Salivary glands and salivary pumps in adult Nymphalidae (Lepidoptera)

The salivary glands and salivary pumps were investigated by means of dissection and serial semithin sections in order to expose the anatomy and histology of Nymphalidae in relation to feeding ecology. The paired salivary glands are tubular, they begin in the head, and extend through the thorax into the abdomen. The epithelium is a unicellular layer consisting of a single cell type. Despite the uniform composition, each salivary gland can be divided into five anatomically and histologically distinct regions. The bulbous end region of the gland lies within the abdomen and is composed of highly prismatic glandular cells with large vacuoles in their cell bodies. The tubular secretion region extends into the thorax where it forms large loops running backward and forward. It is composed of glandular cells that lack large vacuoles. The salivary duct lies in the thorax and also shows a looped formation but is composed of flat epithelial cells. The salivary reservoir begins in the prothorax and reaches the head. Its cells are hemispherical and bulge out into the large lumen of the tube. In the head the outlet tube connects the left and right halves of the salivary gland, and its epithelial cells are flat. The salivary pump lies in the head ventral to the sucking pump and leads directly into the food canal of the proboscis. It is not part of the salivary gland but is derived from the salivarium. Both the thin cuticle of the roof of the salivary pump and the thick bottom are ventrally arched. Paired muscles extend from the hypopharyngeal ridges and obviously serve as dilators for the pump. A functional interpretation of the salivary pump suggests that when not in use, the dilators are not contracted and the pump is tightly closed due to its own elasticity. When the dilator muscles repeatedly contract, the saliva is forced forward into the food canal of the proboscis. The salivary gland anatomy was found to be similar to other Lepidoptera. Furthermore, the histology of the salivary glands is identical in all examined butterflies, even in the species which exhibit specialized pollen-feeding behavior.     

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.     

Biology and morphology of the immature stages of Hermeuptychia atalanta (Lepidoptera: Nymphalidae: Satyrinae)

This paper describes the morphology of the immature stages (egg, larva and pupa) and larval behavior of the butterfly Hermeuptychia atalanta, a common and widespread species of satyrinae in South America. Eggs are white and spherical, laid singly on leaves of grasses. The first instar has a light-green body and dark-brown head; all remaining instars present body and head capsule light green. Larvae have no spines, and general morphology is similar to other species of the subtribe Euptychiina.     

A fake Heliconius (Lepidoptera, Nymphalidae)

Four ‘doubtful species’ are listed in catalogues of the genus Heliconius. All are non–existent: (1) H. Carolina (Herbst) is a forgery made from trimmed fragments of at least three butterflies, the two identifiable ones both beingH. erato; (2) H.cinereo–fuscus (Goeze) is a faded specimen of H.ricini; (3) H.arcuatus (Goeze) is so inadequately described that its identity cannot be determined, although it is apparently some species of heliconiine or ithomiine;(4)H.euclea (Godart) is the valid senior name of an ithomiid in the genus Hypothyris.     

Proboscis sensilla in Vanessa cardui (Nymphalidae, Lepidoptera): functional morphology and significance in flower-probing

 Morphology and distribution of the proboscis sensilla in Vanessa cardui have been investigated in order to contribute to the understanding of flower-probing behaviour in butterflies. The proboscis has a bend region approximately one-third of the length from the base. A short tip region is characterized by rows of intake slits leading into the food canal. Along the dorsal, lateral and ventral sides of the proboscis, sensilla trichodea, sensilla basiconica and sensilla styloconica are distributed in varying patterns depending on their distance from the b ase. The medial food canal bears one longitudinal row of sensilla basiconica only. The bristle-shaped sensilla trichodea are longer in the proximal region of the proboscis and become gradually shorter towards the tip. They are most frequent in number near to the bend region and near the beginning of the tip region. Sensilla basiconica arranged in longitudinal rows increase in number the more distal they are on the proboscis. The tip region is characterized by rows of sensilla styloconica on the dorsal side whereas the sensilla trichodea are mostly restricted to the ventral side. The ultrastructure suggests that the aporous sensilla trichodea function as mechanosensilla while the uniporous sensilla basiconica act as contact chemosensilla. The sensilla styloconica are regarded as bimodal contact chemo/mechanosensilla since their sensory cones are equipped with a single terminal pore and a tubular body at the base. The mouthpart sensilla appear to provide tactile cues on the positioning of the proboscis and on the degree of its insertion into a floral tube. Furthermore, they receive chemical stimuli on the availability of nectar and on the immersion status of the food canal. Accepted: 12 September 1997     

B chromosome variation in Euphydryas colon (Lepidoptera: Nymphalidae)

Cytogenetic analysis of an Idaho population of the checkerspot butterfly, Euphydryas colon, has revealed considerable inter- and intra-individual variation in chromosome number which turns out to be a classic case of B chromosome variation. The basic chromosome complement of the species is n (, )=31. The A chromosomes were aligned equatorially at mitotic metaphase and metaphase II, and axially at metaphase I, indicating a restriction of centric activity at the first meiotic division. No failure of pairing between homologous A chromosomes was observed and, although a marked asynchrony of chromatid separation was found to be characteristic of mitotic telophase and telophase II, the frequency of macrospermatid formation was low. The B chromosomes were at least partly heterochromatic but exhibited some variation in both pycnosity and size. Mitotically stable B-containing individuals showed a preponderance of unpaired Bs at first metaphase and these divided at either first or second anaphase. The presence of Bs was associated with a heightened production of abnormal spermatids particularly in individuals with high numbers of B chromosomes. Among the 25 individuals sampled, 21 carried from 1–6 B chromosomes, and of these 14 were mitotically stable. In all 7 unstable individuals the mean number of B chromosomes per cell exceeded the modal number. Assuming that the modal number represents the zygotic number, these results suggest that a mechanism to boost the number of B chromosomes exists in males of E. colon.     

Phylogenetic relationships among genera of danaine butterflies (Lepidoptera: Nymphalidae) as implied by morphology and DNA sequences

Cladistic relationships among genera and subtribes of Danaini (the milkweed butterflies) were inferred by analysis of data combined from five sources: morphology of adults and immature stages, and DNA sequences from three gene regions. The results corroborate and greatly increase support for prior hypotheses based on morphology alone. A new index summarizing incongruence among data partitions, the Partition Congruence Proportion (PCP), is introduced. The significance of the inferred pattern of phylogenetic relationships for comparative chemical ecology of milkweed butterflies is briefly discussed.     

Simulation of movement and egglaying inDanaus plexippus (Lepidoptera: Nymphalidae)

A model simulating the movement and oviposition of monarch butterflies over a female's life time is presented. The model's predictions compare favourably with observed data and suggest that females who lay eggs in an optimal fashion should have low directionalities in areas with high host plant density (patches and single plants) and high directionalities in areas with low host plant density. The model also provides one means of combining individual animal processes and spatial heterogeneity into population dynamics.     

Immature stages of Adelpha malea goyama Schaus (Lepidoptera: Nymphalidae, Limenitidinae)

The immature stages of Adelpha malea goyama Schaus are described from the Esta??o Ecol6gica do Caiuá, Paraná State, Brazil. The species uses the native liana Arrabidaea mutabilis Bur. & K. Schum. (Bignoniaceae) as larval hostplant in the study site. Isolated eggs are laid on leaf tips, and are similar in shape to those described for other species of Adelpha Hübner. Larvae pass through five stadia during 21 days, and first to third instars construct frass chains. Later instars rest on the upper leaf surface, and their body is covered by rigid branched scoli. The general profile of the pupa is elongated, with short head horns projecting laterally and with segment A2 projecting and curved anteriorly. Comparisons with closely related species of Adelpha are not conclusive, and more species in the genus need to be reared before a clear pattern of the variation in immature morphology becomes evident.     

The species of the genus Zethera Felder (Lepidoptera: Nymphalidae,Satyrinae)

The butterfly genus Zethera Felder, endemic to the Philippines and northern Sulawesi, is shown to include six species, four of which comprise the Z. pimplea (Erichson) superspecies. Type data is given for all nominal taxa, and a key to the species recognised is presented, together with data on their variation and distribution. Attention is drawn to the zoogeographic subregions of the Philippine islands, as originally described by Semper, and illustrated by the Zethera species. Their mimicry is also noted, with particular reference to its sex-limitation in the Z. pimplea superspecies, and non-limitation in Z. incerta (Hewitson) and Z. hestioides C. & R. Felder. The need to demonstrate whether or not Zethera represents a true monophyletic unit is briefly discussed.     

A new species of Cyllopsis (Lepidoptera: Nymphalidae: Satyrinae) from Costa Rica

Cyllopsis emilia Chacón and Nishida, a new satyrine species, is described from a single male specimen from Cerro de la Muerte, San José, Costa Rica. This new species can be distinguished from other species of Cyllopsis by its white coloration.     

Within population variation in the demography of Speyeria mormonia (Lepidoptera: Nymphalidae)

The adult demography of a population of Speyeria mormonia (Lepidoptera: Nymphalidae) near Gothic, Colorado USA was studied for four years. Values of demographic parameters, including survival, dispersal and sex ratio, varied yearly, without major changes in density. Shift in sex ratio with constant population numbers entails variation in effective population size among years. Flight period, catchability, relationship between body size and eclosion date, and significance of correlation between weather and daily survival rates differed between the sexes. Sexual dimorphism in correlation of wheather with survival rate may partially explain the lack of yearly variation in total population size. There was no significant sexual dimorphism in large scale habitat use, or in dispersal, except in 1982.
The demography of this insect is compared to that of related species of Argynnis and heliconiines, from the old world temperate zone and new world tropics, respectively. While some differences are observed which can be ascribed to differences in length of the growing season, many patterns are similar. This implies conservation of demographic traits, even across broad habitat and geographic areas.     

斐豹蛱蝶线粒体基因组全序列的测定和分析(英文)

该研究对斐豹蛱蝶(Argyreus hyperbius)(鳞翅目:蛱蝶科)线粒体基因组全序列进行了测定和初步分析。结果表明:斐豹蛱蝶线粒体基因全序列全长为15156bp,包含13个蛋白质编码基因、22个tRNA和2个rRNA基因以及1个非编码的A+T富集区,基因排列顺序与其它鳞翅目种类一致;线粒体全序列核苷酸组成和密码子使用显示出明显的A+T偏好(80.8%)和轻微的AT偏移(AT skew,?0.019)。基因组中共存在11个2～52bp不等的基因间隔区,总长96bp;以及14个1～8bp不等的基因重叠区,总长34bp。除COI以CGA作为起始密码子外,13个蛋白质编码基因中的其余12个基因是以ATN作为起始密码子。除COI和COII基因是以单独的一个T为终止密码子,其余11个蛋白质编码基因都是以TAA结尾的。除了缺少DHU臂的tRNASer(AGN),其余的tRNA基因都显示典型的三叶草结构。tRNA(AGN)和ND1之间的基因间隔区包含一个ATACTAA结构域,这个结构域在鳞翅目中是保守的。A+T富集区没有较大的多拷贝重复序列,但是包含一些微小重复结构:ATAGA结构域下游的20bp poly-T结构,ATTTA结构域后的(AT)9重复,以及位于tRNAMet上游的5bp poly-A结构等。这项研究所揭示的斐豹蛱蝶的线粒体基因组特征,不仅为认识蛱蝶科的遗传多样性贡献数据,而且对于该物种的保护生物学、群体遗传学、谱系地理及演化研究等具有重要意义。     

Homologies in the colour patterns of the genus Charaxes (Lepidoptera: Nymphalidae)

The phylogenetically and morphologically diverse patterns of Charaxes can be reduced to a simple set of pattern elements which can be homologized throughout the genus. At least five types of correspondence (homologies) exist among pattern elements: those between (1) species, (2) forewing and hindwing, (3) dorsal and ventral wing surface, (4) serial wing-cells, and (5) individual pattern elements within a single wing-cell. Differences in Charaxes colour patterns result from the distortion, elaboration, enlargement, reduction or loss of individual pattern elements. Further variation is often the result of dislocation of pattern elements from their serial homologues in neighbouring wing-cells, and fusion of individual pattern elements to create larger areas of colour. The type of analysis presented in this paper should be broadly applicable within the Lepidoptera and may prove useful in studying the systematics of colour patterns and the evolution of the developmental system that gives rise to them.     

Homologies in the colour patterns of the genus Heliconius (Lepidoptera: Nymphalidae)

The colour patterns of Heliconius butterflies are composed from a relatively simple set of pattern elements whose homologues are recognizable throughout the genus. Although Heliconius colour patterns look quite different from those of most nymphalids, these pattern elements are seen to derive from the generalized nymphalid groundplan. The differences arise primarily from the loss or positional shift of certain pattern elements, a high degree of fusion between individual pattern elements, and, in the forewing, asymmetries of the pattern elements relative to the wing-cell midline. The scheme of homologies we present is consistent with what is currently known about the comparative morphology and developmental physiology of colour pattern formation in Lepidoptera, and provides a framework for the interpretation of developmental, evolutionary and genetic studies in Heliconius.     

Immature stages of the Brazilian crescent butterfly Ortilia liriope (Cramer) (Lepidoptera: Nymphalidae)

We provide the first information on the morphology of the immature stages (egg, larva, and pupa), oviposition and larval behavior, and host plant, for the Brazilian crescent butterfly Ortilia liriope (Cramer), based on material from Santarém Municipality, Pará State, Northern Brazil. Females of O. liriope lay eggs in clusters. After hatching, larvae eat the exochorion and remain gregarious in all but the final instar. The host plant recorded in the study site is Justicia sp. (Acanthaceae). Despite the scarcity of data on the immature stages of Neotropical Melitaeini, we can already say that some morphological and behavioral traits observed in the immature stages of O. liriope are also present in all known genera in this tribe.