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THE BLOOD PROTOZOA of NORTH AMERICAN BIRDS by CARLTONM

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THE BLOOD PROTOZOA of NORTH AMERICAN BIRDS by CARLTONM BIRD-BANDING A Jotm• oF ORNITHOLOGICALINVgSTIGATION VOL. XV JULY 1944 No. 3 THE BLOOD PROTOZOA OF NORTH AMERICAN BIRDS By CARLTONM. HERMAn" INTRODUCTION BLOODprotozoa were first reportedfrom arian hostsby Danilewsky in 1885 from the Russian Ukraine only five years after malaria para- sites were first describedfrom the blood of man. During the next decade several European workers, including Danilewsky, reported further on the occurrenceof organismsin the blood of birds but no investigationswere made in North America until the summerof 1897 when Opie(74)• and MacCMlum(58) studied blood smearsof birds from Maryland and Ottawa. Novy and MacNeM(72, 73) published in 1904 and 1905 on blood parasitesof Michigan birds. No further investigationson wild birds were made until Whitmore(97) isolateda strain of Plasmodiumfrom "sparrows" (probably Passer domesticus domesticus}in New York in 1913. This strain is still being carried in canariesand hasbeen subjected to numerousbiological studies during the intervening years. In the early 1920'sthe Department of Proto- zoologyand Medical Entomology of the Johns Hopkins University Schoolof Hygiene and Public Health beganits extensivestudies on the biologyof avianPlasmodiurn, particularly because of its similarity to the malarias of man. The emphasisof the work of this department has continuedto be on theseparasites, but in recent years related investigations have been undertaken on other blood parasites of birds. Many of the former studentsof this department,now connected with other institutions,have carried on or directedfurther investi- gations on the biology of malaria as well as surveys on the occurrence of parasites in wild birds. The first extensivesurvey in North America was made on ducks in Michiganby O'Roke(79}. From the introductionto his paperit is evident that he receivedmuch help from bird banders. The first extensivesurvey of a variety of wild birdswas conducted in the region aroundSyracuse, New York(33,68, 69). Most of the 652birds studied werebanded and released.During the summersof 1936and 1937the author had an opportunity to study birds trapped for banding at •Parasitologist,Bureau of GameConservation, California Division of Fish and Game. •Numerals in parenthesesrefer to the referencesat the end of the paper. 89 901 BloodProtozoa of North.4roerican Birds JulyBird-Banding the O. L. Austin OrnithologicalResearch Station, Cape Cod, Massa- chusetts,and has publisheda seriesof paperson the blood protozoa of these birds(34,35, 36, 37, 38, 39, 40, 41, 42). Huff was probably the first parasitologistto seethe advantageof cooperationwith bird bandem. The resultsof about 10 yearsof suchcooperative endeavors werepublished in 1939(53). Wood and Herman(103)have just com- pleted an extensivestudy of bloodsmears from a largeseries of birds made possiblethrough the cooperationof bird banders in California and Arizona. The cooperationbetween parasitologists and bird bandersthroughout the country should,in the ensuingyears, add greatly to our knowledgeof bloodparasites of birds. The first check-listwas publishedby Ltihe(57)in 1906. In 1926, Wenyon(94) publisheda completecheck-list of all the protozoan blood parasitesreported from birds in the world up to that time. Coatney(8, 9) and Coathey and Roudabush(14) in 1936 and 1937 publishedcheck-lists of Plasmodium,Leucocytozoon and Haemoproteus. Calkins(3)in 1933,published a list of the bird hosts of Trypanosoma. Lucena(55) in 1939 and Hewitt(47) in 1940 publishedreview mono- graphs on bird malaria and included host lists of arian Plasmodiurn. Lucena(56) in 1941 published a check-list of all protozoan blood parasites of birds in neotropicalAmerica. The present paper is intendedto supplementthis last 'check-listof Lueenaand thus bring up to date the host distribution of blood protozoa in birds of the western hemisphere. PARASITE LIST The classification of the blood protozoa of birds is in need of clarification. Many important phaseshave not been sol;•edand much of the publishedinformation is controversial. All trypanosomesreported from birdsbelong to the genusTrypano- soma.These fiagellated organismsare extracellular and in birds have been recordedonly in the bloodstream. In generalmorphology they are similar to the trypanosomeswhich occur in other vertebrates. These organisms,as they occur in birds, are known to be extremely pleomorphic and therefore are difficult to differentiate on the basis of morphology alone. Most trypanosomesfrom arian hostshave been classifiedas Trypanosomaavium. Many authors, however, have preferred to report arian trypanosomessimply as Trypanosomasp. Other specieshave been described,and even varieties have been set up within species,but the evidencefor their validity is very slim. There has been a scarcity of investigationson the possiblepathogenicity of this genus in birds and most protozoologistsconsider them of no pathological significance. In man these parasites and those of the related genus Leishmania cause severe tropical diseases. They are transmitted by various blood-suckinginsects. In birds the only Vol. XV 1944 BloodProtozoa ofNorth A roerican Birds [91 evidence available incriminates mosquitoesas the intermediate host but the completelife cycle has not been worked out. All other protozoanblood parasiteswhich have been reported from avian hosts apparently belong to the Sporozoa. The chief group is classifiedin this paper as belongingto the family Plasmodiidae,which includes the genera Plasmodium,Haemoproteus and Leucocytozoon. In the early stagesrecognizable in blood smearsit is not possibleto differentiatebetween these genera, and severalworkers have reported such findings simply as Plasmodiidae. Most protozoologistshave consideredthese forms as belongingto two families: Plasmodiidae, including the genus Plasmodium, and Haemoproteidae, including Haemoproteusand Leucocytozoon. Current investigations on the exoerythrocytic stages of Plasmodium are presenting much infor- mation which may settle this controversy. Recently Huff(54) has pointed out that Leucocytozoon,in somerespects, may be more closely related to Plasmodiumthan to Haemoproteus. The genusPlasmodium is observedmost readily within red blood cells. The nature of its developmentwithin the vertebrate host makes possiblethe transmissionof the infection to susceptible birds by blood transfusion. Recent work has demonstratedexoerythrocytic stagesas well as the more commonintracellular forms. More than a dozenvalid speciesare recognized. Manwell(65) has reviewed the methods of classificationof speciesin this genus. Evaluation of speciesand differentiating keys are available in several American publications(47,55, 63, 67). All speciesinvestigated are transmitted by culicine mosquitoes. Very little is known of the pathogenicity of theseparasites in wild birds, but canariesreadily succumbto experi- mental infections in the laboratory and have been known to contract a fatal infection spontaneously(44). The genusHaemoproteus occurs in birds and reptiles. In one stage of its developmentit parasitizesred blood cells but in other stages occursonly in internal organs. It is not possibleto transmit parasites of either this genus or Leucocytozoor•by blood transfusion. Most investigators have assumedthat these parasites are extremely host specificand thus many specieshave been named with hardly more of a basis than the fact of being observedin a previously unreported host. Some investigators have attempted to avoid this discrepancy simply by reporting their findingsas Haemoproteussp. The inability to transmit these parasites by simple blood transfusion makes it difficultto investigatethe host-specificityand thus clarify the validity of namedspecies. Only two specieshave beensuccessfully transmitted, the definitive host in both caseswas hippoboscidblood-sucking flies. Information on pathogenicity is in about the same state as our knowledgeof avian Plasmodium. Bird-Banding 92] H•:RM^N,Blood Protozoa of NorthAmerican Birds ,luly The genus Leucocytozoonoccurs only in birds. Students of this parasite are not agreed on the type of host cellsinfected. The host cellscontaining adult parasitesare so distorted from the infection as to make their identification impossible. For a review and recent information on this controversy the reader is referred to a paper by Huff(54). The validity of reportedspecies is in much the samestatus as speciesof the genusHaernoproteus. Only two specieshave been successfullytransmitted, the definitive host in both caseswas simuliid blood-suckingflies. O'Roke(79) hasreported high mortalities among ducks due to infections with Leucocytozoonsimondi. The other Sporozoa which occur in birds have been reported comparativelyinfrequently from North America. Theseinclude the family Haemogregarinidae(hemogregarines) and other parasitesthe family positionof which is not agreedupon by workersin this field: the genera Toxoplasma,Hepatozoon and SFirogregarina,intra-leuco- cytic parasitesand an unclassifiedparasite of monocytecells. Hemogregarinesare sporozoanparasites primarily infecting red blood cells of reptiles and amphibians. Extra-cellular stages also occur. A few casesof infection with this parasite in birds have been reported. The pathogenicity is unknown. Toxoplasmahas beenreported from mammals,including man, and from reptilesas well as birds. The life cycle is not known and i,t is questionablewhether all organismsclassified in this genusin birds are of this genusor new genera.Wenyon(94) has suggestedthat Novy and MacNeal's Haemoproteusrouxii belongsin
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