(Diptera, Culicidae) VIII. a Prodrome of the Genus Orthopodomyia
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Contributions of the American En tordogiccd Institute Volume 3, Number 2, 1968 MOSQUITO STUDIES (Diptera, Culicidae) VIII. A prodrome of the genus Orthopodomyia. By Thomas J. Zavortink MOSQUITO STUDIES (Diptera, Culicidae) VIII. A PRODROME OF THE GENUS ORTHOPODOMYIA ’ BY Thomas J. Zavortink2 CONTENTS INTRODUCTION .......................... 2 HISTORY ............................. 4 MORPHOLOGY. .......................... 6 BIOLOGYANDECOLOGY ...................... 8 DISEASE RELATIONSHIPS AND ECONOMIC IMPORTANCE ....... 9 DISTRIBUTION. .......................... 10 SYSTEMATICS ........................... 10 TAXONOMIC TREATMENT ..................... 13 Genus Orthopodomyia ...................... 14 Bancroftia Section ..................... 22 Signifera Group. .................... 24 Signifera Subgroup ................. 30 1. Orthopodomyia waverleyi ........... 31 2. Orthopodomyia signifera ’ ........... 3 5 3. Orthopodomyia alba .............. 47 Pulchripalpis Subgroup 4. Orthopodomyia pulchripalpis ......... 52 Kummi Subgroup .................. 58 5. Orthopodomyia species 5 ........... 58 6. Orthopodomyia kummi ............ 60 Albicosta Group 7.‘ Orthopodomyia albicosta ........... 66 Thomasina Section ..................... 71 8. Orthopodomyia fascipes. ........... 75 9. Orthopodomyia sampaioi ........... 82 1 Contribution from project “Mosquitoes of Middle America” supported by U. S. Public Health Service Research Grant AI-04379 and U. S. Army Medical Research and Development Command Research Contract DA-49-193-MD-2478. Based on PhD dissertation submitted to the University of California, Los Ange- les and supported in part by U. S. Public Health Service Training Grant TI-AI- 132 and National Science Foundation Grant GB 3871. 2 Department of Zoology, University of California, Los Angeles, Califor- nia 90024. Zavortink: Genus Orthopodomyia 3 relationships among the species. The purpose, then, of the present study is to revise this genus from the world standpoint on the basis of data from all known stages in the life cycle. To achieve this end, species have been delimited on the basis of morphological, ecological and distributional data and arranged into a classification which, it is hoped, is consistent with the evolution of the group. Descriptions and illustra- tions of all known stages of all species and descriptions of all higher taxa are provided, keys to these taxa are included, data on the distribution and bionom- its of each taxon are summarized, and speculations on the affinities of the var- ious taxa are made. Material for this study was first sorted to geographical origin and larval habitat and then tentatively identified with existing keys or descriptions. I have been fortunate in having either paratypic or topotypic material of a large num- ber of species available for examination. My general procedure for any par- ticular species was to first study this paratypic or topotypic material in consid- erable detail. Then, when they existed and specimens were available, species sympatric with the one being studied at or near its type locality were examined. This process, essentially a reversion to that of the local naturalist dealing with nondimensional species, was necessary to obtain an appreciation of the funda- mental morphological differences between undisputed, sympatric species. Only then were additional specimens, tentatively identified as being the species in question, compared with the paratypic or topotypic material and a decision made on the basis of morphological, ecological and geographical data as to whether or not these individuals came from conspecific populations. This en- tire process, when performed for each species, becomes somewhat self-cor- recting and leads to a general refinement of ones’ concepts of the species in- volved. After species were delimited and diagnosed, they were grouped into higher taxa of various levels on the basis of morphological similarity. Many of the groupings are tentative only; until all the stages in the life cycle are known for most species it will not be possible to construct a classification accurately re- flecting the various affinities within the group. I have examined specimens representing all previously described nominal species except geberti, mcgregori and nkolbissonensis. Material contained in the following institutions or private collections was studied: Bernice P. Bishop Museum; British Museum (Natural History); California Academy of Sciences; Canadian National Collection; Centre Scientifique et Technique, Bondy, France; Cornell University; Fred Harmston; Instituto Oswald0 Cruz; Instituto de Salu- bridad y Enfermedadas Tropicales, Mexico City, Mexico; Liverpool School of Tropical Medicine; Antonio Martinez; Museo Firenze; South East Asian Mos- quito Project, Washington, D. C .; United States National Museum; Universidad de Sao Paulo; University of California at Davis; University of California at Los Angeles; University of Utah. Six of these collections deserve special mention because they have been particularly vital to this study. The collections of the British Museum (Natu- ral History) and United States National Museum have been most important be- cause of their broad representation of species and accumulation of lectotypes and holot ype s. The excelent collection of William W. Macdonald has been in- strumentaf. in the treatment of the Oriental species because of its high quality and great number of adults with associated larval and pupal skins. Important also in the study of the Oriental species was the collection amassed by the South East Asian Mosquito Project; this is the largest collection from that area. The collection at UCLA has been invaluable, particularly in the study of the Neo- 4 Contrib. Amer. Ent. Inst., vol. 3, no. 2, 1968 tropical species. The bulk of this material has been collected for the project on the Mosquitoes of Middle America. This collection is outstanding in itsrep- resentation of topotypic material, large series of specimens and number of in- dividually reared adults. The last collection to be mentioned is my own. This collection was important not because of its representation of species but be- cause I acquired an appreciation of the extent of intra- and inter-population var- iation from it. I wish to thank Dr. John N. Belkin for directing this study. Special thanks are extended to Drs. P.F. Mattingly and J. A. Reid at the British Museum (Nat- ural History) and A. Stone and B. de Meillon at the United States National Mu- seum for the courtesies extended to me when I visited those institutions. I am also grateful to the individuals and institutions which loaned material. I am in- debted to Sylvia Barr, Sally Dieckmann and Joyce Roberts for preparation of the final plates and to Sheila Bernstein for typing the copy for reproduction. HISTORY The history of the classification of the genus Orthopodomyia is both short and simple. The first species of Orthopodomyia was described as Culex pul- chripalpis by Rondani in 1872 from material collected in Italy. In 1896 a sec- ond species, the North American signifera, was described in the genus Culex by Coquillett and in 1903 a third species, from India, was named Mansonia anopheloides by Giles. During the next year 2 species were described and gen- era based on them, Theobald (1904:236-237) creating the genus Orthopodomyia for Leicesters’ Malayan species albipes and Lutz (1904) naming Bancroftia al- bicosta from Brazil. Between 1905 and 1911, 9 additional species or varieties of Orthopodomyia were described in the genera Orthopodomyia, Culex, Man- sonia and Kerteszia and 3 additional generic names were published, Dyar (1905: 46) naming Pneumaculex for signifera, Theobald (1909: 297) proposing Newstead- ina for the Mauritian arboricollis, and Newstead and Carter (1911: 553-554) creating Thomasina for the South American longipalpis. The classification of the family Culicidae was in a chaotic state during the early twentieth century. Species and genera were being described at such a rapid rate by workers throughout the world that many taxa were inevitably giv- en more than one name. Additional confusion occurred because descriptions were often so inadequate that many misidentifications and incorrect usages of both specific and generic names took place. The greatest source of confusion, however, was that many workers, particularly Theobald, created and diagnosed numerous genera primarily on the basis of adult palpal length and scale mor- phology. As a consequence of all these factors, even closely related species were often not recognized as such and were placed in different genera. As an example, in Theobalds’ (1910) A Monograph of the Culicidae, which sum- marized the classification of the family, the 10 included nominal species of Orthopodomyia were placed in the following genera: Orthopodomyia, Bancroftia, Pneumaculex, Newsteadina, Mansonia, and Grabhamia. Even before the initial period of proliferation of generic names was over, a great reorganization and clarification of the classification of the Culicidae started taking place. Genera were based on more fundamental morphological features of adults in conjunction with characteristics of the larvae and male genitalia. Fewer, but more natural, genera resulted. In the case of Orthopodomyia, 4 of the 5 generic names proposed were re- duced to synonymy and those species incorrectly placed in other genera were Zavortink: Genus Orthopodomyia 5 properly realigned. The initial steps toward this clarification were taken by Dyar and Knab; first