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University Microfilms, a Xeroxcompany, Ann Arbor 71.,..12,222 GOJRATI, Hassan Ali Navvab, 1935­ EPIZOOTIOLOGICAL SURVEY Of AVIAN I:iALARIA IN THE HAWAIIAN ISLANDS. ' University of Hawaii, Ph.D., 1970 Entomol,ogy University Microfilms, A XEROX Company, Ann Arbor, Michigan THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED EPIZOOTIOLOGICAL SURVEY OF AVIAN MALARIA IN THE HAWAIIAN ISLANDS A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ENTOMOLOGY SEPTEMBER 1970 Hassan Ali Navvab Gojrati Dissertation Committee D. Elmo Hardy, Chairman Andrew J. Berger Mercedes D. Delfinado Wallace C. Mitchell Minoru Tamashiro ACh~OWLEDGMENT8 I would like to express my sincere appreciation to many individuals and organizations for their assistance and cooperation in various aspects of this study. Mr Jack Throp, Director of the Honolulu Zoo for allowing complete freedom in trapping and examination of birds at the zoo. Dr. Joseph E. Alicata of the University of Hawaii for examination of some of the smears; Dr. Allen Y. Miyahara of the University of Hawaii for very essential technical assistance; Miss Elaine M. L. Chang, technician of the Board of Agriculture; Mr. James K. Ikeda of the State Department of Health for prOViding information on mosquito rearing; Mr. Winston Banko of Hawaii National Park for great assistance in mist-netting the birds at higher elevations; Mr. John L. Sincock of the U. S. Fish and Wildlife Service, Koloa, Kauai, for providing large numbers of slides from birds trapped at high-elevations on Kauai in addition to slides from Nihoa Finch, Nihoa Millerbird, Laysan Finch, Laysan Duck, etc. The Pathology section, Queen1s Medical Center for generously permitting me to use some of their laboratory facilities; Dr. Robert S. Gesowitz of the University of Hawaii for examination of slides and technical assistance; and Dr. Marshall Laird of the University of New Foundland, Canada, for indispensable assistance in diagnosis and identification of species of Plasmodium. TABLE OF CONTENTS Page ACKNOWLEDGMENTS •.•.••••••••.•••.•.••••••.•.•••••• ii ABSTRACT •.••••••.•••.•••••.••.••.•.••••••••...••• v LIST OF TABLES ....•••.....•.•.•......•......•.•.. vii INTRODUCTION ....••.......••..•.......•..•........ 1 HISTORICAL BACKGROUND .....••..••................. 5 TAXONOMIC POSITION OF THE AVIAN MALARIA PARASITES •.....•...•..•••...•......•.•....•...• 9 Class Sporozoa Leuckart 1879 ...•....•.•..... 9 Order Haemosporida Danilewsky 1886 •...•..... 10 Genus Plasmodium Marchiafava and Celli 1885 . 10 DEVELOPMENTAL CYCLE OF AVIAN PLASMODIA . 12 MATERIAL AND METHODS ...•.•.•.•••.•..•.•.•...•.•.. 16 Trapping and Handling the Birds •..• ......... 16 Collecting and Handling the Blood .....•••.•. 18 Collecting and Handling the Mosquitoes •..... 20 Description of the Collecting Area . 20 Rearing and Feeding the Mosquitoes . 21 Transmission Experiment.....••......•... 22 Technical Obstacles...•.....•.....•.•... 24 HISTOLOGICAL METHODS .....•....•..•.....•.•....... 25 Preparation of Slides for Making Blood Smears . 26 Making Blood Smears .......•.....••.....•.... 26 Staining Procedure....•.••.......••......... 27 Buffered \\later....•••..••..•......•......... 27 Technique for Dissecting the Salivary Glands of Mosquitoes .....•.••.....•....... 29 Technique for Preparation of Smear of the Salivary Glands for Sporozoites Study.•...............•...........•....•.. 30 Technique for Dissecting the Mid-gut of the Mosquitoes for Oocysts Study . 31 Technique for Preparation of Smear of the Mid-gut of Mosquitoes for Ookinetes Study.........•...•.....•................. 33 iv Technique for Sectioning of Liver for Exoerythrocytic Study.•...•.•....•.•.•.•.•..•..... 34 RESULTS AND DISCUSSION ••.•••••••.•••.••..••••••.•••••...• 38 Information on Mosquitoes .••.•••.••••••••..•.••.••.• 48 Haemoproteus Infections in Birds . 51 Leucocytozoon Infections in Birds . 53 CONCLUSIONS ..•.•••..•••••.•.••.•••.•.•••.•••.••...•••.••. 55 SUMtv1A.RY •••••••••••••••.••••••••••••..•.•••••••••••••.•••• 58 LITERATURE CITED .....•.•.....••.••.•.•.••••••.••••••.•.•• 60 ABSTRACT Avian malaria is believed to be one of the main factors in the extinction of the native Hawaiian avifauna. An epizoological investigation of the disease was conducted in the Hawaiian Islands primarily to determine whether avian malaria was present in Hawaii. The birds were collected from the islands of Oahu, Maui, Kauai, and Hawaii, at different elevations for a period of three years. Blood samples were taken either from the main wing vein, or from the toe nail. Blood smears were prepared and stained with Giemsa1s stain. Histological preparations for the study of the exoerythrocytic stage of the parasites were also made. A total of 4,988 blood smears from 2,604 birds representing 38 species of native and introduced birds was examined. Plasmodium circumflexum Kikuth, f. gallinaceum Brumpt, f. cathemerium Hartman, and probably f. matutinum (Huff), were found in six species of native and introduced birds. Significantly, these records are the first authenticated reports of these parasites from the Pacific Islands, and this is the first time that f. circumflexum has been reported from this part of the world. The presence of f. gallinaceum indicates that this parasite has accompanied its host to many parts of the tropics. Blood smears taken from pigeons and doves indicated that some of these birds were infected with Haemoproteus and Leucocytozoon. vi It was found that 65 percent of the pigeons were infected with Haemoproteus. The doves, however, did not seem to be infected by this pathogen. Although both pigeons and doves were infected by the Leucocytozoon, the incidence was relatively low. Only four percent of the two species of birds were infected. This is the first record for Leucocytozoon from the Hawaiian Islands. The presence of the larvae of the potential vector of avian malaria, Culex pipiens qUinguefasciatus Say, in a water container at 6511 feet elevation indicates that probably one of the limiting factors in the vertical distribution of Culex is the availability of suitable habitats. LIST OF TABLES TABLE PAGE I BUFFERED WATER FOR USE WITH GIEMSA r S STAIN........................... 27 II OCCURRENCE OF PLASMODIUM IN NATIVE AND INTRODUCED BIRDS IN HAWAII . 39 III PREVALENCE OF PLASMODIUM IN NATIVE AND INTRODUCED BIRDS IN HAWAII . 42 IV DISTIRBUTION OF INFECTION RATES BY ISLAND AND ELEVATION ••••••••••••••••••.•• 44 INTRODUCTION Research on avian malaria has offered to microbiologists, physiologists, biochemists, and many other scientists tools for solving many of their problems and this K~ll continue to be true regardless of whether human malaria is eradicated. Moreover, the growing feeling that eradication of malaria is just around the corner has given some people the idea that malarial research is no longer necessary. Because malaria has not as yet been eradicated, our tools for controlling it are rusting away leaving us unprepared for combatting an unexpected return of this great scourge. In this respect research on avian malaria provides us with a means for solving problems of basic importance to many fields of biology. The discovery that birds are subject to malaria by Danielewsky in 1885 followed closely after the finding of malarial parasites in the blood of man by Laveran (1880). Since then, the course of experimental observation with both organisms has proceeded in a parallel course. Unfortunately, the study of bird malaria, and its possible important role in the extinction of the rich Hawaiian avifauna, has been almost completely neglected. It is believed that, the parasites of aaimals and their vectors have gained entrance with the importation of infected animals from various parts of the world. Because of the mild climate and other favorable factors, these parasites have become established. 2 It is characteristic of the parasitic diseases that they do not cause immediate high mortality, and more often their pathological symptoms develop gradually. This has been the case in the Hawaiian Islands. It seems that parasites did not attract the attention of the Hawaiians until about 1800 when the American and European ships began to call frequently. Since that time, parasites have been found in many species of birds and mammals (Alicata, 1969). According to Hardy (1960), the night-biting mosquito (Culex pipiens quinquefasciatus Say) apparently was imported into the state of Hawaii in water casks on a ship (or ships) from Mexico between 1826 and 1830. There are a number of accounts as to how they first arrived here. Van Dine (1904) says, ITprevious to the year 1826 mosquitoes were unknown in Hawaii. During that year they were brought to the port of Lahaina, on the Island of Maui, in the ship TWp.llington T from San BIas, Mexico. 1T Furthermore, up to the year 1826 there was no word in the Hawaiian language for mosquito. The native term is TMakika T, a corruption of the word mosquito (Hardy 1960). On the basis of available evidence, Hardy (1960) also states that the two species of day-biting mosquitoes, Aedes aegypti (Linnaeus) and ~. albopictus (Skuse), apparently did not reach the islands until a much later date. Avian diseases aid greatly in controlling numbers of birds, but we know very little about these in Hawaii. With the exceptions of reports by Alicata (1939) on Haemoproteus columbae Kruse and of Fisher and Baldwin (1947) on Plasmodium vaughani Navy and Mac Neal, 3 practically no information is available. Warner (1968) reported on
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