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Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 75-26,561 COOPER, Clyde Lawrence, 1943- THE HELMINTH PARASITES OF AN INSULAR AVIAN PASSERINE COMMUNITY: RELATIONSHIP TO LANDSCAPE EPIDEMIOLOGY. The Ohio State University, Ph.D., 1975 Zoology

Xerox University MicrofilmsAnn , Arbor, Michigan 48106 THE HELMINTH PARASITES OF AN INSULAR AVIAN PASSERINE COMMUNITY: RELATIONSHIP TO LANDSCAPE EPIDEMIOLOGY

DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University

By Clyde Lawrence Cooper, B.S,, A.M,

The Ohio State University 1975

Reading Committee: Approved by John L. Crites Tony J . Peterle Loren S. Putnam ACKNOWLEDGMENTS

A large number of individuals have contributed to my development as a biologist and scholar. Most importantly, Professor John L. Crites has given generously of his spiritual and material wealth to provide an atmosphere in which one joyfully acquires a working knowledge of the discipline of parasitology. Human beings spend their lives adopting various models of behavior. A part of the man that Dr, Crites is can be fqund as a part of this writer. This study would not have been possible without the aid of Dr, L, S. Putnam, Director, Franz Theodore Stone Labora­ tory. The assistance of Dr, M. Miskimen was, likewise, invaluable. The ready assistance offered to a parasitology student working on birds by these two ornithologists is, perhaps, the most telling statement that can be made about these kind people. Over the years, a number of graduate students in parasitology and ornithology have aided me in this study in various ways. Since they are too numerous to acknowledge individually, I wish to recognize their contributions collectively. VITA September 2, 1943. . • • .Born - Williamstown, West Virginia 1965 ...... B.S., Marietta College, Marietta, Ohio 1966 ...... A.M., West Virginia University, Morgantown, West Virginia 1965-1968...... Elementary and Secondary Science Teacher, Wood County Board of Education, Parkersburg, W. Va. 1968-1969...... NSF Academic Year Trainee, Graduate School, The Ohio State University, Columbus, Ohio 1969-1974 Teaching Associate, Department of Zoology, The Ohio State University Columbus, Ohio i 1974-1975 Research Associate, Center‘'for Lake Erie Area Research, The Ohio State University, Columbus, Ohio

Publications "New Records of Monopetalonema alcedinis (Rudolphi, 1819) (Nematoda: Filarioidea) from Wisconsin and California.11 The Passenger Pigeon 35(1): 36. 1973. "Gordius sp., A New Host Record," The Ohio Journal of Science 73(4): 228. 1973. "A Check List of the Helminth Parasites of the Blue Jay, Cyanocitta cristata L." The Passenger Pigeon 35(4): 191-193. 1 9 m "Helminth Parasites of the Brown-headed Cowbird, Molothrus ater ater, from Ohio." The Ohio Journal of Science 73(6) 1*76-180. 1973. "Occurrence of Helminth Parasites in Avian Hosts from South Bass Island, Ohio." The Ohio Journal of Science 74(1): 60-62. " Parasites of the Brown-headed Cowbird, Molothrus ater ater, in Ohio." Journal of Medical Entomology IIXT): 233. 1974. "Helminth Parasites of the Common Grackle, Quiscalus cuiscula versicolor, from South Bass Island, Ohio." Proceedings of the Helminthological Society of Washington 41(2): 233-237. 1974. "Helminth Parasites of the Robin from South Bass Island, Ohio." Journal of Wildlife Diseases 10(4): 397-398. 1974. "Helminth Parasites of the Red-winged Blackbirds from South Bass Island, Ohio, Including a Checklist of the Helminths Reported from This Host." Journal of Wildlife Diseases 10(4): 399-403. "The Helminth Parasites of the Blue Jay, Cyanocitta cristata bromia, from South Bass Island, Ohio." The Canadian Journal of Zoology 52(11): 1421-1423. 1974. "Helminth Parasites of Over-wintering Red-winged Blackbirds (Agelaius phoeniceus) from Laurel, Maryland;" The Journal of Parasitology 60(6): 962. 1974. "Helminth Parasitism in Juvenile House Sparrows, Passer domesticus (L.), from South Bass Island, Ohio, Including a List ofHelminths Reported from This Host in North America." The Ohio Journal of Science 74(6): 388-389. 1974. "Arthropod Parasites of the Common Grackle, Quiscalus cuiscula versicolor, in Ohio." Journal of Medical Entomology 12(2): In press. 1975. "Arthropod Parasites of Passerine Birds from South Bass Island, Ohio," Journal of Medical Entomology 12(4): In press. 1975. "Helminth Parasites of the Starling, Sturnus vulgaris, from South Bass Island, Ohio." The Journal of Parasitology 61(1): 161. 1975. "Helminth Parasites of the Common Grackle, Quiscalus quiscula versicolor, from Columbus, OhioT11 The Journal of Parasitology 61. In press. 1975. "A Check List of the Helminth Parasites of the Robin, Turdus migratorius Ridgway." American Midland Naturalist. In press. "Additional Check List of the Helminths of the Starling CSturnus vulgaris L .)." American Midland Naturalist. In press.

Field of Study

Major Field: Zoology Ji Studies in Helminthology. Professor John L. Crites

v TABLE OF CONTENTS Page ACKNOWLEDGMENTS...... ii VITA ...... iii LIST OF T A B L E S ...... vii LIST OF FIGURES...... viii INTRODUCTION...... V...... 1 MATERIALS AND METHODS...... 7 RESULTS...... 10 ...... 11 Cestoda ...... 63 . Nematoda 8 5 ...... 139 DISCUSSION...... 150 A. General Consideration of the Helminth Fauna of Passerine Birds...... 15Q B . The Concept of Landscape Epidemiology and a Modified Concept of the Nidality of Disease...... 16 9 C. The Nidality Index...... 173 D. A Helminth Biocenose Involving Passerine Birds on South Bass Island. . . 177 SUMMARY...... 182 LITERATURE CITED ...... 184 LIST OF TABLES Table Page I 1. Summary of Past and Present Studies of Helminth Parasites in Selected Passerine Birds...... 151 2. Prevalence and Intensity of Helminth Parasites in Fifty Red-winged Blackbirds . 15 9 3. Prevalence and Intensity of Helminth Parasites in Fifty Blue J a y s ...... 16Q M-. Prevalence and Intensity of Helminth Parasites in Fifty Brown-headed Cowbirds ...... 162 5. Prevalence and Intensity of Helminth Parasites in Fifty Common Grackles .... 163 6. Prevalence and Intensity of Helminth Parasites in Thirty House Sparrows .... 16M> 7. Prevalence and Intensity of Helminth Parasites in Fifty Starlings ...... 16 5 8. Prevalence and Intensity of Helminth Parasites in Fifty Robins...... 166

vii !

LIST OF FIGURES

Figure Pag< 1. The -Similarity Index of Helminth Faunas of Seven Avian Host Species from South Bass Island, Ottawa County, Ohio, 1968- 1971. (Sorensen, 1948)...... 170

2 Trellis Diagram of 2 80 Birds Examined. The Nidality Index in Percent is Based on Presence or Absence of 10 pecies of Parasitic Helminths in Passerine Hosts from South Bass Island, Ottawa County, Ohio, 1968-1971. (Dice, 1945) ...... 176

• • • V 1 H INTRODUCTION

Despite a slow accumulation of records of helminth parasites of passerine birds, little attention has been given to the prevalence and intensity of different parasite species in their respective hosts. Most accounts of the helminth parasites of passerine birds are based on small numbers of host examinations. There is scant knowledge of the nature and extent of parasitism in some of North America’s commonest species of birds. The knowledge of how a species of parasite is distri­ buted among various hosts in a biological community is less available than how it is distributed in one species in a single locality. No studies in North America have attempted to elucidate the prevalence and intensity of helminth para­ sites in an association of avian hosts from a defined community. In recent years, the work of Dr, L. S. Putnam and his students have revealed an interesting association of pas­ serine birds on South Bass Island. Goldman (196 9) noted a mixed flock association consisting of Red-winged Blackbirds (Agelaius phoeniceus), Common Grackles (Quiscalus quisoula versicolor), and Starlings (Sturnus vulgaris) on the Island.

1 Petrovic C1972) stated that the principal components of this association which forms mixed feeding flocks on lawns are the Red-winged Blackbird, Common Grackle, Brown-headed Cow- bird (Molothrus ater ater), Starling and Meadowlark (Sturnella magna). Other species foraging in the same habitat to a lesser degree include the Robin (Turdus migra- torius), Killdeer (Charadrius vociferus) and Yellow-shafted Flicker CColaptes auratus). Petrovic (1972) noted that Ring-necked Pheasant (Phasianus colchicus), Barn Swallow (Hirundo rustica), House Sparrow (Passer domesticus) and Eastern Kingbird (Tyrannus tyrannus) forage to a limited extent on the same areas. Conversations with these ornithologists stimulated my interest in this mixed flock association. Since these birds feed and flock together, one might expect their helminth fauna to be similar. Similarities and differences in the helminth faunas would be of considerable heuristic value in discussing the biology of both the avian and parasite popu­ lations. This being the case, I decided to investigate the prevalence and intensity of helminth species in several species of this avian association. The Red-winged Blackbird, Brown-headed Cowbird, Common Grackle, Robin, House Sparrow, and Starling were chosen for study due to their preponder­ ance of numbers in the association. Blue Jays (Cyanocitta cristata bromia) were investi­ gated for helminth parasites in the northeastern United States CBoyd, et al, 1956), Due to their ready availability during the spring migration period and the otherwise limited knowledge of helminth parasitism in this host, Blue Jays were included in this study. The ecology of helminth parasites has received little attention until comparatively recently. Many authors have demonstrated the effects of diet and age of the host on helminths in experimental , but very little is known of such effects in natural populations, While surveying the parasitological literature for information of an ecological nature, the author discovered a considerable body of research relating to this aspect of the study of parasites, All of this work had been performed in Central Europe and the U.S.S.R. It soon became apparent that the conceptual framework for this research was provided by the definition of landscape epidemiology.1 A group of workers in Russia under the direction of General Evgeny N. Pavlovsky studied the prevalence and intensity of parasites which cause disease in men and animals, i,e, zooanthroponoses (Garnham, 1971). They were particularly concerned with spring summer encephalitis, leishmaniasis, tularemia, leptospirosis, rickettsiosis, Q fever, brucellosis, and plague. The canopy concept which developed from these studies was landscape epidemiology. This concept was one developed entirely in reference to parasitic organisms in nature. In essence, it is an ecosystem approach in that climatic conditions and all other abiotic components of an area must be taken into considera­ tion if the distribution and abundance of a parasite popu­ lation in a biological community is to be fully understood. The research which gave rise to this concept was ' initiated in the 1930*s in the U.S.S.R. (Garnham, 1971). Charles Elton (1927) in the U.S.A. published an idea that continues to serve ecologists today. He stated that a good way to solve the problems presented by the distribution and abundance of animals in nature was to study the relationships between animals living together in communities. A community, in his mind, was a complex of animals that are usually found living together in a habitat. The meaning of habitat as presented in his essay on * population interspersion1 (19M-9) was defined as an area possessing a certain uniformity with respect to physiography, vegetation, or some other quality that the ecologist decides is important or easily recognized with rather arbitrary limits which are set in advance. Setting the limits is the first step toward the study of a community. Hence, for a generation, ecologists in the• United States and parasitologists in the U.S.S.R. were evolving a set of concepts using very different vocabularies- both literally and figuratively-to solve very similar problems. Although the concept of landscape epidemiology was developed in the study of zoonoses, it seems a very useful concept to use when thinking about the distribution and abundance of helminth parasites in a natural community, this is to say, a community in the sense of Elton. South Bass Island lies in the western basin of Lake Erie and forms a link in a chain of islands stretching from the Ohio mainland toward Canada. The Island's 2.6 square miles are mainly covered with old fields dominated by stag- horn sumac CRhus typhina), raspberries (Rubus sp.), and grasses. Vineyards cover a large part of the island, as do suburban lawns• Only about one-third of the island still has native forest vegetation, all in second or third growth stages. These old fields, lawns, vineyards and woodlots form the habitat which supports the passerine bird associa­ tion during the spring and summer months (Goldman, 1969). The island itself provides a well defined community with an obvious boundary for this study. Landscape epidemiology is a canopy concept composed of a number of supporting concepts. The principal supporting concept is that of the nidality of disease. These concepts developed in reference to infectious disease agents and must be modified to include the consideration of parasitic helminths in animals. Therefore, the purpose of this study is not only to determine the helminth fauna parasitizing the principal components of a passerine avian association in an insular commun^ ty but to place this information in the context provided by the concept of landscape epidemiology and a modified concept of nidality of disease. MATERIALS AND METHODS

A "total of 330 birds was examined. With the exception of House Sparrows, 50 birds of each species were examined for helminth parasites. Thirty House Sparrows were examined. Robins were shot on lawns and old fields in the central portion of the island. The other species were collected alive from two large decoy traps, operated in the central portion and the south point of the island respectively. Immediately upon collection, the birds were transported to the research building of The Ohio State University's Franz Theodore Stone Laboratory on South Bass Island. All birds were collected during the spring, summer and early autumn months between June, 1968, and June, 1971, Both adult and juvenile birds were collected for autopsy. A juvenile bird is defined as a recently fledged young-of-the-year bird. Banding and observational data indicated that juvenile birds collected from decoy traps in the early summer months probably hatched on South Bass Island. With certainty, all juvenile birds were hatched and reared in the immediate western basin region (Miskimen, 1970). Helminths removed from juveniles serve to indicate infections acquired within the limits of the island. To

7 acquire this information, a high proportion of juveniles was collected. The ratio of juvenile to adult birds in the entire collection was 53:47, Excluding the 50 adult Blue Jays taken during the spring migrations, the ratio was 62:38. Each of the birds brought alive to the laboratory was sacrificed by applying pressure to the thorax or by placing them individually in a 2-gallon jar containing chloroform vapors for no more than 30 seconds. Immediately upon sacri­ fice, the birds were autopsied. The brain, body, and nasal cavities and orbits of the eyes were examined. All organs were separated, teased apart and examined sequentially under a dissecting microscope. All developmental forms, mature and immature, from each tissue site were collected, identi­ fied and counted. Preparation of helminths for identifica­ tion followed standard techniques (Schell, 1962): the trematodes and cestodes were killed in heated Ringerfs fWarmf solution, fixed in Landowskyfs AFA solution, stained with Semicohn’s Carmine and mounted in Piccolyte Medium. The nematodes were placed in a 70% ethanol-glycerine solu­ tion and allowed to dessicate to pure glycerine. The • acanthocephala were either placed in 70% ethanol-glycerine solution which was allowed to dessicate to pure glycerine or were stained with Semicohn's Carmine and mounted in Piccolyte Medium. The line drawing was made with the aid of a Wild M-20 microscope equipped with a drawing tube. Representative specimens of the helminths recovered during this study have been deposited in the USNM Helminthological Collection. RESULTS

Descriptions of a given helminth species other than the original are uncommon. Adequate descriptions of specific identifying characters are of importance if the range of variability of such characters is to be defined for any given species. Forty-one taxa of helminth parasites were removed from the 330 passerine birds examined. Each species of helminth encountered during this study is described in detail utilizing the observations of previous workers as well as incorporating variations which have not been previously noted. Almost every description has been emended in light of the variations in morphology occurring in specimens collected during this study. The descriptions in which considerable new data have been incorporated are noted as ones highly modified from the previous literature descrip­ tions. The presentation of species is arranged systematic­ ally. Records of the occurrence of any given helminth are widely scattered in the literature. Following the original description, such records accumulate slowly. Collection of these records is of considerable importance to the para­ sitologist. Tabulation and study of these records serves to

10 11 indicate the geographic range of a parasite and the variety of host species the parasite can successfully utilize for growth, maintenance and dispersal. This information forms the basis for the parasitological concept of host specifici­ ty. This information is assembled and discussed for each of the helminths collected during this study. Basic information describing the life cycle of each helminth encountered during this study is discussed. Life cycle information provides the basis for understanding the transmission of a given species of helminth. Only when life cycle information is available can one begin to understand how a helminth population is maintained in space and time in any biological community.

Trematoda Leucochloridium variae McIntosh, 1932 Synonym: Leucochloridium pricei McIntosh, 19 32 Site of Infection: Cloaca Classification: Order Strigeatoidea La Rue, 1957 Superfamily Brachylaimoidea Allison, 1943 Family Brachylaimidae Joyeux and Foley, 1930 Subfamily Leucochloridiinae Poche, 1907 Genus Leucochloridium Carus, 18 35 Specific diagnosis (modified after Lewis, 1974): Adult ovate, flattened ventrally and convex dorsally, 1.68 to 12 1,93mm long by 0,81 to 1,04mm wide, cuticle armed. Oral sucker subterminal, 0,43 to 0,53mm in diameter, opens into subglobular pharynx, 0,19 to 0.22mm long by 0,13 to 0.18mm wide. Acetabulum slightly post equatorial, 0,42 to 0.51mm in diameter. Testes oblique, 0,10 to 0,22mm long by 0.11 to 0,16mm wide, located in postacetabular, intercecal zone. Cirrus sac oval, 0,13 to 0.18nun long by 0.10 to 0.16mm wide, situated between cecal ends. Cirrus armed. Ovary oval, 0.10 to 0.19mm long by 0.12 to 0.18mm wide, located anterior to posterior testes, Vittellaria lateral extending from midlevel of oral sucker to posterior margin of cirrus sac. Uterus largely intercecal with anterior loops on each side of oral sucker; traversing acetabulum, terminating near dor­ sal genital pore near posterior end of body. Eggs 0.22 to 0.28mm long by 0.13 to 0,19mm wide. Host records: Host Locality Record Mniotilta varia Michigan Canachites canadensis Alaska McIntosh (1932) Bonasa umbellus N. Hampshire Mueller (1941) frurdus migratorius Wyoming Rausch, In Kagan (1952) Lagopus spp. Alaska Babero (1953) Agelaius phoeniceus Cyanocitta cnstata Ohio Present study Discussion: Records of the occurrence of this helminth, although widespread in North America, are scant and restricted to galliform and passeriform hosts. This study records L. variae from Blue Jays and 13

Red^winged Blackbirds for the first time. Lewis C1974a) stated that further study may reveal Leucochloridium fus co striatum Robinson, 1940 as a synonym of L. variae. I Life cycle: Aspects of the developmental cycle of L. variae were observed experimentally by Lewis (1974a). Branched sporocysts containing meta- cercariae enclosed in cysts were recovered from naturally infected specimens of the land snail, Succinea ovalis. Adult worms were recovered 2 to *+6 days after feeding metacercariae to chicks (Gallus gallus),

Prevalence and Intensity of Leucochloridium variae in Passerine Birds from South Bass Island. Range Infected Birds/No. Examined Birds of Mean Adults Juveniles Exam- Preva-Inten-Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Agelaius 50 2 11 — 1 (22) 0 (28) phoeniceus Cyanocitta 50 4 1-12 6.5 2 (50) 0 (0) cristata

Leucochloridium dryobatae McIntosh, 1932 Synonym: Urogonimus dryobatae (McIntosh, 1932) Site of Infection: Cloaca Classification: Order Strigeatoidea La Rue, 1957 Superfamily Brachylaimoidea Allison, 194 3 Family Brachylaimidae Joyeux and Foley, 19 30 Subfamily Leucochloridiinae Poche, 1907 Genus Leucochloridium Carus, 18 35 1l Specific diagnosis (modified after McIntosh, 1932): Body ovate, flattened ventrally, convex dorsally, 2.19mm long by 1.28mm wide; cuticle unarmed. Oral sucker subterminal, 0,48 to 0.56mm in diameter; opens into oval pharynx, 0.17mm long by 0.25mm wide. Acetabulum slightly postequatorial, 0.5 7 to 0.62mm in diameter. Testes tandem, round, 0.14 to 0.20mm in diameter, located in postacetabular, intracecal zone. Cirrus sac round, 0.06 0mm in diameter, located at posterior end of body. Ovary oval, 0.17mm long by 0.22mm wide, located tandem between testes. Vittellaria extracecal, extending from posterior level of oral sucker to level of ovary. Uterus fills intracecal space with anter­ ior loops on each side of oral sucker; terminates at common genital pore at posterior end of body. Eggs 0.25mm long*by 0.018mm wide. Host records: Host Locality Record Dendrocopus villosus, toolothrus ater, Pipilo erythropthalmus, Pirangea olivacea. Host Locality Record Sphvrapicus varius, Vlreo olivacea Michigan McIntosh (1932) Vireo olivaceus Washington, McIntosh and D.C. McIntosh (1935) Carpodacus purpureus Virginia Reynolds (1936) Spizella pusxlla Michigan Cheatum, In Kagan (1952) Regulus satrapa Wisconsin Rausch, In Kagan C1952) Molothrus ater Manitoba, Hodasi (1963) Can. Toxostoma rufum Iowa Peet and Ulmer (1970) Cyanocitta cristata Ohio Present Study

Discussion: Records of the occurrence of L. dryobatae are largely restricted to woodland dwelling passerine birds collected in eastern North America.' This study records L. dryobatae from the Blue Jay for the first time. Eleven species of the genus Leucochloridium were described by McIntosh (1927, 1932). All of these species were collected from woodland dwelling or woodland frequenting birds, largely passerines. Differ­ ences in body size and body conformation and organ size and organ conformation between these describ­ ed species are slight. In addition, a number of species of this genus hqve been described from avian hosts in Europe. In Russia, Bykhovskaya - Pavlovskaya (1951, 1953) studied over 900 specimens of the genus Leucochloridium and concluded that their 16 individual variability precluded the use of most of the characters commonly used to differeniaate species in this genus. She stated that 10 species are probably valid and named 18 species as synonyms. She named Leucochloridium sp. of Witenberg, 1925; L. vlreonis, McIntosh, 1927; L. certhiae McIntosh, 1927; L. mniotitlae McIntosh, 1927; L. seiuri McIntosh, 1932; L. dryobatae McIntosh, 1932; L. cardis Yamaguti, 1939; and L, witenbergi Skryabin, 1948 as synonyms of L. macrostomum (Rudolphi, 1803). In the same study, sihe named L. insigne of Witenberg, 192 5, nec Looss, 1899; L. pricei McIntosh, 1932; L. variae McIntosh, 1932; L. cyanocittae McIntosh, 1932; L. melospizae McIntosh, 1932; and L. sime CYamaguti, 1935) as synonyms of L. actitis McIntosh, 1932. Several experimental studies by Pojmanska C1967, 1969a, 1969b) provided considerable evidence to reinforce the opinions of Bykhovskaya - Pavlovskaya. In contrast, Kagan (1952) and Lewis C1974a, 1974b) were extremely reluctant to synonomize any of the species described by McIntosh. Both authors state the opinion that until extensive life history studies have been performed, the status of the North American species of Leucochloridium must 17 remain unchanged. Life cycle: Details of the developmental cycle of L. dryobatae are unknown. The life cycle of L. dryobatae is probably similar to that of L. certhiae (McIntosh, 1927). Lewis (1974b) reported aspects of the developmental cycle of L. certhiae. Irregularly branched sporocysts containing encyst­ ed cercariae were recovered from naturally in­ fected land snails, Cionella lubrica. Mature trematodes were recovered from chicks (Gallus gallus) twenty-two days after ingestion of metacercariae. Lewis (1974b) stated that life cycles in the sub­ family Leucochloridiinae exhibit progressive adaptations for reduction of water loss and for facilitation of host transfer. Adaptation for reduction of water loss include the retention of cercariae and/or metacercariae within the sporocyst and development of thick gelatinous metacercarial cysts within the sporocysts. Adaptations for facilitation of host transfer involve the development of motile and pigmented sporocysts broodsacs which alter snail behavior and attract avian predators. Prevalence and Intensity of Leucochloridium dryobatae in Passerine Birds from South Bass Island: Twelve mature 18 specimens were removed from a single Blue Jay of the 50 adult Blue Jays examined during this study.

Leucochloridium vireonis McIntosh, 1927 Site of Infection: Cloaca Classification: Order Strigeatoidea La Rue, 1957 Superfamily Brachylaimoidea Allison, 194-3 Family Brachylaimidae Joyeux and Foley, 1930 Subfamily Leucochloridiinae Poche, 1907 Genus Leucochloridium Carus, 18 35 Specific diagnosis (.modified after McIntosh, 1927) : Body ovate, flattened ventrally, convex dorsally, 2.50mm long by 0.95mm wide; cuticle unarmed. Oral sucker subterminal, rounded, 0.47 to 0,53mm in diameter; opens into muscular, subglobular pharynx, 0.17mm long by 0.21 mm wide. Acetabulum 0.57mm in diameter. Testes oblique, oval 0.22 to 0.23mm long by 0.18 to 0.19mm wide, located in postacetabular, intercecal zone. Cirrus sac large, 0.16 to 0.20mm long by 0.12 to 0.17mm wide, situated posterior and ventral to posterior testes. Ovary round, 0.18 to 0.24mm in diameter, situated to right of median line at level of anterior testes. Vittellaria lateral extending, from level of oral sucker to level of ovary. 19 Uterus fills intercecal zone with anterior loops on each side of oral sucker, terminates at common genital pore near posterior end of body. Eggs 0.23mm long by 0.15mm wide. j Host records: Host Locality Record Vireo g. griseus Mississippi McIntosh (.19 27) Telmatodytes palustris Michigan Cheatum, In Kagan C1952) Cyanocitta cristata Ohio Present Study Discussion: Records of the occurrence of L. vireonis are scant and restricted to woodland dwelling birds from the central portion of the United States. This study records L . vireonis from the Blue Jay for the first time. Life cycle: Details of the developmental cycle of L. vireonis are unknown. Aspects of the life cycle of L. vireonis probably parallel those of L. certhiae (McIntosh, 19 27) reported by Lewis (.1974b) . Prevalence and Intensity of Leucochloridium vireonis in Passerine Birds from South Bass Island: three mature specimens were removed from a single Blue Jay of the 50 adult Blue Jays examined during this study.

Echinostoma revolutum (Froelich, 1802) Looss, 1899 Synonyms: Fasciola revoluta Froelich, 18 02 20 Echinostoma echinatum CZeder, 1803) Nicoll, 1923 Distoma oxvcephalum Rudolphi, 1819 Distoma dilatatum Miran, 1840 Distoma armaturn Molin, 1850 Echinostoma mendax Dietz, 1909 Echinostoma paraulum Dietz, 1909 Echinostoma coalitum Barker and Beaver, 1915 Echinostoma armigerum Barker and Irving, 1915 Echinostoma limicoli Johnson, 1920 Echinostoma cjnetorchis Ando and Osaki, 1923 Echinostoma columbae Zunker, 19 25 Echinostoma mivagawai Ishii, 1932 Echinostoma rubustum Yamaguti, 19 3 5 Echinostoma revolutum tenuicollis Baschkirowa, 1941 Echinostoma multispinosa Perez-Vigueras, 1941 Species inquirende Beaver, 193 5 Echinostoma echinocenhalum Rudolphi, 1819 Echinostoma Sudanese Odner, 1911 Echinostoma acuticauda Nicoll, 1914 Echinostoma callawavensis Barker and Noll, 1915 Echinostoma erraticum Lutz, 1924 21 Echinostoma neglectum Lutz, 1924 Echinostoma nephrocystis Lutz, 1924 Echinostoma microchis Lutz, 1924- Site of Infection: Cloaca 1 Classification: 1 Order Echinostomida La Rue, 1957 Superfamily Echinostomatoidea Faust, 1929 Family Echinostomatidae Looss, 1899 Subfamily Echinostomatinae Faust, 1929 Genus Echinostoma Rudolphi, 1809 Specific diagnosis: Body elongate, 4.60mm long by 1.17mm wide, provided with head collar bearing a single row of 37 spines. Oral sucker, 0.22mm in diameter, situated medially at anterior edge of head collar. Pharynx present, 0.17mm long by 0.15mm wide. Acetabulum large, muscular, 0.6 5mm long by 0.56mm wide. Testes tandem, oval, 0.30mm long by 0.20mm wide, located posterior to ovary. Cirrus sac oval, 0.17mm long by 0.13mm wide, located at anterior edge of acetabulum. Ovary round, 0.24mm in diameter, situated at approxi­ mate midpoint of body length. Vittellaria extend 3.58mm from region posterior to acetabulum to posterior tip of body. Uterus fills mid-third of body with coil extending to median genital pore located- just anterior of acetabulum. 22 Discussion: Echinostoma revolutum has been reported extensively from avian and mammalian hosts throughout the tropical and temperate regions of the world. Few, if any, helminths of wildlife have been reported as frequently from as many host' species. The listing is enormous, even for the United States. Remarkably, this study records E. revolutum from the starling for the first time. Life cycle: The developmental cycle of E. revolutum has been observed experimentally in most of the regions of the world. Studies in the United States include those by Johnson (.1920), Beaver (1937) and Churchill (1950). This life cycle has been summarized by Schell (.1970). Development of miricidia requires three to four weeks after the eggs are passed by the definitive host. Miricidia hatch and penetrate snails of the genera Fhysa, Lymnaea, Stagnicola, Helisoma and Fseudosuccinea, in which a generation of sporocysts and two generations of rediae develop. Daughter redia produce cercariae which, after emerging from the snail, encyst in the same snail, or a different species of snail or in clams or in larval amphi­ bians. Definitive hosts become infected upon ingestion of an infected second intermediate host. 23 Prevalence and Intensity of Echinostoma revolutum in Passerine Birds from South Bass Island: One mature specimen was removed from a juvenile starling of the ^9 juvenile starlings examined during this study. I Plagiorchis noblei Park, 19 36 Synonym: Plagiorchoides noblei Olsen, 19 37 Site of Infection: Cloaca and lower infection Classification: Order Plagiorchioida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Plagiorchiidae Luhe, 1901 Subfamily Plagiorchiinae Pratt, 1902- Genus Plagiorchis Luhe, 1899 Specific diagnosis: Body fusiform, 0.98 to 2,71mm long by 0.2 8 to 1.07mm wide; cuticle armed. Oral sucker subterminal, 0.16 to 0.31mm in diameter; opens into oval pharynx, 0.07 to 0.15mm long by 0.12 to 0.18mm wide. Acetabulum muscular, 0.1M- to 0.23mm in diameter, located in anterior one-half of body. Testes oblique, oval to round, 0.17 to 0.31mm long by 0.16 to 0.29mm wide, located posterior to ovary. Cirrus sac claviform, 0.36 to 0.72mm long by 0.06 to 0.09mm wide, located sub- medially just anterior to acetabulum. Ovary round, submedian, Q.05 to 0.20mm in diameter, 24 located at level of posterior edge of acetabulum. Vittellaria lateral, meeting medially at posterior tip of hody, Q.88 to 1.53mm long. Uterine coils fill mid-section of body; single coil of uterus »I extends anteriorly to terminus near genital pore at anterior edge of acetabulum. Eggs 0.07 to 0.08mm long by 0.0 3 to 0.04mm wide. USNM No. 72692. Host records: Host Locality Record Agelaius phoeniceus California Park (.1936) Xanthocephalus xanthocephalus Iowa Lee (.1961) Agelaius phoeniceus Iowa Ellis (1963) Agelaius phoeniceus, Icterus galbula, Xanthocephalus xanthocephalus Manitoba, Can. Hodasi (1963) Agelaius phoeniceus Ohio Williams (1964) Agelaius pheoniceus Ontario, Can. Bourns (19 66) Agelaius phoeniceus Colorado Wallace and Olsen (1966) Agelaius phoeniceus Ohio Stanley and Rabelais (1971) Agelaius phoeniceus, Eptesicus fuscus Iowa Blankenspoor (19 70) Agelaius phoeniceus, Molothrus a. ater, Quiscalus quiscula, Sturnus vulgaris Ohio Present Study Discussion: Records of the occurrence of P. noblei indicate that the distribution of this helminth is widespread in North America and restricted largely to members of the avian family Icteridae. Blankenspoor (1970) stated that P. noblei is a naturally occurring intestinal parasite of Red­ winged Blackbirds and Yellow-headed Blackbirds (Xanthocephalus xanthocephalus). This study records P. noblei from the Brown-headed Cowbird, Common Grackle and Starling for the first time. Life cycle: Details of the developmental cycle of Plagiorchis noblei have been observed experi­ mentally by Blankenspoor (1970). Blankenspoor summarized the life cycle in the-- fr&*Liewing- manner Two species of aquatic snails are involved, Stagnicola reflexa and Lymnaea stagnalis. Following ingestion of eggs, miricidia hatch, penetrate the snails' intestines and metamorphose to form mother sporocysts on the outer surface of the intestines. Daughter sporocysts, produced by mother sporocysts, migrate to the hepatopancreas of the snail and produce large numbers of cercariae. Cercariae penetrate and encyst in a variety of second intermediate hosts, usually . Definitive hosts acquire their infec­ tions by ingestion of the infected second inter­ mediate hosts. Blankenspoor succeeded in experi­ mentally infecting ten species of passeriform birds, three species of galliform birds, one species of ralliform bird and two species of mammals with P. noblei. Considerable variability 26 in organ development: and body shape was observed in specimens recovered from the various experi­ mental infections. Prevalence and Intensity of Plagiorchis noblei in Passerine Birds from South Bass Island: Range Infected Birds/No.Exam. Birds of Mean Adults Juveniles Exam-Preva- Inten- Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Agelaius phoeniceus 50 6 1-12 1.3 2 (22) 1 (28) Molothrus a. ater 50 4 1-25 13.0 1 (46) 1 (4) Quiscalus quiscula 50 4 2 2 0 (.13 ) 2 (37) Sturnus c vulgaris 50 2 32 0 Cl) 1 (49)

Conspicuum icteridorum Denton and Byrd, 1951 Synonym: Eurytrema icteridorum Lumsden and Zischke Site of Infection: Gall bladder Classification: Order La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Dicrocoeliidae Odhner, 1911 Subfamily Dicrocoeliinae Faust, 1929 Genus Conspicuum Bhalerae, 1936 ^ __ _ Specific diagnosis (highly modified after Denton and Byrd, 1951): Body broadly fusiform, flattened ventrally, strongly convex dorsally, 1.7 to 5.89mm tiS

' 27 long by 0.51 to 2.12mm wide; cuticle thick, wrinkled, and unarmed. Oral sucker subterminal, 0.187 to 0.578mm in diameter; opens into globular pharynx, 0.1U to 0.21mm in diameter. Acetabulum i large, deep, muscular, situated at junction of anterior and middle body thirds, 0.32 to 0.90mm in diameter. Testes opposite, oval to slightly lobed in mature specimens, round to triangular in specimens not fully mature, 0.08 to 0.52mm in diameter, located at level of posterior edge of acetabulum. Cirrus sac median, elongate, 0.179 to 0.81mm long by 0.08 9 to 0.4-2mm wide, located anterior to acetabulum. Ovary median, round to transversely oval, 0.09 to 0.3 9mm in diameter, located posterior to testes. Vittellaria, largely extracaecal, extending posteriorly from testicular zone 0.7 8 to 2,23mm. Uterus appears as dark, brown chain-like thread in posterior half of young specimens. Uterus follows a medial dorsal course anteriorly to median genital pore at • pharyngeal level. Eggs 0.025 to 0.032mm long by 0.015 to 0.020mm wide; contain miricidia. USNM Nos. 72689, 72690 28 Host records: Host Locality Record Cassidix mexicanus Texas ma^ or. Cassidix m. Texas prosopidicola Eu^>hagus carolTnus Texas Quisculus quiscula Ga., Mich., Tenn. Texas Sturnella magna Georgia, Texas Denton and Byrd C1951) Quisculus quiscula New York Patten C19 52) versicolor Ammospiza maritima North Carolina Hunter and Quay C1953) Euphagus cyanocephalus California Bassett (1958) Quiscalus quiscula versicolor Manitoba, Can. Hodasi (1963) Agelaius phoeniceus, Cassidix mesamexicanus Louisiana Lumsden and Zischke (196 3) Passerella iliaca, Quiscalus quiscula, Sturnella magna Iowa Ellis 0963) Agelaiusphoeniceus, Quiscalus quiscula venscolor Ohio Stanley and Rabelais (1971) Molothrus a. ater Ohio Cooper, Troutman and Crites (1974) Agelaius phoeniceus Maryland Cooper and Crites (1974) Quiscalus quiscula Ohio Buck, et al. (1975) Agelaius phoeniceus, Molothrus a. ater, Quiscalus quiscula versicolor, Passer domesticus, Sturnus vulgaris- Ohio . Present Study Discussion: Records of the occurrence of this helminth show a widespread distribution across the conti­ nent. The vast majority of records are confined to the family Icteridae. This study records C. icteridorum from the House Sparrow and Starling 29 for the first time. Specimens collected from the House Sparrow and Starling were quite immature. No specimen resembling a fully mature C. . icteridorum was collected from either host. I Determination was made by comparison of these‘ specimens from juvenile House Sparrows and Starlings with equally immature specimens collected in considerable numbers during this study from juvenile Common Grackles. Several studies of Starling helminths have been made in this country. Based‘on the lack of records of C . icteridorum from this host 9 it seems reason­ able to conclude that icteridorum seldom if ever reaches maturity in Starlings. Conspicuum icteridorum seems adapted to icterids. The tissue response of C. icteridorum infections in Brewerf s Blackbirds (Euphagus cyanocephalus) was studied by Bassett (1958). The pathology attending this infection in the gall bladder was minimal. Bassett concluded that host tissue- ex­ hibits a continued ability to heal in the presence of the fluke after a mild initial in­ flammatory response. Observation of infections in House Sparrows and Starlings revealed no gross pathological changes. Studies by a number of european parasitologists, including Timon-David (1953), Baer (1957), Mettrick (1963), and Macko (1968, 1971) revealed that the range of variability in described organ size, organ displacement, and body shape for a given species of the trematode family Dicro- coeliidae is considerable. Specimens from a single host may exhibit proportions which overlap the limits of several described species. This is. true of specimens of C. icteridorum collected during this study as well. Denton and Byrd (1951) described C. icteridorum and C. macrorchis in the same publication. The original descriptions vary only in some organ measurements and in subtlety of wording. Considerable overlap exists in organ measurements recorded in the original descriptions. It seems reasonable to synonomize the two species. C. icteridorum has page priority. Life cycle: Details of the developmental cycle of £. icteridorum were observed experimentally by Patton (1952). Mature eggs containing miricidia are passed in the feces of the bird host. When eaten by the land snail Zonitoides arboreus, the eggs hatch in the intestine. The released miricidia penetrate the gut wall and initiate . development in the digestive gland of the snail. Daughter sporocysts are released from original 31 mother sporocysts 70 days after infection of the snail* Daughter sporocysts mature in 95-100 days post infection. At maturity, daughter sporocysts, containing 5-14 cercariae, migrate to the mantle cavity of the snail, and into the slime trail during periods of high humidity. The terrestrial isopods Armadillidium quadrifrons and Oniscus acellus become infected with meta­ cercariae upon ingestion of cercariae-containing sporocysts in the slime trails of the snails. Immature flukes were recovered from the gall bladder of a Common Grackle 16 hours after in­ gesting infected isopods. Relatively mature specimens were recovered from a single Common Grackle 12 weeks post infection CPatten, 1952), If a large number of avian hosts had been infected and maintained for lengthly periods of time, the similarity of C . icteridorum and C. macrorchis descriptions would have become evident. In the original descriptions and plates, C. icteridorum resembled relatively young mature specimens while C. macrorchis resembled fully gravid specimens. Patten (1952) depicted an incompletely mature specimen from the 12 week-old Common Grackle. 32 Prevalence and Intensity of Conspicuum icteridorum in Passerine Birds from South Bass Island:

No. Range Infected Birds/No. Examined Birds of Mean Adults Juveniles Exam- Preva-Inten- Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Agelaius 50- 48 1-8 2.4 13 (22) 11 (28) phoeniceus Molothrus 50 4 1-5 3.0 2 (46) 0 (4) a. ater Passer 30 3.3 1 - 0 (5) 1 (25) domesticus Quiscalus 50 46 1-13 4.5 10 (13) 13 (37) quiscula Sturnus 50 8 1-3 1.5 0 (1) 4 (49) vulgaris

Lutztrema microstomum Denton and Byrd, 19 51 Site of Infection: Gall bladder and liver Clas s if ication: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Dicrocoeliidae Odhner, 1911 Subfamily Dicrocoeliinae Faust, 1929 Genus Lutztrema Travassos, 1941 Specific diagnosis (modified after Denton and Byrd, 1951): Body filiform, 3.00 to 4.95mm long by 0.19 to 0.46mm wide, cuticle thin, unarmed. Oral sucker subterminal, 0.07 to 0.13mm in diameter, 33 preceded by liplike projection. Acetabulum large, 0.14 to 0.23mm in diameter, situated about one-tenth of body length from anterior. Pharynx, 0.03 to 0.05mm long by 0.0 3 to 0.06mm wide, opens

j to indistinct esophagus. Single cecum present; passes between anterior testes and posterior testes and ovary, ending at approximate mid-point of body. Testes, oblique to tandem, round to tranversely oval, 0.11 to 0.20mm long by 0.12 to 0.31mm wide, located posterior of acetabulum. Cirrus sac elongated-pyriform, 0.11 to 0.16mm long by 0.06 to 0.07mm wide, located just anterior to acetabulum. Ovary transversely oval, 0.08 to 0.15mm long by 0.08 to 0.18mm wide, located sub- medially posterior to testes. Vittellaria com­ posed of 7-21 large, oval follicles on each side of body; follicles often fuse in median line anteriorly. Uterus fills posterior 1/2 of body; follows looping course between ovary and testes to genital pore which is located midway between oral sucker and acetabulum. Eggs 0.029 to 0.035mm long by 0.019 to 0.24mm wide; contain miracidia. USNM No. 72633. 34 Host records: Host Locality Record Cyanocitta cristata Texas, Ga., N . Denton and Byrd Carolina C1951) Cyanocitta cristata Mass, N.Hamp., Boyd, et al. Vermont C1956) Cyanocitta cristata Iowa Ellis C1963) Toxostoma rufum Iowa Peet and Ulmer (1970) Cyanocitta cristata Ohio Present Study Discussion: The limited number of records of L. microstomum infections indicate that the Blue Jay is most frequently host to this helminth. Surveys of the crow (Corvus brachyahynchos), the other common Worth American corvid, report Brachylecithum americanum CDenton, 1945) as the most frequent parasite of the gall bladder and liver of this avian species. L . microstomum is unreported from the crow. During this study, Lutztrema microstomum was recovered only from Blue Jays. Life cycle: Aspects of the developmental cycle of L. microstomum are unknown. Its life cycle is probably similar to that of Lutztrema monenteron CPrice and McIntosh, 1935) which utilizes land snails and a terrestrial arthropod for develop­ ment . 35 Prevalence and Intensity of Lutztrema microstomum in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva-Iten- Iten-Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Cyanocitta cristata 50 ’ 60 3-199 24.1 30 C50) 0 CO)

Lutztrema monenteron CPrice and McIntosh, 1935) Travassos, 1941 Synonym: Lyperosomum monenteron Price and McIntosh, 1935 Site of Infection: Gall bladder Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Dicrocoeliidae Odhner, 1911 Subfamily Dicrocoeliinae Faust, 1929 Genus Lutztrema Travassos, 1941 Specific diagnosis (modified after Price and McIntosh, 1935): Body lanceolate, 1.9 to 5.2mm long by 0.630 to 0.670mm wide at. the level of acetabulum; cuticle unarmed. Oral sucker subterminal, 0.120 to 0.170mm long by 0.123 to 0.150mm wide; aceta­ bulum 0.170 to 0.320mm wide. Pharynx subglobular, ' 0.042 to 0.060mm long by 0.050 to 0.070mm wide; 36 esophagus present. Single ceca undulates between testes and ovary ending approximately 1/3 of body length from posterior tip. Testes transversely oval, 0.140 to 0.26 0mm long by 0.250 to 0.46 0mm wide; positioned diagonally posterior to aceta­ bulum. Cirrus sac elongate, 0.160 to 0.320mm long by 0.0 65 to 0.10 0mm wide, located anterior to acetabulum. Ovary transversely oval, 0.09 0 to 0.125mm long by 0.100 to 0.220mm wide, located posterior to testes and to right of median line. Vittellaria clumped in relatively large follicles in mid-body; follicles tend to meet in median line. Uterus fills posterior half of body with one uterine loop extending anteriorly to median genital pore at level of pharynx. USNM No. 7 2691. Host records: LocalityHost Record Sialis sialis, Virginia Price and Turdus mxgraterius Quebec, Can., Va., McIntosh(C19 3 5) Washington, D .C. Bonasa umbellus, Tyrannus tyrannus Minnesota Ishii (1942) Turdus migratorius New York Webster (1943) Bonasa umbellus Minnesota Erickson, Highby, and Carlson (1949) Mimus polyglottos, Georgia Toxost'oma rufum Texas, Virginia Turdus migratorius Ga. , N. Carolina, Denton and Byrd Ohio, Tenn., Texas (1951) Cyanocitta cristata Massachusetts Boyd, et al. (1956) Corvus f . frugilegus, 37 Host Locality Record Turdus m. merula, Turdus v. viscivorus England Mettrick Cl956) Ixoreus n . naevis Idaho PipilcT"erythophalmus Washington Turdus migratorius Idaho Schell C1957 Corvus f frugilegus , Turdus m . merula, Turdus pilaris Turdus v . visciyorus England Mettrick (195 8a) Turdus migratorius Michigan Villella (1961) Sturnella magna Iowa Ellis (1963) Turdus migratorius Colorado Slater (1964) Turdus migratorius Montana Carney (1966) Turdus migratorius Alberta, Can. Holmes, In Carney (1966) Turdus merula Czechoslovakia Groschaft (1969) Toxostoma rufum Iowa Peet and Ulmer (1970) Turdus migratorius Illinois Levin (1970) Corvus frugilegus Czechoslovakia Barus, et al. (1972) Turdus migratorius Ohio Present Study

Discussion: Records indicate that L. monenteron is the common dicrocoeliid trematode of the gall bladder of Robins across North America. Both Carney (1966) and Levin (19 70) suggested that infections by this helminth in other species of passerine birds are uncommon. Of all the birds examined during this study, only adult Robins were infected with L. monenteron. Denton and Byrd (1951) stated that the description of this North American form agrees very closely with the description of Lutztrema obliquum (Travassos, 1971), a form described from South American avian hosts. The fact that none of the reported hosts 38 for L. monenteron, with the exception of the Kingbird (Tyrannus tyrannus), migrate to the southern hemisphere seemed a sufficient argument for not synonomizing specific names of the two forms. More .recently, a number of .european workers have reported L. monenteron from several species of passerine birds in Europe. Groschaft C1969) stated that the relatively expansive geographical distribution of birds of the family Turdidae Choloarctic and neotropical) and the far- reaching migration of some representatives of this family could have resulted in these avian species having several helminth species in common. Further studies may reveal L. monenteron to be .a, .sy.npnym of L.. obliguum. Life cycle: Several workers have contributed to our knowledge of the developmental cycle of L. monenteron. The entire cycle has yet to be fully elucidated experimentally. Denton (In Carney, 1966) found four molluscs could serve as first intermediate hosts of L. monenteron: Bulimulus alternatus mariae, Polygyra texasiana, Practicellela berlandieriana .. and Deroceras agrestis. Villella (1961) recovered daughter sporocysts from Vallonia pulchella. Development proceeded only to the mother sporocyst stage in 39 ■the molluscs Anguispira alternata, Deroceras laeve, Retinella indentata, and Triodopsis multilineata. Carney (1966) observed cercariae in sporocysts in the slime trails of five I Anguispira ptychophora 148 days post infectioii. Many of the details of the development•cycle of L. monenteron have apparently1 been observed experimentally by Krissinger and Denton (1972). In their studies, sporocysts developed after . ingestion of eggs by the molluscs Ventridens intertextus, Deroceras reticulatum, and Lehmannia poireri 80 days post infection. A small milliped, Dixidesmus sp., was found to serve as the second intermediate host. Millipeds fed actively on the cercariae-containing slime balls. Encysted metacercariae were recovered from the hemocoel of millipeds within 2 days of slime ball ingestion (Krissenger and Denton, 1972). The details of the development of L. monenteron within the second intermediate host and the • definitive avian host remain to be reported. Prevalence and Intensity of Lutztrema monenteron in Passerine Birds from South Bass Island: 40 Range Infected Birds/No. Exam. Birds of Mean- Adults Juveniles Exam- Preva- Inten- Inten-Infect-/No. Infect-/No. Host ined lence sity sity_____ed Exam. ed Exam.

Turdus 50 20 1-11 5.9 10 (19) 0 (31) migra­ torius

Lyperosomum oswaldoi (Travassos, 1919) Travassos, 1944 Synonym: Oswaldoia oswaldoi Travassos, 1919 Lyperosomum urocissae Yamaguti, 1939 Site of Infection: Gall bladder Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Dicrocoeliidae Odhner, 1910 Subfamily Dicrocoeliinae Faust, 1929 Genus Lyperosomum Looss, 1899 Specific diagnosis (modified after Denton and Byrd, 1951): Body filiform, 5.5 3mm long by 0.65mm wide, flattened dorsoventrally; cuticle unarmed. Oral sucker 0.13mm in diameter. Acetabulum large, 0.43mm long by 0.35mm wide, muscular, located within anterior 1/6 of body. Pharynx globular, 0.15mm in diameter. Ceca slender, terminating midway between end of vittellaria and posterior . tip of body. Testes tandem, oval, 0.20mm long by 0.15mm wide, located posterior of acetabulum. 41 Cirrus sac claviform, 0.20mm long by 0.04mm wide, located anterior to acetabulum. Ovary round, 0.17mm in diameter, situated posterior to caudal testes. Vittellaria lateral, composed of numerous small follicles beginning within zone of anterior testes, extending posteriorily for 2.39mm. Uterine coils fill posterior 2/3 of body length, with coil extending anteriorly between testes to genital pore which is located at level of pharynx. Eggs 0.26 to 0.33mm long by 0.18 to 0.22mm wide; contain miricidia. Host records: Host Locality Record Cyanocitta cristata Texas Toxostoma rufum 6a., Miss., Texas Denton and Byrd C1951) Cape11a stenura USSR Bikhovskaya- Pavlovskaya (1962) Toxostoma rufum Iowa Ellis (1963) Hirundo rustica Poland Jaron (1969) Cacicus h. haemorrhous, Molothrus b. bonariensis Mviospiza h. humeralis. Odontotophorus c.. capueira, Ramphocelus b. bresilius Brazil Travassos, et_ al. (1969) Tachyphomus coronatus Toxostoma rufum Iowa Peet and Ulmer (1970) Cyanocitta cristata , Ohio Present Study Discussion: Records of the occurrence of L. oswaldoi - are infrequent, yet widespread. Its distribution is holoarctic and neotropical. Denton and Byrd (1951) stated that L. oswaldoi is the western hemisphere's equivalent of the eastern hemis­ phere •s Lvperosomum longicauda (Rudolphi, 1&09)* These workers stated that L. oswaldoi would probably fall as a synonym of L. longicauda* Since that opinion was rendered several records of L. oswaldoi from European hosts have been reported. The failure of a number of European workers to synonomize the two indicates the forms may be distinct where their ranges overlap. Specimens of the genus Lvperosomum are quite variable. Macko (1969) studied 50 specimens taken from 11 Crex crex. The variability among specimens of L. pawlowskvi Strom, 192& in his study was considerable. Macko (I960) compared Lvperosomum spp. from European hosts and found them quite similar. He did not include L. oswaldoi. L. longicauda. nor L. pawlowskvi among them. Comparison of descriptions by this author indicate that the latter three species are equally similar. One is lead to a conclusion similar to that of Groschaft (1969) for the genus Lutztrema. To paraphase Groschaft, when the relatively ex­ pansive geographical distribution of birds of the families Corvidae and Turdidae and when the far reaching migrations of some representatives of these families are considered, these birds may be parasitized by a single cosmopolitan species of the genus Lvperosomum* j Life cycles Aspects of the development cycle of Lvperosomum sp* have been briefly outlined by Denton (1941)* Eggs were passed in the feces of the avian host and were eaten by the land snails (Polvgvra texasiana and Praeticallela berlandieri- ana). Sporocyst generations result in cercariae which migrate to the mantle cavity of snail and subsequently are expelled in masses of slime* Gercarial development requires 100 or more days in snails* Metacercariae were recovered from several chrysomelid (Gastroidia cvanea)* Prevalence and Intensity of Lyperosomum oswaldoi in Passerine Birds from South Bass Island: A single mature specimen was removed from a single Blue Jay of the 50 adult Blue Jays examined during this study*

Zonorchis alvevi (Martin and Gee, 1949) Denton and Byrd, 1951 Synonym: Eurvtrema alvevi Martin and Gee, 1949 Site of Infection: Gall bladder Classification: Order Plagiorchiida La Rue, 1957 4 4 Superfamily Plagiorchioidea Dollfus, 1930 Family Dicrocoeliidae Odhner, 1910 Subfamily Dicrocoeliinae Faust, 1929 Genus Zonorchis Travassos, 1944 Specific diagnosis (modified after Martin and Gee, 1949)s Body fusiform, flattened, 2*61 to 5«32mm long by 1.13 to l.£4mra wide; cuticle is unarmed but with small protuberances on the anterior sur­ face. Oral sucker subterminal to terminal, averaging 0.345mm in diameter, opening into pharynx, 0.13 to 0.24mm long by 0.06 to 0.15mm wide. Acetabulum very muscular, 0*32 to 0.45mm long by 0.43 to 0.54mm wide. Bsophagus present. Intestinal caeca approach posterior terminus of body. Testes oval to slightly lobed, 0.19 to 0.32mm long by 0.14 to 0.31mm wide, located at level of acetabulum. Cirrus sac claviform, 0.14 to 0.26mm long by 0.06 to O.OSmm wide, located almost entirely anterior to acetabulum. Ovary smooth is distinctly lobed, oval to round, 0.23 to 0.39mm long by 0.29 to 0.41mm wide. Vittel- laria extend along lateral margins of body, 0.76 to 1.41mm long. Uterine coils fill posterior portion of body, extending anteriorly to median genital which is located at level of pharynx. Eggs oval, operculate, 0.29 to 0.32mm long by O.Old to 0.022mm wide; contain miricidia. USNM Nos. 72694, 72695. Host records: Host Locality Record Junco hvmenalis Indiana Martin and Gee (1949) Melospiza georgiana. Georgia fldelospiza meloaia.~ Georgia Pipilo ervthropthalmus. Virginia, N. Carolina, Ga. Zonotrichia albicollis. Texas Zonotrichia leucophrvs Georgia Denton and Byrd (1951), ^ x Zonotrichia albicollis Manitoba, Hedasi (1963) Can. Molothrus a. ater Ohio Cooper, et al. (1973) Agelaius phoeniceus. Ohio bvanocitta cristata Ohio Present Study

Discussion: Records of Z. alvevi are relatively rare and are restricted to passerine birds. This study records Z. alvevi from the Blue Jay and the Red—winged Blackbird for the first time. A very similar species, Zonorchis petiolatum (Railliet, 1900) was first reported from North American hosts by Denton and Byrd (1951) and by Boyd, et al. (1956). The original description of Z. alvevi was based on the study of 14 specimens from Slate-colored Juneos (Junco hvemalis). Denton and Byrd (1951) contributed further refinements to the descrip­ tion* The description of Z. petiolatum provided 46 by Denton and Byrd (1951) for their specimens closely approximates that of Zm alvevi* Zonorehis petiolatum is widely reported from avian hosts in Europe* Macko (1965) examined 35 specimens of Z. petiolatum taken from the gall bladders of 6 Jays (Garrulus glandarius)• Macko studied the morphological variability between individual specimens taken from the same host and between specimens taken from different hosts of the same species* Comparison of body shapes and organ measurements revealed that the descriptions of nine species of trematodes from four different genera fell within the range of limits of the 35 study specimens* Macko did not include Z. alvevi in his comparative study* This author finds that the measurements of Z* alvevi are equally compar­ able and similar* Mettrick (1963) also conducted a study revealing the highly variable morphology of Z* petiolatum* The study of Macko (1965) and the addition of Z* alvevi to his comparison is not sufficient grounds to synonomize this North American form with the European one* There exists sufficient geographic isolation between North American and European passerine bird populations to warrant the maintenance of Z* alvevi as the specific 4 7 appellation for the North American form of this trematode. Life cycle: Aspects of the developmental cycle of Zonorchis alvevi are unknown. Timon—David (i9 6 0 ) studied the developmental cycle of £• petiolatum experimentally. The eggs of this helminth are passed in the feces of the passerine bird host and are eaten by a land snail (Helicopsis arenosa)• Two generations of sporocysts develop in the digestive gland of the snail. Daughter sporocysts migrate into the mantle cavity of the snail host. Each daughter sporocyst contains from 6 to £ cercariae. These sporocysts migrate from the snail into the slime trail during periods of high humidity. The encysted cercariae are ingested by terrestrial isopods. Two species of isopods, Armadillidium vulgare and Armadillo officinalis, supported the further development of this helminth. Meta— cercariae in the hemocoel of isopods become in­ fective approximately 2 months after ingestion by the isopod. When infected isopods were fed to a sparrow, Passer montanus. immature flukes were found in the bird's gall bladder 66 hours post infection. 4 # Prevalence and Intensity of Zonorchis alvevi in Passerine Birds from South Bass Island: No* Range Infected Birds^o* Exam. Birds of Mean Adults Juveniles Exam— Preva—Inten-Inten— Infect—/No* Infect-/No* Host ined lence sitv sity ed Exam* ed Exam* Aeelaius 50 2 1 1 (22) 9 (26) pnoeniceus Cyanocitta 50 10 2—4 3«0 5 ( 50) 0 (0) cristata

Tanaisia zarudnvi. (Skrjabin, 1924) Byrd and Denton, 1950 Synonyms: Tamerlania zarudnvi Skrjabin, 1924 Tamerlania meruli Nezlobinsky, 1926 Tamerlania panuri Nezlobinsky, 1926 Tamerlania .iaponica Yamaguti, 1935 Tamerlania melospizae Penner, 1939 Tamerlania gallica Dollfus, 1946 Site of Infection: Ureters and renal calyces Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Eucotylidae Skrjabin, 1924 Subfamily Tanaisiinae Freitas, 1951 Genus Tanaisia Skrjabin, 1924 Specific diagnosis (modified after Kingston, 1965)s Body elongate, flattened, 1*75 to 3*94ram long by 0*275 to 0*763mm wide; cuticle with flattened scales measuring 0*015 to 0.027mm long by 0.005 to O.OO&nm wide, disposed in oblique files on ventral and lateral surfaces. Oral sucker anterior, subterminal, 0.162 to 0.297mm long by 0.155 to 0.297inm wide, opens into pyriforai, muscular pharynx, 0.067 to 0.106mm long by 0.60 to 0.105mm wide. Acetabulum is obscure, 0.03# to O.OdOmm long by 0.035 to 0.102mm wide. The pharynx opens directly into the intestine or into an esophagus 0.025mm long. The esophagus bifur­ cates into paired caeca which extend down the lateral margins of the body for a distance of 1.43mm to 2.69mm from the anterior end, where they turn inward and fuse. Testes paired, 0.107 to 0.346mm long by 0.53 to 0.169mm wide, borders smooth to lobed, opposite, located posterior to ovary. Ovary round to oval, with smooth to deeply indented margins, 0.053 to 0.346mm by 0.093 to 0.2l6mm, generally displaced to right of midline. Cirrus pouch transversely elongate, 0.051 to 0.112mm long by 0.051 to 0.134mm wide, located at midline of ovary. Uterus fills posterior portion of body with coils leading to female genital pore next to the cirrus pouch. Vittellaria paired, follicular, extracaecal, ex­ tending 0.65 to 0.69mm, from anterior margin of 50 the testicular zone toward anterior end of body* Eggs are operculate, 0*03# to 0.047mm long by 0.020 to 0.023mm wide. USNM No. 72693* Host records: Host Locality Record Melospiza lincolni Minnesota Penner (1939) Cyanocitta cristata. Hvlocichla guttata Parula americanus. Pipilo ervthropHthalmus Georgia, N. Carolina Zonotrichia albicollis Texas. Virginia Byrd and Denton (195°) ^ x Melospiza melodia Iowa Ellis C1963) 6'onasa umbellus Michigan, Kingston (1965) Ontario; Canada Toxostoma rufum Iowa Peet and Ulmer (1970) Cyanocitta cristata* Molothrusater Ohio Present study

Discussion: North American records of this helminth are relatively rare. Records of this helminth from the palaearctic region, especially the U.S. S.R. are more numerous. This study records £.* zarudnvi from the Brown—headed Cowbird for the first time. Byrd and Denton (1950) reported Tanaisia bragai (Santos, 1934) from the Common Grackle in Georgia. Passerine birds can serve as hosts to all four of the widely acknowledged species of the genus.T. zarudnvi. however, Is the species most often reported from passerine birds. Life cycle: The developmental cycle of T. zarudnvi was observed experimentally by Kingston (1965)* The eggs presumably pass out of the body of the host with the kidney excretions* The eggs remain viable for extended periods, over 3 years, and hatch only in the gut of terrestrial and semi- aquatic gastropods* The miracidium penetrated the’ gut wall and developed to daughter sporocysts and unencysted cercariae in the digestive gland of two species of snails (Anguisnira altemata and Succinea ovalis)* Development prior to this point was not observed* Fully developed cercariae encyst within the daughter sporocyst* The earli­ est encysted raetacercariae were seen about 45 days after infection* Encysted metacercariae from A* altemata were infective to Ruffed Grouse (Bonasa umbellus) when fed to the definitive host orally* Immature flukes reached the ureters of the host 6 hours after infection* The life cycle of T* bragai. a parasite of pigeons and land snails, in Puerto Rico was ex­ perimentally observed by Maldonado (1945) • The details of the developmental cycle of T* bragai and T* zarudnvi are quite similar* These organ­ isms differ only in respect to the time required to reach their respective developmental stages* 52 Prevalence and Intensity of Tanaisia zarudnvi in Passerine Birds from South Bass Island: No* Range Infected Birds/Mo* Exam* Birds of Mean Adults Juveniles Exam-Preva— Inten—Inten- Infect—/No# Infect—/No# Host ined lence sitv sity ed Exam* ed Exam# Cyanocitta 50 2 15 15 1 (50) 0 (0) cristata Molothrus 50 2 4 4 1 (46) 0 (4) ater

Prosthogonimus macrorchis Macy, 1934 Site of Infection: Cloaca Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Prosthogonimidae Nicoll, 1924 Subfamily Prosthogoniminae Luhe, 1399 Genus Prosthogonimus Luhe, 1399 Specific diagnosis: Body pyriform, flattened, 3#97mm long by 2#24aim wide; cuticle entirely aimed# Oral sucker subterminal, 0#17mm in diameter; opens into subglobular pharynx, 0«13mm in diameter# Acetabulum relatively small, 0#35mm in diameter, situated just within anterior 1/2 of body# Eso­ phagus present# Ceca filled with black digested material, extending to a point approximately 1/3 of body length from posterior tip of body# 53 Testes opposite, 0#59mm long by 0*39 to 0#46mm wide, located in midportion of body# Cirrus sac elongate, 0#43mm long by 0#09mm wide, located lateral at level of oral sucker, pharynx, and esophagus# Ovary transversely oval, 0#37mm long by 0#6lmm wide, consists of 10-14 large follicles# Uterus fills posterior 1/2 of body with coil ex­ tending anterior to lateral genital pore at level of pharynx* Eggs 0#015mm long by 0#006mm wide# Vittellaria lateral, composed of small follicles extending for 1.67mm in the center 1/2 of the body# Discussion: A large number of species of the genus Prosthogonimus have been described, largely from anseriform and galliform birds# Each working in­ dependently, two workers, Krasnolobova (1970) in Russia and Boddeke (I960) in the Netherlands have established the existence of 2 valid species in Eurasia, Prosthogonimus cuneatus (Rudolph!, lf$09) and £• ovatus (Rudolphi, 1&03) • The vast majority of the other described species have been placed in synonomy with one or the other of these two species# Both workers stated that the validity of P. macrorchis is uncertain# It may be a synonym of P# cuneatus. Krasnolobova (196S) stated that variations in development and relative size of some diagnostic characters in the genus depend upon the host species and tissue site of infec­ tion* In the U*S*f Macy (1950) has noted a number of developmental anomalies in specimens of P* macrorchis* Anomalies included examples of monorchisn, a specimen with three testesf and a specimen with rudimentary vitelline glands* P* macrorchis most frequently infects the bursa Fabricius of immature birds and the oviducts of mature female birds* Macy reported that infection of the oviduct resulted in curtailment of egg production in domestic turkeys (1939) and pheasants (1940)* This study recorded P* macrorchis from the Common Crackle for the first time* Life cycles Details of the developmental cycle of P* macrorchis were observed experimentally by Macy (1934)* This work has been summarized by Schell (1970)* Eggs containing miricidia are passed in the feces of the host* When eaten by the aquatic snailf Amnicola limosa. miricidia hatch in the snail intestine* Cercariae develop in sporocysts* Upon escape from the snail9 cercariae are drawn into the brachial chamber of dragonfly nymphs of the genera Epicordulia. Leucorrhinia* Mesothenis, and Tetragoneuria. Cercariae penetrate the hemo— coel of the nymphs and encyst in the abdominal musculature. Definitive hosts become infected upon ingestion of infected nymphs or adult dragonflys. Prevalence and Intensity of Prosthogonimus macrorchis in Passerine Birds from South Bass Island: One specimen was removed from one juvenile Common Grackle of the 37 juvenile Crackles examined during this study.

Echinuscodendium echinus (McIntosh, 1936) Synonym: Gyrabascus echinus McIntosh, 1936 Site of Infection: Herniations in Wall of lower intestine Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Lecithodendriidae Odhner, 1910 Subfamily Gyrabascinae Macy, 1936 Genus Echinuscodendrium Yamaguti, 195^ Specific diagnosis (modified after Macy, 1936): Body pyriform to oval, with ventral side somewhat flattened, 1 to 1 .23mm long by 0.325 to 0.960mm wide; cuticle covered with spines. Suckers nearly equal, 0.135mm in diameter, oral sucker slightly subterminal, sucker equatorial. Pharynx spherical 0*070mra in diameter; esophagus long, the crura branching anterior to acetabulum; caeca broad, sacculate extending to level of the testes* Testes nearly oval, dorsal and opposite; 0*200mm long by 0*135mm wide* Genital pore medioventral and posterior to acetabulum near posterior border of testicular zone* Ovary with 3 to 4 lobes at acetabular level* Vittellaria consisting of a broad band of follicles surrounding esophagus* Uterus consisting of dense transverse coils, con­ fined to posterior half of body* Anterior coil of uterus terminates in a short metraterm* Eggs 0*27mm by 0*020mm, ending basally in a short spine- like process* Host records: Host Locality Record Icteria virens Washington, D.C* McIntosh (1936) Cyanocitta cristata Ohio Present Study

Discussion: The trematodes occurred in pairs in herniated pockets in the wall of the lower in­ testine of the bird* The only other record of the occurrence of this parasite is the original description* This record confirms the unusual tissue site of infection. This study records E. echinus from the Blue Jay for the first time* 57 Life cycle: Other aspects of the biology of the E* echinus are unknown. Prevalence and Intensity of Echinuscodendrium echinus in Passerine Birds from South Bass Island: Four specimens were removed from a single Blue Jay of the 50 adult Blue Jays examined ‘during this study.

Omithodendrium imanensis Oshmarin and Dosenko, 1951 Synonym: Parabascus imanensis Baer, 1961 Site of Infection: Cloaca, lower intestine and bursa Fabricius Classification: Order Plagiorchiida La Rue, 1957 Superfamily Plagiorchioidea Dollfus, 1930 Family Lecithodendriidae Odhner, 1910 Subfamily Phaneropsolinae Mehra, 1935 Genus Omithodendrium Oshmarin and Dosenko, 1951 Specific diagnosis (highly modified after Oshmarin and Dosenko, 1951) • Body pyriform, 1.32 to 2.30mm long by 0.7^0 to 0*915mra wide; cuticle armed. Oral sucker subterminal, 0.25 to 0.42mm in diameter; pharynx, 0.13 to O.lSmm in diameter. Esophagus absent. Acetabulum pre-equatorial, 0.23 to 0.31mm in diameter. Initial compartments of ceca oval, distinctive * stomach—shaped* widenings located dorsally and to the sides of the body 5* mid—line* Cecal inflations distinctly set apart, with thick glandular walls. In young specimens, widenings less obvious, located at level of acetabulum, appear as inflated cecal zone. Cecae terminate 0.25 to 0.40mm from posterior tip of body* Testes oval, opposite, 0.20 to 0.45mm long by 0.25 to 0.7&mra wide. Cirrus sac elongate, 0.16 to 0.28mm long by 0.10mm wide. Ovary round, 0.14 to 0.21mm in diameter, located dorsal to testes and at level of acetabulum. Vittellaria fill anterior portion of body in space anterior to ovary. Uterus fills posterior portion of body, terminating near genital pore immediately an­ terior to acetabulum. Eggs operculate, 0.31mm long by 0.014mm wide. USNM Nos. 72630, 72634* 72635* Host records: Host Locality Record Coryus corone. Gallus alius. §riolus sinensis Primorsk Prov.. Oshmarin and USSR Dosenko (1951) Rinaria rioaria Moldavian. Schmilo and USSR Tixon (1971) Melaneroes erythrocenhalus Ohio Cooper (1974) Cyanocitta cristata. (jmiscalus qulscala. Sturnus vulgaris Ohio Present Study 59 Discussion: This is the first specific description to be prepared in a language other than Russian. 0. imanensis was described from three species of avian host collected in Far Eastern Siberia. Prior to its description, Oshmarin C1950) examined several hundred specimens from the Black Crow (Corvus corone) and reported considerable varia­ tion in body size. Trematodes varied from 0.9 5 5mm to 2.30mm in length. In most cases, the trema­ todes had strongly developed sexual organs and fully developed uteri completely filled with eggs. The small flukes were young but had fully develop­ ed genital organs. The trematode infecting Blue Jays and Starlings collected during this study revealed similar size variation and development. Oshmarin hypothesized that such accelerated development of the genital system may explain the origin of the phenomenon of progenesis in some trematodes in which metacercariae develop to sexual maturity. The hypothesis is untested. Ellis (196 3) reported Ornthithodendrium sp. from a single Kingbird (Tyrannus tyrannus) collected in Iowa. This was the first report of the genus in North America. Cooper C197M-) reported 0. imanensis, sensu strictu, for the first time from a North American host. This study records 0. imanensis from the Blue Jay, Common Grackle, and Starling for the first time. Life cycle: Other aspects of the biology of 0^ imanensis are unknown. Prevalence and Intensity of Omithodendrium imanensis in Passerine Birds from South Bass Island:

No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten- Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam.

Cyanocitta 50 26 1-18 10.4 13 (50) 0 (0) cnstata Quiscalus 50 4 1-3 2.0 0 (13) 2 (37) quiscula Sturnus 50 8 3-161 80.5 0 (1) 4 (49) vulgaris PLATE I

Omithodendrium imanensis

61 CM CO

PLATE 1 uj

uj S O Cestoda Hymenolepis farciminosa (Goeze, 1782) Railliet, 1899 Synonyms: Taenia farciminosa Goeze, 17 82 Piplacanthus farciminosa (Goeze, 1782) Weinlandia farciminosa (Goeze, 1782) : Wardium variable Mayhew, 1925 Variolepis variable (Mayhew, 1925) Hymenolepis variablis (Mayhew, 1925) Variolepis farciminosa (Goeze, 1782) Site of Infection: Intestine Classification: Order Cyclophyllidea Braun, 1900 Superfamily Hymenolepidoidea Spasski, 1960 Family Hymenolepididae Fuhrmann, 1907 Subfamily Hymenolepidinae Perrier, 1897 Genus Hymenolepis Weinland, 185 8 Specific diagnosis (modified after Mettrick, 1958): Long, elongate cestodes, maximum length of 82mm, maximum width of l-2mm. Scolex 0.18 to 0.2 5mm in diameter. Rostellum armed with a single row of ten hooks, 0.020 to 0.02411011 long. Diameter of four suckers is 0.08 5 to 0.090mm. Neck, 0.3mm long by 0.085mm wide. Excretory system follows the usual pattern. Dorsal vessel overlies the ventral which is 0.03mm in diameter. Genital pores unilateral, opening marginally on the left 6*f of the strobila in the middle of each proglottid. Arrangement of the genitalia follows the usual hymenolepid pattern. Testes are 0.09 to 1 mm in diameter. Cirrus sac is 0.20mm by 0.04-5mm, in­ ternal vesicula comparatively small. Seminal receptacle quite well developed, completely covers the external seminal vesicle. Large ovary lobed, follicular and ventral in position, extending beyond the lateral margins of the testes when fully developed. The compact vitelline gland is 0.09mm by 0.1mm, lies between the two posterior testes. Embryos are 0.048mm by 0.03mm, surrounded by an outer envelope 0.06mm by 0.065mm. Embryonic hooks are 0.02mm long. Two aporal testes lie slightly obliquely to each other. External seminal vesicle small, covered by the seminal receptacle. North American host records: Host Locality Record Sturnus vulgaris USA Salmon Cl696) Corvus brachyrhynchos Illinois Mayhew C1925) Corvus brachvrhvncnos' Washington, D.C. Jones (1934) Sturnus vulgaris Illinois Sommer (1936) Sturnus vulgaris USA Boyd (1951) Corvus brachyrhynchos Virginia Daly (1959) Sturnus vulgaris Colorado Ballard and Olsen (1966) Turdus migratorius Colorado Slater (1967) Pica pica hudsonia Utah Todd, et al. (1967^5 Corvus brachyrhynchos Ohio Jones (1968) Sturnus vulgaris Newfoundland, Threlfall (1968) Can. 65 Host Locality Record Corvus brachyrhynchos N. Carolina Hendricks, et^ al. C1969) Pica pica hudsonia Colorado Wacha and Schmidt C1971) Sturnus vulgaris California Vincent (197 2) Quiscalus quiscula Ohio Buck et al.■ (19757 ~ Agelaius phoeniceus, Cyanocitta cristata, Quiscalus quiscula, Sturnus vulgaris, Turdus migratonus Ohio Present Study

Discussion: Records of the occurrence of H. farcimi­ nosa indicate a holarctic geographical distri­ bution. This observation was emphasized by Spasski and Spasskaya (1956). Passerine birds of the families Corvidae and Sturnidae are most frequently infected in both Europe and North Ameri.ca,.. Hym,enolepis farciminosa is reported from the Red-winged Blackbird, Blue Jay and Common Grackle the first time during this study. Life cycle: Aspects of the developmental cycle of H . farciminosa were studied by Jones (1934) in the United States, Dutt and Mehra (1962) in Indip., and Mourad (1967) in Czechoslovakia. Eggs are passed in the feces of the avian host. Eggs are subsequently ingested by grasshoppers (Acrotylus humbertianus, Acrida exalta, Oedaleus abruptus, Crotogonus sp. and Aiolopus sp. in India; Melanop- lus bivittatus in the USA) and beetles ' 66 CPterostichus madidus and hortensis in Czechoslovakia; Aphodius granarius in the USA). Cysticercoids were recovered from all inter­ mediate hosts 7-14 days post infection. Defini­ tive hosts become infected upon ingestion of ! infected intermediate hosts. Spasski C1965) stated the opinion that Dutt and Mehra were not working with Hymenolepis farcimi- nosa. Spasski argued that Dutt and Mehra were working with eggs from Passerilepis crenata (Goeze, 1782). This opinion is based on the observation that H. farciminosa has not been reported in the tropical and steppe zones in i Asia. However, the work of Mourad (1967) support­ ed that of Dutt and Mehra. Prevalence and Intensity of Hymenolepis farciminosa in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam-Preva- Inten- Inten-Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exa Agelaius 50 2 3 - 1 (22) 0 (28) phoeniceus Cyanocitta 50 16 1-35 7.0 8 (50) 2 - (0) cristata Quiscalus 50 8 1— 3 2.0 1 (13) 3 C37) quiscula Sturnus 50 38 1-15 5.0 1 Cl) 18 (49 ) vulgaris Turdus 50 10 1-2 1.2 1 (19) 4 (31) migratorius Dilepis undula (Schrank, 1788) Weinland, 1834 Synonyms: Taenia angulata Rudolphi, 1810 Taenia angulata Dujardin, 1845 Hymenolepis pyramidalis (Sinitzin, 1896) Dilepis angulata (Dujardin, 1845) 1 Hymenolepis undulata Parona, 1899 Dilepis undulata Volz, 1900 Drepanidotaenia undula CSchrank, 1788) Southwellia ransomi Chapin, 1926 Dilepis turdi CYamaguti, 1935) Erschovilepis vulpis Spasski, 1965 Site of Infection; Intestine Classification: Order Cyclophyllidea Braun, 19 0 0 Superfamily Dilepiodoidea Matevosyah, 1962 Family Dilepididae Fuhrmann, 1907 Subfamily Dilepidinae Fuhrmann, 19 07 Genus Dilepis Weinland, 1834 Specific diagnosis (modified after Mettrick, 1958): Medium sized worms varying considerably in size, with a maximum length of 70mm, maximum width 3.5mm. Well developed scolex, diameter of 0.45- 0.93mm, bears rostellum armed with 48-64 hooks arranged in a double row. There is no constant size or number of hooks associated with any particular host. Size of the hooks in the first 68 row varies from 0.091-0.116mm, and those in the second from 0.70-0,088mm. The neck is 0.55- 0.75mm long, merges into a fairly long immature region with evident segmentation. Excretory system consists of two pairs of lateral excretory canals, the dorsal pair lying slightly inside the ventral pair. The dorsal vessels are 0.008-0.014mm in diameter, and the ventral ones 0.28-0.042mm. The transverse vessel between the ventral pair in each segment is 0.Ql-0.0 2mm in diameter. The genital pores are unilateral, opening on the right side of the strobila, situated marginally in the anterior half of each segment. In a mature proglottid, 2 8-36 testes situated behind the vitelline gland and the ovary. Testes, 0.048- 0.063mm wide by 0.06-0.07mm long. Cirrus sac long and narrow, 0.28-0.42mm by 0.0 32mm in diameter. Cirrus is armed with tiny spines, diameter 0.012- 0.014mm. The vagina passes inwards parallel with and either antero-dorsal or postero-dorsal to the cirrus sac. The ovary of two scattered groups of follicles lying anterior and ventral to the vitelline gland. Follicles round, 0.028-0.04mm in diameter, vary in number from 7-12 in the right group, and from 12-24 in the left group. Vitel­ line gland compact, slightly lobed organ situated 69 in the raid-line and in the centre of each proglottid. It is 0.04-0.07mm long by 0.097- 0.17mm broad. Mehlis* gland lies dorsal to the other female organs, and is 0.08 by 0.03mm broad. Uterus sac-like extends beyond the lateral ex­ cretory vessels when it becomes distended with eggs. The embryos are 0.036-0.04mm in diameter, surrounded by an envelope 0.004-0.048mm wide by 0.048-0.056mm long. The embryonic hooks are 0.018 -Q.02mm long. USNM No. 72 711.

North American host records: Host Locality Record Turdus migratorius Washington, D . C. Chapin (19 26) Turdus migratorius New York Webster (1943) Turdus migratorius Illinois Ogren (1958) Turdus migratorius Colorado Slater (1967) Pica pica hudsonia Montana Todd and Worley (1967) Turdus migratorius Newfoundland, Threlfall Can. (1968) Melanerpes erthrocephalus Ohio Cooper (1974) Turdus migratorius Ohio Present Study

Discussion: Records of the occurrence of D. undula indicate a geographical distribution limited primarily to passerine birds in Europe and North America. By far, members of the family Turdidae are the most frequently infected. In Europe, small mammals are infrequently infected with this helminth (Rysavy, 1968). In North America, in­ fection is confined almost exclusively to Robins. 70 Life cycle: Embryonic development of D. undula within mature proglottids has been observed by Ogren .£1958, 1959a, 1959b, 1962). Ogren concluded that the embryonic development of D. undula was similar to that of other invertebrates. . Maturation of the oocyte including growth and meiotic prophase occurs in the ovary of each mature proglottid, penetration of the spermatazoon in the oviduct and completion of meiosis in the uterus. Cleavage is unequal. Embryonic hooks developed in cortical cells of the morula stage. Contractile parenchyma originated from embryonic mesenchyme. Mature eggs are apparently passed in the feces of the host. Harper (.193 3) reported cysticercoids in earthworms CAllolobophora longus, Lumbricus terrestris) from Scotland. Rysavy £196^, 197 3) reported D. undula cysticercoids from earthworms CEiseniella tetraedra, L. terrestris) in Czechoslovakia. Definitive hosts become infected upon ingestion of earthworms containing mature cysticercoids (Rysavy,.1968). Prevalence and Intensity of Dilepis undula in Passerine Birds from South Bass Island: 71 Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten-Inten-Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Turdus 50 »+2 1-8M- 15.3 8 (19) 13 (31) migratorius

Choanotaenia iola Lincicome, 19 39 Site of infection: Intestine Classification: Order Cyclophyllidea Braun, 1900 Superfamily Dilepidoidea Matevosyan, 1962 Family Dilepididae Fuhrmann, 1907 Subfamily Dipylidiinae Stiles, 1896 Genus Choanotaenia Railliet, 1896 Specific diagnosis (modified after Lincicome, 1939): Small cestodes, average length about 2.M-mm, maximum width 0.5 57mm, anterior proglottids broader than long, with average maximum wide at anterior end, O.M-2Mmm. Approximately 21 segments present in strobila, with one or two gravid. Mature segments about as long as broad, gravid ones longer than wide. Proglottids telescoped, overlapping. Genital pores irregularly alterna­ ting, occasional strobila showing regularly alternating pores. Scolex prominent, average width 0.356mm with four cup-shaped suckers. Average width of suckers 0.114mm with inner surface armed throughout with cuticular spines 0.003-5mm long. Rostellum bears crown of 17-20 hooks. Average width of crown and body of rostellum 0.068 and 0.06 5mm, respectively. Rost'ellar hooks approximately 0.032mm long. Rostellar sac 0.202mm long. Vagina in ventral position along anterior half of proglottid margin, and dilated at distal end near center of proglottid forming seminal receptacle. Ovary 0.036mm in diameter arises dorsally receiv­ ing first the vitelline duct, secondly, the spermatic duct from seminal receptacle. Vitelline gland lobated, pyramidal in outline, apex directed posteriorly, greatest width 0.063mm at base. Union of oviduct and spermatic duct forms ootype into which shell gland empties. Uterus, a laterally branching organ, arises on poral and antiporal sides of ootype at approximately the level of shell gland. Uterus ultimately breaks up into individual egg capsules. Testes 13-17 in number posterior and lateral to female complex, not confined to single plane, but scattered throughout depths of proglottid. Average diameter of testes 0.050mm. Vas defereus 73 coiled before entering cirrus pouch, Cirrus heavily armed minute spines. Host records: Host Locality Record Turdus migratorius Illinois Lincicome (1939) Agelai'us phoeniceus Colorado Wallace and Olsen (1966) Turdus migratorius Colorado Slater (1967) Quiscalus quiscula Ohio Buck, et al.(1975) Cyanocxtta crxstata, Sturnus vulgaris, Turdus"~mxgratorxus Ohio Present Study

Discussion: Records of the occurrence of C. iola indicate its geographical distribution is re­ stricted to the mid-western United States. Future investigations of helminth parasitism in passerine birds should enlarge both the host range and geographical limits of this helminth. Choanotaenia iola is reported from the Blue Jay and Starling for the first time during this study. Life cycle: Aspects of the life cycle and development of C. iola are unknown. Morphologically, <2. iola differs only slightly from C. infundibulum (Bloch, 1779), a cosmopolitan parasite of galliform birds. The principal difference between the two species is the presence of minute spines on the walls of the suckers of iola. Further investigations may reveal that C. iola is the form that C. in­ fundibulum takes when this species infects 74 passeriform birds. Such an event would not be unprecedented, a number of helminth species are known which infect both galliform and passiform birds. Enigk and Sticinsky (1959) list Musca i domestica, 2 spp. of Locustidae, and 92 spp. of beetles reported from various countries as being suitable intermediate hosts of C. infundibulum. In addition, these workers observed successful experimental infections in a number of species of beetles, flies, ants, Orthoptera, Collembola, Myriapoda, Acarina and Crustacea. Dutt and Sinha (19 61) successfully infected 15 spp. of beetles and 15 spp. of grasshoppers under experimental conditions in India. A wide range of potential intermediate hosts may exist for C. iola as well, assuming it is a species distinct from C. in- fundibulum. Prevalence and Intensity of Choanotoenia iola in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam-Preva- Inten-Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exa Cyanocitta 50 6 1-4 2.0 3 (50) 0 (0) cristata Sturnus 50 42 1-14 4.3 0 (1) 21 (49) vulgaris Turdus 50 10 1-7 2.6 0 (19) 5 (31) migratorius 75 Choanotaenia musculosa (Fuhrmann, 1896) Fuhrmann, 1899 Synonyms: Monpylidium musculosum Fuhrmann, 1876 Anomotaenia musculosa (Fuhrmann, 1896) Site of infection: Intestine I Classification: ' Order Cyclophyllidea Braun, 1900 Superfamily Dilepidoidea Matevosyan, 1962 Family Dilepididae Fuhrmann, 1907 Subfamily Dipylidiinae Stiles, 1896 Genus Choanotaenia Railliet, 1896 Specific diagnosis (modified after Joyeux and Baer, 1936): Small, fragile cestodes. Strobila 30 to 70mm long by 1 to 2mm wide. Scolex measures 0.260 to 0.270mm in diameter, the suckers 0.090mm in diameter, the rostellum 0.070 to 0.085mm in diameter. Rostellum armed with a double crown of 22 hooks, measuring 0 .02 8 and 0.02lhnm in length, respectively. Testes occupy the posterior portion of the prog­ lottid; testes number 21-38. Vas deferens is sinuous; lacks a seminal vesicle. Cirrus pouch measures 0.160 to 0.270mm long by 0.022 to 0.050mm in diameter. Cirrus is armed with very small spines. Female genital complex occupies the anterior portion of the mature- proglottid. Vagina is very muscular and measures 0.270 to 0.2 80mm 76 long* Seminal receptacle measures 0.170mm in diameter. Uterus branches laterally and breaks into capsules containing a single egg when ripe. Egg capsules measure O.OSOmm in diameter; egg proper is 0.050mm in diameter; embryo is 0.30ram in diameter. Embryonic hooks measure O.Ol&mm long. North American host records; Host Locality Record Stumus vulgaris USA Ransom (1909) Stumus vulgaris USA Boyd (1951) kturnus vulgaris Colorado Ballard and Olsen (1966) Sturnus vulgaris Newfoundland 9 Threlfall (1963) Can. Stumus vulgaris Kansas Rodrick and Johnson (1971) Quiscalus quiscula Ohio Buck, et ali (19751 Agelaius phoeniceus. Ohio fouiscaTus quiscula. Ohio Sturnus vulgaris Ohio Present Study

Discussion: Records of the occurrence of this hel­ minth indicate its geographical distribution is limited to Western Europe and North America. The principal host in both regions is the Starling. Other passerine birds are infected infrequently. Choanotaenia musculosa is reported from the Red­ winged Blackbird and Common Grackle for the first time during this study. Life cycle: Aspects of the life cycle and development of C. musculosa are unknown. Utilization of one or more species of Insecta as intermediate hosts seems probable. Prevalence and Intensity of Choanotaenia musculosa in Passerine Birds from South Bass Island: No. Range Intected Birds/No. Exam* Birds of Mean Adults Juveniles Exam— Preva— Inten- Inten-Infect-/No. Infect-/No. Host ined lence sity sitv ed Exam. ed Exam. Aaelaius 50 6 1-9 4.7 1 (22) 2 (2d) onoeniceus Quiscalus 50 10 1-3 2.2 3 (13) 2 (37) auiscula Sturnus 50 12 3-10 6.7 0 (1) 6 (49) vulgaris

Anonchotaenia globata (von Linstow, 1&79) Synonyms: Taenia loxiae recurvirostrae Blumenback, 1779 Taenia clavata Marchi, 1B69 Taenia rudolphiana von Linstow, 1&79 Taenia breviceps von Linstow, 1&79 Anonchotaenia clava Cohn, 1900 Amurina inermis Fuhrmann, 1901 Amerina alaudae Cerutti, 1901 Amerina inermis Clerc, 1902 Site of infection: Intestine Classification: Order Cyclophyllidea Braun, 1900 Superfamily Paruterinoidea Matevosyan, 1962 73 Family Anonchotaeniidae Matevosyan, 1965 Subfamily Anonchotaeniinae Matevosyan, 1950 Genus Anonchotaenia Cohn, 1900 Specific diagnosis (modified after Mettrick, 1953): Small to medium sized cestodes, maximum length 66mm, maximum width l*0mm* Well developed scolex 0 *53-0 *62mm in diameter* No rostellum, scolex unarmed* Four suckers have a diameter of 0*20— 0*2Z»jnm. Neck 1*2—1*5mm by 0.25-O*26mm wide* Excretory system follows the usual pattern* Dorsal vessel is 0*006-0.003mm in diameter, ven­ tral 0.012-0*01ifinm* Genital pores alternate ir­ regularly, open marginally by a deep atrium in the middle of each segment. In mature proglottid there are 5 testes arranged in a transverse row 0*02S—0*035mm in diameter* Cirrus sac is clavi— form, 0•07-0*06imn long by 0.024-0*03mm wide* Vagina passes inwards parallel with the cirrus sac* Ovary is compact round organ 0*05-0.053mm in diameter, ventral in position in the posterior part of the segment* It lies between the vitel­ line gland, which is in the mid—line, and the genital atrium* Uterus is at first a sac—like organ which develops in front of and dorsal to the ovary* Eggs, when in the uterus, are 0*013— 0*017mm in diameter* No embryonic hooks* 79 North American host records: Host Locality Record Dendroica striata. Melospiza melodia Maryland Ransom (1909) Passer domesticus Quebec, Can. Rayner (1932) Passer domesticus Indiana Kintner (1938) 5ay o m i s phoebe Virginia Jones (1945) Melospiza georgiana. Ohio, Mich. Melospiza meloaia. Ohio Passercuius sandwichensis. Ohio Passerina cvanea. Michigan Poecetes g. gramineus. Ohio, Mich. RichmoncTea cardinal:! s. Ohio 5pizella arborea. Ohio SpizellsT passerina. Michigan gpizella pusilla. Ohio IZonotrTc'hia albicollis. Wisconsin Agelaius phoeniceus. Ohio, Wisconsin Molothrus ater. Ohio Dendroica striata. Wisconsin Dendroica coronata. Wisconsin Raush and Morgan (1947) Melazonle rieneri California Voge and Davis (1953) „ x Molothrus ater Manitoba, Can. Hodasi (1963) Agelaius phoeniceus Ohio Spory (19o5) Sturnus"vulgaris Kansas Rodrick and Johnson (1971) Molothrus ater Ohio Cooper, et al. (1973) Agelaius phoeniceus. Molothrus ater. Quiscaius quiscula Ohio Present Study

Discussion: Records of the occurrence of A* globata indicate a cosmopolitan geographical distribution in passerine birds. In North America, this hel­ minth is one of the two most frequently reported cestodes, along with Hymenolepis farciminosa. in­ fecting passerine birds. Anonchotaenia globata ■ is reported from the Common Grackle for the first time during this study. Life cycles Aspects of the life cycle of A. globata are not known* Life cycle information concerned with other members of this genus is likewise lacking* Aspects of the development of the eggs of A* globata have been observed in mature and gravid proglottids (Woodland, 1929)* The embryo first elongates and grows until it is about 0*llmm long* It then coils itself in the mid—region, the investing nuceleated membrane becomes detached from the coils and forms a loose sac around them* Finally the embryo is tightly coiled in this sac and measures 0*026 by O.OlBmm. The embryos pass into the paruterine organ which meanwhile has been i developing at the anterior end of the uterus. The two organs, i*e*, uterus and paruterine organ to­ gether form a rather ovoid structure lying in the middle of the segment, and slightly diagonal in position when the segments are well relaxed* The embryos in the paruterine organ are few, and are spindle shaped due to the two surrounding membranes which are prolonged into slender poles* The outer membrane is 0*030-0*035mm long (Woodland, 1929).

Prevalence and Intensity of Anonchotaenia globata in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam— Preva— Inten—Inten— Infect—/No. Infect—/No. Host ined lence sity sity ed Exam, ed Exam. Agelaius 50 22 1-11 4.0 3 (22) 3 (23) onoeniceus Molothrus 50 22 1-12 4.5 10 (46) 1 d ) ater Quiscalus 50 3 1-15 7.3 1 (13) 3 (37) auiscula

Orthoskr.iabinia rostellata (Rodgers, 1941) Spassky, 1947 Synonyms Anonchotaenia rostellata Rodgers, 1941 Classifications Order Cyclophyllidea Braun, 1900 Superfamily Paruterinoidea Matevosyan, 1962 Family Biuterinidae Matevosyan, 1965 Subfamily Orthoskrjabinae Matevosyan, 1965 Genus Orthoskr.iabinia Spassky, 1947 Specific diagnosis (modified after Rodgers, 1941)J Description. Strobila 400-700mm long by 0.7mm in maximum width. Scolex 0.523mm wide with a dis­ tinct, but minute, unarmed spheroidal rostellum 0.0375mm in diameter. Average diameter of suckers 0.0157mra between segments of entire strobila rather indistinct. Margins of strobila acrasped— ote; immature proglottids 3 times as wide as long. Mature proglottids 0.350 to 0.450ram wide, 6—7 times as wide as long. Ripe segments 82 slightly longer than vri.de 0*47 by 0.41mm. Strobila attains its maximum width at about two—thirds the length from the front end, becoming somewhat nar­ rower farther back. Gonads become mature at 85- i 100 segments from the neck. Two pairs of excretory canals, comprising the usual large straight ventral and minute sinuous dorsal ones, the former being connected at the back of each proglottid by a small transverse vessel. Vas deferens and vagina ventral to ex­ cretory canals. Vagina behind cirrus pouch. Testes usually 9 in number, 4 poral and 5 aporal; broadly oval, average 0.031 to 0.022rom in mature proglottid. Ovary ellipsoidal, somewhat larger than testes, centrally located in mature proglot­ tid in front of the smaller compact vitellarium. In partially gravid segments, ovary is still recognizable as a small spheroidal body postero— centrally located between the two principal arms of the uterus. Uterus, at first a small simple single sac, becomes transversely elongate, dis­ tinctly double when partially gravid; still later it becomes spheroidal, no longer double. Para­ uterine organ appears first as a small dense transversely striated anteromedian mass that gradually enlarges and develops an equatorial flange which spreads out in a vertical plane; the peripheral edges of the flange curve backward along the uterus. Posterior surface of the para­ uterine organ is closely applied to the anterior

i surfaces of the uterus. Eggs in the paruterine organ only in the hindmost segment of the strobila. Host records: Host Locality Record Richmondea cardinal!s Oklahoma Rodgers (1941) Fringilla montifringiila USSR Belopolskaya (1963) ^ Corvus brachvahvnchos Ohio Jones (1963) Cvanocitta cristata. Molothrus ater. Quiscalus quiscula Ohio Present Study

Discussion: Records of the occurrence of 0. rostellata indicate its geographical distribution is re­ stricted to the mid—western United States. The record of 0. rostellata from the USSR must be regarded as anomalous. Several cestodes of passerine birds exhibit a holarctic geographical distribution. Those that exhibit a holarctic distribution are always reported with greater frequency from the palaearctic region than from the nearctic region. The latter fact is due to the larger number of investigations conducted over a longer period of time in the palaearctic 64 zone# The report of Belopolskaya (1963) must be regarded as a misidentification for the present# Further investigations may eventually reveal 0# rostellata in a wider range of hosts and wider geographic distribution than presently known# Orthoskr.iabinia rostellata is reported from the Blue Jay, Brown-headed Cowbird and Common Grackle for the first time during this study. Life cycles Aspects of the life cycle and development of 0. rostellata is unknown. Details of the life cycle of other members of this genus are also unknown# Prevalence and Intensity of Ortho skr.i abinia rostellata in Passerine Birds from South Bass Island: No# Range Infected Birds/No# Exam# Birds of Mean Adults Juveniles Exam- Preva-Inten— Inten— Infect-/No# Infect-/No. Host ined lence sity sity ed Exam# ed Exam# Cvanocitta 50 6 1-4 2.5 4 (50) 0 (0) cristata Molothrus 50 4 2-6 4.0 2 (46) 0 (4) ater Quiscalus 50 4 1-3 2.0 0 (13) 2 (37) auiscula 65 Nematoda Svngamus trachea (Montague, l3ll) Chapin, 1923 Synonyms: Fasciola trachea Montagu, lSll Svngamus tracheal!s Siebold, 1&36 Strongvlus trachealis (Siebold, 1636) Sclerostoma tracheale (Siebold, 1336) Svngamus primitivus Molin, 1360 Svngamus nasicola von Linstow, 1699 Svngamus king! Leiper, 1913 Svngamus gracilis Chapin, 1925 Svngamus parvus Chapin, 1925 Svngamus merulae Baylis, 1926 Svngamus tenuispiculum Manter and Pinto, 1936 Svngamus skr.iabini Elperin, 1936 Svngamus anterogonimus Ryzhikov, 1949 Site of infection: Trachea Classification: Order Strongylata Superfamily Strongyloidea Weinland, IS56 Family Syngamidae Leiper, 1912 Cenus Svngamus Siebold, 1936 Specific diagnosis (modified after Cram, 1927): Cylin­ drical red worms* Head enlarged and truncated* Mouth orbicular, with a hemispherical capsule, at the base of which are usually 3, occasionally 9# 86

sharp teeth; the mouth is surrounded by a plate, the outer margin of which is incised to form 6 lobes opposite each other, with 4 smaller lobes between them in opposed pairs; a lateral papilla is between each pair of small lobes and 4 sub- median papillae are present. The male is per­ manently attached in copula to the female, forming a Y. Male: Length 2 to 6mm long by approximately 0.200 mm wide. The bursa is obliquely truncated, pro­ vided with rays, sometimes with asymmetrical dorsal rays. The male is permanently attached to the female about the vulva. Two equal, slender, short spicules, -0.057 to 0.064mm long, present. Female: Length 5 to 20mm long by 0.310 to 0.350mm wide, somewhat more slender anteriorly and ir­ regularly swollen when fully gravid. Conical tail bears a pointed process. Vulva 1/4 to 1/6 of body length from anterior end. Ellipsoidal eggs operculate, 0.085 to 0.090mm long by 0.050mm wide. USNM Nos. 72696, 72697, 72698. North American host records: Host Locality Record Nebraska Walker (1886) B U X U b f ivjeleagrls gallopavo USA Cram (1927b) Turdus migratorxus Nebraska Manter and Pinto (1928) 87

Host Locality Record Meleagris eyiHnpavo Washington, D*C* Wehr (1937b) Turdus migratorius Nebraska Ripple (1941) Turdus migratorius New York Webster (1943) Alectoris graeca. Lophortvx California California Herman (1945) dolinus virginianus Mississippi Ward (1945) Soiinus" virginianus Texas Webster and Addis (1945) Bonasa umbellus. dorvus brachyrhvnchos. Phasianus colchicusT" Quiscalus quiscula. Sturnella magna New York Goble and Kutz (1945b) Phasianus colchicus Conn* Anderson and Shapire (1955) Cvanocitta cristata Mass* Boyd, et al* (1956^---- Phasianus colchicus Canada McGregor, et al. ?19SlT Phasianus colchicus Canada McGregor (1963) ftica pica hudsonia Montana Todd and Worley (1967) Golinus virginianus Maryland Rose and Hwang (1967) Gorvus brachyrhvnchos Ohio Jones (1963) Cluiscalus quiscula Ohio Stanley and Rabalais (1971) Quiscalus quiscula Ohio Buck, et al* (19751 Agelaius phoeniceus. dvanocitta cristata. Molothrusater. Passer domestlcus. duiscalus quiscula. Turdus migratorius Ohio Present Study

Discussion: This species of helminth exhibits an almost cosmopolitan pattern of geographical distri­ bution* The majority of records indicate a holoarctic geographical distribution* The range • of definitive hosts is extremely large and com­ prises numerous species of wild birds of 10 orders* In North America, records from galliform and passeriform hosts predominate* This helminth is recorded from the Red**winged Blackbird and the Brown-headed Cowbird for the first time* In Europe, Lewis (1925* 1926), Morgan (1931), and Campbell (1935) have stated that Starlings serve as an important host for S* trachea* but this helminth has yet to be reported from naturally infected Starlings in North America* The absence of infection in North American Starlings is dif­ ficult to explain* Barus (1946b) stated that the wide range of hosts for this species of helminth is due to the stable environment, physiologically and biochemically, in which the nematode exists* He further stated that no physiological races of trachea exist* The original stock of Starlings introduced into North America may have been re­ sistant to infection* Life cycle: The development cycle of £>• trachea may be direct or may utilize invertebrate intermediate hosts* A large number of workers, including Ortlepp (1923), Clapham (1934, 1935) and Nolst (1971, 1973) in Europe and Walker (1SS6), Waite (1920), and Wehr (1937b) in the United States have studied the development of this helminth in earthworms (Eisenia foetida* Lumbricus terrestris)* Wehr (1937b) and Barus (1966b) have studied the overall development of the helminth* In addition, Clapham (1939)# Hwang (1961) and Barus (1964a) have, respectively, observed that Diptera, cock­ roaches, and fresh-water snails may successfully serve as intermediate hosts* Barus (1964b) observed that eggs are passed in the feces of the host* Third—stage infective larvae occur within the egg 40 days later* The larvae moult twice within the egg shell* When ingested by an intermediate host, the third—stage larvae penetrate the intestine and persist in the hemo— coel or ventral blood vessel* If ingested directly by an avian host, the larvae penetrate the intestinal wall, migrate in a blood vessel through the liver and heart to the lungs, pene­ trate the lung wall and accumulate in the alveoli where males and females become attached* Mated pairs subsequently migrate to the trachea (Wehr, 1937a). Barus (1966b) noted that adults persist for 92-126 days in galliform birds* Female worms produce $5$ of their eggs during the first 35 days of patency* Clapham (1939), Guilford and Herrick (1954) and Fernando, et al* (1971) have observed considerable pathogenesis and subsequent pneumonic infections induced by migrating Sm 90 -trachea larvae in the lungs of the avian host. Prevalence and Intensity of Svngamus trachea in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam—Preva- Inten—Inten— Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam.

Agelaius 50 4 1 — 1 (22) 1 (23) loemceus Cyanocitta 50 16 1-7 3.3 3 (50) 0 (0) cristata lolothrus 50 2 1 - 0 (46) 1 (4) ater Passer 30 6.7 3 3 0 (5) 2 (25) omesticus uiscalus 50 72 1-3 3.2 5 (13) 31 (37) uiscula Turdus 50 42 1-3 3.2 5 (19) 16 (31) m gratorius

Porrocaecum ensicaudatum (Zeder, 1300) Baylis, 1920 Synonyms: Fusaria ensicaudatum Zeder, 1300 Ascaris ensicaudata (Zeder, 1300) Spiroptera turdi Molin, .1360 Filaria turdi (Molin, 1360) Ascaris fissilabium von Linstow, 1373 Yiguiera turdi Seurat, 1913 Site of infection: Gizzard and upper intestine Classification: Order Ascaridida 91 Superfaraily Ascaridoidea Railliet and Henry, 1915 Family Ascarididae Blanchard, 1396 Genus Porrocaecum Railliet; and Henry, 1912 Specific diagnosis (modified after Cram, 1927)s Large worms; large labia, wider than long; each labia with 2 large, oval, closely set papillae* Inter- labia 3/5 the length of labia* Intestinal cecum small, almost rudimentary. Male: Length 23 to 32mm long by 0*92 to l*03mm wide* Esophagus 1/10 the length of body* Six pairs of post anal papillae, 5 of which are near the caudal extremity, and 1 pair of double papil­ lae near anus* Preanal papillae in a single row, numbering 13-19#' Female: Length 50 to 53mm long by 1*0 to l*3mm wide* Vulva somewhat anterior to middle of body

c# length in ratio of 4*5# Eggs 0.110 by 0.035mm wide, the outer shell with delicate lattice--work markings* North American host records: Host Locality Record Turdus migratorius N* Jersey Cram (1933) Turdus migratorxus Quebec, Can* Rayner (1932) jFurdus mi gratorlus New York Webster (1943) 3turnus vulgaris USA Boyd (1951) Corvus ~bracnyrhyn cho s * Molothrus ater. Oulscallus quiscula* Turdus migrafcorius Canada Mawson (1956a) Corvus brachvrnynchos Virginia Daly (1959) Host Locality Record Stumus vul garls Colorado Ballard and Olsen (1966) Turdus migratorius Colorado Slater (1967) Corvus brachyrhvncho s Ohio Jones (1965) Quiscalus aulscu-La Ohio Stanley and Rabalais (1971) Quiscalus ouiscula Ohio Buck, et al. (1975) Agelaius^phoeniceus. CyanocItTta cristata, uis c alusqui scula« §_tuivius~yulgarls, Turdus"migratorius Ohio Present Study

Discussion: Records indicate that the geographical distribution of P# ensicaudatum is holoarctic in passerine birds# Hartewich (1959) observed that this nematode is most frequently a parasite of birds of the family Turdidae in Europe# It is less frequent in Corvidae, Motacillidae and Stumidae# Perusal of North American records indicated that this observation is true for this region as well# The family Icteridae can be added as a frequent host in North America# This hel­ minth is recorded from the Blue Jay and Red-winged Blackbird for the first time during this study# Life cycle: Details of the developmental cycle of P# ensicaudatum were observed experimentally by Rysavy (1959) in Czechoslovakia and by Levin (1961) in the United States# Eggs are passed in the feces of the avian host# Development of first stage larvae within the egg requires 12—14 days# 93 When eaten by earthworms (Lumbricus terrestris), larvae hatch, penetrate the intestine and lodge in the ventral aorta. Infective third-stage larvae are found in the ventral aorta 7-10 days post infection (Rysavy, 1959)* Definitive hosts be­ come infective upon ingestion of infected earth­ worms* Levin (1961) stated that larvae are re­ leased from the earthworm in the proventricuius of the avian host* Larvae migrate and moult be­ neath the lining of the gizzard* Fourth-stage larvae enter the lumen of the intestine, penetrate and moult again in the intestinal mucosa* Fifth- stage worms subsequently re-enter the lumen of the i intestine and mature* All of the latter develop­ mental stages have been observed in naturally in­ fected birds during this study* Prevalence and Intensity of Porrocaecum ensicaudatum in Passerine Birds from South Bass Island: No* Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam—Preva— Inten- Inten— Infect-/No* Infect-/No* Host ined lence sitv si tv_____ed Exam* ed Exam* 50 4 2-6 4.0 0 (22) 2 (23) Dhoeniceus Cvanocitta 50 14 1-4 1.7 7 (50) 0 (0 ) cristata Quiscalus 50 46 1-10 2.3 3 (13) 20 (37) auiscula Sturaus 50 32 1-14 2.5 1 (1) 15 (49) vulgaris 50 34 1-3 2*2 3 (19) 14 (31) mxgratonus 9 4 Oxvspirura petrowi Skrjabin, 1929 Synonym: Oxvspirura lumsdeni Addison and Anderson,

1969 Site of infection: Orbit of eye Classification: Order Spiruida Diesing, 1661 Superfamily Spiruroidea Family Thelaziidae Genus Oxvspirura Drasche, in Stoss, 1697 Specific diagnosis (modified after Pence, 1972): Body slender, yellow to cream color, bluntly rounded anteriorly, sharply attenuated posteriorly* Cervical alae present* Cuticle transversely i striated* Mouth with four submedian pairs and three circumoral pairs of cephalic papillae* Amphids present* Cuticularized buccal capsule undivided* Esophagus not discemibly divided* Deirids lateral* Male: Length 6*27 to 6.63mm, maximum width 0*165 to 0*330mm* Esophagus 0*530 to 0*660mm long* Right spicule 0*121 to 0*320mm long and boat­ shaped* Left spicule slender with sharp tip, 0*264 to 0*517mm long* Tail sharp—pointed* Pre— anal papillae 4 to 6 in number, asymmetrically arranged* Postanal papillae 4 to 6 in number, asymmetrically arranged. 95 Females Length 7*70 to 12.35mm, maximum width 0*200 to 0*455mm* Buccal capsule 0*01d to 0*02&mn long and 0*015 to 0*026mm wide* Esophagus 0*550 to 0*730mm long. Vulva and anus in posterior quarter of body* Vulva 0*500 to 0*700mm from posterior end* Anus 0*242 to 0*400mm from posterior end. Eggs contain larvae, 0*035 to 0.044mm long by 0.015 to 0.031mm wide* North American host records: Host Locality Record Tympanuchus cupido Michigan Cram (1927a) Pnasianus colcnicus Nebraska McClure (1949) ^onasa umbelius Minnesota Erickson, et al* (1949) Ammospiza maritima N* Carolina Hunter and Quay (1953) Geothlypis trichas S* Carolina Wells and Hunter (I960) Bonasa umbellus. Ontario, Can* gentrocercus urophasianus Sask*, Can* Pedioecetes phaslanellus. S. D., USA; Sask*, Can* Tvmpanuchus pallidicinctus Oklahoma Addison and Anderson (1969) Agelaius phoeniceus. Anthus splnoietta* Bubulcus ibis, ButoriHes virescens, Cassidlx mexicanusT Iridoprocne bicolor, J^olotnrus ater. Mviarchus crinitus. Quiscalus quiscuTa. IgLchmondea cardinal!s« Sturnella magna* Turdus migratorius* Tvrannus tvrannus. Louisiana Pence (1972) gyanocitta cristata. Ivioiotnrusater Ohio Present Study ' 9 6 Discussion: Records of the occurrence of 0* petrowi indicate a holoarctic geographical distribution in galliform and passeriforra birds* This hel­ minth is recorded from the Blue Jay for the first time during this study* Pence (1972) noted that even in heavy infections with as many as 30 worms per eye, no gross or histopathology has been demonstrated as a result. Life cycle: The life cycle of only one species of the genus Oxvspirura has been observed experimentally namely 0* mansoni (Cobb, 1&79)• Fielding (1926, 1927) in Australia, Kobayashi (1927) in Formosa, Saunders (1929) in Florida, Hutson (1936) in Antigua and Schwabe (1951) in Hawaii found that a cockroach (Pvcnoscelus surinamensis) was a suitable host for the development of 0^ mansoni* Eggs are deposited in the orbits of the definitive host* The eggs pass down the naso—lacrymal ducts, are swallowed and passed in the feces of the avian host. The intermediate host is infected by eating the eggs* Infective third—stage larvae were removed from cockroaches 51 days post in­ fection* After ingestion of an infected roach, the larvae are released in the crop of the avian host, migrate up the esophagus, along the roof of the mouth and finally up the naso—lacrymal ducts 9 7 into the orbits* The life cycle of 0* petrowi probably parallels that of 0. mansoni* utilizing an orthropteran intermediate host* Prevalence and Intensity of Oxvspirura petrowi in Passerine Birds from South Bass Island: ' No* Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam- Preva- Inten- Inten—Infect—/No* Infect-/No* Host ined lence sitv sitv ed Exam* ed Exam* Cvanocltta 50 4 2—4 3*0 2 (50) 0 (0) cnstata Molothrus 50 2 1 1 1 (46) 0 (4) ater

Tetrameres americana Cram, 1927 Synonym: Tetrameres pattersoni Cram, 1933 Site of infection: Proventriculus Classification: Order Spirurida Diesing, 1861 Superfamily Spiruroidea Family Tetrameridae Travassos, 1914 Genus Tetrameres Travassos, 1915 Specific diagnosis (modified after Cram, 1927)s Mouth with 3 small labia* Slender muscular pharynx present* Female: Length 3•9mm long, by 3mm wide; body globular, blood—red in color, with 4 longitudinal furrowso Anterior end protrudes from globular 93 body for a length of 0.9mm; the protruding posterior part of body is 0.3mm long* Buccal capsule 0*03 5mm long by 0.010mm wide; pharynx 0.300mm long by 0.027mm wide; esophagus 1.4mm long by 0.040mm wide. Intestine sacculate, filled with black detritus. Anus 0.330mm from posterior end. Eggs 0.042 to 0.050mm long by 0.025mm wide, Host records: Host Locality Record Gallus gailas. USA Cram (1927a) Colinus virginianus USA Cram (1931) Colinus* virginianus. Meleagris gallopavo. Gallus gallus New Jersey Beaudette, et al. (1933) Richmondea cardinal!s Missouri Durant and Knight (1941) Columba livia California Raggi and Baker (1957) Colinus virginianus Florida Kellogg and Prestwood (1963) Columba livia Florida Flatt and Nelson (1969) Molothrus ater Ohio Present Study

Discussion: Most of the species of the genus Tetra— meres are parasites of aquatic and shore birds. Tetrameres americana is exceptional in that records indicate it is exclusively a parasite of upland birds. This helminth is recorded for the first time from the Brown—headed Cowbird. Life cycle: A number of terrestrial have been identified or implicated as intermediate 99 ho sis s of T# americana. Eggs were fed to two species of grasshoppers (Melanoplus femurrubrum and Chorotophaga viridifasciata) and to cock­ roaches (Blattella germanica)• Infective third- stage larvae were found encysted in body muscles 21+ days post infection (Cram, 1934)* Cram (1937) similarly studied the development of T» americana in a grasshopper (Scvllina cvanipes) in Puerto Rico* Raggi and Baker (1957) and Flatt and Nelson (1969) have suggested that terrestrial isopods (Armadillidium sp# and Porcellio sp#) may serve as intermediate hosts# .Prevalence and Intensity of Tetrameres americana in Passerine Birds from South Bass Islands One female specimen was removed from one adult Brown-headed Cowbird of the 1+6 adult Cowbirds examined during this study#

Microtetrameres centuri Barus, 1966 Synonym: Microtetrameres xiphidiopici Barus, 1966 Site of infections Proventricuius Classifications Order Spirurida Diesing, lS6l Superfamily Spiruroidea Family Tetrameridae Travassos, 1914 Genus Microtetrameres Travassos, 1915 Specific diagnosis (modified after Ellis, 1969b): Female: Body bright red, tightly coiled, approximately 1.3mm in length. Transverse striations present. Longitudinal furrows or flanges absent* Buccal capsule width at bulbous midportion 0*010mm* Muscular portion of pharynx 0.025mm in length; glandular portion 0.370mm in length. Intestinal diverticulum present. Dis­ tance between anus and vulva 0.125mm. Vulva 0.300mm from end of body. Anus 0.175mm from end of body. Eggs 0.034 to 0.036mm long by 0.051mm wide. Host records: Host Locality Record Centurus superciliaris Cuba Barus (1966a) Stumella magna. Sturnella neglecta Iowa Ellis (1969a) Centurus florentinoi. Centurus superciliaris. Xiphidiopi cus percussus Cuba Barus (1971) Ageiaius phoeniceus Maryland Cooper and Crites (1974c) Agelaius phoeniceus Ohio Present Study

Discussion: Ellis (1969b) stated that nematodes of this genus parasitize the proventricuius of at least 10 orders of birds. Fourteen species of Microtetrameres have been described from avian hosts in the western hemisphere. Comparative measurements, host ranges and geographical distribution of the described species of this genus were compiled by Ellis (1971). Anatomical 101 differences used in separating certain species were minute. In addition, many of the dimensions reported for several species overlap. Ellis (1969b) observed that any worker attempting to determine the species of a female Microtetrameres experiences difficulty in deciding which of the reported dimensions match most closely. This worker agrees. Ellis (1969b) concluded that small variations in the buccal capsule, spicule, and egg are less subject to change at time of tissue fixation and are of greater significance than are large variations in other dimensions. Experimental observation of the development cycle of M. centuri indicated that eggs were remarkably constant in size as was the buccal capsule. There­ fore, until redecriptions are available, it is best to use only dimensions of structures con­ sidered to be most stable and which change least under fixation. Life cycles Details of the developmental cycle of Microtetrameres centuri have been observed experi­ mentally by Ellis (1969a). Eggs are passed in feces of the definitive hosts. Third—stage in­ fective larvae were recovered from cysts within the perivisceral sinus of the hemocoel of grass­ hopper nymphs (Melanoplus spp.) nine days post 102 infection* Definitive hosts are infected upon ingestion of infected grasshoppers. Prevalence and Intensity of Microtetrameres centuri in Passerine Birds from South Bass Island: One female specimen was removed from a single adult Red—winged Blackbird of the 22 adult Red-winged Blackbirds examined during this study.

Microtetrameres helix Cram* 1927 Site of infection: Proventricuius Classification: Order Spirurida Diesing, l£6l Superfamily Spiruroidea Family Tetrameridae Travassos, 1914 Genus Microtetrameres Travassos, 1915 Specific diagnosis: Following the conclusion of Ellis (1969b) only egg and buccal capsule dimensions are of value for specific determination* Eggs 0.042mm long by 0.033mm wide; buccal capsule 0 .0223mm long. Host records: Host Locality Record Corvus brachyrhvnchos Washington, D.C, Cram (1927a) 1CorvidaeT USSR Spasski and Oshmarin (1939) Corvus brachyrhvnchos Iowa, Wise. Morgan and Waller (1941) Corvus frugilegus USSR Skrjabin, Shikobalova, and Soboley (1 9 4 9 ) 103

Host Locality Record Sturaus vulgaris Mass* Boyd (1951) Corvus^corone. Cvanopica cvana. garrulus glandarius. Pica pica USSR Oshraarin (1956) Corvus levaillanti. Cvanopica cvana. karrulus glandarius USSR Bashkirova (I960) Corvus frugilegus Roumania Stoican (I960) Corvus frugilegus^ Pica pica USSR Ablasov and Chibichenko (1962) Tockus birostris India Sultana (1962) Corvus brachyrhvnchos Manitoba, Can* Hodasi (1963) dorvus corone Bulgaria Stoimenov 0.963) _ Corvus brachvrhvncho s Ohio Jones (196S) dvanocitta cristata Ohio Present Study

Discussion: Records indicate that the geographical 1 distribution of M* helix is holarctic, principally in birds of the family Corvidae* Life cycle: The development cycle of M* helix is unknown* The life cycle of M* helix probably parallels that of M* centuri * Prevalence and Intensity of Microtetrameres helix in Passerine Birds from South Bass Island: No* Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam—Preva— Inten—Inten— Infect—/No* Infect—/No* Host ined lence sitv sitv ed Exam* ed Exam* Cvanocitta 50 2-4 3.0 (50) (0) cristata 104 Microtetrameres spiculata Boyd, 1956 . ... Site of infections Proventricuius Classification: Order Spirurida Diesing, l36l Superfamily Spiruroidea Family Tetrameridae Travassos, 1914 Genus Microtetrameres Travassos, 1915 Specific diagnosis: Following the conclusion of Ellis (1969b), only egg and buccal capsule dimensions are of value for specific determination. Eggs 0.04$ to 0#050mm long by 0*031mm wide; buccal capsule 0.022 to 0.023mm long. Host records: Host Locality Record Cvanocitta Mass. Boyd, et al. (1956) Aphelocoma coerulescens Florida KinselTa T^974) Cvanocitta cristata Ohio Present Study

Discussion: Records of the occurrence of M. soiculata are scant and restricted to passerine birds of family Corvidae. Further study may reveal that passerine birds are parasitized by one or two cosmopolitan species of the genus Microtetrameres. Life cycle: The developmental cycle of M. spiculata is unknown. The life cycle of M. spiculata probably parallels that of M. centuri. 105 Prevalence and Intensity of Microtetrameres spiculata in Passerine Birds from South Bass Island: No Range Infected Birds/Mo. Exam* Birds of Mean Adults Juveniles Exam— Preva— Inten—Inten— Infect—/No* Infect—/No* Host ined lence sity sitv ed Exam* ed Exam* Cvanocitta 50 22 1— 5 2*5 11 (50 ) 0 (0) cristata

Microtetrameres sp* Site of infection: Proventricuius and under lining of Gizzard* Classification: Order Spirurida Diesing, l£6l Superfamily Spiruroidea Family Tetrameridae Travassos, 1914 Genus Microtetrameres Travassos, 1915 Specific diagnosis: Following the conclusion of Ellis (1969a), the identity of male Microtetrameres sp* specimens from the Western Hemisphere must be questioned* Males cannot be specifically de­ termined due to the absence of recognized specific identifying characters* A single male specimen of Microtetrameres sp* was re­ moved from each of two Blue Jays of the 50 adults examined during this study* Acuaria nebraskensis Williams, 1929 Site of infections Under the lining of the gizzard Classification: Order Spirurida Diesing, 1&61 Superfamily Spiruroidea Family Acuariidae Seurat, 1913 Genus Acuaria Bremser, lSll Specific diagnosis (modified after Williams, 1929)s Body tapering anteriorly and narrowing abruptly posteriorly. Cordons extending posteriorly about 1/3 of body length, the posterior termination either abrupt or gradual; usually double, but may be single for as much as half their length. Pseudocordons well developed. Males Length 7#05mra by 0.26lmm wide. Cordons 2.56mm long; pseudocordons 0.011 long. Cervical papillae 0 .260mm from anterior end of body, slightly oval. Transverse body striations 146 to 192 per millimeter. Pharynx 0.175mm long; eso­ phagus 2.270mm long, maximum width 0.122mm. Caudal alae O.&Lmm long. Four pair preanal papillae, eight pair postanal papillae. Left spicule 0.274mm long by 0.043mm wide. Right spicule 0.234mm long by 0.039mm wide. Both spicules strongly curved ventrally. 1 0 7 Host records: Host Locality Record Corvus brachyrhvnchos. Nebraska Williams (1929) Cvanocitta cristata Ohio Present Study

Discussion: These thin, transparent worms are dif­ ficult to perceive in situ. The dearth of records of this organism is probably due in part to its location in a tissue site that is often not examined and to the difficulty of observing the infection when it is present. Life cycle: Details of the life cycle of A. nebrasken— sis are unknown. Development may be similar to that of A. anthuris (Rudolphi, 1&19), a similar helminth frequently reported from corvid birds throughout Eurasia. Quentin, et al. (1972) observed the developmental cycle of A. anthuris experimentally. Eggs passed in the feces of the avian host are ingested by the migratory locust (Locusta migratoria) • Three days post infection, first stage larvae were recovered from the hemo— coel of the . Seven days post infection, the larvae were localized in the abdominal muscles. Infective third stage larvae were re­ covered from grasshoppers 15 days post infection. Definitive hosts become infected upon ingestion of intermediate hosts containing infective third— 10d stage larvae. Prevalence and Intensity of Acuaria nebraskensi s in Passerine Birds from South Bass Islands One male specimen was removed from a single Blue Jay of the 50 adults ex­ amined during this study.

Acuaria auiscula Williams, 1929 Site of infection: Under the lining of the gizzard Classification: Order Spirurida Diesing, 1361 Superfamily Spiruroidea Family Acuariidae Seurat, 1913 Genus Acuaria Bremser, l&ll i Specific diagnosis (modified after Williams, 1929)s Long, slender worms, body tapers rapidly an­ teriorly and posteriorly. Cordons very short; pseudocordons undeveloped. Male: Length 5«22 to 10.34mm by 0.100 to 0.175mm wide. Cordons 0.146 to 0.2d7mm wide. Cervical papillae 0.137 to 0*227mm from anterior end of body, slightly oval. Transverse body striations 200 to 2d6 per millimeter. Pharynx 0.116 to 0.1^5mm long; esophagus 1.164 bo 1.535mm long, maximum width 0.070mm. Caudal alae 0.300 to 0.512mm long. Four pair pre- anal papillae, six pair postanal papillae. Left 1 09 spicule 0*157 to 0*136 long by 0*014 to 0.017mm wide, slightly grooved* Right spicule 0*109 to 0.136mm long by 0.015 to 0.017mm wide, deeply grooved. Both spicules slightly curved ventrally. Host records: Host Locality Record Quiscalus auiscula Nebraska Williams (1929) Aohelocoma coerulescens Florida Kinsella (1974) Cvanocitta cristata Ohio Present Study

Discussion: The infrequency of reports recording the presence of A. ouiscula may well have the same explanation as that of A. nebraskensis. A more plausible explanation is that an intensive study of specimens of Acuaria parasitizing North American passerine birds may reveal these speci­ mens best refer to A* anthuris (Rudolphi, 1319) or A. depressa (Schneider, 1366). The distribution of most of the helminths of corvid birds seems to be holarctic* Life cycle: Details of the developmental cycle of A. auiscula are unknown* The life cycle of A. auiscula probably parallels that of A. anthuris (Rudolphi, 1319) and (Quentin, et al* 1972)*

Prevalence and Intensity of Acuaria ouiscula in Passerine Birds from South Bass Island: 110 No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam—Preva— Inten—Inten— Infect—/No. Infect—/No. Host ined lence sitv sitv ed Exam. ed Exam. Cvanocitta 50 4 1*2 1.5 2 (50) 0 (0) crxstata

Acuaria sp. Site of infection: Under the lining of the gizzard Classification: Order Spirurida Diesing, 1361 Superfamily Spiruroidea Family Acuariidae Seurat, 1913 Genus Acuaria Bremser, l&ll

Discussion: A single immature specimen of Acuaria sp. was removed from each of seven Blue Jays of the 50 adults examined during this study. Identifying structures were insufficiently developed to permit specific referral of these specimens.

Disoharvnx nasuta (Rudolphi, 1&L9) Stiles and Hassal, 1920 Synonyms: Spiroptera nasuta Rudolphi, 1&19 Dispharagus nasutus (Rudolphi, l£l9) Dispharagus spiralis Molin, 1&53 Filaria nasuta (Rudolphi, 1&19) Dispharagus tentaculatus Colucci, 1393 Acuaria nasuta Railliet. Henry and Sisoff, 1912 I l l Cheilospirura nasuta (Rudolphi, 1&L9) Pispharynx notoi Smit, 1927 Acuaria resticula Canavan, 1929 Pispharynx stonae Harwood, 1933 Pispharynx emberizae Yamaguti, 1935 Pispharynx galli Hsu, 1959 Site of* infection: Proventricuius Classification: Order Spirurida Diesing, l£6l Superfamily Spiruroidea Family Acuariidae Seurat, 1913 Genus Pispharynx Railliet, Henry and Sisoff, 1912 Specific diagnosis (modified after Cram, 1927)s Four wavy cuticular cordons on anterior end, origina­ ting at the 2 papilliform lips, not anastomosing, but recurrent, the distal extremity of the cordons turning forward and extending anteriorly. Post- cervical papillae small, bicuspid, situated be­ tween the recurrent branches of the cordons. Body usually coiled in a spiral. Male: Length 7 bo 5mm long by 0.230 to 0.315mm wide. Cordons 0.415 bo 0.515mm long, slender and curved; shorb spicule 0.150mm long, navicular. Ventral surface of body in cloacal region covered with small, elongated shields which give the appearance of a longitudinal striabion. 1X2 Females Length 9 to 10*2mm long by 0*360 to 0* 565mm wide* Cordons 0*9 to l*lmm long* Anus 0*120mm from tip of tail* Small mucron on tip of tail* Vulva in posterior portion of body, 2 to 2*45mm from tip of tail* Cylindrical ovijector curved, S-shaped at its origin, and then directed anteriorly; the limiting boundary of the vestibule and sphincter is clearly marked by an annular cuticular fold* Eggs 0.036 to 0*040mm long by 0*021mm wide; contain larvae when laid* USNM No* 72712. orth American host records: tost Locality Record Gallus gallus Oregon, Wash* Purvis, In Stiles and HassaL (1920) ■uiscalus auiscula Illinois Walton (1923) onasa umbellus New York Allen (1924) Sonasa umbellus Conn*, Maine Gross (1925) Sonasa umbellus Mass*, N* J*, Allen and Gross R* I. Columba livia Texas Cram (1923) j-jonasa "umbellus N* Hampshire Gross (1930) Colinus virginianus Virginia Cram (1931a) Bonasb"*umbellus * Nova Scotia, Can* Pedio^cetes phasianellus Wisconsin Gross (1931) Perdix perdix Wisconsin Cram (1931b) Passer domesticus. Washington, D*C* lurdus mlgratorius New Jersey Cram (1932a) Dumetelia carolinensis New Jersey Cram (1932b) Inrvothorus ludovicianus Texas Harwood (1933) Colinus virginianus N* Carolina Moore (1933) Colinus* virginianus Ohio Venard (1933) Perdix Verdlx Mich., Ohio Yeatter (1934) Bonasb. umbellus Michigan Fisher (1939) Chrvsolophus pTctus. 113 Host Locality Record Phasianus colchicus New York Goble and Cheatum (1943) , Turdus migratorius New York Webster (1943) Columba ixvia California Wehr (1943) Bonasa umbellus# coiinus virginianus, Pavo cristatus• PeHix" perdixT Phasianus' colchicus, Corvus brachyrhynclios, Dumetelia caroilnensis, Molothrus ater, Passer domesticus, flails siaiis, Sturnus vulgaris, Ll‘urdus'"migratorius New York Goble and Kutz (1945a) % Sturnus vulgaris USA Boyd (1951) Ammospiza marxtima N, Carolina Hunter and Quay (1953) Dendragapus obscurus Brit, Columbia, Can, Bendell (1955) Cvanocitta cristata Mass, Boyd, et al, (19567 Columbia livia Florida Hwang, et al, (1961)7“ 7" Dendragapus obscurus Ontario, Can, Jensen (1964) ilxrclus migratorius Colorado Slater (1967) Colinus virginianus Georgia, Kellogg and Florida Prestwood (196S) Sturnus vulgaris California Vincent (1972) Molotnrus ater Ohio Cooper, et al, (1973) ~ “ Melanerpes ervthrocephalus Ohio Cooper (1974) Apneiocoma coerulescens Florida Kinsella (1974) Cmiscaius quTscuia Ohio Buck, et al,(1975) Ageiaius^phoeniceus, Cvanocitta cristata, Jtolohnrus ater. Passer domesticus, Quiscalus quiscula, sturnuETvui gari s, Turdus migratonus Ohio Present Study

Discussion: The pattern of distribution of this nematode is cosmopolitan. This writer noted fifty—two citations recording the occurrence of D# nasuta in addition to the North American records listed above# The wide range of defini­ tive hosts comprises birds of the order Charadriiformes, Columbiformes, Coraciadiformes, Galliformes and Passeriformes, principally the latter two# This study records D# nasuta from the Red-winged Blackbird for the first time# Nescos (1954) reported that D# nasuta infections killed 1/3 of the pigeons in an Italian aviary# Infection of the proventri cuius re suited in papilliform hyperplastic involvement of the glandular layer# Bendell (1955) stated that D# nasuta infections are the principal factor limi­ ting Blue Grouse (Dendragapus obscurus) recruit­ ment in British Columbia# In an experimental study of the pathology of D. nasuta infections in Blue Grouse, Jensen (1964) found that the severity of the pathological signs was directly proportion­ al to the size of the infection and the age— class of the host# Infected juvenile Blue Grouse gained less weight than control birds and exhibited a retarded development of juvenile feathers# Chronic hematological signs in Jensen’s study were anemia and leucocytosis# 115 Life cycle: Piana (1&97) first observed aspects of the life cycle of Pispharynx nasuta. Eggs are passed in the feces of the avian hosts and are subsequently eaten by terrestrial isopods. The larvae hatch and penetrate the intestine of the isopod. First-stage larvae were found in the hemocoel 4 days post infection. Infective third- stage larvae were present in the hemocoel 26 days post infection (Chabaud, 1954)• Cram (1931) ex­ perimentally infected terrestrial isopods (Porcellio scaber and Armadillidium vulgare)• Barus, al. (1970) experimentally infected an isopod (Porcellionides pruinosis) in Cuba. De­ finitive hosts become infected upon ingestion of infected terrestrial isopods. Prevalence and Intensity of Pispharynx nasuta in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam-Preva— Inten—Inten— Infect—/No. Infect—/No. Host ined lence sitv sitv ed Exam. ed Exam. Apelaius 50 ' 3 1-7 4*5 1 (22) 3 (2$) pnoeniceus Cvanocitta 50 12 1-5 3*0 6 (50) 0 (0) cristata Molothrus 50 12 1-2 1.7 4 (46) 2 (4) ater Passer 30 3*3 11 — 0 (5) 1 (25) domesticus No* Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam— Preva— Inten—Inten— Infect—/No* Infect—/No* Host ined lence sitv si tv ed Exam* ed Exam* O.uiscalus 50 23 1-13 6.7 1 (13) 13 (37) auiscula Sturnus 50 24 1-3 3*2 0 (1) 12 (49) vulgaris Turdus 50 23 1-26 7.0 1 (19) 13 (31) migratorius

Diplotriaena bargusinica Skrjabin, 1917 Synonyms: Diplotriaena henrvi Blanc, 1919 Diplotriaena isabellina Koroliowa, 1926 Diplotriaena roveglial Caballero, 1941 Diplotriaena conceptionis Caballero, 1943 Site of infection: Air sacs Classification: Order Spirurida Diesing, 1361 Superfamily Filarioidea Weinland, 1353 Family Diplotriaenidae Anderson, 1953 Genus Diplotriaena Henry and Ozoux, 1909 Specific diagnosis (modified after Anderson, 1959)i Trident medium-sized; apex sharply pointed in form of arrow head* Esophagus divided into short, anterior, muscular part and long, pos­ terior, glandular part* Cuticle smooth. Female: Length 47 to 54mm, maximum width 0,55 to 0,6lmm wide. Trident 0,14mm long; delicate 1 1 7 tubercles on proximal end* Muscular esophagus 0.45 to 0.54mm long. Vulva on slight protuberance, 0.35 bo 0.39mm from anterior end. Vagina approxi­ mately 1.2mm long. Eggs 0.046 to 0.053mm long by 0.031 to 0.035mm wide. North American host records: Host Locality Record Cassidix mexicanus Mexico Caballero (1941) bives cfTves Mexico Caballero (194&) J* IJ ocichla guttata. Ontario, Can. ocichla ustuiata, USA tospiza lincolnil. Ontario,- Can. Oenanthe oenanthe V Alaska Passerculus sandwishensis. Ontario, Can. 6uiscaJ-iis quiscula Ontario, Can. Anderson (1959) Hylocichia fuseescens. Qulscaluswrlocichla quisculaustuiata. Ontario, Can. Anderson (1961) HviocicHra fuscescens Ontario, Can. Anderson (1962) Molothrus ater Ohio Present Study

Discussion: Records of the occurrence of D. bargusi— nica indicate a holoarctic and neotropical geo­ graphical distribution in passerine birds. Over­ all, records of the occurrence of this helminth are scant. The paucity of records is probably due, in part, to a failure of workers to report the occurrence of diplotriaenids removed from birds. Prior to the work of Anderson (1959), specific determination of specimens was nearly impossible due to scattered literature sources and inadequate specific descriptions. Diplotriaena 118 bargusinica is reported from the Brown-headed Cow- bird for the first time during this study# Life cycle: Details of the developmental cycle of D# bargusinica were observed experimentally by i Anderson (1962)# Thick-shelled eggs deposited by female worms pass through the respiratory tract, are swallowed, and passed in the feces of the avian host# When eaten by grasshoppers (Melanoplus bilituratus. M# fasciatus. and Camnula nollucida), larvae hatch, penetrate the midgut, and develop further in the fat body# In­ fective third-stage larvae were removed from cysts in the fat body of grasshoppers 18-19 days post infection# Definitive hosts are infected upon ingestion of infected intermediate hosts# Migration to the air sacs is apparently via the circulatory system. The developmental cycle of D# agelaius (Walton, 1927) was studied by Anderson (1957) • The cycle of D# tridens (Molin, 1858) was studied by Bain and Vaucher (1973)# The develop­ mental cycles of the latter two species closely parallel that of D# bargusinica. utilizing grass­ hoppers as intermediate hosts# Prevalence and Intensity of Diplotriaena bargusinica in Passerine Birds from South Bass Island: Two female speci­ mens were removed from a single adult Brown—headed Cowbird of the 46 adult Brown-headed Cowbirds examined during this study#

Diplotriaena tricuspis (Fedtschenko, 1^74) Henry and Ozoux,

1909 Synonyms; Filaria attenuata (Rudolphi, 1&03) Filaria attenuata (Fedtschenko, 1&74) Anrocta tricuspis (Fedtschenko, 1^74) Filaria ninni (Stossich, 1J$91) Diplotriaena bifidus Canavan, 1931 Diplotriaena manipoli Chu, 1931 Site of infections Air sacs Classification: Order Spirurida Diesing, 1&61 Superfamily Filarioidea Weinland, 1£$5& Family Diplotriaenidae Anderson, 195# Genus Diplotriaena Henry and Ozoux, 1909 Specific diagnosis (modified after Anderson, 1959): Large nematodes# Trident small, apex flat to slightly concave in lateral view and pointed in dorso-ventral view, arms rather far apart# Eso­ phagus divided into short, anterior, muscular part and long, posterior glandular part# Cuticle with fine transverse striations# Male: Length 45 to 6lmm# Maximum width 0#55 to 0#72mm near middle of body# Trident 0#09 to 120 0.12mra in length* Muscular esophagus 0.32 to 0.63mm, glandular part 3*1 to 3*3ram in length* Nerve ring 0*20 to 0.30mm from anterior extremity* Tail broad, with prominent lateral wing-like ex— • pensions bearing at least nine pairs of minute papillae, one pair of which are dorsal in position. Right spicule 0*96 to 1.1mm in length, bent three times and with a narrow ala on much of its length* Left spicule long and slender, 1*6 to 2*0mm in length* Anus 0*07 to 0*llmm from caudal extremity* Lateral line 0*13 to 0* 01mm, canal 0.009 to 0.014mm in width. Female: Length 123 to 153mm. Maximum width 0.90 to 1.0mm near middle of body* Trident 0.09 to 0*12mm in length* Muscular esophagus 0.27 to 0*34mm, glandular part 3.7 to 4 •3mm in length. Nerve ring 0.19 to 0.20mm, and excretory pore 0.12mm from anterior extremity. Vulva opening on prominent protuberance 0.39 to 0.76mm from an­ terior extremity. Vagina 1.9 to 3»3mm in length. Eggs 0*056 to 0.063mm by 0.033 to 0.041mm in size. Lateral line 0.17 bo 0.22mm and canal 0.022mm in width. USNM No. 72700. North American host records: Host Locality Record Corvus brachvrhvnchos Wisconsin Morgan.and Waller 121 Host Locality Record Cvanocltta cristata Mass* Boyd, et al* (19567“ ~ „ Corvus brachvrhvnchos New York;Quebec Anderson (1959) Can* Cvanocltta cristata Ontario, Can* Anderson (1961) Vermivora ruficapilla Ontario, Can* Judd (1959) Corvus brachvrhvnchos Ohio Jones (1958) 6orvus brachvrhvnchos N* Carolina Hendricks, et al* (1969) Cvanocltta crxstata Ohio Present Study

Discussion: The pattern of geographical distribution of D* tricuspis is cosmopolitan* Records of occurrence of this helminth indicate it is limited almost exclusively to species of the passerine family Corvidae* Life cycle: The life cycle of D* tricupis has not been observed experimentally or in nature. It probably parallels that of D. bargusinica* utilizing grasshoppers as an intermediate host for development* Prevalence and Intensity of Diplotriaena tricuspis in Passerine Birds from South Bass Island: No* Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam- Preva— Inten-Inten- Infect—/No. Infect-/No* Host ined lence sitv sitv ed Exam* ed Exam* Cvanocitta 50 14 1-3 3.$ (50) (0) cristata 122 Splendidofilaria algonguinensis (Anderson, 1955) Anderson, 1961 Synonyms Omithofilaria algonguinensis Anderson, 1955 Site of infection: Aorta, auricles of heart Classification: Order Spirurida Diesing, 1S61 Superfamily Filarioidea Weinland, 135# Family Onchocercidae Chabaud and Anderson, 1959 Subfamily Splendidofilariinae Chabaud and Choquet, 1953

Genus Sp Iendidofilaria Skrjabin, 1923 Specific diagnosis (modified after Anderson, 1955)s Small worms tapering markedly at the extremities. Oral opening a dorsoventral slit bordered by a delicate, circumoral, cuticular ring. Oral papillae consist of 4 submedian pairs. Amphids lateral. Esophagus not divided. Intestine broad and transparent. Cuticle with delicate transverse striations. Male: Length 9.2 to 12.3mm, maximum width 0.059 to 0.063mm. Esophagus 0.52 to 0.65mm long, 0.013 to 0.019mm wide. Spicules subequal. Right spicule 0.031 to 0.033mm long; left spicule 0.029 to 0.030mm long. Caudal papillae prominent, conical, consisting of 2 preanal and 3 postanal papillae. Tail with 2 prominent terminal 123 protuberances# Phasmids lateral, O.OlOmra from end of tail# Anus surrounded by a slightly raised cuticular plate, 0.042 to 0.059mm from end of tail# Female: Length 19#1 to 22#6mm long, maximum width 0.031 to 0#042mm# Esophagus 0.55 to 0.56mm long# Tail bluntly pointed, with pair of small terminal protuberances# Anus on slightly raised protuberance# Vulva 0#19 to 0#26mm from anterior end# Vagina convoluted, 0.42 to 0 #66mm long, filled with microfilaria, 0.076 to 0#0S4mm long# Eggs indistinct# Host records: Host Locality Record Hirundo ervthrogaster Ontario, Can< Anderson (1955) K^olothrus ater Ohio Cooper, et al# (19735 ” Agelaius phoeniceus Maryland Cooper and Crites (1974c) Molothrus ater Ohio Present Study

Discussion: Records of the occurrence of S# algon- quinensis are scant# Failure to closely in­ spect the heart musculature and blood vessels may partially explain the paucity of records# Only by teasing apart the blood vessels and heart tissues does the infection become evident to the investigator. The nematodes lie embedded in the musculature. 124 Life cycles Aspects of the developmental cycle of S. algnnmiinensis are unknown. Aspects of the developmental cycles of two members of the genus Splendidofilaria are known* Splendidofilaria sis (Anderson, 1954) utilizes several

species of blackflies (Simulium s p p .) for de­ velopment (Anderson, 1956). Hibler (1964a) re­ ported S. picacardina Hibler (1964) and utilized several species of biting midges (Culicoides spp.) for development. Either blackflies or biting midges serve as intermediate hosts for S. algon- auinensis. • Prevalence and Intensity of S. algonguinensis in Passerine Birds from South Bass Island: Eight specimens were re­ moved from a single adult Brown-headed Cowbird of the 46 adult Brown-headed Cowbirds examined during this study.

Chandlerella auiscali (von Linstow, 1904) Robinson, 1971 Synonyms: Filaria auiscali von Linstow, 1904 Splendidofilaria auiscali (von Linstow, 1904) Site of infection: Under pia-mater of cerebral hemispheres Classification: Order Spirurida Diesing, 195^ Superfamily Filarioidea We iniand, 1&5& 1 25 Family Onchocercidae Anderson and Chabaud, 1959 Subfamily Splendidofilariinae Chabaud and Choquet, 1953 Genus Chandlerella Torke and Maplestone, 1926 Specific diagnosis (modified after Robinson, 1971)• Male: Length 23 to 34mm, maximum width 0* 172mm* Esophagus 1*00 to l*5mm long, maximum width 70mm* Tail coiled, bluntly rounded, with 4 pairs of postanal papillae* Spicules dissimilar, curved, smooth, blunt, graod in the middle* Left spicule 0*076 to 0*105mm long by 0*015 to 0*020mm wide; right spicule 0*064 to 0*095mm long by 0*015 to 0*020mm wide* Female: Length 50 to SOmm, maximum width 0*200mm* Esophagus 0*35 to l*13mm long, maximum width 0*090mm* Tail slightly tapered, bluntly rounded* Vulva anterior to esophage—intestinal junction, 0*409 to 0*924mm from anterior end of body* Microfilarioid embryos in uterus with thin, clear envelope* Host records: Host Locality Record htiscalus quiscula USA von Linstow (1904) luiscalus quiscuLa Iowa Odetoyinbo and Ulmer (1959, I960) Quiscalus quiscula Indiana Welker (19o2) Jorvus "‘TSrachyrhyncho s Ohio Jones (1963) Quiscalus quiscalus Minnesota Robinson (1971) Moxotnrus ater Ohio C?l§?3J — — ’ 126 Host Locality Record Cyanocitta cristata, Molothrus ater, Quiscalus quiscula Ohio Present Study

Discussion: Records of the occurrence of C. quiscali indicate its geographical distribution is limited to the midwestern United States. Definitive hosts are passerine birds of the families Corvidae and Icteridae. In an endemic area, the incidence in a single avian species may be quite high. Welker C1962) found adult £. quiscali in 100% of the Common Grackles examined for this helminth. Odetoyinbo and Ulmer (1959, 1960) observed that the microfilariae demonstrate a highly nocturnal periodicity in the avian hosts* peripheral blood stream. The microfilariae of C. quiscali displayed considerable variation in density in the peripheral blood for the same sampling time on successive nights. Schmid and Robinson (1972) observed the distri­ bution of larvae in experimentally infected gnats CCulicoides crepuscularis). The variation in den­ sity of microfilariae in the peripheral blood was reflected in the clumped pattern of occurrence of larvae in the gnats observed after feeding on an infected avian host. Schmid and Robinson concluded 127 •that the microfilariae may have been distributed randomly throughout the peripheral circulation, but their density may have varied during the periods of exposure to the gnats. Life cycle: Aspects of the developmental cycle of C . quiscali have been observed experimentally by Robinson (1961). Ceratopogonid gnats (Culicoides crepuscularis) were exposed to an infected Common t Grackle. Gnats ingest microfilariae randomly and > * exsheathment and migration to thoracic muscles occurs within minutes of ingestion. Larvae moult twice at about 84- and 120 hours, respectively, post infection. The infective third-stage larvae migrate to the head of the insect in less than 12 hours after the second moult. Other aspects of the developmental cycle of this helminth are unknown. Prevalence and Intensity of Chandlerella quiscali in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten-Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Cyanocitta 50 2 4 1 1 (50) 0 (0) cristata Molothrus 50 4 1 1 2 (46) 0 (4) ater Quiscalus 50 12 1-4 2 5 (13) 1 (37) quiscula Capillaria contorta CCreplin, 1839) Travassos, 1915 Synonyms: Triehosoma contortum Creplin, 18 39 Thominix contorta (Creplin, 1839) Trichosoma longicolle Rudolphi, 1819 Eucoleus contorta (Creplin, 1839) Eucoleus corvicola Vasilkova, 19 30 Capillaria perforans Kotlan and Orosz, 1931 Capillaria lophortygis Baylis, 1934 Eucoleus raillieti Lopez-Neyra, 1947 Site of infection: Esophagus Classification: Order Trichinellida Chitwood, 1969 Superfamily Trichinelloidea Family Trichuridae Railliet, 1915 Subfamily Capillariinae Railliet, 1915 Genus Capillaria Zeder, 180 0 Specific diagnosis Cmodified after Barus, e£ al_., 1972) White to yellowish worms; cuticle with transverse striations. Bacillary bands extending along entire length of body. Muscular portion of eso­ phagus 0.36 to 0.77mm -long. Stichocytes number 33 to 4 2 . Male: Length 11.38 to 12.57mm, width 0.073 to 0.088mm. Anterior end of body 0.0l t o wide. Length of esophagus 4.0 3 to 4.68mm. Ratio of esophagus to posterior portion of body, 1:1.6-1.8. 129 Spicules indistinct. Spicule sheath with small spines distributed most densely in its central ampulla-shaped portion. Posterior end of body surrounded by a small caudal vesicle supported at each side with a rounded papilla and one pair of ventral papillae. Cloaca subterminal. Female: Length 11.70 to 15.44mm, width 0.11 to 0.16mm. Anterior end of body 0.015 to 0.016mm wide. Length of esophagus 4.21 to 4.86mm. Ratio of eso­ phagus to posterior portion of body, 1:1.6-2.5. Vulva at 0.079 to 0.29mm from end of esophagus. Vulva transverse slit, borders unraised. Eggs

4 0.058 to 0.067mm long by 0.026 to 0.030mm wide. Anus subterminal. North American host records: Host Locality Record Colinus virginianus New Jersey Cram (1930) Meleagris gallopavo Virginia Wehr (1948) Bonasa umbellus Minnesota Erickson, et al. (1949) Sturnus vulgaris Ohio Boyd (1951) CyanocTtta cnstata Mass. Boyd, et al. (19567” Querquedula discors Quebec, Can. Mawson (1956b) Capella gailinago Colorado Schmidt (1964) Colinus virginianus Georgia, S. Kellogg and Carolina Prestwood (1963) Corvus brachyrhyncho s Ohio Jones (1968) CapelXa gailinago USA Threlfall (1968)- Colinus virginianus Georgia, Davis, et al. Florida (1969) Colinus virginianus Georgia, Dawe, et al. Florida (19697” 130 Host Locality Record Cyanocitta cristata Conn. Helmboldt9 et al. ------C1971) Cyanocitta cristata, Turdus migratorius Ohio Present Study

Discussion: The pattern of geographical distribution of this helminth is cosmopolitan. Barus, et^ al. (1972) noted that the numerous findings of eso­ phageal capillarids reported from birds of 6 different orders indicartes "their -location in the same organ of the host results in an apparent uniform specialization of these ‘forms. The exist­ ence of other valid taxa of esophageal capillarids specific to hosts of the orders Anseriformes and Lariformes is problematic and■requires a detailed study of specimens collected from hosts of these two orders. C . contorta is reported from the Robin for the first time during this study. Helmboltj et al. (1971) observed a diphtheritic membrane formation in the esophagus and oral cavity of Blue Jays infected with C^. contorta. The oral . and pharyngeal cavities and esophagus down to the proventriculus were covered by a whitish membrane. The birds were emaciated and dehydrated. The effect of the membrane was mechanical as it pre­ vented normal esophageal activity. No gross pathology was observed in the birds examined by 131 the writer during this study. Life cycle: Aspects of the development of contorta have been observed experimentally by Cram C19 36). Uncleaved eggs are passed in the feces of the avian host. Eggs develop in the soil, requiring approximately 30 days. Upon ingestion by an avian host, the larvae hatch and the worms become sex­ ually mature in 30-60 days. The closely related species, C. annulata (Molin, 1858) requires an earthworm CEisenia foetida, Allobophora caliginosa and Lumbricus terrestris) intermediate host to complete its larval development CWehr, 19 36; Allen, 19lt9). Wehr C1952) demonstrated that the red- banded earthworm (Eisenia foetida) serves as a suitable intermediate host for the transfer of the infective stage of C. contorta to its avian defini- tive host as well as confirming its direct trans­ mission. Prevalence and Intensity of Capillaria contorta in Passerine Birds from South Bass Island: No Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten- Inten-Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Cyanocitta 50 6 1 1 3 C50) 0 (0) crxstata Turdus 50 2 1 - 0 (19) 1 (31) migratorius Capillaria exilis (Dujardin, 1845) Travassos, 1915 Synonym: Trichosomum exile Dujardin, 1845 Liniscus exilis Dujardin, 1845 Site of infection: Intestine Classification: Order Dorylaimida Superfamily Trichinelloidea Family Trichuridae Railliet, 1915 Subfamily Capillariinae Railliet, 1915 Genus Capillaria Zeder, 1800 Specific diagnosis (modified after Wakelin, 1966): Male: Length 7.27-to 17.72mm. Width at base of esophagus 0.040 to 0.059mm. Length of esophagus 3.39 to 8.04mm. Length of spicule 0.7 8 to 1.50mm. The body tapers considerably to the caudal end, where it is bordered by lateral alae 0.035 to 0.080ram in length. The posterior tip bears a small bursa 0.027 to 0.035mm wide, supported by two ventro-lateral projections from the body, each of which bears a small terminal papilla. The spicule is slender and flexible, circular in cross- section and 0.00 3 to 0.008mm wide at the mid­ point. Proximally it is only slightly expanded; at the distal tip it tapers to a fine point and

. often shows a constriction before the tip. Sclerotization is often irregular at the 133 extremities and the distal end may present a beaded appearance• The spicule sheath is smooth and shows transverse striations. Female: Length 10*00 to 21*14mm* Width at the vulva 0.04& to 0«064mm. Length of esophagus 4»77 to ?*96mm* The vulva is situated a little less than half of the body length from the anterior end and bears a small rounded appendage 0*020 to 0.087mm long and 0*032 to 0*056mm wide* The presence and appearance of the vulval appendage was constant in all the females examined* The vagina is 0*07 to 0*i3mm long* At the caudal end, the body terminates bluntly; the anus is sub­ terminal* Egg length 0*053 to 0*069mm, width 0*021 to 0*032mm* The outer egg shell layer has a characteristic lattice pattern with prominent striae at each pole; the inner layer forms a collar at each pole and is slightly reflexed* North American host records: Host Locality Record Turdus migratorius Quebec, Can* Rayner (1932) 5turnus vulgaris USA Boyd (1951) Aix sponsa* flails s* sialis N* America Wakelin (1967) Cyanocitta cristata, quiscalus qulsculaT Sturnus vulgaris* turdus migratonus Ohio Present Study 134 Discussion: Records of the occurrence of C,* exilis indicate that the geographical distribution of this helminth is limited to Africa, Europe and North America* Overall, records of this helminth are scant* It is reported most frequently in passerine birds* Capillaria exilis is reported from the Blue Jay and Common Grackle for the first time during this study* Life cycle: Aspects of the developmental cycle of C* exilis are unknown* The life cycle of this hel­ minth may parallel that of Capillaria caudinflata (Molin, 1353). Details of the developmental cycle of JC* caudin— flata* a frequently reported intestinal helminth of galliform and passeriform birds, were observed experimentally by Morehouse (1944)* Eggs are passed in the feces of the avian host* Direct feeding of "eggs to birds failed to produce new infections* Upon ingestion by earthworms (Eisenia foetida. Allolobophora oAlj fiin-Lsa’) f larvae hatch from the eggs, penetrate the intestine, and develop to infective forms within 11 days* Upon ingestion of infected earthworms, development to sexually mature adults in chickens requires 22-24 days* Prevalence and Intensity of Capillaria exilis in Passerine Birds from South Bass Island: No. Range Infected Birds/No* Exam* Birds of Mean Adults Juveniles Exam— Preva— Inten— Inten—Infect—/No. Infect—/No. Host ined lence sity sitv ed Exam. ed Exam. Cvanocitta 50 12 1-2 1.3 6 (50) 0 (0 ) cristata Quiscalus 50 6 2-14 6.0 0 (13) 3 (37) auiscula

Sturnus 50 IS 1-S 4.0 1 ' (1 ) 8 (49) vulcarls Turdus 50 70 1-35 S. 6 9 (19) 26 (31) micratorius

Capillaria ovopunctatum (von Linstow, 1S73) Travassos, 1915 Synonyms: Trichosoma ovopunctatum von Linstow, 1S73 Trichosomum ornatum Dujardin, 1&43 Calodium ornatum Dujardin, 1S45 Capillaria quiscali Read, 1949 Site of infection: Intestine Classification: Order Dorylaimida Superfamily Trichinelloidea Family Trichuridae Railliet, 1915 Subfamily Capillariinae Railliet, 1915 Genus Capillaria Zeder, 1S00 Specific diagnosis (modified after Wakelin, 1966): Male: Length 4.77 to 10.00mm. Width at base of esophagus 0.043 to 0.53mm. Length of esophagus 3.57 to 5•72mm. Length of spicule 0.56 to 0«97mm. The body tapers slightly to the caudal end, which is bilobed and bears a spatulate bursa 0.053 to 0.067mm wide. The bursa is supported by two ventro—lateral projections from the body. The spicule is well developed, circular in cross section and 0.004 to 0.009mm at the mid-point. Proximally it is expanded to 0.011 to 0.027mm and occasionally has a triangular section. Distally' the spicule ends in a claviform tip 0.006 to 0.016mm wide. The spicule sheath is smooth and shows extensive transverse foldings when retracted. Female: Length 6.36 to 14.77mm. Width at the vulva 0.04& to 0.069mm. Length of esophagus 3«41 to 6.25mm. The vulva is situated approximately one—half of the body length from the anterior end and bears a broad—based, funnel-shaped appendage 0 . 0 9 to 0.16mm long. Frequently the shape of the appendage was distorted by the prolapse of body contents through the vulva. The vagina is muscu­ lar, 0 . 1 7 to 0.24mm in length. At the posterior end the body tapers bluntly; the anus is sub­ terminal. Egg length 0.056 to 0.69mm, width 0 . 0 2 4 to 0.032mm. The outer egg shell layer is markedly punctate, giving the egg a characteristic appearance. The inner shell layer forms a collar 137 at each pole but is not, or* is only slightly, reflexed. North American host records: Host Locality Record Quiscalus quiscula Wisconsin Read (191*9) Sturnus vulgaris USA Boyd (1951) Quiscalus quiscula, Sturnus vulgaris Quebec, Can. Mawson (1956b) Quiscalus quiscula Manitoba, Can, Hodasi C1963) Quiscalus quiscula Ohio Stanley and Rabalais (1971) Sturnus vulgaris California Agelalus phoeniceus, Molothrus ater, Quiscalus quiscula, Sturnus vulgaris, Turdus migratonus Ohio Present Study

Discussion: Records of the occurrence of C. ovo­ punctatum indicate that the geographical distri­ bution of this helminth is limited to North America and Western Europe. The most frequently recorded host in Europe is the Starling. One can speculate that this common helminth of Starlings was introduced into North America with its host and is now maintained in this region largely by Common Grackles and Starlings. This helminth is reported from the Red-winged Blackbird, Brown­ headed Cowbird, and Robin for the first time during this study. Life cycle: Aspects of the developmental cycle of C. ovopunctatum are unknown. The life cycle of C . ovopunctatum probably parallels that of Capillaria 138 caudinflata (Morehouse, 1944). Prevalence and Intensity of Capillaria ovopunctatum in Passerine Birds from South Bass Island: No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten-Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam Agelaius 50 4 3-9 6.0 1 (22) 1 (28) phoeniceus Molothrus 50 4 1 2 (.46) 0 (4) ater Quiscalus 50 44 1-25 8.4 4 C13) 18 (37) quiscula Sturnus 50 46 1-40 9.7 0 (1) 23 (49) vulgaris Turdus 50 12 2-15 4.7 0 (19) 6 (31) migratorius Acanthocephala Mediorhynchus grandis Van Cleave, 1916 Synonyms: Heteropus grandls CVan Cleave, 1916) Empodius grandis CVan Cleave, 19162. Site of infection: Lower intestine > Classification: Order Archiacanthocephala Meyer, 1931 Family Gigantorhynchidae Hamann, 1892 Genus Mediorhynchus Van Cleave, 1916 Specific diagnosis Cmodified after Moore, 1962): Mature females 27 to 51mm in length by 0.8 to 1.4mm in width. Mature males 7 to 12mm in length by 0.6 to 0.8mm in width. Anterior and posterior ends of body flexed ventrally. Body slightly ex­ panded just posterior to proboscis. Fully ex­ tended proboscis, cone-shaped, 0.57 to 0.86mm in length, divided into anterior and posterior regions by insertion of proboscis receptacle. Anterior region 0.28 to 0.42mm long, posterior region 0.29 to 0.44mm long. Diameter of proboscis near apex Clevel of most anterior hooks) 0.14 to 0 .21mm; at level of insertion of proboscis re­ ceptacle 0.28 to 0.43mm; at base 0,35 to 0.79mm. Anterior region of proboscis possesses 18 more or less longitudinal rows of hooks with 4 or 5 hooks per row, each hook provided with posteriorly 140 directed root. Average total number of hooks anterior to insertion of proboscis receptacle 70. Anterior three or four hooks in each longitudinal row 0.025 to 0.0 3 5mm long Cmeasurement taken at largest straight line from tip of hook to top of angle between hook and root). Length of root 0.03 to 0.05mm Cfrom base of root to top of angle between root and hook). These hooks strongly recurved, tapering to fine point. Posterior one or two hooks in each longitudinal row much stouter, with longer root and blunt point; length 0.032 to 0.036mm, root 0.077 to 0.09mm. Diameter about 0.01mm. Posterior region of proboscis possesses about 30 longitudinal rows of small hooks with 4 to 6 hooks per row. Average total number 130. These hooks are simple except for the first two in each row which are slightly recurved, lack posteriorly directed roots but possess small protuberance at base of each hook, length 0.029 to 0.038mm. Proboscis receptacle thin-walled, sac-like, with narrower thick-walled posterior tip. Anterior thin-walled portion 0.28 to 0.3mm by 0.19 to 0.25mm (width measured at level of cervical ganglion). Thick-walled posterior portion 0.08 to 0.2 5mm by 0.04 to 0.14mm. Lemnisci emerge from receptacle anterior to thick-walled 141 portion. Cervical ganglion centrally located in anterior region of receptacle between the lemnisci. Five to 8 pyriform cement glands. Testes oval, 0.98 to 1.03mm by 0.26 to 0.35mm located in tandem in posterior half of body. Lemnisci filiform with 6 nuclei each. Eggs elliptical, provided with two shells and three accessory membranes, outer shell 0,058 to 0.064mm by 0.0 35 to 0.0 38mm, inner shell rigid and highly . refractive, acanthor within the egg 0.041 to 0.054mm by 0.025 to 0.027mm. Host records: Host Locality Record Quiscalus quiscula, Maryland Sturnella magna N . Carolina Van Cleave (.1916 ) Corvus brachyrhyncho s Maryland Van Cleave (1918) Sturaus vulgaris Illinois Sommer (19 36) Agelaius phoeniceus Ohio Corvus brachyrhynchos, Ohio Euiphagus carolmus Illinois Quiscalus quiscula Ken., 111., Kan., N.J., Ohio Sturnella magna Illinois, Ohio Van Cleave (1947) Sturnella magna Oklahoma Bachman and Berry (1948) Sturnus vulgaris USA Boyd (1951) Corvus brachyrhynchos Virginia Daly (1959) Meleagris gallpavo South Dakota Huggins and Daumann (1961) Agelaius phoeniceus, Texas Molothrus ater, Texas ^assernerbulus caudacutus, Texas 14-2

Host Locality Record Serinus canarius , Expt. infection Sturnella magna, Texas Turdus migratorius Ohio Moore (196 2) Quiscalus quiscula, Xanthocephalus xanthocephalus Manitoba, Can. Hodasi (1963) Sturnus vulgaris Colorado Ballard and Olsen (.1966 ) Quiscalus quiscula Colorado Sturnella neglecta Schmidt and Neiland (.1966 ) Corvus brachrhynchos Ohio Jones (1968) Corvus brachyrhynchos North Carolina Hendricks, Harkema, and Miller (196 9) Centurus Carolina, Sturnella magna Louisiana Nickol (1969) Agelaius phoeniceus, Sturnella magna Georgia Byrd and Kellogg (.1971) Sturnus vulgaris Kansas Rodrick and Johnson C1971) Aphelocoma coeaulescens Florida Kinsella C1974) Passer domesticus Ohio Present Study

Discussion: Records of the occurrence of M. grandis indicate that the distribution of the helminth is largely limited to members of the family Icteridae in the eastern and central portion of the United States. With the exceptions of Crows (Corvus brachyrhynchos) and Starlings, records from non- icterid passerine birds are infrequent. The feeding habits of crows and starlings often over­ lap those of icterid birds. Study of the inci­ dence of M. grandis indicates that the principal host species of this helminth are meadowlarks - (Sturnella magna and S^. neglecta). This study records M. grandis from the House Sparrow for the 143 first time. Life cycle: Details of the developmental cycle of M. grandis were observed experimentally by Moore C1962). Eggs containing acanthors are passed in l the feces of the avian host. When ingested by grasshoppers CChortophaga viridifasciata, Orphulella pelidna, Arphia luteola and Schisto- cerca americana) the acanthor escapes and pene­ trates the wall of the midgut. The larvae remain for twelve days post infection on the outer wall of the midgut. Subsequently, the acanthors drop free into the hemocoel and develop into acan- tella stage larvae 25 days post infection. Further development results in encysted cyst- acanths within 2 8 to 30 days post infection. Definitive hosts become infected upon ingestion of infected grasshoppers. Prevalence and Intensity of Mediorhynchus grandis in Passerine Birds from South Bass Island: Four specimens were removed from a single juvenile House Sparrow of the 25 juvenile House Sparrows examined during this study.

Plagiorhvnchus formosus Van Cleave, 1918 Synonym: Prosthorhvnchus formosus CVan Cleave, 1918) Site of infection: Lower intestine 144 Classification: Order Meyer, 1931 Family Plagiorhynchidae Golvan, 19 59 Subfamily Plagiorhynchidae Meyer, 1931 j Genus Luhe, 1911 ' Specific diagnosis Cmodified after Schmidt and Olson, 1964): Mature female 9 to 15mm long, 2 to 3mm wide, mature male 8 to 13mm long, 1.5 to 2.5mm wide. Body elliptical to slightly ovoid, more or less flattened dorsally; trunk spines absent; color in life milky-white; hypodermal nuclei numerous, distributed evenly throughout trunk. Fully extended proboscis nearly cylindrical 0.80 to 1.10mm long, 0.0 25 to 0.029mm wide in female, slightly smaller in male. Armature identical in both sexes, consisting of 15 to 18 longitudinal, regularly alternating rows of 11 to 15 hooks which measure 0.075 to 0.08 3mm long except first and last two hooks in each row which are 0.055 to 0.065mm long Cmeasurement of longest straight line from tip of hook to top of angle between hook and root). Roots of hooks simple, occasionally ir­ regularly knobbed or lobed, directed posteriorly, 0.063 to 0.066mm long; last two or three hooks in each row with no roots. Proboscis receptacle double-walled, cylindrical, rounded at base, 1.70 1 4 5 to 1.90mm long, 0.410 to 0.5 32mm wide, inserted in base of proboscis at level of last three or four hooks in each row. Cervical ganglion located in middle third of proboscis receptacle. i Conspicuous retinacula penetrate receptacle walls laterad and insert on wall. Lemnisci slightly longer than proboscis receptacle, claviform, flattened, containing numerous fragments of nuclei; rarely split along part or all of their length. Testes tandem or slightly oblique, oval, 0.080 to 0.090mm long, 0.040 to 0.050mm wide, about one-third body length from anterior end; anterior testes commonly somewhat smaller than posterior. Three to six tubular cement glands extend from rear margin of posterior testes to region of bursa. Genital pore dorsoterminal. Copulatory bursa lacking, rays. Completely formed, living egg elliptical, averaging 0.075mm long by 0,03 0mm wide, with two shells and inner membrane; outer sculptured with ridges and grooves, most prominently al poles, near each of which a residual body lies between inner membrane and second shell. Acanthor within egg averages 0.076 by 0.026. USNM Nos. 72684, 72701, 72702. 146

Host records: Host Locality Record Colaptes auratus Maryland Van Cleave (1918) Corvus brachyrhynchos, Gallus gallus, Turdus migratorius New Jersey Jones (1928) , Dumella carolinensis, New Jersey r Hvlocichla sp. , Washington, Cuvillier (1934) D.C. Hvlocichla guttata, New York Pipilo ervthropthalamus, New Jersey Quiscalus quiscula, Kentucky Sturnus vulgaris, New Jersey Turdus migratorius Mich., Penn. ,Van Cleave (1942) Va. Toxostoma rufrum Ohio Chlandler and Rausch (1948) Sturnus vulgaris USA Boyd (1951) Turdus migratorius Alaska Van Cleave and Williams (19 51) Turdus migratorius Washington Werby, In Van Cleave and Williams (1951) Ammospiza maritima N . Carolina Hunter and Quay i C1953) Cvanocitta cristata Washington, Jones, In Boyd e±_ al, D.C. (1956) Turdus migratorius Colorado Schmidt (196 3) Colaptes cafer, Colorado Meleagris gallopavo Expt. in­ Schmidt and Olsen * fection (1964) ■ Sturnus vulgaris Colorado Ballard and Olsen (1966) Sturnus vulgaris, Turdus migratorius Virginia Holloway (1966) Turdus migratorius New York Howell, In Holloway (1966) Ixoreus naevius Oregon, Schmidt and Neiland Wash. (1966) Pica pica hudsonia Utah Todd, Ernst and Hammond (1967) Blarina brevicauda New York Nickol and Oetinger (1968) Sturnus vulgaris, Turdus migratorius Newfoundland,Threlfall (1968) Can. Corvus brachyrhynchos N . Carolina Hendricks, Hartiema, and Miller (1969) 147

Host Locality Record Crex crex Czechoslovakia Macko (1969) Sturnus vulgaris Kansas Rodrick and John­ son C1971) Agelaius phoeniceus, Quiscalus quiscula Ohio Stanley and

•i Rabelais (1971) Sturnus vulgaris California Vincent (1972) Dumatella carolinensis Ohio Cooper (1974) Quiscalus quiscula Ohio Buck, et al. (1975) Agelaius phoeniceus, Cyanocitta cristata, Molothrus ater, Quiscalus quiscula, Passer domesticus, Turdus migratorius Ohio Present Study

Discussion: Plagiorhynchus formosus is the most frequently recorded acanthocephalan parasite of North American passerine birds. The distribution i of this helminth spans the continent. Schmidt and Kuntz (1966) stated that P. formosus is a parasite of North American representatives of the avian family Turdidae. A perusal of the records indicates a much wider host range. The recording of P. formosus from a rail (Crex crex) in Czechoslovakia is anomalous and impossible to explain. The large number of records from Robins (family Turdidae) is probably a reflection of their obvious availability. This availability is illustrated by the fact that more accidentally killed Robins from the Ohio State University campus have been brought to this author for 1M-8 examination than any other host species. Star­ lings (family Sturnidae) rank second. Hosts other than Robins seem equally susceptible to infection. This study records P. formosus from Brown-headed Cowbirds and House Sparrows;for the;first time. Life cycle: Details of the developmental cycle have been observed experimentally by Schmidt and Olsen (196H). Eggs are passed in the feces of the host. When eaten by terrestrial isopods (Armadillidium vulgare and Porcellio scaber) the enclosed acanthor escapes and penetrates the intestinal wall. The acanthor remains dormant in the intestinal wall of the isopod for 15 to 25 days. The acanthor subsequently migrates into the hemocoel and develops within 2 days to form the acantella stage. In the hemocoel, the larval acanthocephalan becomes encysted and continues development to form the cystacanth stage within 30 to HO days post infection. The definitive host becomes infected when infected isopods are ingested.

Prevalence and Intensity of Plagiorhynchus formosus in Passerine Birds from South Bass Island: 1 149 No. Range Infected Birds/No. Exam. Birds of Mean Adults Juveniles Exam- Preva- Inten-Inten- Infect-/No. Infect-/No. Host ined lence sity sity ed Exam. ed Exam. Agelaius 50 10 1-4 1.8 0 (22) 5 (28) phoeniceus l Cyanocitta 50 30 1-20 5.9 15 (50) 0 (0)_ cnstata Molothrus 50 2 1 - 1 (46) 0 (4) ater Quiscalus 50 56 1-10 3.2 4 (13) 24 (37) quiscula Passer 30 2 1 - 0 (.5) 1 C 2 S ) domesticus Sturnus 50 42 1-8 1.9 0 (1) 21 C49) vulgarTs Turdus 50 62 1-47 5.0 12 (19) 19 (31) migratorius DISCUSSION

A. General Consideration of the Helminth Fauna of Passerine Birds I. Comparison of Previous Studies and Present Study. Most of the reports of helminth parasites of passerine birds have resulted from surveys for selected parasites and from regional surveys of avian helminths. The records of helminth parasites reported from Red-winged Blackbirds were tabulated by Cooper and Crites (1974b), from Blue Jays by Cooper (1973), from Brown-headed Cowbirds by Cooper, et^ al. (19 73), from Common Grackles by Cooper and Crites (1974a), from House Sparrows by Cooper and Crites (1974d), from Starlings by Hair and Forrester (1970) and Cooper and Crites (1975b), and Robins by Cooper and Crites (1975a). This writer's introductory statement that helminth parasitism in these birds has been little investigated is illustrated by the numbers of new host records that have resulted from this study. The accompanying Table 1 summarizes previous studies, the present, and the resulting number of new host records. As a result of this study, the diversity of helminths which can utilize these species of

150 1 5 1 birds as definitive hosts is considerably greater than previously known.

Table 1. Summary of Past and Present Studies of Helminth Parasites in Selected Passerine Birds

New Host Trema- Acantho- Records toda Cestoda Nematoda cephala

Red-winged Blackbirds Previous Studies - 10 4 9 4 Present Study 9 4 3 5 1 Blue Jays Previous Studies - 13 2 11 1 Present Study 17 9 3 12 1 Brown-headed Cowbirds . Previous Studies 7 4 6 1 Present Study 8 3 2 8 1 Common Grackles Previous Studies - 9 2 10 2 Present Study 9 4 4 6 1 House Sparrows Previous Studies - 29 12 13 5 Present Study 3 1 0 2 2 Starlings Previous Studies 27 11 30 11 Present Study 5 4 3 4 1 Robins Previous Studies — 8 10 11 5 Present Study 2 2 3 6 1

II. Sites of Infection. . Helminths parasitize practically all the organ 152 systems of passerine birds. However, the vast majority parasitize the alimentary canal and its associated organs. Cestodes and acanthocephala are restricted to the intestines of these birds. Trematodes are almost entirely restricted to the intestine, gall bladder and liver. The only ex­ ception in this study is a species of trematodes infecting the ureters of the kidney. Nematodes infect the widest range of tissue sites. Primarily, nematodes infest tissue sites in the alimentary canal. Other tissue sites utilized by nematode parasites in these birds include the circulatory and nervous systems and the orbits of the eyes. III. Infections in Juvenile and Adult Birds. The species of helminths removed from avian i hosts during the present study may be arranged into five groups according to their distribution and abundance in juvenile and adult birds: 1. Species rare in juveniles and absent in adults. (3) Echinostoma revolu±um Prosthogonimus macrorchis Mediorhynchus grandis 2. Species frequent or fairly frequent in both juveniles and adults. (8) Conspicuum icteridorum Hvmenolepis farciminosa Capillaria exilis Capillaria ovopunctatum Dispharynx nasuta Porrocaecum ens icaudatum Syngamus -trachea Plagiorhynchus formosus Species frequent or fairly frequent in a single host species and rare or absent in other host species. (5) Lutztrema microstomum (Blue Jays) Lutztrema monenteron (Robins) Choanotaenia iola (Starlings3 Robins) Dilepis undula (Robins) Microtetrameres spiculata (Blue Jays) Species relatively infrequent or rare in both juveniles and adults. (7) Plagiorchis noblei Ornithodendrium imanensis Anonchotaenia globata Choanotaenia musculosa Orthroskjabinia rostellata Capillaria contorta Chandlerella quscali Species infrequent or rare in adults and absent in juveniles. (16) Echinuscodendrium echinus Leucochloridium dryobatae 1 5 4 Leucochloridium variae Leucochloridium vireonis Leucochloridium sp. Lyperosomum oswaldoi Tanaisia zarudnyi Zonorchis alveyi Acuaria nebraskensis Acuaria quiscula Acuaria sp. Diplotriaena bargusinica Diplotriaena tridens Microtetrameres centuri Microtetrameres helix Microtetramere s sp. Oxyspirura petrowi Splendidofilaria algonquinensis Tetrameres americana Thirty-nine species of helminth parasites were removed from the 330 birds, 174 juveniles and 156 adults, examined during this study. Fourteen species of trematodes were removed. Of the 14, two species of trematodes occurred as single specimens from separate juvenile hosts (Group 1). Nine species of trematodes were removed from adult birds only (Groups 3, 5). Three species of trematodes occurred in both juvenile and adult birds (Groups 2, 4). The 6 species of cestodes removed from the birds examined 155 during this study were common to both juvenile and adult birds (Groups 2, 3, 4). Ten species of nematodes were common to both juvenile and adult birds (Groups 2, 4). One of the two species of acanthocephalan was encountered as

i four specimens in a single host (Group 1). The other species of acanthocephalan was common to both juveniles and adults (Group 2). Overall, 17 species of helminth occurred in both juveniles and adults, 19 in adults only and 3 in juveniles only. Naumov (1963) stated that the parasitic fauna of the definitive host undergoes substantial change as the host ages. With increasing age, the usual pattern involves an increase in the number of parasite specimens per indi­ vidual host, an increase in the number of host individuals infected, and an increase in the number of species of parasites utilizing the definitive host population(s). In contrast, separate studies of helminth parasites in passerine birds from the United Kingdom, Owen and Pemberton (196 2) and James and Llewellyn (1967) observed greater intensities of helminths in juvenile than in adult birds. In the eight (group 2) species of helminths which occurred frequently or fairly frequently in the birds examined during this study, juvenile birds displayed higher intensities of infection of a given helminth parasite than adult birds. The greater susceptibility of young animals and an increased resistance to infection with age has been demonstrated experimentally by Dobson (1962). In general, young birds seem susceptible to a wider range of helminths than adults, because of a less well-developed physiological resistance and perhaps because they have slightly wider feeding habits. The results of this study indicate that Naumov’s conclusion of the prevalence and intensity of helminth infections increasing with the age of the host is not true for passerine birds. Naumov’s conclusion that the number of species of helminth parasite to be found utilizing adults is larger than that utilizing juveniles is borne out to some extent in this study. This conclusion is evident in the observation that 16 species of helminths were infrequent or rare in adults and absent in juveniles i (Group 5). It is also reflected in the species that were relatively infrequent or rare in both juveniles and adults (Group MO. In this latter instance, intensities of in­ fection in both adults and juveniles were low. Apparently a variety of helminth infections accumulate in these avian hosts resulting in a greater variety of helminth species occurring in older birds. A large proportion of the initial infection in a juvenile is lost as the host ages. The infections in adults are of low intensity. These low in­ tensity infections in adults probably result from two sources. One source is the long-lived residual infection that persists from the initial large infection in the juvenile bird. The other source is the occasional ingestion 1 5 7 or exposure to infective dispersal agents of parasites which frequently infect other definitive hosts but which have the capacity to successfully utilize a broad range of host species. One can attribute all of the infections that occurred with rarity in juveniles and that were absent in adults (Group 1) to the latter source of infection. The 7 species that occurred infrequently in both juveniles and adults (Group 4) are impossible to attribute absolutely to one or the other of the stated sources of infection. The low intensity of infection in juveniles indicates that contact with the dispersal agent is limited. In these instances, species of definitive hosts other than those

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Juveniles Parasite % sity sity N =20 N = 02 N =16 N = 12

Treraatoda Conspicuum icteridorum 48 1-8 2,4 11 02 04 07 Leucochloridium variae 2 (11) 11 01 00 00 00 Plagiorchis noblei 6 1-2 1.3 01 01 00 01 Zonorchis alveyi 2 1 0 01 00 00 00

Cestoda Anonchotaenia globata 22 1-11 4 08 00 01 02 Choanotaenia musculosa 6 1-9 4.7 01 00 00 02 hymenolepis farciminosa 2 3 3 01 00 00 00

Nematoda Capillaria ovopunctatum 4 3-9 6 01 00 00 01 Dispharynx nasuta 8 1-7 4.5 00 01 02 01 Microtetrameres centuri 2 1 1 01 00 00 00 Porrocaecum ensicaudatum 4 2-6 4 00 00 01 01 Syngamus trachea 4 1 Pair 1 Pair 01 00 00 01

Acanthocephala Plagiorhynchus formosus 10 1-4 1.8 00 00 05 00

Total number of Birds Infected in Each Class 84 •* 17 02 12 11

M cn ID TABLE 3. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN FIFTY BLUE JAYS

Range Infected Birds _____ Preva- of Mean Adults Juveniles lence Inten- Inten- .Males Females Males Females Parasite % sity sity N =25 N = 25 N =00 N = 00

Trematode E ch inuscodendrium 2 2 04 4 4 01 01 00 00 00 00 00 echinus Leucochloridium 2 2 12 12 12 00 00 01 01 00 00 00 dryobatae Leucochloridium variae 4 4 01-12 06.5 06.5 02 02 00 00 00 00 00 Leucochloridium vireonis 2 03 3 3 00 00 01 01 00 00 00 Lutztrema microstomum 60 60 03-19903-199 24.1 24.1 13 13 17 17 00 00 00 Lyperosomum oswaldoi 2 2 01 1 1 00 00 01 01 00 00 00 Ormthodendrium imanensis 26 01-18 10.4 10.4 07 07 06 06 00 00 00 Tanaisia zarudyni 2 2 15 15 15 01 01 00 00 00 00 00 Zonorchis alveyi 10 10 02-04 03.0 03.0 03 03 02 02 00 00 00

Cestoda Choanotaenia iola 6 6 01-04 01-04 02.0 02.0 03 03 00 00 00 00 00 Hymenolepis~farciminosa 16 16 01-35 01-35 ■■ 07.007.0 05 05 03 03 00 00 00 Orthoskrp abinia 8 8 01-04 01-04 02.5 02.5 02 02 02 02 00 00 00 rostellata

Nematoda Acuaria nebraskensis 2 01 1 00 01 Acuaria quiscula 4 01-02 01.5 01 01 Acuaria sp. 14 01 1 04 03 Capillaria contorta 6 01 1 00 03 — H (7) o TABLE 3. (Cont'd.)

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N =25 N = 25 N =00 N = 00

Nematoda (con.) Capillaria exilis 12 01-02 01.3 04 02 Chandlerella quiscali 2 04 4 01 00 Diplotriaena tricuspis 14 01-Q8 03.8 03 04 Dispharynx nasuta 12 01-05 03.0 04 02 Microtetrameres helix 4 02-04 03.0 01 01 Microtetrameres spiculata 22 01-05 02.5 06 05 Microtetrameres sp. 4 01 1 02 00 Oxyspirura petrowi 4 02-04 03.0 01 01 Porrocaecum 14 01-04 01.7 03 04 ensicaudatum Syngamus trachea 16 01-07pr. 03.8 pr. 05 03

Acanthocephala Plagiorhynchus formosus 30 01-20 05.9 05 10

Total Number of Birds Infected in Each Class 98 *■ 24 25

H CD H TABLE 4. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN FIFTY BROWN-HEADED COWBIRDS

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N =43 N = 03 N =04 N = 00

Trematoda Conspicuum icteridorum 4 01-05 03.0 02 00 00 00 Plagiorchis noblei 4 01-25 13,0 01 00 01 00 Tanaisia zarudnvi 2 04 - 01 00 00 00

Cestoda Anochotaenia elobata 22 01-12 04.5 10 00 01 00 Orthoskiabihia rostellata 4 02-06 04.0 02 00 00 00

Nematoda

Capillaria ovopunctatum 4 01 — 02 00 00 00 Chandlerella auiscali 4 01 - 01 01 00 00 Diplotriaena bareusinica 2 02 - 01 00 00 00 Dispharvnx nasuta 12 01-02 01.7 03 01 02 00 Oxvspirura petrowi 2 01 - 01 00 00 00 Splendidof ilaria 2 08 - 01 00 00 00 algonquinensis Svngamus trachea 2 01 pr. - 00 00 01 00 Tetrameres americana 2 01 — 01 00 00 00

Acanthocephala Plaeiorhvnchus formosus 2 01 - 01 00 00 00 Total Number of Birds Infected in Each Class 48 - - 18 02 04 oo S TABLE 5. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN FIFTY COMMON GRACKLES

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N = 5 N = 8 N =21 N = 16

Trematoda Conspicuum icteridorum 46 01-13 4.5 03 07 08 05 Ornithodendrium imanensis 4 01-03 02.0 00 00 02 00 Plagiorcnis noblei 4 02 02.0 00 00 00 02 Prosthogonimus macrorchis 2 1 1 00 00 00 01 Cestoda Anonchotaenia globata 8 01-15 07.8 00 01 02 01 Choanotaenia musculosa 10 01-03 02.2 01 02 01 01 Hyraenolepis farciminosa 8 01-03 02.0 01 00 02 01 Orthoskrjabinia rostellata 4 01-03 02.0 00 00 02 00 Nematoda Capillaria exilis 6 01-14 06.0 00 00 01 02 Capillaria ovopunctatum 4 01-25 08.4 02 02 07 11 Chandlerella quiscali 12 01-04 02.0 02 03 01 00 Dispharynx nasuta 28 01-13 06.7 00 01 09 04 Porrocaecum ensicaudatum 46 01-10 02.3 01 02 11 09 Syngamus trachea 72 1-8 pr-. 03.2 pr. 01 04 19 . 12 Acanthocephala Plagiorhynchus formosus 56 01-10 03.2 01 03 12 12

Total Number of Birds Infected in Each Class 96 05 08 21 14 163 TABLE 6. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN THIRTY HOUSE SPARROWS

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N =03 N = 02 N =14 N = 11

Trematoda Conspicuum icteridorum 3.3 01.0 00 00 01 00

Nematoda Dispharynx nasuta 3.3 11.0 - 00 00 01 00 Syngamus trachea 6.7 3 pr. 3 pr. 00 00 01 01

Acanthocephala Mediorhynchus grandis 3.3 04.0 - 00 00 01 00 Plagiorhynchus formosus 3.3 01.0 - 00 00 01 00

Total Number of Birds Infected in Each Class 13.3 00 00 03 01 9I t + TABLE 7. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN FIFTY STARLINGS

Range Infected Birds Preva­ Of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N =00 N = 01 N =24 N = 25

Trematoda ■ Conspicuum sp. 8 1—3 1.5 00 00 03 01 Echinostoma revolutum 2 1 - 00 00 00 01 Ornithodendrium imanensis 8 3-161 80.5 00 00 03 01 Plagiorchis noblei 2 32 - 00 00 00 01 Cestoda- Choanotaenia iola 42 1-14 4.3 00 00 12 09 Choanotaenia musculosa 12 3-10 6.7 00 00 03 03 Hymenolepis farciminosa 38 1-15 5 00 01 08 10

Nematoda Capillaria exilis 16 1-08 4 00 01 03 05 Capillaria ovopunctatum 46 1-40 9.7 00 00 12 11 Dispharynx nasuta 24 1-08 3.2 00 00 04 08 Porrocaecum ensicaudatum 32 1-14 2.5 00 01 07 08

Acanthocephala Plagiorhynchus formosus 42 1-08 1.9 00 00 10 11

Total Number of Birds Infected in Each Class 96 •* 00 01 23 24

H cn tn TABLE 8. PREVALENCE AND INTENSITY OF HELMINTH PARASITES IN FIFTY ROBINS

Range Infected Birds Preva­ of Mean Adults Juveniles lence Inten­ Inten­ Males Females Males Females Parasite % sity sity N =09 N = 10 N =13 N = 18 • Trematoda Leucochloridium sp. 2 01 1 01 00 00 00 Lutztrema monenteron 20 1-11 5.9 05 05 00 00 Cestoda Choanotaenia iola 10 1-07 2.6 00 00 02 03 Dilepis undula 42 1-84 15.3 04 04 05 08 Hymenolepis farciminosa 10 1-02 1.2 00 01 00 04 Nematoda Capillaria contorta 2 01 1 00 00 00 01 Capiliaria exilis 70 1-35 8.6 04 05 10 16 Capillaria ovopunctatum 12 2-15 4.7 00 00 02 04 Dispharynx nasuta 28 1-26 7 00 01 06 07 Porrocaecum ensicaudatum 34 1-08 2.2 03 00 06 08 Syngamus trachea 42 1-08 pr. 3.2 pr. 03 02 05 11 Acanthocephala Plagiorhynchus formosus 62 1-47 5 05 07 09 10

Total Number of Birds Infected in Each Class 96 - 08 09 13 18

H o> O) 167 Most of the helminths with a prevalence of less than 10% similarly utilize invertebrates for development and dispersal. In these instances, one must conclude that contact between intermediate host and these species of i birds is minimal. The invertebrates being utilized by this group of helminths are either not present on South Bass Island or are seldom encountered and/or selected as a food source by these species of birds. Among’free-living organisms, the pattern of a relatively few common species associated with many rare species is characteristic of biological community structure everywhere (Odum, 1971). In this study, the total number of helminth specimens collected from the 330 birds was 4333. i Of the 41 taxa identified, 27 were represented by 30 or fewer specimens. Of the latter 27 taxa, 18 were represented by fewer than 10 specimens. The pattern of a relatively few common species associated with more rare species is reflected in the species of helminth parasites utilizing a portion of the biological community on South Bass Island. The quantitative descriptions of species abundance relationships vary widely. One approach is comparisons based on diversity indices, which are ratios or other mathematical expressions of species importance relationships. Sorensen's (1948) Index of Similarity is best suited for comparing the helminth faunas of the birds examined during this study. To investigate the degree of similarity of the 168 helminth faunas of each pair of host species, all possible combinations of pairing were tested. The system of notation is as follows:

2C . . . S = -r—z— r—x 100 where A = number of helminth species m host species A B = number of helminth species in host species B C a number of helminth species common to both host species.

The obtained values for every two species compared were arranged in a trellis diagram (Fig. 1) so that the host species having the most similar parasites faunas are placed together. The results reveal a number of interesting re­ lationships. The Blue Jay is not a member of this feeding association. Blue Jays have a relatively low index of similarity with all of the other species examined during this study. Although observed in the same areas as the mixed feeding flocks, the low level of parasitism in the House Sparrow is reflected in its low index of similarity with all other host species. The three species of icterids show relatively high levels of similarity, especially the Red-winged Blackbird and the Common Grackle. Their para­ site faunas are similar, but differ in degree of prevalence and intensity. The three widely acknowledged omnivores in this association show interesting relationships. The index 169 reveals that the helminth faunas of the Common Grackle and Starling are quite similar and that both differ to a marked degree from the Robin* This can be attributed to the Robin's differential feeding on earthworms* The two former species select earthworms as a food resource less commonly and are parasitized by helminths utilizing this invertebrate host to like degree (see Tables 5# 7, and &).

B* The Concept of Landscape Epidemiology and a Modified Concept of the Nidalitv of Disease The conceptual framework of ecological parasitology in the USSR was established in the 1930's* Its main aim was and continues to be the study of the dependence of a para­ site fauna on changes in the external environment of the host, on changes in the physiological state of the host it­ self and the consideration of the host as the habitat of the parasite (Bykhovskaya-Pavlovskaya, 1967)* The rudiments of the concept of landscape epidemiology were first presented by Pavlovsky in a course on human parasitology in 1934* It reached its fullest statement in Pavlovsky's (1964) monograph on the Natural Nidalitv of Transmissible Diseases* In essence, the canopy concept of landscape epidemiology and the concepts which compose it are quite simple and straight forward* It is difficult to understand initially in that it possesses a distinct vocabu­ lary that must be assimilated if one is to think effectively Common Crackle 74 Starling 64 71 Red-winged Blackbird 48 58 52 Robin 31 52 38 62 Brown—headed Cowbird 36 43 37 37 45 Blue Jay 20 42 35 44 35 40 House Sparrow v O£ . rt >» F-t tj ‘ a*H J-j G ® t* - 3 *H & *H *H P* U ,0 O r O r* O rta d Q) o £ 05 ^u *3cd G Fi co r-4 O aj -P « PQ rH CO

O Ti u a) PQ 05

Figure 1* The Similarity Index of Helminth Faunas of Seven Avian Host Species from South Bass Island, Ottawa County, Ohio, 1968-1971* (Sorensen, 194&) in terms of the concept# Nonetheless, the concept is most useful to the parasitologist conducting field work. It provides a vocabulary and conceptual framework which makes organization of one's thoughts and communication in terms of the prevalence and intensity of parasite populations 1 within a biological community much easier than any other in the discipline of parasitology. The concept developed as part of the study of trans­ missible zooanthroponoses, i.e. the study of diseases which are transferred from animals to man by vectors. Vectors are various invertebrate organisms which can acquire the disease agent of the corresponding disease while feeding on an infected animal and transmit it to another host, including man, later. In contrast, nontransmissible diseases do not require a vector and anthroponoses are diseases characteristic of man alone. The essence of the concept in Pavlovsky's (1964) words is: It is characteristic of a series of trans­ missible diseases that their sources (nidi) exist in nature either untouched by man or more or less altered (modified) by conscious or unconscious human activity in past his­ torical times or in the contemporary period. The nidi of such diseases are called natural ones, while the corresponding diseases are called natural nidal. The concept of the natural nidality of transmissible diseases is characterized by the sequential transmission of infective agents from infected definitive hosts to inter­ mediate invertebrate hosts or vectors. Subsequently the Intermediate hosts transmit the infective agent to yet another definitive host. The new definitive host may be the same or another host species which may or may not have been previously infected in the same manner. More im­ portantly, this kind of transmission occurs only in areals where abiotic factors are favorable. The disease nidus exists in those areas in which invertebrate vectors, in­ fected definitive hosts and potential definitive hosts aggregate under favorable climatic, edaphic, etc., condi­ tions. Such a favorable landscape acts as a biogeocenose for the establishment and continuation of the nidus of infection. The term biogeocenose was coined by Molius in 1377 (Audy, 195#)• Ecologists in the United States have developed a variety of terms quite similar in meaning. Elton and Miller,s (1954) discussion of a biological com­ munity is the best example to use for comparative purposes. The difference is that a biocenose, i.e. a naturally linked assemblage of species, is a term historically associated with disease (Audy, 195#)» lb is a concept frequently used by epidemiologists. The concept of nidality is modified when limited exclusively to endoparasitism in animals other than man. Endoparasites for the most part have either direct life cycles in which the infective agents in the form of eggs or larvae are transmitted directly from animal to animal or 17 3. indirect life cycles which use invertebrate intermediate hosts for development and dispersal of infective stages* Vectors* per se. are rare but not totally absent in most situations of this nature* Audy (195&) has reviewed the various modifications of the concept of nidality which Pavlovsky pioneered and widely applied in reference to zooanthroponoses*

C* The Nidalitv Index It was concluded that factors which control or deter­ mine the spatial distribution of a helminth species in this avian association are principally dependent on the feeding activities of each species of bird* The existing methods attaching numerical value to the found association all have their drawbacks* Sometimes the methods used do not give any distinction between complete and absolute association (Pielou, 1969)* Sometimes the methods contain means like Sorensen*s QS (194$) which are unsuitable when the differ­ ences of the absolute frequencies are large* A standard way to examine the association between two species of free-living animals is to study their presence or absence in a number of samples (Dice* 1945)* In the present analysis* use was made of every bird examined* Blue Jays are not a part of the association and were not in­ cluded in the data analysis* To investigate whether some parasite species were more often associated than others* the 174 10 species considered to have natural nidi of infection on South Bass Island were tested. They are listed in an association diagram (Figure 2.). The system of notation is as follows: Species B Presence Absence onQ^ ae a Presence a b a + b = m Species A Absence 0 d c + d = n

All data in the 2x2 table above have not been used to construct an index. No consideration was taken to the negative association, to the term d. This has no effect on the positive side when as in this case N always is 2S0. The remaining three terms formed the association index, M* here­ after referred to as the nidality or association index, in the following manner (Dice, 1945)•

M = luxaMax. Im * § x x 100 or M = min 1 ^m,rj----- X 100

The denominator is not the same as Dice, but as at the 2x2 table above, the formula is also multiplied by 100. The obtained values for every two helminth species compared were arranged so that species which have high association with one another are placed together. Regroup­ ings must be done until the highest possible associations of the groups were found (Agrell, 1945)* The result (Fig. 2) showed all species had a low 1 7 5 mutual association# The low association between helminth species utilizing common species of intermediate hosts indicates that a single intermediate host species is likely to harbor infective stages of only one species of parasitic t helminth# The highest mutual association value (41$) is1 between Capillaria exilis and Dilepis undula. This value indicates that the same earthworms may be utilized by both species of helminths# The life cycle of C# exilis is un­ known# Other species of Capillaria are known to utilize earthworms as transport hosts# Robins serve as the principal hosts for both helminths# The relatively high mutual association indicates Robins select earthworms as a food source to a much greater extent than do the other avian species of the association of definitive hosts# This study is the first to develop an index of simi­ larity or an index of association for host-parasite data from natural populations# The problems of host collection, parasite collection and preparation, and resolution of systematic questions which arise in a study of this nature have been more than adequate to occupy the minds of previous workers conducting similar extensive studies# The index of association values produced in this study indicate an important parasitological phenomena# Although more than one species of helminth may utilize a given species of in­ vertebrate intermediate host for development and dispersal in a community, any one specimen of such an intermediate 176

Plagiorhvnchus formosus 15 Dispharvnx nasuta 12 24 Svngamus trachea 12 4 4 Conspicuum icteridorum

5 25 15 13 Porrocaecum ensicaudatum 32 17 29 9 33 Capillaria ovopunctatum 34 5 3 1 10 6 Hvmenoleois farciminosa 3 4 14 0 11 11 20 Capillaria exilis 4 1 4 2 0 1 1 6 Choanotaenia iola 4 2 4 5 0 9 3 3 Dileois undula

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Figure 2* Trellis diagram of* 230 birds examined* The nidality index in percent is based on presence or absence of 10 species of parasitic hel­ minths in passerine hosts from South Bass Island, Ottawa County, Ohio, 1963—1971• (Dice, 1945). 177 host species is likely to harbor the larval forms of but a single species of helminth* This phenomena is revealed by the low association values of C. icteridorum. D. nasuta and P. formosus which utilize terrestrial isopods and the values of P. ensicaudatum, S. trachea, and D. undula which utilize terrestrial oligochaetes*

D. A Helminth Biocenose Involving Passerine Birds on South Bass Island A natural nidus is a well developed complex of defini­ tive hosts, intermediate hosts and parasitic agents in a defined biological community. The theory of natural nidality states that the infective dispersal agents of any parasite population are not distributed randomly in a community. On the contrary, if a parasite population is to be maintained through time, the infective agents must be aggregated at points where probability of host contact is high. Foci of infection are those points in the space-time continuum where hosts and infective agents are aggregated and transmission is maximized. An assemblage of parasite populations is being supported by the.association of avian hosts examined during this study. This assemblage of parasite populations make up a biocenose. The species of helminths which occur fre­ quently or fairly frequently in both juvenile and adult birds (Group 2) have, in all likelihood, foci of infection on South Bass Island# The two species of helminths which occur frequently in Juveniles and adults of a single or two host species likewise have foci of infection on the Island (Group 3: C. iola. D. undula)# Each parasite population has a distinct, although often overlapping, nidus in the biological community formed within the limits of the Island# This argument is based on the observation that the preva­ lence and intensity of infection of these 10 helminth species was much greater in recently fledged Juveniles from the Island than in adult birds# The 7 species of helminths which occurred infrequently in both Juvenile and adult birds (Group k) have foci of infection in the western basin region of Lake Erie but hot on South Bass Island# After fledging, Juveniles move between the Bass Islands and the mainland with increasing frequency as the summer months progress (Miskimen, 1970)# Those species of parasites which occurred infrequently or rarely in adult birds and were absent in Juvenile birds must have been acquired in foci of infection in other parts of the definitive hosts geographical ranges. The three species of helminths which occurred rarely in Juveniles only must have foci of infection in the western basin region but not on South Bass Island. Two of the latter three species are commonly reported from non-passeriform hosts# The indiscriminate feeding of Juvenile birds on infected intermediate hosts rarely utilized as a food 179 resource by these birds probably resulted in these rare in­ fections* A total of 10 species of helminth parasites has been identified as having foci of infection, i.e* established life cycles, on South Bass Island* Host species for each parasite population are- listed below in descending order of importance* Prevalence of infection in each host species if3 included* In most instances, one or two host species are primarily responsible for the maintenance of a given helminth species* The remaining host species exhibit relatively low levels of parasitism for a given helminth* In the latter instance, the accumulative effect of these low infection levels is con­ siderable* Trematoda Conspicuum icteridorum — Red—winged B'.lackbird (4#&) Grackle (465S), Brown—headed Cowbi]rd (4#)t Starling (immature specimens 6$), House Sparrow (immature specimens 3*39&) Cestoda Choanotaenia iola — Starling (425S), ffcobin (10?6), Blue Jay (6fa) Dilepis undula — Robins (427S) Hvmenolepis farciminosa — Starling (3(Wo 8?S),, Blue Jay (16$), Robin (10?S), Grackle (#5&)r Red—winged Blackbird (2?o) Nematoda Capillaria exilis — Robin (70$) , Starling (1.8$) § Blue Jay (12$) 9 Grackles (6$) Capillaria ovopunctatum — Starling (1+6$) $ Grackle (l+l+$)f Robin (12$) f Brown-headed Cowbird (1+$), Red—winged Blackbird (l+$) Dispharvnx nasuta — Grackle (28$), Robin (28$) 9 Starling (21+$)9 Blue Jay (12$) 9 Brown-headed Cowbird (12$), Red-winged Blackbird (8$), House Sparrow (3*3$) Porrocaecum ensicaudatum — Grackles (1+6$), Robins (3k$), Starlings (32$), Blue Jays (11+$), Red­ winged Blackbirds (t+$) < Svngamus trachea — Grackle (7255), Robin (U2$), Blue Jay (16$), Red-winged Blackbird (i+$), House Sparrow (3*3$), Brown—headed Cowbird (2$)* Acanthocephala Plagiorhvnchus formosus — Robin (62$), Grackle (56$), Starling (1+2$), Blue Jay (30$), Red- winged Blackbird (10^5), House Sparrow (3*3$) 9 Brown—headed Cowbird (2$)»

The natural nidus of definitive host, intermediate host, and parasite species have been deduced for 6 of these com­ plexes* The trematode Conspicuum icteridorum. the nematode Dispharvnx nasuta. and the acanthocephalan Plagiorhvnchus formosus utilise terrestrial isopods (Armadillidium 181 quadrifrons. A. vulgara. Oniscus acellus. Porcellio jg.gabffjJ for development and dispersal* These are common isopods in open areas such as lawns on South Bass Island (Beatty, 1 9 7 0 ) * The cestode Dilepis undula and the nematodes Porrocaecum ensicaudatum and Svngamus trachea utilize terrestrial oligochaetes (Lumbricus spp*) for development and dispersal* Common food resources in the form of terrestrial iso­ pods and oligochaetes are being utilized by several helminth populations for development and dispersal of intermediate life-cycle stages* Wynne—Edwards (1962) stated that mixed species flocks such as those that form on South Bass Island form expressly because there is competition for the same food resources between them* The distribution and abundance of parasite populations which utilize the same invertebrate hosts for development and dispersal confirms the contention that the avian species comprising this association are utilizing common food resources* The competition between these avian species for common food resources helps to maximize the transmission of those helminth populations which utilize the invertebrate food resources* In addition, the aggregation of birds into mixed feeding flocks insures a heavy infection of the feeding grounds with agents (ova) infective for the invertebrate populations* SUMMARY

Red-winged Blackbirds (Agelaius nhoeniceus). Blue Jays (Cvanocitta cristata), Brown—headed Cowbirds (Molothrus ater), Common Grackles (Quiscalus quiscula). House Sparrows (Passer domesticus), Starlings (Stumus vulgaris). and Robins (Turdus migratorius) were collected on South Bass Island, Ottawa County, Ohio, and examined for helminth parasites. A total of 4333 specimens of helminth parasites was re­ moved from the 330 passerine birds collected. Forty—one taxa of helminth parasites were identified. Fifty—three new host records resulted. Five distinct groups of helminth species were discerned in the pattern of prevalence and intensity of helminths in juvenile and adult birds examined. Sorensen's Index of Similarity quantitatively described the species abundance relationships of the helminths encountered. The helminth fauna of the Common Grackle and the Starling was most similar, according to the Index. The Nidality Index constructed quantitatively describes the biocenose revealed. This index indicates that most specimens of infected intermediate hosts harbor the larval forms of a single species of helminth. The concept of landscape epidemiology is applicable to the study of helminth populations in biological com­ munities. A modified concept of nidality of disease delineates' a biocenose of six nidi of infection involving the species of passerine birds examined. The biocenose discerned on South Bass Island is composed of ten species of helminths. LITERATURE CITED

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* 1974b* The helminth parasites of the red— winged blackbird from South Bass Island, Ohio, in­ cluding a check list of the helminths reported from this host* J* Wildl* Dis* 10: 399-403* * 1974c* A study of the helminth parasites of overwintering red—winged blackbirds (Agelaius phoeniceus) from Laurel, Maryland. J. Parasit* 60: 962* . 1974d* Helminth parasitism in juvenile house sparrows, Passer domesticus (L*), from South Bass Island, Ohio, including a list of helminths reported from this host in North America* Ohio J* Sci. 74:388-389* * 1975a* A check list of the helminth parasites of1 the robin, Turdus migratorius Ridgway* Am* Midi* Nat* In press* * 1975b* * Additional check list of the helminths starling (Sturnus vulgaris L*)* Am* Midi* Nat* In press* Cram, E* B. 1927a* Bird parasites of the nematode sub­ orders Strongylata, Ascaridata, and Spirurata* U.S*N* M. Bull. 140: 237-244* * 1927b* New records of distribution for various nematodes. J. Parasit. 14s 70. * 1928. Nematodes of pathological significance found in some economically important birds in North America* U.S.D.A* Tech* Bull* 49* 10 pp* * 1930* Sowbug found to be host of parasite of ruffed grouse, a game bird* U*S*D*A*, Off* Rec* 965. * 1931* Nematodes (Roundworms) in quail* In Stoddard* Bobwhite Quail. New York: Scribner's Sons* 240- 2 9 6 . * 1932a* New host records for Dispharvnx spiralis* J* Parasit* 18: 303* . 1932b* Additional observations in bird hosts for Dispharvnx spiralis* J. Parasit* 18: 310* * 1932c. Recent findings in connection with parasitism of game birds. Trans* Am. Game Conf* 1 8 : 243-247* 190

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