Phylogeny of the Limnophilinae (Limoniidae) and early evolution of the Tipulomorpha (Diptera)
Guilherme Cunha RibeiroDepto. Biologia, Faculdade de Filosofia, Ciências e Letras de Ribeirão Preto, Universidade de São Paulo, Avenida Bandeirantes 3900, 14040–901. Ribeirão Preto, SP, Brazil. Email: ribeirogc@hotmail.com
Invertebrate Systematics 22(6) 627-694 https://doi.org/10.1071/IS08017
Submitted: 11 April 2008 Accepted: 14 November 2008 Published: 22 December 2008
Abstract
Tipulomorpha (crane flies) comprise one of the largest subgroups of Diptera, but its phylogeny at different levels has been poorly explored. This study presents the most comprehensive cladistic analysis of the group ever made, with emphasis on the genera and subgenera of the subfamily Limnophilinae (Limoniidae), assumed to include some of the earliest lineages of Tipulomorpha sensu stricto and therefore important for the understanding of the early patterns in the evolution of the crane flies. Eighty-eight characters of the male imago were scored for 104 exemplar species. The most parsimonious trees were searched using implied weighting, in the framework of a sensitivity analysis with different values of k (2 to 6). The dataset based on the characters of adult male morphology showed high levels of homoplasy and yielded very incongruent and unstable phylogenetic results, which are very sensitive to changes in analytical parameters. In the preferred and most parsimonious phylogenetic hypothesis, the Pediciidae is the sister-group of all other Tipulomorpha sensu stricto. The results indicate the paraphyly of the Limoniidae with respect to the Cylindrotomidae and Tipulidae, which are considered sister-groups. The Limoniidae subfamilies Limnophilinae, Limoniinae and Chioneinae are considered non-monophyletic. The study allowed a reconstruction of the possible ground plan condition of selected features of the adult male morphology of crane flies. The genera/subgenera Epiphragma (Epiphragma), Acantholimnophila, Shannonomyia, Limnophila (Arctolimnophila), Eloeophila, Conosia, Polymera, Polymera (Polymerodes), Prionolabis, Eutonia, Phylidorea (Phylidorea), Metalimnophila, Gynoplistia (Cerozodia), Gynoplistia (Dirhipis), Nothophila, Pseudolimnophila (Pseudolimnophila), Pilaria and Ulomorpha are considered monophyletic, but in general are defined by combinations of very homoplastic character states. Two Temperate Gondwanan clades, (Tonnoirella + (Edwardsomyia + (Tinemyia + (Rhamphophila + (Nothophila))))) and ((Notholimnophila + Bergrothomyia) + (Mesolimnophila + (Chilelimnophila + Ctenolimnophila))) are recovered. The genera Limnophila, Neolimnomyia, Gynoplistia (sensu lato) and Hexatoma (sensu lato) are considered non-monophyletic. The systematic position and some morphological characters of ‘problematic’ taxa, such as Dactylolabis, Elephantomyia, Helius and Atarba are discussed on the light of the proposed phylogeny and the analysis of the characters. Character states are richly illustrated. A detailed study of the morphology of the male genitalia is made, and several genera and species have the morphology of the male genitalia illustrated for the first time.
Additional keywords: comparative morphology, Cylindrotomidae, implied weighting, Limnophilinae, Limoniidae, Pediciidae, Tipulidae.
Acknowledgements
During the years spent in conducting this research, many different people were important, in different phases, to make it possible. This paper is a synthesis of my PhD thesis, made under the advisement of Dr Dalton de Souza Amorim, to whom I am especially indebted. Special thanks to Dr Wayne Mathis for the great support, and for hosting me at the Department of Entomology of the National Museum of Natural History, Smithsonian Institution, Washington D.C. in 2004. Many thanks to Dr Christian Thompson, for his unconditional support to my work, and for the donation of several reprints and books. I thank Dr Pjotr Oosterbroek for the exchange of ideas and literature, loan of specimens, and for allowing access to his world catalogue of crane flies years before its publication. Many thanks to Dr Jaroslav Starý and Dr Jon Gelhaus, the former for the donation of specimens, and both for sharing their critical views and opinions that greatly improved the final version of this work. I thank my colleague Matthew J. Petersen, also for our exchange of ideas and for the donation of some specimens. I thank Peterson Lásaro Lopes, for his help with the calculation of the Consistency and Retention indexes in TNT. Jon Gelhaus and another anonymous reviewer are very much thanked for the detailed reading of an early version of this text, and for raising deep and significant critiques that greatly improved the quality of its final version. This research was financially supported by a PhD and Post Doc fellowships from FAPESP. My studies at the National Museum of Natural History were funded with a fellowship by CAPES.
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