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.