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Xxxiv. a Revision of the Albimanus Section of The 107773 107773 ^ ^ Contributions of the Amenccm Entomological Institute Volume 15, Number 7, 19:80 MOSQUITO STUDIES (Diptera, Culicidae) XXXIV. A revision of the AlbimanusSection of the subgenus Nyssorhynchusjof; Anopheles '/.'"" ;7. .'By .'. I..": ^... ; Michael E. Faran 11JUN Contributions of the American Entomological Institute Volume 15, Number 7, 1980 MOSQUITO STUDIES (Diptera, Culicidae) XXXIV. A revision of the Albimanus Section of the subgenus Nyssorhynchus of Anopheles By Michael E. Faran CONTENTS INTRODUCTION 1 MATERIAL AND METHODS 3 TAXONOMIC HISTORY 5 TAXONOMIC CHARACTERS 9 SYSTEMATICS 14 BIONOMICS 16 MEDICAL IMPORTANCE 17 TAXONOMIC TREATMENT 18 Albimanus Section 18 Keys to Groups and Species. 27 Albimanus Group 35 1. Anopheles (Nys.) albimanus 35 Oswaldoi Group 50 Oswaldoi Subgroup 52 Oswaldoi Complex 53 2. Anopheles {Nys.) oswaldoi 55 3. Anopheles (Nys.) galvaoi 64 4. Anopheles (Nys.) noroestensis 67 5. Anopheles (Nys.) aquasalis 75 6. Anopheles (Nys.) ininii 89 7. Anopheles (Nys.) anomalophyllus 95 8. Anopheles (Nys.) rangeli. 99 9. Anopheles (Nys.) trinkae 106 10. Anopheles (Nys.) nuneztovari 112 Strodei Complex 122 11. Anopheles (Nys.) strodei 123 12. Anopheles {Nys.) random 132 13. Anopheles (Nys.)benarrochi 136 Triannulatus Subgroup 141 14. Anopheles (Nys.) triannulatus HI REFERENCES 152 FIGURES 178 TABLE OF DISTRIBUTIONS 211 CONSPECTUS OF TAXONOMIC CHANGES 212 INDEX TO SCIENTIFIC NAMES 213 MOSQUITO STUDIES (Diptera, Culicidae) XXXIV. A REVISION OF THE ALBIMANUS SECTION OF THE SUBGENUS NYSSORffYNCHUS OF ANOPHELES1 By Michael E. Faran2 INTRODUCTION Zavortink (1973:4), when he was reviewing the subgenus Kerteszia of Anopheles, stated, ". the systematics of Nyssorhynchus are too poorly known and hopelessly confused." In 1942, L. E. Rozeboom (1942b) used only one couplet in a key to the adult females for 9 different species in the Albimanus Section. The reason for this, as the pioneering work of W. H. W. Komp (1942:25-26) and Rozeboom recognized, is that many of the characters that have been used to distinguish the adult females in the Albimanus Section are extremely variable and unreliable for species identifica- tion. On the basis of one character alone it is often impossible to identify with any confidence an adult female as belonging to a particular species in this section, which includes several important vectors of malaria. The purpose of this revision is (1) to describe and illustrate, in detail, all the currently known species in the Albimanus Section, (2) to reconcile some of the systematic problems that have resulted in this aura of confusion and (3) to develop effective keys that use more than one character, when possible, for all stages of the life cycle for the species in this section, I am subdividing the subgenus Nyssorhynchus into 2 sections, the Albimanus Sec- tion and the Argyritarsis Section. These sections are very closely allied and form a tight, well-defined unit. The entire subgenus is restricted to the Neotropics except for albimanus which extends into the Nearctic. The Albimanus Section is distin- guished from the Argyritarsis Section in the adults primarily by the basal dark band on hindtarsal segment 5, and in the male genitalia by the variously developed fused ventral claspette. Only on the basis of these 2 characters can these sections be readi- ly differentiated. The Myzorhynchella group has been excluded from consideration because of the paucity of material available for study and because of its uncertain taxonomic position. In the present revision 14 species are recognized in the Albimanus Section. The nominal species sanctielii is not included because of the lack of material. An. gorgasi This research was supported by the Medical Entomology Project, Smithsonian Institution, U. S. Army Medical Research and Development Command Research Contract DAMD-I7-74C-4086; the Mosquitoes of Middle America project. University of California, Los Angeles, supported by U. S. Army Medical Research and Development Command Research Contract DA-49-193-MD-2478 and U. S. Public Health Service Research Grant AI-04379; and U. S. Public Health Service Predoc- toial Trainee Grant 5T01 AI00070-14. Captain, Medical Service Corps, U. S. Army, Department of Entomology, Walter Reed Army Institute of Research, Washington, DC 20012. 2 Contrib. Amer. Ent. Inst., vol. 15, no. 7, 1980 is removed from synonymy with albimanus and relegated to status of a nomen du- bium as is evansi which was formerly considered to be the senior synonym of strodei. The synonymy of metcalfi is changed from oswaldoi to tliat of noroestensis. An. tri- annulatus is treated as a single, although variable, form, not as being composed of 2 separate subspecies, t. triannulatus and t. davisi (or t. bachmanni). An attempt has been made to assemble the species into distinct monophyletic groups on the basis of correlated characters in the adults and immatures. The charac- ter state, ancestral or derived, of the taxonomically important features has been de- termined whenever possible and evolutionary trends stated in formulating the phylo- genetic relationships within the section- I am dividing the section into 2 groups, the monotypic Albimanus Group and the Oswaldoi Group. An. albimanus is the least de- rived species in the section, possessing several ancestral features which it shares with some of the ancestral species in the Argyritarsis Section. An. albimanus is easily dif- ferentiated from the Oswaldoi Group by several correlated unique features in the adult, male genitalia and larva. I am separating the Oswaldoi Group into 2 subgroups, che monotypic Triannulatus Subgroup and the Oswaldoi Subgroup composed of 12 species, further separated into the Oswaldoi Complex and the Strode! Complex. The Oswaldoi Complex consists of 9 species. Within the complex, 2 separate phy- letic lines are discernible on the basis of the structure of the ventral claspette of the male genitalia. One line is composed of oswaldoi, galvaoi, noroestensis, aquasalis, in- inii and possibly the relict anomalophyllus, and the other by rangeli, trinkae and nu- neztovari. The Strode! Complex contains strodei, rondoni and benarrochi. An. strodei and random are very closely related, but their relationship to benarrochi is difficult to de- termine as benarrochi is the most derived species in the section. An. benarrochi has been placed in the Strodei Complex only because of the similarity of its male genita- Ha with those of strodei and rondoni. The larvae and pupae of the Albimanus Section occur primarily in ground water habitats. All the species breed in fresh water, except for aquasalis and often albima- nus which are often found in brackish water. The females feed predominantly on large mammals. The adults are active either crepuscularly or noctumally; triannula- tus is the only species reported to bite occasionally during the day. Two species, albimanus and aquasalis, are major vectors of malaria in Central and South America and the islands of the Caribbean. An. nuneztovari is the primary vec- tor of malaria in western Venezuela and northern Colombia and possibly a vector in Suriname. An. triannulatus, strodei and noresfensis have been implicated in the transmission of malaria, since they have been found naturally infected- An. rangeli has been suspected of transmitting malaria in Ecuador. Two viruses, Venezuelan Encephalitis virus and Tiacotalpan virus, have been isolated from aquasalis and albi- manus respectively. Nothing is known about the medical importance of the remain- ing species. I would like to express my deep gratitude to John N. Belkin for his constant sup- port and guidance. 1 thank the other members of my doctoral committee, A. Ralph Barr, George A. Bartholomew, Walter Ebeling and Peter Vaughn, for their encourage- ment and helpful advice. I am greatly indebted to Ronald A. Ward of Walter Reed Army Institute of Re- search and Oliver Flint of the Smithsonian Institution for the loan of material; to P. Cova Garcia and A. Gabaldon, J. F. Reinert and D. R. Roberts, J- B. Kitzmiller, R. F. Darsie, Jr., and R. S. Panday for valuable specimens from Venezuela, Brazil, Colom- bia, El Salvador and Suriname respectively. Also, I would like to thank the following Faran: Albimanus Section of Anopheles (Nyssorhynchus) 3 individuals and institutions that have so generously loaned specimens for this study: Paul H. Arnaud, California Academy of Sciences, San Francisco; Henry S. Dybas, Field Museum of Natural History, Chicago; Peter F. Mattingly and Graham B. White, the British Museum (Natural History), London; L. L. Pechuman, Comell University, Ithaca; Selwyn S. Roback, Academy of Natural Sciences, Philadelphia; Milan Trpis, Johns Hopkins University, Baltimore; and Pedro W. Wygodzinsky, American Museum of Natural History, New York. I am very grateful to my friends and colleagues George K. Bryce and Kenneth Lin- thicum for reading and criticizing the manuscript and to J. Hal Amell, 0. G. W. Ber- lin and Thomas J. Zavortink for suggestions, advice and assistance whenever sought. To Sandra Heinemann, William Powder, Thomas Gaffigan, Ellen Paige, Ruby Sims and Melvin Stave I wish to express my thanks for the preparation of material and miscellaneous assistance. During my field studies I have been assisted by many people. I would especially like to thank Nelson Papavero of the Museo de Zoologia, Sao Paulo, Oscar de Souza Lopes and his staff of Institute Adolfo Lutz, Sao Paulo, and Herbert C. Barnett and Ricardo Iglesias Rios of Institute de Microbiologia, Rio de Janeiro. I wish to extend my sincerest appreciation to Sharon Toalson, Trinka Faran and Cynthia Lomax for the difficult and frustrating Job of typing the preliminary drafts of the manuscript. I thank Sandra Heinemann for criticizing and editing the manu- script, as well as preparing the text copy for lithoprinting. Also, I thank Nobuko Ki- tamura, L. Margaret Kowalczyk, Young Sohn and Vichai Malikul for preparation of the preliminary and final drawings. MATERIAL AND METHODS MATERIALS. This taxonomic revision is based on a study of 14,792 specimens: 2816 males, 774 male genitalia, 6078 females, 2088 pupae and 3036 larvae, including 1682 individual rearings (755 larval, 741 pupal, 186 incomplete) and 30 progeny roarings.
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