Asymmetric larval head and mandibles of Hydrophilus acuminatus (Insecta: Coleoptera, Hydrophilidae): Fine structure and embryonic development.

The larvae of a water scavenger beetle, Hydrophilus acuminatus, have strongly asymmetric mandibles; the right one is long and slender, whereas the left one is short and stout. The fine structure and embryonic development of the head capsule and mandibles of this species were examined using light and scanning electron microscopy, and asymmetries in shape were detected in these structures applying an elliptic Fourier analysis. The larval mandibles are asymmetric in the following aspects: whole length, the number, structure and arrangement of retinacula (inner teeth), and size and shape of both the molar and incisor regions. The larval head is also asymmetric; the left half of the head capsule is larger than the right, and the left adductor muscle of the mandible is much thicker than the right. The origin and developmental process of asymmetric mandibles were traced in developing embryos whose developmental period is about 270 h and divided into 10 stages. Mandibular asymmetries are produced by the cumulative effects of six stepwise modifications that occur from about 36% of the total developmental time onward. The significance of these modifications was discussed with respect to the functional advantages of asymmetries and the phylogeny of members of the Hydrophilidae.

Embryonic development of Carabus insulicola (Insecta, Coleoptera, Carabidae) with special reference to external morphology and tangible evidence for the subcoxal theory.

The egg morphology and successive changes in the developing embryos of the carabid ground beetle Carabus insulicola (Carabidae) are described based on light and scanning electron microscopy observations. Newly laid eggs of this species are ellipsoid and measure approximately 6.1 × 2.9 mm, before increasing to 6.6 × 3.4 mm at hatching. The egg period is about 11 days at 23°C. The egg shell is characterized by a thin fragile chorion covering a hard serosal cuticle. The embryo forms on the ventral egg surface, where it develops for the duration of the egg period. During the process of thoracic leg formation, two subcoxal rings, subcoxae-1 and 2, are clearly discernible at the basalmost region of the leg rudiments, and these subcoxae participate in the formation of the larval pleura and sterna. The result thus provides tangible evidence for the subcoxal theory, that is, that thoracic pleura and sterna are derived from subcoxal regions. Despite the complete absence of abdominal appendages in the larvae of this species, two pairs of appendage-like swellings, the medial and lateral ones, temporarily arise in the first eight abdominal segments during the middle of embryonic development. The medial swellings are assumed to be serially homologous with the coxal part of the thoracic leg, and they later flatten out and participate in the formation of the larval pleura (hypopleurites). In the light of the serially homologous relationships among gnathal appendages, thoracic legs, and abdominal appendage-like swellings, we identified the subcoxal regions in both the gnathal and abdominal segments. Although, the lateral swellings soon degenerate and disappear, it is considered that the swellings originate in the abdominal subcoxae-2 and may be homologous to the tracheal gills of larvae of Gyrinidae. Based on the embryological results, new interpretations for the constituent of gnathal appendages are proposed.

Embryonic development of a whirligig beetle, Dineutus mellyi, with special reference to external morphology (insecta: Coleoptera, Gyrinidae).

The egg morphology and successive changes of developing embryos of the whirligig beetle, Dineutus mellyi (Adephaga: Gyrinidae) are described from observations based on light and scanning electron microscopy. The egg surface is characterized by minute conical projections covering the entire egg surface, a stalk-like micropylar projection at the anterior pole of the egg, and a longitudinal split line along which the chorion is cleaved during the middle embryonic stages. The germ band or embryo is formed on the ventral egg surface, and develops on the surface throughout the egg period; thus, the egg is a superficial type, as is the case in most coleopteran species. A pair of lateral tracheal gills (LTGs) of the first abdominal segment originates from appendage-like projections arising at the lateral side of pleuropodia, and the LTGs of the second to ninth abdominal segments are arranged in a row with that of the first segment. Therefore, LTGs are structures with serial homology. The paired dorsal tracheal gills (DTGs) of the ninth abdominal segment are formed on the regions just latero-dorsal to the LTGs of this segment. Regarding the pleuropodia as the structures being homologous with thoracic legs, neither the LTGs nor DTGs are homologous with thoracic legs, but originate in the more lateral region corresponding to the future pleura of the thoracic segments. The last (10th) abdominal segment in the larva is formed by the fusion of the embryonic 10th and 11th abdominal segments. Four terminal hooks at the end of the last abdominal segment originate from two pairs of swellings on the posterior end of the embryonic 11th abdominal segment. It is proposed that the terminal hooks possibly correspond to the claws of medially fused cerci of the embryonic 11th abdominal segment.

Embryogenesis of the glowworm Rhagophthalmus ohbai Wittmer (Insecta: Coleoptera, Rhagophthalmidae), with emphasis on the germ rudiment formation.

The early embryonic development and features of the developing embryo of the glowworm Rhagophthalmus ohbai are described chiefly by light microscopy, with emphasis on the germ rudiment formation and its phylogenetic implication. The egg period is 30-34 days at about 23 degrees C. The newly laid egg is a short ellipsoid, 1.09 by 0.78 mm in size, and the size increases to 1.15 by 0.95 mm by 17 days after oviposition. Cleavage is of the typical superficial type. The germ disk is formed by cell aggregation of the embryonic area at the anterior end of the egg. The central part of the germ disk then sinks into the yolk and the spherical germ rudiment is formed by fusion of the amnioserosal folds extended from all margins of the germ disk. The inner region of the germ rudiment soon becomes slender and develops into the short embryo, whereas the outer region facing the anterior end is extended to form the thin amnion. The embryo then rapidly elongates, the elongation being accompanied by embryo segmentation and formation of appendages. The submerged condition of the embryo persists until about 17 days after oviposition (about 1 day before embryonic revolution) and thereafter the embryo becomes superficial in position. The presence of the following embryonic characters in R. ohbai supports the molecular data placing it within the Lampyridae: 1) formation of a spherical germ rudiment near the anterior end of the egg, and 2) the submerged condition of the developing embryo persists until shortly before revolution.

Early embryonic development and external features of developing embryos of the caddisfly, Nemotaulius admorsus (trichoptera: Limnephilidae).

Formative processes of the blastoderm, germ disk, embryonic membranes, and germ band in the trichopteran Nemotaulius admorsus are described. Successive external changes of developing embryos are also described, chiefly on the basis of SEM data. The blastoderm is very thin, and a circular germ disk forms in its ventro-posterior region about 32 hr after oviposition. Embryonic membranes are completed by fusion of amnio-serosal folds extending from the margin of the germ disk. The germ disk then differentiates into a long germ band in situ. This process is similar to that of other trichopterans of the suborder Integripalpia (e.g., Neophylax concinnus and Neoseverinia crassicornis), but differs from that of Stenopsyche griseipennis (suborder Annulipalpia) in which the germ band and embryonic membranes are formed by deep invagination of the germ disk. The germ band or embryo, as in other trichopteran species, assumes a completely superficial position throughout development. The following description summarizes its external differentiation. An egg tooth develops at the top of the head capsule. Antennal rudiments become small as development advances. In the maxillary segment, the lacinial and galeal lobes are indistinguishable and united into one laciniogaleal complex. Thoracic appendages develop considerably during the embryonic period. Pleuropodia are formed in the first abdominal segment. Two pairs of appendage-like swellings, a ventral and a medial pair, arise in each of the first eight abdominal segments, and homologies of these swellings and pleuropodia are discussed. Terminal hooks originate from the bilateral projections of the telson. Tracheal invaginations appear in the mesothoracic, metathoracic, and the first eight abdominal segments, but their openings or spiracles close later. Embryonic membranes rupture after revolution, and thus the secondary dorsal organ is formed.

Mesodermal organogenesis in the embryo of the primitive moth, Neomicropteryx nipponensis Issiki (Lepidoptera, Micropterygidae).

The formation of somites, coelomic sacs, splanchnic mesoderm, fat bodies, circular system, gonads, and musculature in the embryo of the primitive moth, Neomicropteryx nipponensis Issiki, is described. The following paired somites are formed: the labral, antennal, intercalary, mandibular, maxillary, labial, three thoracic, and 11 abdominal. Small but distinct coelomic cavities appear in all these somites. Labral somites differentiate into the labral muscles, stomodaeal muscles, and dorsal dilator muscles of the pharynx. Antennal somites differentiate into the antennal muscles, aorta, and the ventral dilator muscles of the pharynx. Intercalary somites are short-lived, disintegrating to liberate many free cells into the yolk. The suboesophageal body is not formed. Mandibular somites differentiate into the mandibular flexor and extensor muscles. Maxillary and labial somites differentiate into the splanchnic mesoderm, fat bodies, and into muscles of the maxillolabial region. Three pairs of thoracic and ten pairs of abdominal somites split into the splanchric and somatic mesoderm. The 11th abdominal somites merge into the proctodaeal mesoderm, and differentiate into the musculature of the hindgut. The heart is formed by the fusion of the cardioblasts derived from the first thoracic to the tenth abdominal segment. The aorta arises from the antennal median mesoderm. Blood cells are derived from the median mesodermal cells of the maxillary to the tenth abdominal segment. Germ cells appear at the mediodorsal corner of each somite in the fifth abdominal segment. They become enclosed with a mesodermal sheath to form a pair of rudimentary gonads in this segment. Major muscles in the head, thorax, and abdomen of the fully grown embryo are described.

Embryonic development of the alimentary canal and ectodermal derivatives in the primitive moth, Neomicropteryx nipponensis issiki (Lepidoptera, Micropterygidae).

The formation of the alimentary canal, nervous system, and of other ectodermal derivatives in the embryo of the primitive moth, Neomicropteryx nipponensis Issiki, is described. The stomodaeum is formed from an invagination in the medioposterior portion of the protocephalon. The proctodaeum arises as an extension of the amnioproctodaeal cavity. The midgut epithelium orginates from anterior and posterior rudiments in blind ends of the stomodaeum and proctodaeum. The decondary dorsal organ is formed in developing midgut. The development of the brain is typical of insects. The ventral nerve cord originates in large part from neuroblasts arising in 3 gnathal, 3 thoracic, and 11 abdominal segments. Intrasegmental median cord cells probably differentiate into both ganglion cells and glial elements of the ventral nerve cord; intersegmental cells appear not to participate in the formation of the nervous system. The stomatogastric nervous system develops from three evaginations in the dorsal wall of the stomodaeum, and consists of the frontal, hypocerebral, and ventricular ganglia, the recurrent nerve, and corpora cardiaca. Five stemmata arise from the epidermis on each side of the head. Five pairs of ectodermal invaginations are formed in the cephalognathal region to produce the tentorium, mandibular apodemes, corpora allata, and silk glands. Prothoracic glands orginate in the prothorax. Mesothoracic spiracles shift anteriorly to the prothorax during development. Oenocytes arise in the first seven abdominal segments. Invaginated pleuropodia are formed in the first abdominal segment.

The early embryonic development of the primitive moth, Neomicropteryx nipponensis lssiki (lepidoptera, micropterygidae).

This paper describes the early embryonic development of Neomicropteryx nipponensis from oviposition to the formation of the inner layer. Newly laid eggs are covered with many hygroscopic, gelatinous masses. The chorion is composed of a porous, spongy exochorion of variable thickness and of a thin endochorion. The eggs have a very thin periplasm and contain large amounts of yolk made of proteid and fatty yolk droplets. The processes of maturation division, fertilization, and cleavage are similar to those commonly found in lepidopteran eggs. The blastoderm of Neomicropteryx is very thin in comparison with that of other lepidopteran eggs. The small circular germ disk is formed on the ventral egg surface. It then invaginates deeply into the yolk to form a sac-shaped germ rudiment. The formation of the germ rudiment and of the embryonic membranes resembles that of swift moths, Endoclita (suborder Monotrysia) and of the caddisfly, Stenopsyche (Trichoptera), but differs from that of ditrysian Lepidoptera. As in other lepidopteran insects, the formation of the inner layer begins after completion of the germ band, which has a bilobed protocephalon and a slender protocorm. Unlike the situation in most lepidopteran eggs, yolk segmentation does not occur in N. nipponensis. During formation of the germ band, hydropyle cells are formed in the dorso-posterior region of the serosa; these are here reported for the first time in the eggs of holometabolan insects.

The embryonic development of the primitive moth, Neomicropteryx nipponensis Issiki (lepidoptera, micropterygidae): Morphogenesis of the embryo by external observation.

The micropterygid moth Neomicropteryx nipponensis belongs to the most primitive suborder Zeugloptera of the Lepidoptera. During embryogenesis the small circular germ disk formed on the ventral egg surface invaginates deeply into the yolk. It finally separates from the egg periphery or rudimentary serosa, and becomes a sac-shaped germ rudiment. Its anterior part later develops into the germ band, while its posterior part is the future amnion. Just before revolution of the embryo, the embryo assumes a completely superficial position beneath the yolk. Neither amnion nor serosa rupture during revolution; by completion of dorsal closure they have been incorporated into the yolk to form the secondary dorsal organ. The formation of the germ rudiment and embryonic membranes in N. nipponensis resembles those of swift moths, Endoclyta (suborder Monotrysia) and of the caddisflies, Stenopsyche (Trichoptera), but differs from those of ditrysian Lepidoptera. The secondary dorsal organ has never been found in any other lepidopteran embryos; however, it is formed in N. nipponensis and in the Trichoptera. The results of the present study strongly support the general phylogenetic views that the Zeugloptera have a close affinity to the Trichoptera.