Neurobiological constraints and fly systematics: how different types of neural characters can contribute to a higher level dipteran phylogeny

Much uncertainty still exists regarding higher level phylogenetic relationships in the insect order Diptera, and the need for independent analyses is apparent. In this paper, I present a parsimony analysis that is based on details of the nervous system of flies. Because neural characters have received little attention in modern phylogenetic analyses and the stability of neural traits has been debated, special emphasis is given to testing the robustness of the analysis itself and to evaluating how neurobiological constraints (such as levels of neural processing) influence the phylogenetic information content. The phylogenetic study is based on 14 species in three nematoceran and nine brachyceran families. All characters used in the analysis are based on anatomical details of the neural organization of the fly visual system. For the most part they relate to uniquely identifiable neurons, which are cells or cell types that can be confidently recognized as homologues among different species and thus compared. Parsimony analysis results in a phylogenetic hypothesis that favors specific previously suggested phylogenetic relationships and suggests alternatives regarding other placements. For example, several heterodactylan families (Bombyliidae, Asilidae, and Dolichopodidae) are supported in their placement as suggested by Sinclair et al. (1993), but Tipulidae and Syrphidae are placed differently. Tipulidae are placed at a derived rather than ancestral position within the Nematocera, and Syrphidae are placed within the Schizophora. The analysis suggests that neural characters generally maintain phylogenetic information well. However, by "forcing" neural characters onto conventional phylogenetic analyses it becomes apparent that not all neural centers maintain such information equally well. For example, neurons of the second-order visual neuropil, the medulla, contain stronger phylogenetic "signal" than do characters of the deeper visual center, the lobula plate. These differences may relate to different functional constraints in the two neuropils.