The Pupae of Culicomorpha—Morphology and a New Phylogenetic Tree
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Zootaxa 3396: 1–98 (2012) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Monograph ZOOTAXA Copyright © 2012 · Magnolia Press ISSN 1175-5334 (online edition) ZOOTAXA 3396 The Pupae of Culicomorpha—Morphology and a New Phylogenetic Tree Art Borkent Research Associate, Royal British Columbia Museum, American Museum of Natural History and Instituto Nacional de Biodiversidad, 691-8th Ave. SE, Salmon Arm, British Columbia, V1E 2C2, Canada. (e-mail: [email protected]) Magnolia Press Auckland, New Zealand Accepted by J.K. Moulton: 22 Mar. 2012; published: 23 Jul. 2012 Art Borkent The Pupae of Culicomorpha—Morphology and a New Phylogenetic Tree (Zootaxa 3396) 98 pp.; 30 cm. 23 Jul. 2012 ISBN 978-1-86977-957-3 (paperback) ISBN 978-1-86977-958-0 (Online edition) FIRST PUBLISHED IN 2012 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/ © 2012 Magnolia Press All rights reserved. No part of this publication may be reproduced, stored, transmitted or disseminated, in any form, or by any means, without prior written permission from the publisher, to whom all requests to reproduce copyright material should be directed in writing. This authorization does not extend to any other kind of copying, by any means, in any form, and for any purpose other than private research use. ISSN 1175-5326 (Print edition) ISSN 1175-5334 (Online edition) 2 · Zootaxa 3396 © 2012 Magnolia Press BORKENT Table of contents Abstract . 3 Introduction . 4 Materials And Methods . 5 Acknowledgments . 6 Glossary of the Structures of the Pupae of Culicomorpha . 6 Adult Emergence . 21 Pupal Key to the Families of Culicomorpha . 22 Description of Ptychopteridae and Families of Culicomorpha . 23 Ptychopteridae . 23 Culicomorpha . 24 Chironomidae . 24 Ceratopogonidae . 26 Thaumaleidae . 27 Simuliidae . 28 Dixidae . 30 Corethrellidae . 31 Chaoboridae . 32 Culicidae . 33 Phylogenetic Analysis . 34 Eggs . 35 Larvae . 36 Pupae . 39 Adults . 48 Misinterpreted or Questionable Synapomorphies . 52 Conflicting Synapomorphies . 55 Phylogenetic Conclusions . 55 Bionomic Divergence . 60 References . 63 ABSTRACT The pupae of each of the families of the Culicomorpha are described and, for the first time, their structures homologized. A glossary provides a standard set of terms to be applied to each structure, including a common chaetotaxy. A cladistic analysis incorporates information from each life stage, including a number of new features discovered from the pupal stage, to provide a new phylogenetic hypothesis, as well as indicating autapomorphies for each family. Analysis included states for one egg, 21 larval, 33 pupal, and 37 adult characters. The Chironomidae is the sister group of all remaining Culicomorpha, the Ceratopogonidae is the sister group of Thaumaleidae + Simuliidae and these three are newly recognized as members of the re-defined superfamily Simulioidea. The superfamily Culicoidea are the sister group of the Simulioidea and include, as previous work has already demonstrated, the Dixidae as the sister group of Corethrellidae + Chaoboridae + Culicidae. Corethrellidae is the sister group of Chaoboridae + Culicidae. The superfamily Chironomoidea now includes only Chironomidae. Analysis of the fossil record shows that the Chironomidae (and the Culicomorpha) originated in the Triassic and both Simulioidea and Culicoidea were present by 176 million years ago in the Jurassic. Phylogenetic patterns are used to interpret bionomic features such as differences in the nature of blood-feeding by adult females, daytime or nighttime feeding by adult females, and occurrence of immature stages in aquatic habitats. Chironomidae do not feed on blood as adults and have likely diversified by invading virtually all aquatic habitats as larvae. Its sister group is more than twice as diverse and feeding on vertebrate blood is strongly correlated with high diversification within the Simulioidea + Culicoidea (likely because a reliable source of protein was available to dispersing females since the Triassic from terrestrial vertebrates). Families with blood-feeding females have larger numbers of species than do those without this behaviour. Each family in the Simulioidea + Culicoidea have specialized larval habitats or specialized habits, largely in aquatic habitats where Chironomidae are either not, or are marginally present, suggesting a level of competitive exclusion by the Chironomidae. Key Words: Chironomidae, Ceratopogonidae, Thaumaleidae, Simuliidae, Dixidae, Corethrellidae, Chaoboridae, Culicidae, aquatic, phylogeny, pupal homologies, key, glossary, blood-feeding, diversification, egg, larva, pupa, adult. PUPAE OF CULICOMORPHA Zootaxa 3396 © 2012 Magnolia Press · 3 INTRODUCTION One of the joys but also one of the great challenges of being an entomologist is the remarkably vast remaining ignorance regarding our cherished organisms. Diptera is a case in point. Largely due to the remarkable contributions of the three volumes of the Manual of Nearctic Diptera (McAlpine et al. 1981; McAlpine et al. 1987; McAlpine & Wood 1989), we have a consistent set of morphological terms applied to the larvae and adults of the order, based on careful study of homologous structures. The pupae of Diptera, however, are another matter. They were not included in that massive effort (other than within some family treatments) and this stage has been largely ignored by subsequent major treatises (e.g. Papp & Darvas 1997, 1998, 2000a, 2000b; Brown et al. 2009). As such, there has never been a concerted effort to homologize the morphological details of pupae at the familial level; nor do we have a good key to the pupae of the families. In this paper, I have carved out a slice of that diversity and interpret the pupae of the Culicomorpha. At present, there is a great disparity in the vocabulary applied to the pupal structures of the included families. Researchers of the most diverse families or groups of Culicomorpha, namely the Simuliidae, Chironomidae, Ceratopogonidae and the superfamily Culicoidea have each developed independent sets of terms. The homologies and associated terms shared by Culicidae, Chaoboridae, Corethrellidae and Dixidae are primarily due to the historical inclusion of all four in the Culicidae and their subsequent treatment by Belkin and his associates (e.g. Belkin et al. 1970), who described the pupae of the latter three families as part of their primary studies of Culicidae. This study originated, in part, from my continuing quest to better understand the phylogenetic relationships between the genera of Ceratopogonidae. For the past number of years I have gathered and studied the immature stages of this family as a basis for a comprehensive analysis. An important barrier to the phylogenetic interpretation of pupal features was the question of outgroup comparisons with other families of Culicomorpha. Therefore I examined material of each of the other families of Culicomorpha in an attempt to interpret homologies. As a consequence, I provide a single set of terms for the pupae of Culicomorpha, based on my understanding of their homology and also provide a new key for their identification to the family level. As I studied the pupae of other culicomorph families, it rapidly became clear there were significant morphological differences between the families and this provided the impetus to study even a broader outgroup. This broader survey resulted in the discovery of new and important synapomorphies grouping various families within the infraorder that challenge earlier conclusions regarding relationships. When Wood & Borkent (1989) published their study of the phylogenetic relationships between the families of nematocerous Diptera they wrote that "The monophyly of the Culicomorpha, along with the relationships between its component families, is one of the least contentious issues in the phylogenetic interpretation of the nematocerous Diptera." It is true that the composition of the Culicoidea.