Gelechioidea: Cosmopterigidae) from Papua New Guinea Associated with Rubiaceae

ABSTRACT. Two new species, Syntomaula xanthofasciata n. sp. and S. flavoangulata n. sp. (Lepidoptera: Gelechioidea: Cosmopterigidae) are described from Papua New Guinea and are associated with Neonauclea obversifolia (Valeton) Merr. & L.M. Perry, Uncaria appendiculata Benth., and Uncaria cordata (Lour.) Merr. (Rubiaceae). Macromorphological features and DNA barcodes provide corroborative evidence to diagnose these similarly patterned species. Illustrations of the wing pattern, wing venation, and male and female genitalia are provided. The distinctive yellow and brown forewing coloration is discussed with regard to potential mimicry.

A new species of Aboilus (Insecta, Orthoptera) from the Jurassic Daohugou beds of China, and discussion of forewing coloration in Aboilus

Wang, H., Li, S., Zhang, Q., Fang, Y., Wang, B. & Zhang, H., 13.02.2015. A new species of Aboilus (Insecta, Orthoptera) from the Jurassic Daohugou beds of China, and discussion of forewing coloration in Aboilus. Alcheringa 39, xxx–xxx. ISSN 0311-5518He Wang* [wanghe0701@163.com], Sha Li* [lisha.paleo@gmail.com], Qi Zhang* [zhqi1105@126.com], Yan Fang [yanfang@nigpas.ac.cn], Bo Wang† [bowang@nigpas.ac.cn] and Haichun Zhang [hczhang@nigpas.ac.cn], State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, Nanjing 210008, PR China.*Also affiliated with University of Chinese Academy of Sciences, Beijing 100049, PR China. †Also affiliated with Steinmann Institute, University of Bonn, Bonn 53115, Germany.A new species of Aboilinae (Orthoptera: Prophalangopsidae), Aboilus perbellus, is described and illustrated based on three well-preserved forewings recovered from the Middle–Upper Jurassic Daohugou beds of Inner Mongolia, China. The new species differs from all congeneric forms in its special forewing coloration and features of its wing venation. To date, three types of forewing coloration have been found among different species of Aboilus at Daohugou, suggesting that these taxa inhabited different ecotopes.

Forewing pigmentation predicts migration distance in wild-caught migratory monarch butterflies

Surprisingly, little is known about how the environment influences the production of the iconic orange coloration of the monarch butterfly (Danaus plexippus). Previous research under controlled laboratory conditions has shown that the temperature during larval development influences the color of monarch wings, where females raised in warm conditions had a greater proportion of melanization, whereas males raised in warm conditions had a lower proportion of melanization. These melanin-based colors have been found to increase flying ability in Lepidoptera, and recent experiments have found that monarchs with redder forewings flew greater distances than monarchs with less intense coloration. We examined whether wild-caught monarchs captured in the Great Lakes region exhibited geographic polyphenism by using stable isotopes to estimate natal origin, and hence rearing temperature, spectrophotometry to measure forewing coloration, and image analysis to estimate shape. We found that monarchs from the Gulf Coast were more melanized than monarchs from the Great Lakes, and southern male monarchs were more saturated than northern male monarchs. This supports previous research suggesting that colors that absorb more solar energy allow for greater flying ability but contradicts the patterns we expected based on natal temperature. Interestingly, this effect of color on migration distance was independent of wing shape. We provide the first evidence that the coloration of wild monarchs influences their migration ability over a continental scale, and we suggest that these differences in color may benefit the cohort of monarchs destined for long-distance migration to their wintering ground.

Epinotia cinereana (Haworth, 1811) bona sp., a Holarctic tortricid distinct from E. nisella (Clerck, 1759) (Lepidoptera: Tortricidae: Eucosmini) as evidenced by DNA barcodes, morphology and life history

DNA barcodes of European tortricid moths identified as Epinotia nisella (Clerck, 1759) were found to comprise two ge-netically distinct clusters. These coincided with E. nisella and E. cinereana (Haworth, 1811) (sp. rev.), the latter havingbeen considered a synonym of the former for several decades. Comparing these DNA barcodes with those of North Amer-ican Epinotia showed that both species are Holarctic. The North American Proteopteryx criddleana Kearfott, 1907 is anew junior synonym of E. cinereana (syn. nov.). The two species also show distinct differences in male and female gen-italia. North American populations of both species show moderate differences in barcodes from their respective Europeanpopulations but there are no morphological differences correlated with the intraspecific barcode clusters. Tortrix petranaHübner, 1813 is considered as a junior synonym of E. cinereana (syn. rev.). Epinotia nisella has several synonyms reflect-ing its highly variable forewing colour pattern. We discuss its variation and the status of the names associated with it. The biology of both species in Europe and North America is summarized. Adults and genitalia are illustrated.

A new leaf-feeding species of Neohydatothrips from southwestern China (Thysanoptera: Thripidae)

Neohydatothrips magnoliae sp. n., is described from leaves of Parakmeria yunnanensis (Magnoliaceae) in SouthwesternChina. This thrips is similar to N. samayunkur, N. elaeagni, N. trypherus and N. plynopygus, but can be distinguished fromthese by details of the fore wing colour, the body colour, and shape of the pronotal blotch. An illustrated key is providedto distinguish the 13 described species of Neohydatothrips recorded from China. Type specimens of the new species are preserved in Yunnan Agricultural University.

Description of 33 new species of Calamoceratidae, Molannidae, Odontoceridae and Philorheithridae (Trichoptera), with detailed presentation of their cephalic setal warts and grooves

Additions to the taxonomy of the leptoceroid families Calamoceratidae, Molannidae, Odontoceridae, and Philorheithridae are given, and 33 new species in the superfamily are described based on characters in the male genitalia, head and thoracal setose warts, and groove patterns, wing venation, and forewing colour and pattern. In the Calamoceratidae Ulmer, the Anisocentropus latifasciata diagnostic species-group and Anisocentropus brevipennis diagnostic species cluster are described for the first time. The following 17 species are described as new: Anisocentropus bungus (Vietnam), A. csorbai (Vietnam), A. dvaupadhah (Malaysia), A. fridae (Fiji Islands), A. hannahae (Fiji Islands), A. hoisat (Laos PDR), A. malaisei (Myanmar), A. maralus (Malaysia), A. mjoebergi (Borneo), A. samuh (Madagascar), A. tapenan (Solomon Islands), A. thinlin (Laos PDR), A. thonmihn (Vietnam), A. vanuensis (Fiji Islands), A. vitiensis (Fiji Islands), Ganonema malickyi (Thailand) and G. rong (Vietnam). In addition, Ganonema fuscipenne (Albarda) is synonymized with G. ochraceellum (McLachlan), and Ganonema pallidum Martynov is synonymized with Anisocentropus kawamurai (Iwata). In the Molannidae Wallengren, a single new species, Molanna gamdaha, is described from Myanmar. In the Odontoceridae Wallengren, the genus Inthanopsyche Malicky is synonymized with Psilotreta Banks. The following 14species belonging to previously described genera, are described as new: Lannapsyche birathena (Myanmar), L. suksma (Myanmar), L. kamba (Myanmar), Marilia enikiana (Laos PDR), M. jonssoni (Laos PDR), M. katakaha (Malaysia), M. malickyi (Laos PDR, Malaysia, Vietnam), M. mendolonga (Malaysia), M. namha (Laos PDR), M. tuyetmira (Laos PDR), Phraepsyche pectinata (Vietnam), P. yitungshana (China: Hong Kong), Psilotreta enikoae (Vietnam), and P. malickyi (Myanmar). In addition, the following 3 diagnostic species-groups are described for the first time: Psilotreta japonica diagnostic species-group, Psilotreta trimeresuri diagnostic species-group, and Psilotreta frontalis diagnostic species group. The species Ganonema odaenum Kobayshi is synonymized with Psilotreta locumtenens Botosaneanu, and Psilotreta pyonga Oláh is synonymized with P. falcula Botosaneanu. In the Philotheithridae Mosely, a single new species, Psilopsyche granda, is described from Chile. In addition to describing new taxa, new species records for 34 species in the superfamily are presented.

Integrating morphology and mitochondrial DNA for species delimitation within the spruce budworm (Choristoneura fumiferana) cryptic species complex (Lepidoptera: Tortricidae)

Species in cryptic complexes tend to be very difficult, if not impossible, to identify using morphological characters. One such complex is the spruce budworm (Choristoneura fumiferana Clemens, 1865) species group, an economically important group of Nearctic forest pests. Morphological, ecological, behavioural and genetic characters have been studied to try to understand the taxonomy of this group better, but diagnostic character states differ in frequency rather than being complete replacements between each species. We used mitochondrial DNA (mtDNA), together with a new morphology-based character system that focuses on forewing colour components, to determine if one or a combination of character sources can be used for species diagnoses within the spruce budworm complex. We characterized 47 forewing morphometric measurements and sequenced a 470 bp region of cytochrome c oxidase I mtDNA for 111 ingroup individuals comprising five taxa within the complex. Larval host association and coloration or adult pheromone attraction were used as the prior method for grouping individuals. Our results showed that linear discriminant analysis of morphometric wing characters gave unique clusters for all species on the first and second canonical axes, except for a partial overlap between C. fumiferana and C. biennis, which are not sympatric in nature. In contrast, mtDNA distinguished C. fumiferana, C. pinus pinus Freeman, 1953 and a group of western species, but the three western species (C. occidentalisFreeman, 1967, C. biennisFreeman, 1967 and C. lambertiana Busck, 1915) shared mtDNA haplotypes. On the basis of the linear discriminant analysis of the combined character set, this study supports the application of both morphology and mtDNA within a framework of integrative taxonomy as the most accurate method for species identification. Furthermore, it demonstrates the utility of quantitative colour analysis, which may be particularly helpful for groups in which colour characters are difficult to divide into discrete units due to intergrading hues.

No genomic mosaicism in a putative hybrid butterfly species

Recent descriptions of hybrid animal species have spurred interest in this phenomenon, but little genomic data exist to support it. Here, we use frequency variation for 657 amplified fragment length polymorphism (AFLP) markers and DNA sequence variation from 16 genes to determine whether the genome of Heliconius pachinus, a suspected hybrid butterfly species, is a mixture of the putative parental species, Heliconius cydno and Heliconius melpomene. Despite substantial shared genetic variation among all three species, we show that the genome of H. pachinus is not a mosaic; both AFLP and DNA sequence data overwhelmingly associate H. pachinus with just one of the potential parents, H. cydno. This pattern also applies to the gene wingless, which is tightly linked to the locus that determines forewing colour-one specific H. pachinus trait that has been hypothesized to have originated from H. melpomene. As a whole, the data support a traditional, bifurcating model of speciation in which H. pachinus split from a common ancestor with H. cydno without a genetic contribution from H. melpomene. However, comparison of our data to DNA sequence data for another putative hybrid Heliconius species, Heliconius heurippa, suggests that the H. heurippa genome may be a mosaic.

Asymmetrical colour and wing-folding inTithrone roseipennis(Saussure 1870) a neotropical praying mantis (Mantodea Hymenopodidae)

The frequency and type of forewing colour asymmetry present in Tithrone roseipennis (Saussure 1870) (Mantodea Hymenopodidae) was examined and 29 out of 35 specimens had red/green forewing colour asymmetry. The insects, from both field-collected material and museum specimens, contained both left and right fore wing asymmetry morphs. This colour asymmetry was found to be linked to wing-folding. Asymmetrical mantids always appeared cryptically green at rest, decreasing the chance of predator detection. It is suggested that red and green colour pigments are genetically encoded for, and genetic epistatic interactions result between forewing colouration and wing folding mechanisms. The forewing colouration of T. roseipennis is an example of antisymmetry.

Morphological and Genetic Variation in the Speckled Wood Butterfly (Pararge Aegeria L.) Among Differently Fragmented Landscapes

We compared morphological and genetic variation between populations of the speckled wood butterfly (Parage aegeria L.) from four landscapes differing in degree of fragmentation: a continuous woodland, a set of woodland fragments, another, more isolated set of woodlots and a highly fragmented area with very small woodland fragments and hedgerows scattered in an intensively used agricultural landscape. Male butterflies were collected, weighed and their wing features (size and colour) measured by means of an image analysis system. We used Polyacrylamide Gel Electrophoresis (PAGE) to investigate genetic variation. The clearest morphological differences were found between the large, continuous woodland and the three other areas. Males of this woodland had a paler basal and distal forewing colour and larger yellow patches on the dorsal side of the forewing. Since these features are relevant to theroregulation and anti-predation, this result suggests differences with respect to microclimate and predation between the study areas. Relative thorax size also increased with degree of fragmentation. Since the thorax contains the flight muscles, this result may relate to differences in patterns of mobility or different costs for the same mobility pattern. The genetic diversity of the butterflies of the large, continuous woodland was significantly higher than in the other study areas. There were also indications for genetic differentiation. Hence, this study may point at adaptive morphological variation in the speckled wood butterfly among differently fragmented landscapes.