In this review, we use the wing veins of dipteran insects as potential models for understanding the evolution of development. We briefly discuss previous work in this field and examine the genetic complexity of wing formation, discussing the genes involved in wing formation and their roles in Drosophila wing development and vein formation. Furthermore, patterns of wing vein formation, addition, and reduction are discussed as they occur throughout the Diptera. Using the phyletic phenocopy paradigm, we draw attention to many wing vein morphologies that phenocopy various wing mutants in Drosophila melanogaster. The systematic issues of the nature of characters, homology, and the role of modern developmental approaches to evolutionary studies, which has recently become important, can be addressed from the perspective of the wing. We argue that further developmental evolutionary studies, and the interpretation of data therefrom, must be conducted within the context of a well-supported phylogeny of the organisms under study.
The Hawaiian Drosophilidae possess spectacular diversity in male foreleg modifications, many of which are unknown in other Diptera. The greatest diversity in foreleg morphology is in the antopocerus, modified tarsus, and ciliated tarsus clade (AMC Clade), a group of 95 species. The modified tarsus flies are divided into the bristle, ciliated, split, and spoon tarsus subgroups. The bristle tarsus species feature one or two rows of thickened setae on the basitarsus. The split tarsus species are characterized by only having four tarsal segments, in contrast to five tarsomeres in the remainder of Diptera. Based on comparisons of the apparent ground state of ventral setal rows across the Hawaiian Drosophila, we suggest that it is the second tarsal segment which has been lost. The spoon tarsus species are characterized by having the second tarsomere modified into a setae-filled, concave-shaped spoon. The ciliated tarsus species, all of which possess one or more elongate setae on the tarsus of males, are probably not monophyletic with respect to the bristle tarsus subgroup. The antopocerus flies are characterized by a long basitarsus, with extensive setation on the tibia and basitarsus of some species. The use of these foreleg modifications in courtship behavior has been previously described and it is suggested that they represent the results of sexual selection. The current work expands on previous morphological analyses, presenting a level of detail not previously possible without SEM images. The new characters revealed will figure prominently in future cladistic studies.
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