Comparative study of larval head structures of Megaloptera (Hexapoda)

Abstract

External and internal head structures of the larva of Neohermes are described in detail. The results are compared to conditions found in other representatives of Corydalidae, in Sialidae, and in Raphidioptera and Neuroptera. Corydalidae and Sialidae are mainly characterised by plesiomorphic features such as distinct frontal and coronal sutures, six stemmata, a movable labrum with a full set of muscles, a thin tentoriomandibular muscle, a distinct maxillary groove, 4-segmented maxillary palps, an oblique arrangement of the extrinsic maxillary muscles, a labium with all components except for the glossae and paraglossae, 3-segmented labial palps, and a largely complete muscle system. The partly reduced maxillary groove, the strongly elongated stipes, the apical membranous stipital collar, the close connection of the palp and galea, the subdivision of the galea, the strongly shortened palp, the bipartite tentoriocardinal and tentoriostipital muscles, the anterolateral submental notch, the lateral tentoriopharyngeal muscle, and the postgular plate are autapomorphies of Corydalidae. An additional antennomere is present in large corydalid species. The monophyly of the subfamilies Corydalinae and Chauliodinae is not supported by features of the larval head. The reduced condition of the anterior and dorsal tentorial arms and the antennal muscles, the transverse labial muscle, the loss of muscles of the salivary duct, and possibly the lateral origin of M. frontopharyngalis posterior are autapomorphies of Sialidae. The monophyly of Megaloptera is suggested by the insertion of a peg-like or spine-shaped sensillum on the antepenultimate antennomere, the vestigial salivary duct, and a verticopharyngeal muscle composed of several bundles. The distinct neck region, the parietal ridge, and the anterior position of the posterior tentorial grooves are features shared by Corydalidae and Raphidioptera. Arguments in favour of a clade comprising Megaloptera and Raphidioptera are the presence of a circular ridge anterad of the neck region, an increased number of Semper cells and retinula cells in the stemmata, the presence of a gula in adults, a similar cleaning behaviour, and molecular data. Potential autapomorphies of Neuropterida are the prognathism of the larvae and the absence of a mandibular mola. However, the polarity of these characters is unclear. A derived condition found in most groups of Endopterygota, but not in Hymenoptera, is the presence of one or two sensorial appendages on one of the intermediate antennomeres, usually the penultimate. Larval autapomorphies of Endopterygota suggested in earlier studies are confirmed for Corydalidae.