Larvae are a peculiar developtmental stage of holometabolous insects, occupy a considerable part of the insect life history, and contribute significantly to the prosperity of
Holometabola. The insect larvae are important to the study of insect taxonomy, ecology, and evolutionary biology, and cause direct injury to crops and other valuable materials of humans. However, since larvae are more difficult to obtain compared with adults, the most reliable way to get larvae is to rear the identified adults. For these reasons, the knowledge of insect larvae is far from satisfactory, especially for some small groups like Mecoptera.
Mecoptera is one of the most primitive lineages of Holometabola, remarkable for bearing
a pair of compound eyes on the larval head capsule, and is considered important to the study of the evolutionary origion of holometabolous insects. The Mecoptera contains more than 600 extant speceis across the world, assigning to nine families. The larvae of these families exhibit dramatically morphological and biological diversity, including the aquatic campodeiform larvae of Nannochoristidae, the herbivorous scarabaeiform larvae of Boreidae, and the edaphic eruciform larvae of Bittacidae and Panorpidae. These larvae are even more diverse morphologically and biologically on familial or generic levels, and may provide valuable characters for phylogenetic analysis.
In this study, the larvae of 33 species in Panorpidae, two species in Bittacidae, and one species in Panorpodidae were obtained through rearing in the past six years. The larvae were examined and compared morphologically using light and scanning electron microscopy. The morphology of larval mouthparts in related to their feeding behaviour, and the morphological diversity associated with habitat divergence were discussed. The phylogenetic analysis of Panorpidae was conducted on the basis of these larval morphological characters.
The larval feeding habits vary among Bittacidae, Panorpidae, and Panorpodidae, and the
larval mouthparts are also diverse morphologically among these families. In Panorpidae the larvae are saprophagous, consuming both the soft tissues and chitinous cuticle of dead insects. The molar regions of panorpid larvae bear numerous tuberculate teeth for grinding solid food. In Bittacidae the larvae are also saprophagous, feeding especially on the inner semi-fluid tissues of dead insects, and discard the empty exoskeleton on the ground. The molar regions of bittacid larvae are furnished with long spines for preventing larger particles from entering the pharynx. In Panorpodidae, however, the larval feeding habits are unknown. The molar regions of panorpodid larvae are glabrous, likely unsuitable for ginding solid food. It is more likely that these edaphic larvae of Panorpodidae feed on tissue of juice of the tender roots of
some specific host plants.
The larvae of Panorpidae are usually edaphic, living in/on the soil. However, based on
our current investigations, the specific living habits of larvae are diverse significantly on the generic level. The larvae of Dicerapanorpa are epedaphic, living predominantly on the soil surface. The larvae of Cerapanorpa are semi-epedaphic, staying either on the soil surface or beneath the soil horizon. The larvae of Panorpa are nocturnally active, staying in the soil during the day and crawling on the soil surface in the night. The larvae of Neopanorpa are euedaphic, living concealedly in the soil. The larval morphological features in these genera are also diverse associated with their living habits. In order to increase their survival opportunities, the larvae of Dicerapanorpa and Cerapanorpa all have darkish dorsal integuments, simulating the coloration of the soil as camouflage. The larvae of Panorpa are even nocturnally active to decrease the predation risk from the visually hunting predators. The larvae of Dicerapanorpa, Cerapanorpa, and Panorpa, all possess well-developed visual organs (prominent compound eyes) and locomotive organs (thoracic legs and prolegs). However, the euedaphic larvae of Neopanorpa adopt no camouflage strategy, but have additional shallow furrows on their head capsules to enhance the mechanical strength during locomotion beneath the soil horizon.
The phylogenetic analysis of Panorpidae was conducted on the basis of the larval
morphological characters for the first time. The phylogeny of 33 panorpid species was
reconstructed with maximum parsimony based on 52 larval morphological chararcers,
including 27 numeric characters and 25 continuous characters, employing Panorpodes
kuandianensis (Panorpodidae) and Bittacidae as the outgroups. The result shows that the
larval morphological characters support the current taxonomical system of Panropdiae. Most of the panorpid genera, including Neopanorpa, Dicerapanorpa, Sinopanorpa, and
Cerapanorpa, were reconfirmed to be monophyly. Panorpa, however, is supposed to be a
paraphyletic group and may need further split. The topology of the phylogenetic tree shows that the Panorpidae are basally splited into two clades, Neopanorpa is the sister group of all the other species. The monotypic genus Furcatopanorpa is the basal lineage of the latter. The genus Cerapanorpa is the sister group of the monophy comprising Dicerapanorpa and Sinopanorpa.