Primitive Lepidoptera Research Articles

Courtship and mating behavior of Endoclita signifer (Hepialidae: Lepidoptera) and the male sex pheromones in hairbrushes.

Endoclita signifer Walker is the most destructive wood-boring pest of Eucalyptus in China, causing significant economic and ecological damage. As an insect of the primitive Lepidoptera family Hepialidae, E. signifer fly and mat for only 10-20 min at dusk. The courtship and mating behavior of E. signifer adults and whether male moths release sex pheromones are still unknown, especially since transitory flight survival strategies in primitive moths differ from advanced moths like noctuids. In this study, we first observed the courtship and mating behavior of E. signifer by considering the effects of space and then analyzed extracts of male hairbrushes using gas chromatography-electroantennogram detection. Our results indicated that during the courtship period, flying males form courtship fields, lekking, and chase flying females before mating with them; E. signifer were more successful in mating in larger spaces (Length × Width × Height = 9.6 × 7 × 4 m); 5 compounds in the hairbrushes of the male moths which elicited antennal responses of 2 sexes, despite at high concentrations. Combined with it, indicating that communication between male and female may rely on male sex pheromones. These findings can serve as a basis for studying the mechanisms of sex communication in E. signifer and developing sex pheromone-based trapping techniques.

An annotated catalogue of nonditrysian Lepidoptera of India

The present catalogue comprises 88 species in 24 genera and 10 families of nonditrysian Lepidoptera from India. Information on type localities, records from India including their diversity in different biogeographic zones of India and from outside India is included. Among all, Hepialidae is the dominant family, consisting 25 species in four genera whereas, the least dominant families are Palaeosetidae and Incurvariidae with one species each. Biogeographically, North-East is home to maximum species (37 in eight families) of primitive Lepidoptera, followed by Himalaya (31 species in five families), Western Ghats (24 species in seven families) with Hepialidae as a dominating family in all the three zones. No nonditrysian is reported from Desert and Islands. Among the states of India, Meghalaya is the most diverse with 27 species in seven families.

First Report of the Immature Stages of the Leaf-Mining Genus Subclemensia Kozlov, 1987 (Lepidoptera: Incurvariidae), with a Re-Illustration of the Type Species and a Generic Concept Discussion Based on Immature Characters.

Simple SummaryCaterpillars and pupae are important sources of evidence for the identification of primitive moths, and they can often provide quite different but useful morphological characters from those of adults. Incurvariidae is one of the most primitive groups of extant Lepidoptera, but half of the genera lack any information on caterpillars and pupae. It is important to increase the knowledge on the immature stages of Incurvariidae. Subclemensia Kozlov, 1987 is one of the monotypic genera in Incurvariidae. In this study, the caterpillar and pupa of the type species of Subclemensia are reported for the first time. The leaf mine, host plant and its biological characteristics are also provided. DNA barcodes were generated to help with the species delimitation. The adult male and female genitalia are re-illustrated by color photography to supplement the original line drawings.The immature stages of primitive Lepidoptera can provide quite different but often useful morphological evidence and synapomorphies from those of adults. Incurvariidae is one of the most primitive lineages of extant Lepidoptera, which is species-poor but highly diverse, but half of the genera lack any information on immature stages. New knowledge on the immature stages of the family is expected to provide useful morphological evidence and synapomorphies to stabilize the generic nomenclature. Subclemensia Kozlov, 1987 is one of the monotypic genera in Incurvariidae. In this study, the immature stages of the type species of Subclemensia are reported for the first time. The leaf mine, host plant and its biological characteristics are also provided. DNA barcodes were generated to aid the species delimitation. The adult male and female genitalia are re-illustrated by color photography to supplement the original line drawings. The generic concepts of Subclemensia and other related genera are discussed based on immature characters.

On the systematic position of a highly derived amphiesmenopteran insect from Burmese amber (Insecta, Amphiesmenoptera)

ABSTRACTA small fossil insect with scales on the wings and body was identified as a representative of Aphiesmenoptera from Burmese amber. The species is introduced here as Tarachocelis microlepidopterella (†). The insect is described in detail, and photos and line drawings are provided for wing venation, head, mouthparts, scales, legs and abdomen. All characters shared with primitive Lepidoptera and Trichoptera are symplesiomorphies or groundplan traits of Amphiesmenoptera. In addition, the Burmese fossil has a number of remarkable autapomorphies, giving it an appearance that deviates clearly from known families of Lepidoptera and Trichoptera. The species, representing a family of its own, Tarachocelidae, is considered a separate and unique taxon in the stem-group of Amphiesmenoptera and is provisionally placed as Amphiesmenoptera incertae sedis.**NB: See Note Added in Proof (Section 4).

Phylogenetic and biogeographical analysis of the New Zealand Helicopsyche von Siebold (Trichoptera: Helicopsychidae)

AbstractThe phylogenetic relationship between the seven known New Zealand Helicopsyche von Siebold, 1856 species is outlined. The New Zealand Helicopsyche comprises Helicopsyche albescens Tillyard, 1924, Helicopsyche poutini McFarlane, 1964, Helicopsyche howesi Tillyard, 1924, Helicopsyche zealandica Hudson, 1904, Helicopsyche haurapango Johanson, 1999, Helicopsyche torinoJohanson, 1999, and Helicopsyche cuvieri Johanson, 1999 and the cladistic analysis revealed a single most parsimonious tree, ((albescens, (poutini, howesi)), (zealandica, (haurapango, (torino, cuvieri)))). The basal species in the two species groups formed, H. zeczlandica and H. albescens, are widely distributed in both North Island and South Island, while the derived species have more restricted distribution ranges and sister clades form non-overlapping distributions. New Zealand was divided into four and five separate geographical entities based on a) distribution patterns from the Helicopsyche and primitive Lepidoptera and b) distribution patterns from the Helicopsyche alone, respectively. To map the history of Helicopsyche in New Zealand reconciliation using COMPONENT 2.0 and Dispersal-Vicariance analysis by DIVA 1.1 was applied in the search for a reduced area cladogram and ancestral areas, respectively. The results indicate that the southernmost part was part of the ancestral area, and that the division of the earliest ancestor has a minimum age of 10 million years.

Sex chromatin and sex chromosome systems in nonditrysian Lepidoptera (Insecta)

Eleven representatives of the superorder Amphiesmenoptera (Trichoptera + Lepidoptera) were examined for sex chromatin status. Three species represent stenopsychoid, limnephiloid and leptoceroid branches of the Trichoptera; eight species belong to the primitive, so-called nonditrysian Lepidoptera and represent the infra-orders Zeugloptera, Dacnonypha, Exoporia, Incurvariina, Nepticulina and Tischeriina. The female-specific sex chromatin body was found in the interphase somatic nuclei of Tischeria ekebladella (Bjerkander 1795) (Lepidoptera, Tischeriina). The sex chromatin was absent in all investigated Trichoptera species as well as in all representatives of the nonditrysian Lepidoptera except Tischeria ekebladella. The sex chromosome mechanism of Limnephilus lunatus Curtis 1834 (Trichoptera, Limnephilidae) is Z/ZZ. The sex chromosome mechanism of Tischeria ekebladella (Lepidoptera, Tischeriina) is ZW/ZZ including the W chromosome as the largest element in the chromosome set. The data obtained support the hypothesis that the Z/ZZ sex chromosome system, the female heterogamety and the absence of the sex chromatin body in interphase nuclei are ancestral traits in the superorder Amphiesmenoptera. These ancestral characters are probably kept constant in all the Trichoptera and in the most primitive Lepidoptera. The W sex chromosome and the sex chromatin evolved later in the nonditrysian grade of the Lepidoptera. It is proposed that the sex chromatin is a synapomorphy of Tischeriina and Ditrysia.

The preoral cavity of lower Hymenoptera (Insecta): comparative morphology and phylogenetic significance

The skeletal and muscular morphology of the preoral cavity, including the labrum, hypopharynx and labium, was examined in the imago in representatives of all the ‘symphytan’ families as well as the apocritan families Stephanidae, Megalyridae and Trigonalyidae.Xyelidae have complex modifications for masticating pollen, remarkably similiar to those of primitive Lepidoptera. These modifications, collectively termed the triturating basket complex, include an asymmetrical distal epipharyngeal wall with a microtrichial brush and an enlarged infrabuccal pouch with heavy cuticular armature that interacts with the mandibles during feeding. There were striking structural differences between the two subfamilies of Xyelidae in the ligular region; the reduced glossa and clubshaped paraglossae of Macroxyelinae resembles those of primitive Lepidoptera, while the well developed, flattened glossa and paraglossae in Xyelidae are similiar to those of most other ‘Symphyta’.A putative transformation series, leading from a relatively large labrum with unsclerotised distal epipharyngeal wall lying anterior to the mandibles, as seen in Xyelidae and enthredinoidea, to a small and heavily sclerotised labrum and distal epipharyngeal wall lying posterior to the mandibles, as seen in ‘Siricoidea’, Orussidae and the Apocrita, was revealed. These modifications may be adaptations to enable the adult of the families pupating in wood to emerge from the pupal chamber.The Anaxyelidae, Orussidae and Apocrita have similiar configurations of the glossa and nsertions of the ventral premental adductors. This indicates a close affinity of the Anaxyelidae to Orussidae + Apocrita, a hypothesis that is in conflict with other character systems. The Orussidae and Stephanidae share a unique condition in the development of a pair of large apodemes attached to the labrum; this renders the groundplan state of the labrum in the Apocrita uncertain.Twentyfive characters were defined in an attempt to eludicate the ‘Symphyta’–Apocrita transition. A numerical cladistic analysis of the characters was undertaken, resulting in 522 minimum length trees. The characters are also discussed with reference to a cladogram which resulted from an analysis of the characters derived from the present study and a survey of characters from literature.

Presence of only eupyrene spermatozoa in adult males of the genus Micropterix hübner and its phylogenetic significance (Lepidoptera : Zeugloptera, Micropterigidae)

The morphology of the male genital organs and sperm ultrastructure were studied by light and transmission electron microscopy in adult males of two species of the genus Micropterix (Micropterigidae; Zeugloptera; Lepidoptera). The results are compared with findings from other primitive Lepidoptera and Trichoptera, and evaluated from a phylogenetic viewpoint. Both Micropterix species examined agree with regard to essential characters of the male genital organs. The paired testes are separate and show no internal compartmentalization. The male genital organs contain only mature nucleate (eupyrene) spermatozoa. Anucleate (apyrene) spermatozoa characteristically found in all other Lepidoptera were not observed. The eupyrene spermatozoa are filiform, measure 100 μm in length, and contain an elongated nucleus, 2 mitochondrial derivatives without paracrystalline materials, and a 9 + 2 axoneme without accessory tubules; the nucleus extends for almost the entire length of the spermatozoa. The absence of apyrene spermatozoa in Micropterix is in contrast to their presence in another species of the same family.

Local or global? Biogeography of some primitive Lepidoptera in New Zealand

Abstract Distribution maps are presented for 30 species of small forest moths representing two genera of Micropterigidae and one genus of Mnesarchaeidae. Evolutionary patterns are interpreted by means of the panbiogeographic method in order to seek a relationship between dispersal patterns within New Zealand and the overseas links of the taxa in question. It is established that different evolutionary lines indeed exhibit different geographic dispersal patterns within New Zealand. Moreover, certain characteristics of these patterns can be attributed to the overseas affinities of the groups, i.e., groups of organisms with New Caledonian or Australian affinities have a western disjunct pattern, whereas those with circum-Pacific affinities have a different pattern in which the taxa occupying a basal position in the phylogeny are to be found in the eastern South Island. However, both groups have overlapping centres of diversity, particularly in NW Nelson.

Phylogeny of Pupation in Trichoptera, with Proposals on the Origin and Higher Classification of the Order

Analysis of modes of pupation in Trichoptera reveals two fundamental types of pupal enclosures and concomitant systems for water circulation. In one (Rhyacophilidae, Hydrobiosidae, Glossosomatidae, Hydroptilidae-infraorder Spicipalpia Weaver), pupating larvae construct a closed cocoon of parchment-like silk, usually discrete from the pupal enclosure of small stones; water currents bathe the external surface of the cocoon during metamorphosis. In the other type (most families in the suborders Annulipalpia Martynov s.s. and Integripalpia Martynov s.s.), larvae con- struct a pupal cell with open meshes or holes at each end, permitting water currents to bathe the surface of the pupa directly during metamorphosis. Exceptions in the Philopotamidae, Stenopsy- chidae, Ecnomidae, and Phryganopsychidae are considered. The function of trichopteran cocoons during metamorphosis is considered, indicating that osmotic relations in closed cocoons of parchment-like silk in the Rhyacophilidae and allied families impede the efficiency of respiration mediated solely by diffusion of oxygen across the semipermeable wall of the cocoon. Because ovoid, closed cocoons of parchment-like silk also occur in primitive Lepidoptera, the sister group of Trichoptera, cocoons of this type are proposed as part of the groundplan of Tri- choptera. Consequently, the open pupal cells of Annulipalpia (retreat-makers) and Integripalpia (case-makers) are interpreted as derived. It follows that the habitat common to the families con- structing closed cocoons-cool, flowing waters-would likely have been the habitat in which Tri- choptera originated. The hypothesis proposed for the phylogeny of pupation in Trichoptera is that the closed cocoon of semipermeable silk in the ordinal groundplan required Trichoptera to become aquatic in cool, lotic waters; and that evolutionary innovation through subsequent derivation of cocoons of perme- able silk with ventilatory openings enhanced the efficiency of respiration, enabling Trichoptera to invade warmer waters of reduced current, and opening the way for radiation of the major lineages now extant-the retreat-making (Annulipalpia s.s.) and the case-making (Integripalpia s.s.) families. An alternative form of the higher classification of Trichoptera is proposed and discussed, elevating the Spicipalpia, termed the cocoon-making Trichoptera, to the rank of a third suborder, co-ordinate with the Annulipalpia and Integripalpia.

Floral Biology of Myristica fragrans Houtt. (Myristicaceae), the Nutmeg of Commerce

The flowering and pollination of Myristica fragrans were studied on plants cultivated in southern India. The staminate flowers are borne in indeterminate inflorescences and function only for a single night. Pistillate flowers are borne solitarily and appear receptive for 2-3 days. Staminate plants may produce over 50 times as many flowers as pistillate plants. Both types of flowers are strongly fragrant. A single species of beetle, Formicomus braminus (Anthicidae, ant-mimicking flower beetles), appears to be the best candidate for an effective pollinator of nutmeg in the area of the study, although other species of small beetles may be involved. The beetles are strong fliers, are active nocturnally around staminate flowers, and carry significant pollen loads. Pollen is the only evident pollinator reward. Pistillate flowers produce no reward, and spatial constraints of the perianth and stigma appear to prevent beetles from entering the flowers. Pollen carried on the beetle's head or body would be deposited on the stigma when the insect attempts to enter the flower. This is the first report of a food-deception automimicry in a cantharophilous pollination system. The pollen/ ovule ratio for M. fragrans is estimated to be 801,000/1, a very high P/0 ratio for an insect-pollinated plant. THERE ARE SEVERAL REASONS TO SUSPECT that the four Southeastern Asian genera of Myristicaceae may be beetle pollinated. These include (1) the predominance of white or cream to light-yellow flowers, (2) strong nocturnal fragrance, (3) lack of well-defined nectaries or other secretory structures (Wilson & Maculans 1967, Armstrong & Wilson 1978) (although reports of nectaries persist, e.g., Cobley 1976), and (4) the prevalence of beetle pollination among other members of the Annonales (Thien 1974, 1980; Faegri & van der Pijl 1979). However, flowers of Myristica, Horsfieldia, Knema, and Gymnacranthera differ morphologically from typical beetle-pollinated, annonalean flowers in several important respects. These genera are dioecious, as are all Myristicaceae, although there are reports of monoecious species in Iryanthera (Smith 1937). The flowers are small and do not form a broad cup or saucer. Pistillate flowers have one simple pistil containing a single ovule. Androecia consist of few to many, variously fused stamens rather than numerous, free, leaf-like stamens. In several genera, including Myristica, the perianth is fused into an urn-shaped enclosure. These differences suggest that Myristicaceae have either a nonbeetle pollinator or a different type of cantharophilous syndrome. Most accounts of nutmeg biology (e.g., Mcllroy 1967, Purseglove 1968, Cobley 1976) assume that pollination is effected by small insects, although wind pollination also is considered possible (Mcllroy 1967), and one researcher suggested that moths (Lepidoptera) are pollen vectors of M. fragrans (Deinum 1949). Pollen vectors of other annonalean families (Winteraceae, Degeneriaceae) include Diptera, Thysanoptera, primitive Lepidoptera, and Coleoptera (Thien 1980). In this study of M. fragrans we present preliminary information about the floral biology and pollination ecology of Myristicaceae. Specifically we made field observations and manipulated plants to determine (1) the phenology of staminate and pistillate flowers, (2) the relative abundance of staminate and pistillate flowers, (3) the nature of floral attractants and pollinator rewards, and (4) the identity and behavior of the effective pollinator.

Early embryonic development of the primitive moths, Endoclyta signifer walker and E. excrescens butler (Lepidoptera : Hepialidae)

Little information is available on the embryology of primitive moths. This paper describes and discusses the early embryogenesis of the monotrysian moths, Endoclyta signifer and E. excrescens. The small round germ disk is formed in the posterior ventral part of the egg. It invaginates into the yolk, and becomes the sac-like germ rudiment. Soon after, it separates from the rudimentary serosa, and completely sinks into the yolk. This sac-like germ rudiment develops into the germ band. In both species the embryonic envelopes are formed from the amniotic folds. The formation of the germ rudiment and embryonic envelopes in Endoclyta resembles those of Neomicropteryx nipponensis in the Zeugloptera and Stenopsyche griseipennis in the annulipalpian Trichoptera, and differs from those of the ditrysian Lepidoptera. The close affinity between the primitive Lepidoptera and the Trichoptera is supported by the present embryological evidences.

MORPHOLOGY OF THE FEMALE TERMINALIA IN LEPIDOPTERA, AND ITS TAXONOMIC SIGNIFICANCE

AbstractIn the phylogenetic classification of the Lepidoptera, the position of the genital opening and its modifications help to clarify the relationship of the Zeugloptera to the lepidopterous suborders. Three different systems of the suborder classification of the Lepidoptera, into Homoneura and Heteroneura (Tillyard, Imms, etc.), Monotrysia and Ditrysia (Borner), and Dacnonypha, Monotrysia, and Ditrysia (Hinton), are not supported by evidence obtained in the study of the female genitalia.The development of the female terminalia is closely associated with the mode of oviposition. Several types of female terminalia are found in the primitive Lepidoptera (Hepialidae, Neopseustidae, Tischeriidae, Lyonetiidae, Agathiphagidae, Tineidae, Nepticulidae, and Eriocraniidae). They are divided into four categories: hepialid type — the eggs are dropped randomly; typical lepidopterous type — the eggs are laid on the surface of food plants; eriocraniid type — the eggs are laid inside the food plants; tineid type — the eggs are laid in crevices of food. Each of the types is derivable from the types occurring in Neopseustidae, Tischeriidae, Agathiphagidae, and Eriocraniidae, respectively. These types are still far removed from the type of female terminalia in Micropterygidae of the Zeugloptera.The modes of egg-laying as well as the morphological characters of the female genitalia must be taken into consideration in suborder classification of the Lepidoptera.A genealogical tree based on the female terminalia suggests four main branches: hepialid branch — includes Neopseustidae and Prototheoridae; typical lepidopterous branch — includes all Ditrysia and Nepticulidae; tineid branch — includes only Tineidae; eriocraniid branch — includes Incurvariidae, Prodoxidae, Adelidae, and Heliozelidae.

A contribution towards an understanding of the morphology and phylogeny of the Ditrysian Lepidoptera

Summary 1. A morphological study of five body regions of the Lepidoptera has been made, these being (a) the radial system of the mesothoracic wing, (b) the mesothoracic sternopleural region, (c) the ventral thoraco-abdominal articulative (and adjacent) structures, including the integumental components of the tympanum where present, (d) the dorsal thoraco-abdominal articulation, etc., and (e) the metathoracic furca or endosternite. These regions were selected because the nature of the variation they exhibit suggested that, of the much larger number of regions primarily investigated, they are the most likely to provide information on the evolution of the higher taxa in the order. None of these regions had been investigated at all fully previously, and therefore a considerable amount of basic morphological study was necessary as a preliminary to phylogenetic discussion. This morphological work has been extended to include the primitive Lepidoptera (with the Trichoptera) in the case of the mesosternopleural r...

Morphological observations on the wing scales in some primitive lepidoptera (insecta)

Scanning electron micrographs show that the ordinary wing scales in the families Micropterygidae and Eriocraniidae are not transversely ribbed as in higher Lepidoptera, whereas some of the scales in the family Mnesarchaeidae are. In the two former families the wing surface scales are seen in thin section to be solid, while in the latter solid scales occur together with hollow scales resembling those of higher Lepidoptera. In Micropterygidae and Eriocraniidae hollow scales are present only along the wing margins. Longitudinal ridges are present on the outer surface of all scales; only occasionally are they seen on the surface facing the wing and then close to the margin of the wing scale.

The Comparative Morphology of the Male Genitalia of the Primitive Lepidoptera.*

The primitive Lepidoptera discussed in this paper include the Jugate Lepidoptera, (Hepialidae and Prototheoridae), the Jugo-frenate Lepidoptera, (Mnesarchaeidae, Micropterygidae, and Eriocraniidae), and the Frenate superfamilies Tineoidea, Euceleoidea, and Bombycoidea.[†][1] The object in this discussion is to compare the male genitalia of these groups of Lepidoptera with each other and with those of nearly related orders of insects, and to ascertain from this comparison the value of these organs in gaining a more complete knowledge of the natural relationships of the insects treated. [1]: #fn-2