This dissertation has been 62—3609 microfilmed exactly as received
WELKER, George William, 1923- HELMINTH PARASITES OF THE COMMON GRACKLE QUISCALUS QUISCULA VERSICOLOR VIEILLOT IN INDIANA.
The Ohio State University, Ph.D., 1962 Zoology
University Microfilms, Inc., Ann Arbor, Michigan HELMINTH PARASITES OP THE COMMON GRACKLE
QUISCALU5 QUISCULA VERSICOLOR VIEILLOT
IN INDIANA
DISSERTATION
Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University
By
George William Welker, B. S., M. A.
u _ u u u
The Ohio State University 1962
Approved by: 1'XJijdJi ~7 Adviser aenturtmeht of Zoology and Entomology Dedicated as a tribute of appreciation and admiration to ELLEN ANN, my wife, for her help and for the sacrifices which she made during the
four years covered by this study.
ii ACKNOWLEDGMENTS
The author wishes to express his sincere appreciation for all the help and cooperation which he has received from many people during the course of this study: Dr. Joseph Jones, Jr. of St. Augustine's College,
Raleigh, North Carolina; Dr. Donal Myer, Southern Illinois university;
Dr. E. J. Robinson, Jr., Kenyon College, Gambier, Ohio; Dr. Martin J.
Ulmer, Iowa State University, Ames, Iowa; and Dr. A. Carter Broad and
Dr. Carl Reese of the reading committee who helped in checking the paper for errors.
Special acknowledgment goes to two persons whose help and influence are most deeply appreciated. To Professor Robert H. Cooper, Head of the
Department of Science at Ball State Teachers College, whose sincere and continuous interest, encouragement and help made possible the completion of the work; and to Professor Joseph N. Miller, Department of Zoology, the Ohio State University, who served as my adviser. Without his help, kindness and cooperation this study could not have been done.
iii CONTENTS
Introduction ...... 1
Historical Review ...... k
Materials and Methods ...... 6
Classification and Description of Parasites ...... 10
Acanthocephala ...... 10
Platyhelminthes ...... 13
Cestoda...... 13
Trematoda...... 17
Aschelminthes...... 36
N e m a t o d a ...... 36
Observations and Results ...... ^7
Interpretation of Data ...... 56
General Discussion ...... 69
S u m m a r y ...... 7^
Appendix...... 76
Bibliography...... 9V
Autobiography...... 104
iv TABLES
Table Page
1. Published records of helminth parasites from the
Common Grackle...... ^9
2. Helminth parasites obtained from the Common Grackle
in Indiana...... 50
3« Percentage of infection found in the Common Grackle
with different species of helminth parasites in relation
to geographical regions of Indiana ...... 51
Percentage of infection of male and female grackles
with different classes and speciesof helminths ...... 52
5. Totals of Common Grackles taken in the spring and
autumn in the three regions of Indiana...... 53
6 . A composite table of the number of Common Grackles
infected with different species of helminths ...... 5^
7 . A comparison of helminth parasites in four passerine
b i r d s ...... 55
v APPENDIX ILLUSTRATIONS
Plates
I......
Map of Indiana
II......
Mediorhynchus grandis
III...... '......
Paricterotaenia parina
Orthoskrjabinia quiscali
IV......
Bracbylecithum americanum
Conspicuum icteridorum
V ......
Conspicuum macrorchis
Echinostoma revolutum
VI......
Crepidostomum cooperi
Capillaria quiscali
VII......
Splendidofilaria quiscali
Dispharynx spiralis INTRODUCTION
This is a study of helminth parasites of the Common Grackle,
Quiscalus quiscula versicolor Vieillot. It was made to determine what species of helminths infected this bird; the geographical distribution of the parasites; their relative frequencies in the spring and autumn; and their distribution in relation to the sex of the host in the state of Indiana. The grackle is a common bird in' Indiana. In the 5th edition of the A. 0. U. Check-List of North American Birds (1957) three sub-species of grackles were grouped under the name Common Grackle.
These were the Bronzed Grackle, the Purple Grackle, and the Florida
Purple Grackle. The host used in this study is the one previously known as the Bronzed Grackle. This bird is considered highly undesirable because it has become so numerous in the past few years. No previous study has been made of the helminth parasites of it, although individual reports of helminths of this bird are found in the literature.
The Common Grackle is omnivorous; however, it feeds primarily upon seeds, which include the cereal grains of the farmers of the state.
These birds do much damage to fields of grain. The author has observed fields of corn in which flocks of these birds had destroyed practically
the entire crop, often feeding on c o m while it was still green. Records
indicate that c o m makes up the greatest percentage, of the diet of this bird.
During the autumn of both i960 and 1961 the author observed flocks
of grackles flying to their night roosts in numbers estimated to be in
the thousands. Large, tall trees are favored as roosts. Residents near 2
such roosts are plagued hy the incessant chatter and odor and filth of the droppings of the birds. Because of the large numbers of these birds even in urban areas, they have come to be abhorred by city dwellers as well as the rural farmer.
Although the Common Grackle is classified as a song bird, and is therefore protected, the residents of some areas have illegally shot or
otherwise destroyed many of them. However, all efforts to route them from their chosen sites have met with little or no success. Certain
cities in Indiana have "open season" on these birds. At predetermined
times, residents, under supervision of the police department, have been
allowed to shoot them. Since it is a migratory bird, and therefore
protected by law, it was necessary for the author to obtain scientific
collecting permits from both the state of Indiana and the Federal Bureau
of Wildlife Conservation. Annual reports were made to the Regional
Director of the Bureau of Sport Fisheries and Wildlife.
In order to determine whether the parasites or their relative
frequencies varied with the season, spring and autumn were selected as
the times in which collections were made. This bird generally arrives
in early spring (March) and leaves Indiana with the onset of cold
weather, migrating into the southern states and wintering from South
Carolina to Texas. Parasites taken from birds in the early spring
likely represent infections obtained during the winter when the host was
out of the state. On the other hand parasites found in the autumn, but
not present in the spring, evidently had to be obtained locally by the
birds. 3
The author believed that a study of the geography of Indiana Justi fied a three way division of the state into north, central, and south parts (Fig. 1). In this report, the north area is referred to as the lakes region, being characterized by many lakes of various sizes. The j central region of the state, primarily an agricultural area, has few natural lakes. The southern part of Indiana is an unglaciated area and extremely hilly. Because of the nature of the terrain, this part of the state is used mostly for forestry and grazing. While some crop farming is done in the southern part, this does not occur nearly so extensively as in central Indiana.
Since it was proposed to determine whether the parasites or their relative frequencies varied with the sex of the host, it was intended that approximately equal numbers of each sex of bird would be examined, thus allowing for a check on the distribution of parasites according to season, geographical distribution, and sex. HISTORICAL REVIEW
Many studies have been conducted on avian parasites; however, only three species of passerine birds have had surveys made of their helminth parasites. These included the Blue-jay, crow, and starling.
A review of the literature has revealed only scant information con cerning the parasites of grackles, especially those of this species. In fact, no study of helminth parasites has been undertaken heretofore for the Common Grackle, although from time to time a total of seven species of worms has been reported in the literature. These come from isolated reports of parasites obtained by workers checking this host for a particular species of parasite.
Van Cleave (1918), in a study of the acanthocephala of North
American birds, reported Mediorhynchus grand!s as a new species from the
Common Grackle in Kansas, Illinois, Ohio, and Kentucky.
Denton and Boyd (1951) reported a new species of trematode,
Conspicuum icteridorum from this host in Texas, Tennessee, Ohio, Georgia, and Michigan.
Byrd (1950) reported Tanaisia bragai Santos, 193^ from the Common
Grackle in Georgia. This parasite infects the ureters near the kidneys.
Of twelve grackles examined by him, one was found to be infected with this trematode. Biis is the only record of this species in this host.
Denton (19^5)> in a study made in Georgia, described Brachylec ithum americanum, a new species of trematode, reporting an 18 percent infection in the Common Grackle. He also recovered the metacercariae of this fluke from the 12-spotted cucumber beetle, Diabrotica 5 duodecimpunctata, which he had experimentally infected.
Read (19**9) reported a new species of nematode, Capillaria quiscali, from grackles collected in Madison, Wisconsin.
Walton (1923) reported Dispharynx spiralis (Molin, 1858) Skrjabin,
1916b, a nematode from the Common Grackle in Illinois.
Von Linstow (190*0 described, as a new species, Filaria quiscali, a nematode from the cerebral hemisphere of the grackle, Quiscalus versicolor versicolor. Odetoylnbo and Ulmer (i960) reported a species, which conformed to von Linstow*s description, as Splendifofilaria quiscali (von Linstow, I90U) new combination. This was obtained from the Common Grackle in Iowa, and appears to be identical with a nematode,
Splendidofilaria ohioensis, which Jones (i960) reported as a new species from the Eastern Crow, Corvus brachyrhynchos brachyrhynchos. MATERIAL AND METHODS
This study was conducted over a period of four years. Collections were made during the spring and fall months for comparisons.
The birds were killed either with a 22 calibre rifle or a 16 gauge shotgun using maximum range birdshot. In some cases the shotgun destroyed the bird, making it of no use in the study; however, it eventually proved to be more effective with maximum kill and minimum waste of birds.
The birds were posted as soon as possible following their col lection. Some injured birds were taken into the laboratory and examined, while yet alive, for ectoparasites. Ohe eyes, ears, feathers, and anal region were examined with a hand lens. Dead birds were placed in a container with water and a detergent (Surf), being doused up and down in the detergent water. While this was being done the feathers were ruffled with a fine tooth comb. The detergent solution was then strained through a double layer of cheese cloth and the cheese cloth then was examined for ectoparasites using the binocular dissecting microscope.
Next, the feathers were removed from the abdominal region of the bird, and an incision was made in the abdomen in order to remove the entire digestive tract, which then was placed in a physiological salt solution
(Ringer's warn). The liver, gall bladder, heart, trachea, and lungs were removed and examined separately. The lungs were teased and doused
in a saline solution. Other organs such as the liver, gall bladder and heart were examined by teasing them under a binocular dissecting, micro
scope, using sub-stage lighting. The brain was removed from the cranial 7 cavity and examined.
Blood smears were made from the heart and veins. Giemsa's stain and Wrights stain were "both employed. In Giemsa's method the blood was smeared thinly on a slide and allowed to dry in the air. The dried smear was then immersed in Methyl alcohol for 1 - 2 minutes. Without drying, the slide was then placed in a container of diluted Giemsa's stain (one drop of stain to each mi. of distilled water) for 1+5 minutes.
The slide then was rinsed in tapwater and allowed to dry while standing on one end.
Cestodes and trematodes were fixed under pressure in either
Lavdowsky's formula of A F A fixing reagent or in 10 percent formalin.
The A F A fixing reagent proved to be more effective in fixation as well as less irritating to the worker.
The acanthocephala were relaxed in the refrigerator for 1 - 2 hours, after which they were fixed in 70 percent alcohol. The nematodes were relaxed and fixed in warm 70 percent alcohol.
All specimens were removed from the fixing reagents and stored in
75 percent alcohol until ready for staining. Several staining pro cedures were employed during the course of the study. Delafield's
Hematoxylin, Meyer's Paracarmine, and Semichon's stain were the ones used. The author obtained best results with Semichon's stain. After being stained, the specimens were dehydrated in alcohol, cleared in xylene, and mounted in piccolyte under cover glasses. Some nematodes were cleared in petri dishes containing a 10 percent solution of glycerine-alcohol, made by mixing 90 ml. of 75 percent alcohol and 8
10 ml. of glycerine. This solution was then placed in a dessicator, allowing the alcohol to evaporate and thus leaving the worms in pure
glycerine which cleared them.
Birds collected in the northern part of Indiana were examined in a field laboratory set up by the author. Most of the birds from this part
of the state were collected from the areas of Silver Lake, Hogback Lake,
and Lake James.
A H birds taken from Central Indiana were brought into the labora
tory at Ball State Teachers College, Muncie, Indiana, where they were
examined.
A field laboratory was set up also in Southern Indiana, but, as a
survey of the collections will show, most of the birds were brought into
the laboratory at Ball State Teachers College for examination. Birds in
Southern Indiana were very difficult to obtain. Several that were
collected from this area were brought into the laboratory by students.
These birds came principally from the general vicinity of Salem and
Bedford, Indiana. Birds collected personally by the author in this area
came from near Pekin, Indiana. Many persons contacted in Southern
Indiana were unfamiliar with this bird either by name or description-
The Common Grackle does not seem to be a frequent inhabitant of this
part of the state.
Birds were numerous in North and Central Indiana and could be
collected in numbers for short periods of time during the spring and
autumn. In order to make examination of fresh material, no more birds
were taken than could be examined that day. Each bird required approximately one hour for the examination.
All drawings were made by using either a camera lucida or micro- projector. Measurements are indicated either in millimeters or microns.
Where sufficient material was available the average measurement as well
as the extremes are given. The average measurement is given in paren
theses following the extremes. In some instances there were not enough
good, specimens to justify the use of an average measurement. Information
obtained in this study is summarized in the specific description for
each species. CLASSIFICATION AND DESCRIPTION OF PARASITES
Acanthocephala
Van Cleave (1948) established the phylum Acanthocephala, including two classes and four orders. The spiny headed worms infect fish, reptiles, birds, and mammals as definitive hosts. Both larval and adult stages are completely lacking a digestive tract. While several species parasitize birds, only one, Mediorhynchus grand!s Van Cleave, 1918 was found in the grackle.
Mediorhynchus grandis Van Cleave, 1918. (Figs. 2-6).
Synonyms: Heteroplus grandis Van Cleave, 1916.
Empodius grandis Travassos, 1924.
Site of infection: Intestine.
Classification according to Chandler (1955)*
Order - Archiacanthocephala Meyer, 1931*
Family - Gigantorhynchidae Hamann, 1892.
Genus - Mediorhynchus Van Cleave, 1916.
Synonyms: Heteroplus Kostylev, 1914.
Empodius Travassos, 1916.
Micracanthorhynchus Travassos, 1917*
Leiperacanthus Bhalerao, 1937*
Generic Diagnosis according to Van Cleave (1916): Acanthocephala of medium size, reaching sexual maturity in the alimentary canal of birds. Proboscis receptacle inserted near the middle of the proboscis wall. Receptacle a single walled muscular sac, with invertors of proboscis passing through its wall some distance anterior to the posterior tip of the receptacle. Central nervous system near the center of the proboscis receptacle between the invertor muscles. Cement glands of male a
10 11
compact mass of rounded or pear shaped cells, usually eight in number. Proboscis hooks of two distinct types; the hooks on anterior proboscis with flask shaped roots. The hooks on posterior portion of proboscis, without reflexed roots.
Specific description; Mediorhynchus grandis with the character istics of the genus Mediorhynchus. The proboscis ends bluntly, having hooks on the anterior part arranged in diagonal rows, each of which contains six to ten hooks. The body of both male and female specimens is smooth, without spines. Body length of female 22 - 30 mm (24).
Maximum width 1.4 mm. Eggs 0.040 - 0.043 mm in length. Body of male
8 - 10.5 mm (8 .5) in length. Maximum width 1.0 mm. Lemnisci extending to level of anterior testis. Two testes,.in tandem, located in the posterior half of the body. Cement glands containing eight cells, and located posterior to testes.
Discussion: The genus Mediorhynchus was established by Van Cleave
(1916). He obtained specimens of this worm from passerine birds in -the
Mid-west United States. There are several synonyms for the genus.
Van Cleave (1947) proposed that the generic name Heteroplus, which
Kostylev (1914) used, was not available, since it had been us.ed by
Mulsant and Rey as a name for a sub-genus of beetles. The date of their work is uncertain, but the evidence indicates that it preceded
Kostylev's work by several decades, the latest date given by Van Cleave being 1870. Two genera, Empodius and Micracanthorhynchus were proposed by Travassos (1917) for species which later were recognized as belonging to the genus Mediorhynchus. Van Cleave (1947) placed Leiperacanthus
Bhalerao, 1937 in synonomy with Mediorhynchus. 12
Van Cleave (1916) proposed the name Mediorhynchus grandis for the specimens taken from the sub species Quiscalus quiscula stonei in
Maryland and New Jersey, and from the Meadowlark, Stumella magna, in
North Carolina, Ohio, and Illinois. Van Cleave (1918) took this species from the crow in Maryland and Ohio; from the Common Grackle in Kansas,
Illinois, Kentucky, and Ohio; and from the Red-Wing Blackbird, Agelaius phoenicus phoenicus, in Ohio. Mediorhynchus grandis has not been reported outside of the United States.
Life History: The life history of Mediorhynchus grandis is not completely known. Moore (19^1) fed eggs of this species to the grass hoppers, Chorotophaga viridifaciata, Orphuella pelidna, and Arphia luteola. Infective cystocanths were recovered from the body cavity of these hosts after 25 days. Grackles and meadowlarks would appear to be the natural definitive hosts. Other vertebrates become infected by eating infective arthropods accidentally or otherwise. Cestoda
Many species of cestodes are found in birds. There were two species of cestodes found infecting the Common Grackle. These belonged to the order Cyclophyllidea, which is the most important order to man and domestic animals.
Orthoskrjabinia quiscali (Rodgers, 19*H) Spassky, 19^7- (Figs.
10-12).
Synonyms: Anonchotaenia quiscali Rausch and Morgan, 19^7•
Site of infection: Small intestine.
Classification according to Yamaguti (1959)*
Order - Cyclophyllidea Braun, 1900.
Family - Dilepididae Railliet and Henry, 1909*
Genus - Orthoskrj abinia Spassky, 19^7*
Synonyms - Skrjabinerina Matevosian, 19^8 .
Anonchotaenia Rodgers, 19*11.
Generic Diagnosis according to Wardie and McLeod (1952): With prominent suckers, but with neither rostellum nor armature of any kind. Segmentation apparent only when the segments are mature. Segments short. Genital apertures alternating irregularly. Genital ducts ventral to poral canals. Ovary and yolk gland ovoid, without lobulation. Uterus small, ovoid or spherical, showing when gravid a few eggs; joined to it is a fingerlike paruterine organ which receives the eggs.
Specific Description: Orthoskrj abinia quiscali is a small fragile worm. Stobila 30 to 35 mm in length. Rostellum rudimentary or absent, unarmed, uterus single, testes nine, no special development of a sphincter muscle at mouth of cirrus pouch. Gravid uterus with few eggs.
Discussion: Orthoskrjabinia quiscali was originally described by
13 Rausch and Morgan (19^7) from some material taken from the Purple
Grackle. There are about fifteen valid species in the genus
Orthoskrj abinia according to Wardle and McLeod (1952). Most of the specimens found in North American birds belong to .the species
Orthoskrj abini a globata. Rodgers (19^1) found Orthoskr .1 abinia rostellata in the cardinal. However, according to Wiardle and McLeod, the characteristic difference between the two species is that
Orthoskrj abinia rostellata has a double uterus and Orthoskrj abinia quiscali has a single uterus. Jones (i960) reported Orthoskrjabinia rostellata from the Eastern Crow in Ohio. He gave as the species* characteristic the possession of a single uterus. It is theref&re possible that the cestode taken by Jones from the Eastern Crow in Ohio is the same as that reported here as Orthoskrjabinia quiscali from the
Common Grackle in Indiana.
Life History: There are no life histories known for this genus.
In 1957 Freeman investigated a related species, Paruterina candelabaria, and succeeded in infecting mice with the bladderworm stage. Some similar vertebrate may serve as the intermediate host of Orthoskrjabinia quiscali.
Arthropods, annelids, and mollusks serve as intermediate hosts for the immature stages of tapeworms, while reptiles, birds and mammals harbor the adult worms. 15
Paricterotaenia parina (Dujardin, l8I4-5) Fuhrmann, 1932. Figs. 7-9*
Synonyms: Taenia parina Dujardin, 18U5.
Drepanidotaenia parina (Dujardin, I8U5 ) Stossich,
1898.
Choanotaenia parina (Dujardin, 18^5 ) Clerc, 1906.
Icterotaenia parina (Dujardin, 18^5) Baer, 1925.
Site of infection: Small intestine.
Classification according to Yamaguti (1959)*
Order - Cyclophyllidea Braun, 1900.
Family - Dilepididae Railliet et Henry, 1909*
Genus - Paricterotaenia Fuhrmann, 1932.
Synonyms: Choanotaenia Fuhrmann, 1907 nec Railliet, 1896.
Icterotaenia Railliet et Henry, 1909# partira.
Parachoanotaenia Luhe, 1910, partim.
Generic Diagnosis: After Yamaguti (1959)* With a single crown of rostellar hooks. Testes numerous, posterior to female glands. Genital glands "between excretory stems. Cirrus pouch long, usually over reaching excretory stems medially. Genital pore alternating irregularly. Ovary lobate, pre- equatorial, median, with compact vitelline gland behind. Uterus saclike. Vagina posterior to cirrus pouch, seminal receptacle present. Parasite of birds.
Specific Diagnosis: Paricterotaenia parina is a small cestode with a maximum iength of ^5 mm and a maximum width of 1.5 mm. The scolex is small with a rostellum armed with 18 - 20 hooks arranged in a single row. A short neck follows the scolex. The genital pores, which alternate irregularly, are situated on the lateral margin of the anterior third of the proglottid. There are 18 - 23 testes situated posterior to 16 the ovary and vitelline gland. The vagina opens into the genital atrium posterior to the cirrus and passes inward parallel with the cirrus sac.
The ovary is slightly bilobed and compact with short projections. The vitelline gland is a compact organ and is situated equatorially in the proglottid posterior to the ovary. Mehlis' gland, less apparent, lies just in front of the vitelline gland.
Discussion: Fuhrmann (1932) established the genus Paricterotaenia, including this form in the family, Dilepididae. Lopez-Neyra (1951) proposed new generic definitions for two genera, Anomotaenia and
Choanotaenia, of this family, dropping the genus Paricterotaenia. As a result there were ^3 species of this genus, which had to be placed in either related genera or in new genera. However, Mettrick (1958) was unable to confirm the findings of Lopez-Neyra, and therefore rejected the emended classification in favor of Fuhrmann's earlier work.
Mettrick contended that Lopez-Neyra's work in 1951 was based on various species described in the literature rather than on a study of fresh material. Yamaguti (1958) supported Mettrick in retaining the genus
Paricterotaenia. He designated as the genotype, Paricterotaenia porosa, which Lopez-Neyra (1951) had transferred to the genus Choanotaenia.
This had left Paricterotaenia without a genotype, thus invalidating the genus. The life history of Paricterotaenia is unknown. A single specimen of Paricterotaeni a was taken from the Bob-white Quail, Colinus virginianus texanus, in Texas by Webster and Addis (19^5), who examined two hundred-ninety of these birds. Hhesy did not identify the species represented. Trematoda
This class includes the non-strobilated parasitic flat worms which have rudimentary digestive systems. Dawes (19^) listed 90 species of digenetic trematodes from birds. Birds, next to fishes, have the heaviest infections of fluke parasites. The digenetic trematodes are the most common. They require one or more intermediate hosts for completion of their life cycles. There were five species of the
digenea obtained from the Common Grackle. The tissue sites of infection
represented were the intestine and biliary ducts of the liver.
. Conspicuum icteridorum Denton, 1951* (Figs. 16 and 17).
Site of infection - Gall bladder
Classification according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Dicrocoeliidae Odhner, 1911.
Subfamily - Dicrocoeliinae Looss, 1899.
Genus - Conspicuum Bhalerao, 1936.
Generic Diagnosis according to Yamaguti (1958): Body flattened, fusiform with maximum width just behind middle. Cuticle distributed irregularly with conical papillae. Suckers subequal, esophagus three or four times as long as pharynx. Ceca terminating short of posterior extremity. The acetabulum well apart from anterior end. Testes symmetrical, immediately postacetabular, separated, from one another by the uterus. Cirrus pouch plump, containing winding seminal vesicle, prostate complex, and cirrus. Receptaculum seminis and Lauer's canal present. Vitelline follicles small, extending in extracecal fields from testicular level as far as, or nearly to, cecal ends. Uterus occupying most of hind part of body. Excretory
17 18
vesicle tubular, arms bifurcating in front of testes. Parasitic in gall bladder of birds, rarely mammals.
Specific Description: Conspicuum icteridorum, with characters of the genus Conspicuum. Body thick, muscular, elongate, oval, tapering to posterior end. Body length, 2.20 to 5*97 mm (3•5)• Body width, 0.70 to
1.97 mm (1.5) with widest region between the testes and ovary. Cuticle smooth or with transverse wrinkles when slightly contracted. Oral sucker strongly muscular, sub-terminal; 0.35 to O .50 mm in diameter.
Acetabulum large and muscular, with deep cup-shaped lumen situated at junction of anterior and middle body thirds. Acetabulum, 0.45 to O .79 mm in diameter. No prepharynx; pharynx globular. Esophagus narrow, bifurcating one-half to two-thirds the distance from oral sucker to acetabulum. Ceca of medium width, straight or slightly wavy, passing outside of lateral margins of testes and terminating about midway between testes and posterior end of vitellaria. Excretoiy pore terminal.
Excretory vesicle simple, tubular, extending anteriorly through approxi mately one-third of body length, receiving a common collecting tubule from each side. Genital pore median, at level of pharynx. Testes rounded, 0.15 to 0.25 mm in diameter, arranged bilaterally with their fields widely separated. Cirrus sac, anterior to acetabulum, elongated,
0.24 to 0.50 mm in length by 0.09 to 0.25 mm in diameter, containing a convoluted seminal vesicle surrounded by prostate gland cells. Ovary slightly lobed to transversely oval, 0.10 to 0.35 mm in greatest diameter, situated submedially, posterior to the testes. Vitellaria composed of numerous median-sized follicles, extending posteriorly from 19
testicular zones for a distance of approximately one-fourth the body length. Uterus convoluted, filling most of posttesticular region of body, and continuing an undulating course to genital pore. Mature ova dark brown.
Discussion: Bhalerao (1936) in considering the genus Eurytrema
recognized five sub-genera in the group, including Conspicuum. Strom
(19^0 ), re-examining the material, removed all of the bird parasites
from the genus Eurytrema. He then elevated the sub-genera to generic
rank. Travassos (19^+), working with the genus Eurytrema, elevated two
of the subgenera, including Conspicuum to generic rank. Since Strom's
work antedates that of Travassos, it must be given priority. Denton and
Byrd (1951) obtained two new species, Conspicuum icteridorum and
Conspicuum macrorchis from the Common Grackle and Eastern Crow
respectively. They obtained Conspicuum icteridorum from several hosts
from Texas, Tennessee, Georgia, Ohio, and Michigan. However, of all the
bird hosts that have been reported for this species, it has been found
to be most prevalent in the Common Grackle. Of fifty specimens of this
bird that were examined by Denton and Byrd, forty-two were found to be
infected with this fluke. Their eighty-four percent infection in the
Common Grackle is slightly less than the ninety percent obtained by the
present author.
Bassett (1958) observed that infections of Conspicuum icteridorum
resulted in marked changes in the gall bladder of the host, the walls
becoming much thinner with the epithelium missing. On the other hand,
the tissue was much thicker at the point of attachment of the parasite, 20 resembling a mushroom-like projection. Each layer of tissue was thicker and the epithelium was present. Necrotic tissue was found along with , apparent inflammation. This follows the general pattern of tissue response of other hosts to fluke invasion.
Life History: Patten (1952) succeeded in infecting the snail,
Zonitoides arboreum from which, later, he recovered cercariae. He fed these to two isopods, Armadillium quadrifrons and Onsicus asellus, later recovering metacercariae. The metacercariae were fed next to uninfected grackles, after which he recovered young worms from the gall bladder in sixteen hours and adult flukes in twelve weeks. Denton and Byrd (1951) listed the snail, Deroceros laeve as the first intermediate host.
Recorded definitive hosts of Conspicuum icteridorum are:
Quiscalus quiscula versicolor; Quiscalus quiscula quiscula; Cassidix mexicanus prosopidicola; Cassidix mexicanus major; Euphagus carolinus;
Stumella magna; and Ammospiza maritima macgillivraii. All of the recorded hosts of this parasite belong to the family Icteridae except
Ammospiza which is in the family Fringillidae. 21
Conspicuum macrorchis Denton and Byrd, 1951. (Figs. 18 and 19).
Site of infection: Gall bladder.
Classification according to Yamaguti (1958).
Order - Digenea Van Beneden, 1885.
Family - Dicrocoeliidae Odhner, 1911*
Genus - Conspicuum Bhalerao, 1936.
Generic Diagnosis: From Yamaguti (1958). Body flattened, fusiform, with maximum width Just behind middle. . Cuticle distributed irregularly with conical papillae. Suckers subequal, esophagus three or four times as long as pharynx. Ceca terminating short of posterior extremity. The acetabulum well apart from anterior end. Testes symmetrical, immediately post- acetabular, separated from one another by the uterus. Cirrus pouch plump, containing winding seminal vesicle, prostate complex, and cirrus. Receptaculum seminis and Lauer's canal present. Vitelline follicles small, extending in extracecal fields from testicular level as far as, or nearly to, cecal ends. Uterus occupying most of hind part of body. Excretory vesicle tubular, arms bifurcating in front of testes. Parasitic in gall bladder of birds, rarely mammals.
Specific Description: Conspicuum macrorchis, with characters of
the genus Conspicuum. Body elongated, widest in region between testes
and ovary, 4.25 to 5*57 mm long (5*25) by 1.40 to 2.24 mm wide (2 .20 ).
Cuticle without spines. Oral sucker subterminal and muscular.
Acetabulum large, 0.50 to 0*95 mm in diameter, situated at Junction of
anterior and middle thirds of the body. Pharynx globular. Esophagus
short, bifurcating one-third to one-half the distance from the oral
sucker to the acetabulum. Ceca terminate short of posterior end of body. Genital pore median, approximately midway between oral sucker and bifurcation of esophagus. Testes round, large, 0.35 to O .65 mm in 22 diameter, directly opposite each other. Cirrus sac entirely anterior to acetabulum. Ovary oval, near median plane of the body. Seminal receptacle small; just posterior to the ovary. Vitellaria composed of many follicles, marginal to ceca. Uterus convoluted, filling most of posttesticular region of body. Eggs dark brown.
Discussion: Conspicuum was established as a genus by Bhalerao in
1936 from flukes taken from the robin, Turdus migratorius, in Brazil.
The genus has five valid species of which four are parasites of birds.
Conspicuum macrorchis was described by Denton (1951) from material '4 1 taken from crows in Texas. Jones (i960) obtained this fluke from the
Eastern Crow in Ohio. These were the only host records for this species until the present, which therefore constitutes a new host record. This species has never been reported outside the United States.
Life History: The life history of Conspicuum macrorchis is not completely known. Denton and ByrcL (1951) listed the molluscan host as
Bulimulus alternatus mariae. Patten (1952) infected the snail,
Zonitoides arboreus, with Conspicuum icteridorum. The cercariae from this infection were fed to two isopods, Armadillidium quadrifrons and
Oniscus asellus, from which he was able later to recover metacercariae.
These metacercariae then were fed to grackles, after which adult flukes were recovered in twelve weeks. Denton and Byrd (1951) listed the
snail,* Derocera laeve, as the first intermediate host of Conspicuum
icteridorum. Since Conspicuum icteridorum and Conspicuum macrorchis
are very similar in appearance as well as in site of infection, it is
likely that they have similar life histories. 23
Brachylecithum americanum Denton, 19^5* (Figs. 13, 1^, and 15).
Site of infection: Gall bladder and bile ducts.
Classification according to Yamaguti (1958)*
Order - Digenea Van Beneden, 1885.
Family - Dicrocoeliidae Odhner, 19H.
Genus - Brachylecithum Strom, 19^0.
Synonyms: Qrthorchis Travassos, 19^.
Olssoniella Travassos, 19^, partim.
lyperosomum Looss, 1899, partim.
Generic Diagnosis according to Yamaguti (1958)* Body slender, suckers usually equal or subequal, close to each other. Esophagus short, ceca terminating short of posterior extremity. Testes tandem or somewhat diagonal. Cirrus pouch pre-acetabular. Genital pore usually bifureal. Ovary median, a short distance in front of mid body. Receptaculum seminis present. Uterus filling up most of hindbody. Vitellaria occupying the small lateral area posterior to ovary sometimes commencing at level of ovary. Parasites in bilary ducts and gall bladder of birds and mammals.
Specific Diagnosis: Body thin, transparent, elongated, with almost parallel sides, having a rounded anterior end and a tapering posterior end. Length, 2.5 to 5*6 mm (U.6). Width, in region of the acetabulum
0.25 to O.56. Cuticle without spines. Oral sucker, subterminal.
Acetabulum within anterior fourth of body. Pharynx small; esophagus narrow, bifurcating anterior to the acetabulum. Ceca wavy, lateral to genital organs, terminating posterior to vitelline follicles. Excretory vesicle tubular, narrow, and extending anteriorly almost to vitellaria.
Testes large, lobed, in tandem, almost equal in size. One loop of the 2b uterus passes between the testes. Ovary sma.13.er than the testes, round to oval, situated posterior to the testes. Vitellaria lateral, compact, posterior to ovary. Uterus convoluted and filling most of posterior portion of body. Seminal receptacle smaller than ovary and posterior to same. Eggs, dark brown.
Discussion; Denton and Bird (1951) iu describing Brachylecithum americanum from the crow and Red-shouldered Hawk, reported that the size of the adult was larger in these hosts than in smaller birds such as the grackle. They believed that there was some correlation in size of the host and its bile ducts with the size of the parasites. Dawes
(19^6) listed Brachylecithum as a subgenus of Lyperosomum as proposed by
Strom (19^0).
Life History: Denton (19^5) succeeded in infecting two species of snails, Polygyra texasiana and Polygyra berlandierianna, recovering cercariae after 106 days. He then infected two beetles, Gastroidia cyanea and Diabrotica duodec impunctata, from which he was able to recover metacercariae. Therefore, this cycle was experimentally established. Apparently certain beetles serve as the main source of infection for birds.
Recorded definitive hosts are; Cassidix mexicanus prosopidicola;
Quiscalus quiscula versicolor; Stumella magna; Corvus brachyrhynchos, and Cyanocitta cristata. 25
Echlnostoma revolutum (Froelich, 1802) Looss, 1899. (Figs. 18, 19,
and 20).
Synonyms: Fasciola revoluta Froelich, 1802.
Distoma echinatum Zeder, 1803.
Echlnostoma mendax Dietz, 1909*
Echlnostoma paraulum Dietz, 1909*
Echlnostoma coalitum Barker and Beaver, 1915*
Echlnostoma armigemm Barker and Irvine, 1915*
Echlnostoma limicoli Johnson, 1920.
Echlnostoma cinetorchis Amdo and Ozaki, 1923»
Site of infection: Small intestine.
Classification according to Dawes (19^).
Order - Digenea Van Beneden, 1858.
Family - Echinostomatidae Looss, 1902, emend. Poche, 1926.
Genus - Echlnostoma Rudolphi, 1809. /
Generic Diagnosis; according to Yamaguti (1958). Body elongate, spinose. Head colar well developed, with double dorsally uninterrupted crown of spines. Acetabulum near anterior extremity. Testes usually in posterior half of body. Cirrus pouch rather rounded anterior to acetabulum or somewhat over lapping it. Genital pore postbifureal. Ovary median or a little to one side of median line in front of anterior testis. Vitellaria surrounding ceca in hind part of body. Uterus long, winding in intercecal field between ovary and acetabulum.
Specific Diagnosis: With characters of the genus Echlnostoma.
Body elongate, 4 - 7 mm long by 1.5 - 2-5 mm broad. Widest in region between ovary and testes. Cuticle spinous anteriorly and ventrally as far back as the ventral sucker. Collar bearing thirty-seven spines. 26
Ventral sucker twice as large as the oral sucker. Gopads variable in form, one behind the other, with the ovary foremost. Testes, 0.3 - O.U mm in diameter. Ovary, 0.66 - 0.79 mm. Vitellaria extremely developed,
especially posteriorly, reaching to the posterior end, but not extending
anterior of ventral sucker. Uterus with only an ascending limb, which
is formed into transverse fold. Eggs large, 0.095 - 0.120 by 0.050 -
0.07 mm.
Discussion: Johnson (1920) reported Echinostoma revolutum from
water birds frequenting lakes containing the snail, Physa occidentalis.
This agrees in part with findings in the present report in that
Echinostoma revolutum was taken only from grackles in the northern part
of Indiana which includes the major lakes of the state. The birds from
which this parasite was obtained were taken on the shore of Silver Lake,
near Angola, Indiana. A check revealed that the snail Physa
occidentalis is found in the lakes in this region. The present author
has collected Physa occidentalis at various times from Silver Lake
where the birds were parasitized with Echinostoma revolutum.
Beaver (1937) restudied the descriptions of all closely related
species and possible synonyms of Echinostoma revolutum, but after
working on the older literature he decided it was impossible to deter mine from the figures and descriptions of most of the earlier authors
whether they had studied this species or some other. He placed all but
two of the eighteen forms which have thirty-seven cephalic spines either
in synonomy with Echinostoma revolutum or as questionable species.
Johnson (1920) found that adult worms probably live for several 27 years in the intestine of ducks and geese. Over a period of years the incidence of infection would likely increase. If the Common Grackle harbors this parasite for several years, it would seem reasonable to expect a higher frequency of infection in birds in the lake area. How ever, the low incidence of this parasite might indicate that the grackle is only an accidental host.
Beaver (1937) reported marked variations in body size, organ size, and body spination in specimens of Echinostoma revolutum taken from different hosts. He demostrated that the arrangement, number, and relative size of the cephalic spines were least variable. For this reason these features are considered to be the most reliable in diagnosing the species of Echinostoma. The variations in size of specimens of this parasite in different hosts might account for the great number of Synonyms that have been listed for this species.
Beaver (1937) placed eight species of echinostomes in synonomy with
Echinostoma revolutum. Leidy (1888) reported specimens up to 25 mm, and Barker and Beaver (1915) give 22 - 30 mm as the length for this species. Beaver (1937) observed that this species reached maturity at about 4 mm. Thus the range for mature specimens of Echinostoma revolutum is from 4 - 3° mm. The size range in this present repdrt is small, in part due to the few specimens obtained.
Life History: Many studies have been made of the life history of this genus. Johnson (1920) was able to recover cercariae from the snail Physa occidentalis, which had been experimentally infected. Krull
(1935) experimentally infected the snail Pseudosuccinea columella with 28
cercariae. Natural infections with cercariae of Echinostoma revolutum have been reported by Tubangui (1932) in Lymnaea pervia; by Vevers
(1923) in lymnaea stagnails; by Fallis (193*0 in Stagnicola palustris;
and by Lutz (192*0 in Physa rival!s. Metacercariae have been recovered
from snails, Fossaria modicella and Pseudosuccinea columella, experi
mentally infected by Beaver (1937)* Dollfus (1925) obtained
metacercariae from experimentally infected tadpoles of Bufo americana.
Miki (1923) obtained metacercariae from tadpoles of Rana japonica, and
Dollfus (1925) from tadpoles of Rana rugosa. Beaver (1937) reported
metacercariae from both tadpoles and adults of Rana pipiens.
Echinostoma revolutum is cosmopolitan in geographical distribution
as well as in choice of hosts. Beaver (1937) stated that this species
has been reported from every temperate region in the world with the
exception of South Africa and inland China. He believes that the adult
worm may be found in almost any bird or mammal, with natural infections
limited to those hosts that feed upon aquatic animals. He included as
"natural" hosts animals in which the parasite has been found in nature.
"Normal" hosts were those in which the parasite completed its life
cycle only after experimental infection. Thus many birds and mammals
experimentally infected act as "normal" hosts if not "natural" ones. 29
Crepidostomum cooperi Hopkins, 1931. (Figs. 23, 2k', and 25).
Synonyms: Crepidostomum ambloplitis Hopkins, 1931*
Crepidostomum solidum Van Cleave and Mueller, 1932.
Crepidostomum fausti Hunninen and Hunter, 1933*
Site of infection: Gall bladder.
Classification according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Allocreadiidae Stossich, 190f-
Subfamily - Crepidostominae Dollfus, 1951.
Genus - Crepidostomum Braun, 1900.
Synonyms: Acrodactyla Stafford, 190U.
Stephanophiala Nicoll, 1909*
Acrolichanus Ward, 1917*
Generic Diagnosis: after Yamaguti (1958). Body elongate oval to subcylindrical, unarmed. Oral sucker terminal, surmounted anterodorsally by a half crovm of six head papillae or lobes, of which the two outermost are formed by the anterolateral corners of the sucker projecting prominently. Mouth aperture ventroterminal or practically ventral. Prepharynx present, pharynx well developed. Esophagus short or of moderate length. Ceca terminating near posterior extremity. Acetabulum in anterior half of body. No vesicula seminalis externa. Cirrus pouch more or less elongated claviform, overreaching acetabulum containing seminal vesicle, prostate complex, and ductus ejaculatorius. Genital pore median, pre-acetabular. Ovary submedian, between acetabulum and anterior testis. Receptaculum seminis and Laurer1s canal present. Uterus coiled between anterior testis and acetabulum. Vitellaria circumcecal, extending from forebody to posterior extremity, com mencing at varying levels in different species. Excretory vesicle reaching beyond anterior testis. Parasites of the intestine of freshwater fish. 30
Specific Diagnosis; The following characteristics were hased upon the study of two stained whole mounts. Crepidostomum cooperi with the characters of the genus Crepidostomum. Width varying from one-fourth to one-half the body length according to state of contraction. Greatest width at the level of the testes. Acetabulum one-fifth to one-fourth the body length from anterior end, slightly larger than oral sucker.
Oral papillae all of practically equal size; ventral pair, a part of the oral sucker, directed laterad; four dorsal papillae attached on anterior margin projecting anteriad beyond the oral sucker. Pharynx ovoidal, length one-half that of the oral sucker. Esophagus as long as pharynx.
Genital pore ventral, posterior to bifurcation of ceca. Common genital atrium obliterated under cover glass pressure. Cirrus sac long, tapering anteriorly, reaching dorsally over ventral sucker to level of ovary or to anterior testis, convoluted, with thin walls. Seminal vesicle convoluted, within posterior half of cirrus sac. Testes close together, slightly diagonal, in posterior half of body, about one and one-half times as long as wide, margin entire; distance from posterior testis to posterior end of body one-fourth to one-third of body length.
Ovary pear-shaped, dorsal and posterior to acetabulum, median or slightly to one side. Seminal receptacle separated from ovary by uterus. Ovary and cirrus sac pretesticular. Uterus mostly within area between acetabulum and anterior testis. Uterus extends back to posterior level of the anterior testis, wall thin and becoming thicker near edge of acetabulum. Intrauterine eggs few, varying in number from ^0 to 45.. Spermatazoa with rod-shaped heads and slender tails. 31
Vitellaria extending from near posterior end nearly to pharynx. In the area from the acetabulum to the posterior testis the follicles are lateral and ventral to intestinal ceca. Posterior to this the follicles extend across the ventral surface. Anterior to the acetabulum the follicles are lateral, ventral, and dorsal to ceca. A few follicles lie median to the ceca anterior to the testes.
Excretory bladder, a narrow tubular sac close to dorsal surface, extending forward to anterior edge of anterior testis.
Pigmented eyespots, one on each side of the esophagus, clearly evident in adults.
Discussion: Crepidostomum cooperi was originally described by
Hopkins (1931) from the intestine of a fish, Perea flavescens, collected at Go-Home Bay, Ontario by A. R. Cooper. It had previously been listed by Cooper as Crepidostomum laureatum. In the same journal in which
Hopkins described Crepidostomum cooperi as a new species he described
Crepidostomum ambloplitis also as a new species. At the same time
Van Cleave and Mueller (1932) had at press the description of a new species, Crepidostomum solidum. After comparisons were made, Van Cleave and Mueller appended a footnote to their description of Crepidostomum solidum, concluding that Crepidostomum ambloplitis and Crepidostomum cooperi were variants of the same species and that Crepidostomum ambloplitis and Crepidostomum solidum were synonyms of Crepidostomum cooperi. Hopkins (193*0 accepted this view and gave priority to
Crepidostomum cooperi, since it had appeared first in the original publication. Hopkins (193*0 also placed Crepidostomum fausti in 32 synonomy with Crepidostomum cooperi, believing that Hunninen and Hunter
(1933) were misled by lack of knowledge concerning the variability of
Crepidostomum cooperi.
Hunninen and Hunter (1933) reported Crepidostomum fausti as a new species from the trout. They based their identification on size and shape of papillae and testes "and other minor points." However, these have been shown by Hopkins (193*0 to be insufficient to justify separation into a new species.
Hopkins (193*0 expressed an opinion that Crepidostomum metoecus, which has been reported only from bats, is probably a variant of the species, Crepidostomum cooperi. The only material available is that originally described by Braun (1900). This consisted of specimens from some vials, found in the Vienna Museum, labeled as having been obtained from bats. It has been assumed that Crepidostomum metoecus is a para site of bats, although it has been reported only in the original description. Hopkins (193*+) stated, "It seems most probably that fish are the natural hosts and that bats are only occasional or accidental hosts." No attempt was made by Braun or Hopkins to explain how this apparent parasite of a cold-blooded host could become established in a warm-blooded host. It would seem that Crepidostomum cooperi is a very versatile parasite.
The genus Crepidostomum, which was established by Braun (1900), was included by Looss (1902) with the genus Bunodera in the subfamily
Bunoderinae. Odhner (1905) placed Crepidostomum in the subfamily
Allocreadiinae. Nicoll (1909) pointed out that the.differences in the 33 two genera justified this greater separation, "but he proposed a new generic name, Stephanophiala for Crepidostomum, forming a new subfamily,
Stephanophialinae to accommodate this newly formed genus. Faust (1918) supported Nicoll's classification. On the other hand, Brown (1927) placed priority on Crepidostomum and made Stephanophiala a synonym of
Crepidostomum. Dollfus (1951)> accepting Crepidostomum, established the subfamily Crepidostominae for this genus, which according to
Yamaguti (1958) is the only one belonging to this subfamily.
There are seven valid North American species of Crepidostomum according to Yamaguti, of which six are restricted to North American hosts. One species, Crepidostomum farionis, seems to be cosmopolitan in its distribution. The genotype, Crepidostomum metoecus, described from bats by Braun (1900) has not been reported from North America. 1516 six
North American species now considered in this genus are: Crepidostomum cooperi Hopkins, 1931» Crepidostomum corautum Osborn, 1903J
Crepidostomum illinoiense Faust, 1918; Crepidostomum isostomum Hopkins,
1931; Crepidostomum lintoni Pratt in Linton 1901; and Crepidostomum brevivitellatum Hopkins, 193^•
Life History: The complete life history of Crepidostomum cooperi was demonstrated by Hopkins (193*0 • This was the first known life history of this genus. According to Hopkins the eggs do not undergo cleavage while within the uterus of the adult fluke. Eggs placed in pond or tapwater soon begin to cleave. Mature miracidia develop within seven days. The posterior two-thirds of the body of the "miracidium"
(his term) is filled with germ cells. The redia of Crepidostomum 3^ cooperi was found in the clam, Musculium transversum. There was no apparent order of mother and daughter rediae, the same parent contained both cercariae and^rediae.
Bie cercariae, which are unpigmented and transparent, are very rapid swimmers, progressing sometimes with the tail and sometimes with the head in the lead. When coming in contact with a hard object, the suckers attach and a leech-like creeping begins, after which the tail is cast off.
The metacercariae were found encysted in the muscles and body cavity of nymphs of Hexagenia. The precocious metacercaria was enclosed in a thin transparent cyst, surrounded by an orange-brown granular wall laid down by the host. The adult fluke was found in the intestine of several hosts, including sunfish, perch, and catfish. The above life history, worked out by Hopkins, was from natural infections rather than experimental feedings.
Crawford (19^3) reported on the life history of a related species,
Crepidostomum farionis. A species of clam of the genus Pisidium was found to be the molluscan host, and mayfly naiads were second inter mediate hosts, harboring the metacercariae. The rainbow trout, as a definitive host, contained the adult fluke in the intestine. This part of the cycle was determined both naturally and experimentally.
According to Crawford (19^3) the life cycle of Crepidostomum farionis closely parallels that of Crepidostomum cooperi.
The definitive hosts from which Crepidostomum has been reported are
Perea flavescens; Leucichthys artedi tullibeej Ameiurus nebulosus; 35
NotemiRonus crysoleucas; Cyrpinus carpio; Necturus maculosus; and Amyda rautica. The present report is the first of this species in a warm blooded host. Nematoda
This class contains the unsegmented round or thread worms, ftiere are many more free-living forms than parasitic ones. Nematodes may "be found as parasites in most groups of animals, including even other nematodes. Life cycles may be direct or require an intermediate host, often an arthropod. Many species of nematodes parasitize birds. Three species are reported in this paper.
Dispharynx spiralis (Molin, 1858) Skrjabin, 1916. Figs. 31* and
35).
Synonyms: Dispharagus spiralis Molin, 1858.
Dispharagus nasutus Piana, 1897.
Dispharagus spiralis columbae Bridre, 1910.
Acuaria spiralis (Molin, 1858) Railliet, Henry
and Sisoff, 1912.
Site of infection: Mucosa of digestive tract.
Classification according to Cram (1927).
Suborder - Spirurata Railliet and Henry, 1915*
Family - Acuariidae Seurat, 1913*
Subfamily - Acuariinae Railliet, Henry, and Sisoff, 1912.
Genus - Dispharynx Railliet, Henry, and Sisoff, 1912.
Synonyms: Acuaria Bremser, 1811, part.
Spiropotera Rudolphi, 1819, part.
Dispharagus Dujardin, 18^5# part.
Generic Diagnosis: From Cram (1927). Cordons are present, recurrent, not anastomosing. Lateral
36 37
papillae postcervical, small. Males with spicules unequal and dissimilar. Usually with five pairs of postanal papillae. Parasitic in esophagus, gizzard and proventricuius of birds.
Specific Diagnosis: Dispharynx spiralis. Nematodes with four wavy cordons, the distal extremity of the cordons turning forward and extending anteriorly a short distance. Cordons recurrent, not anastomosing. No_yes-icular swelling anteriorly. Mouth with two simple lips, followed by a vestibule or phaprrax with wall usually striated, and an esophagus differentiated into two parts, the anterior part muscular and the posterior glandular. Lateral papillae postcervical, small. Males with spicules. Body usually rolled in a spiral.
Life History; Piana (1897) worked out the life history of a new species, Dispharagus nasutus, having found the intermediate host to be sowbug, Porcellia laevis. The embryo, which hatches from the egg in the alimentary canal of the sowbug, develops into an infective larva in either the body cavity or anterior portion of the digestive canal. When infective sowbugs are eaten by a suitable definitive host the larvae develop to adults, becoming fixed in the musoca of the digestive canal, principally in the esophagus, proventriculus, and small intestine.
Piana's work on the life history of Dispharagus nasutus was based on comparative studies and not on experimental feedings. Cram (1927) placed this species in synonomy with Dispharynx spiralis.
Discussion; Dispharynx spiralis. This species was first described as Spiroptera nasutus by Rudolphi (1819) from the House Sparrow in
Vienna. Dujardin (18^5 ) found the same species in the sparrow in France.
He established the genus Dispharagus, and placed Spiroptera in synonomy 38 with it. Spiroptera, at that time, was in synonomy with Acuaria
Bremser, 1811. Molin (1858) found this worm in domestic birds in Italy and described it as a new species, Dispharagus'spiralis. Stossich
(1891) obtained it from the Guinea Fowl in Italy, and Bridre (1910) from the pigeon in Tunisia.
Railliet, Henry, and Sisoff (1912) revised the genus Acuaria
Bremser, l8n and established the sub-genus Dispharynx to include
Acuaria spiralis, placing Acuaria nasutus in synonomy. Seurat (1916) reported Acuaria nasutus from the partridge in Algeria and listed the pheasant as the definitive host. Skrjabin (1916) elevated the sub genus r Dispharynx to generic rank.
'Walton (1923) reported Dispharynx spiralis from the Common Grackle in Illinois. Other host records for this species include such game birds as the Sharp Tailed Grouse in Wisconsin; the Bob-white Quail in
Virginia, North Carolina, and Ohio; the Hungarian Partridge in Wisconsin,
Ohio, and Michigan; and both the Ringnecked and Golden Pheasants in New
York. Also included are such passerine birds as the House Sparrow, robin, catbird, Carolina Wren, pigeons, and the Common Grackle. Levine and Goble (19^2) found about one-third of 2800 Ruffed Grouse in New York infected with Dispharynx spiralis. Goble (19^5) reported a high incidence of it in gallinaceous birds, but a relatively low incidence in domestic birds. Capillaria quiscali Read, 19^9- (Figs. 26, 27, 28, and 29).
Site of infection: Lower intestine.
Classification according to Chitwood and Chitwood (1950)*
Supervamily - Trichuroidea Railliet, 1916.
Family - Trichuridae Railliet, 1916.
Genus - Capillaria Zeder, 1800.
Synonums: Trichosoma Rudolphi, 1819.
Trichosomum Creplin, 1829.
Liniscus Dujardin, I8U5.
Thominx Dujardin, I8U5 .
Calodium Dujardin, 18^5.
Generic Diagnosis from York and Maplestone (1926): Body capillary, mouth simple, cuticle with bacillary hands, dorsal, ventral or lateral in position; esophagus long and gradually increasing in size posteriorly. Male: Anus terminal or subterminal, small membranous caudal alae or bursa-like structures present or absent: Spicule long and slender, surrounded by a sheath, with or without spines on its surface. Female: Vulva near termination of esophagus. Oviparous. Eggs lemon shaped with the usual opercular plugs at the poles.
Specific Diagnosis: Capillaria quiscali, with the characters of the genus Capillaria. Mouth simple. Female: 8.5 to 10 mm long (9*5) by 65 to 75 microns wide at the region of the vulva. Maximum width
0.1 mm. Vulva located just posterior to termination of esophagus. Anus subterminal. Male: Maximum length 8 mm, maximum width J2 microns.
Spicule smooth with a blunt tip. Without caudal, lateral, cuticular alae. Bursa-like structure with two papillae. Cloaca subterminal with papillae on each side. ko
Discussion: Capillaria quiscali. Read (19^9) examined eighteen
Common Grackles in Wisconsin, finding two of them infected with specimens of Capillaria. From this material he described, as a new species, Capillaria quiscali. Although it closely resembles Capillaria caudinflata, it differs from the latter in having larger eggs with rough outer shells; however, as an adult it is smaller than Capillaria caudinflata.
Read (19^9) reported Capillaria quiscali from the Common Grackle taken in Madison, Wisconsin. Clark's unpublished survey of the parasites of the grackle in Maryland lists Capillaria quiscali as a parasite obtained from this bird. Capillaria quiscali has not been reported for any host other than grackles.
Life History: The life history of Capillaria quiscali has not been determined. Cram (1936) was able to infect Bob-whites with a related species, Capillaria contorta, by feeding them infective eggs of this worm. Wehr (1936) showed that still another species, Capillaria annulata, could be transmitted by earthworms. According to Noble and
Noble (1961) the eggs of Capillaria annulata are deposited by the female in the mucosa of the crop of poultry, being freed when this layer sloughs off. Eggs are fully embryonated in a few weeks. If such eggs are eaten by earthworms, poultry become infected by eating the infected earthworms. The larval nematodes, which are liberated in the crop of the bird, penetrate the mucosa, mature, mate, and produce a new batch of eggs.
On the other hand, it would seem that an intermediate host may not kl
"be necessary for Capillary quiscali to complete its life history. Waite
(1920), Clapham (1935), and Cram (1936) contend that Capillaria does not always need an intermediate host. If, however, the earthworm serves as an intermediate host for Capillaria quiscali, it certainly provides a widely distributed intermediate host. The low incidence of Capillaria quiscali in the Common Grackle in Indiana might be indicative that earthworms were not on the highly selective food list of the Common
Grackle. Splendidofilaria quiscali (Von Linstow, 1904) Odetoyinbo and Ulmer,
I960. (Pigs. 30 - 3*0*
Synonyms: Filaria quiscali Von Linstow, 1904.
Splendidofilaria ohioensis Jones, i960.
Site of infection: Sub-pia mater of cerebral hemisphere.
Classification according to Adderson and Chabaud (1959)*
Superfamily - Filarioidea Weinland, 1958*
Family - Onchocercidae Anderson and Chabaud, 1959*
Subfamily - Splendidofilarinae Chabaud and Choquet, 1953*
Genus - Splendidofilaria Skrjabin, 1923*
Synonyms: Filaria Mueller, 1787, part.
Chandlerella York and Maplestone, 1926.
Vagrifilaria Augustine, 1937*
Parachandlerella Caballero, 1948.
Generic Diagnosis: According to Skrjabin (1923). Medium-sized worms. Cephalic and without lips and epaulet-shaped structures. The head bears four sub median and perhaps two lateral head papillae. The esophagus is not divided into a muscular and glandular., pqrtion. Male: Posterior extremity digitiform. Caudal alae absent. Two equal or approximately equal apicules.- Female: Posterior extremity digitiform. Vulva in anterior third of the esophagus. Viviparus.
Specific Diagnosis: Splendidofilaria quiscali, with characters of
the genus Splendidofilaria. Male: Length, 20.0 to 24.0 mm (23.0); bearing three pairs of anal papillae. Female: Length, 45.0 to 57*5 nun
(49.0). Ovary and oviducts coiled. Uteri parallel, not crossing each
other. Worms long and slender, cuticle smooth, spineless, and unstriated.
Tail digitiform, that of male recurved, bluntly rounded, having ^3 relatively long spicules. Parasites of the cranial cavity of "birds.
Discussion: This species was first described by Von Linstow (190^) from material taken from the grackle Quiscalus versicolor. Although he had only fragments of females to work on, Von Linstow described the species as Filaria quiscali.
Odetoyinbo and Ulmer (1959) in a study of a nematode from the brain of the Common Grackle in Iowa concluded that the species was Filaria quiscali Von Linstow, 190^. They removed it from the genus Filaria and placed it in the genus Splendidofilaria. This new combination was first reported in an abstract by Odetoyinbo and Ulmer (i960).
The original description of this species was given by Von Linstow
(1904), and now that the new combination Splendidofilaria quiscali has appeared in print, it is a valid designation of the species.
Jones (i960), in an unpublished doctoral dissertation (The Ohio
State Uhiversity) described Splendidofilaria ohioensis as a new species, taken from the cerebral hemispheres of the Eastern Crow, Corvus brachyrhynchos brachyrhynchos. Comparative studies of these specimens indicate that the species from the Eastern Crow in Ohio is the same as the one taken from the grackle, and therefore the species Splendidofilaria ohioensis must now be placed in synonomy with Splendidofilaria quiscali.
Anderson (1961), in a revision of the genus Splendidofilaria does not mention either Splendidofilaria quiscali or Splendidofilaria ohioenais.
Splendidofilaria quiscali is similar to Splendidofilaria sinensis
Li, 1933* In both species the anterior end of the intestine is wider than the posterior end of the esophagus. However, the female and the male alike of Splendidofilaria quiscali are from one and one-half to two times as long as Splendidofilaria sinensis. In Splendidofilaria quiscali the uteri are parallel and do not cross whereas the uteri do cross each other in Splendidofilaria sinensis. Hie esophagus of
Splendidofilaria quiscali is not separated into muscular and glandular portions as is the case in Splendidofilaria sinensis. Li (1933) listed the "divided" esophagus as a species character, although Skrjabin (1923) listed the undivided esophagus as a character of the genus
Splendidofilaria.
Wehr and Hwang (1957) in a study of a species of nematode from the
cranial cavity of the Snake Bird, Anhinga anhinga, removed the species from the genus Filaria and placed it in the genus Splendidofilaria.
Realizing that the present taxonomic position of these brain nematodes
is questionable, they made this statement concerning this group,
"...which has been seen only rarely by helminthologists. It is hoped
that more and more helminthologists who are interested in the study of
filaroid nematodes will..., unravel some of the confusion surrounding
the systematic position and relationship of other species of nematodes
of this group."
Jones (i960) obtained adult specimens of Splendidofilaria in the
cranial cavity from between and within the cerebral hemispheres of the
crow. In the Common Crackle the nematode was found between and upon the
cerebral hemispheres, always beneath the pia mater, but never within the
brain tissue, a fact substantiated by the work of Odetoyinbo and Ulmer. In a preliminary report, Dr. Gordon Clark, who is conducting a survey of the parasites of the grackle in Maryland, listed a species of nematode from the sub-pia mater as Filaria quiscali. Biis probably is the same species as that of the Common Grackle.
Splendidofilaria quiscali was the most common species of worm taken from the grackle in Indiana. Of 184 cranial cavities examined, each contained several worms. Although this is the only species for which one hundred percent infection can be given, it must be understood that all other percentages are based on examinations of Uo6 birds rather than
184. Moreover, it should be pointed out that examinations of the brain were not made in either the summer or winter months. It seems unlikely that once acquired the worms would be lost from the cranial cavity of the bird. However, Jones (i960) indicated that this might occur in the crow.
Odetoyinbo and Ulmer (1959) obtained 3 - 32 adult worms per host, finding living adults even three days after death of the birds.
Odetoyinbo and Ulmer (i960) reported that the nematode
Splendidofilaria quiscali produced sheathed microfilariae which show a marked noctural periodicity, with maximum numbers present about mid night. Exposure to artificial light from 8 p.m. to 8 a.m. reversed the normal periodicity, with more microfilariae found in the period of “ forced darkness. Since the present author did not have living captive birds, blood smears were not made after dark. The birds were all taken during daylight hours, which probably accounts for the absence of microfilariae. k6
Life History:. The life history of Splendidofilaria quiscali is unknown. Work done on related species indicate various intermediate hosts or vectors. Anderson (1956) has suggested that the blackfly might he the intermediate host; Robinson (1955) believes gnats are possibly vectors of this worm; and Boyd (1956) suggests a blood-sucking louse as the vector. In either case the young birds while partially devoid of feathers likely become infected in the nest. OBSERVATIONS AND RESULTS
Four hundred and six Common Grackles were examined for helminth parasites. These birds were collected from three geographical areas of
Indiana (Fig. l). Collections were made in the spring and autumn of
1958, 1959, I960, and 1961.
Early spring collections revealed birds with gizzards void of all food material. Lawns are the frequent feeding places for these birds in the spring. By autumn, when the young birds are mature, flocks of birds feed in gardens, grain fields, etc. While the choice food of the bird is seeds, they do not refuse a meal of insects. Pieces of beetles, grasshoppers, and other unidentified insects were found in the gizzards of the birds.
During the summer this bird is rarely seen in flocks, for it is during this time that it feeds in fields and roosts near its feeding places, but when cool weather sets in the birds flock together in great numbers at feeding and roosting times. With the advent of cool weather in the autumn they move into the city or near dwellings at night to roost. Again, choice roosts are tall trees which are close together.
A pine grove on the Ball State Teachers College Campus became the roost for thousands of these birds. As cold weather with frost arrived, the grackles disappeared almost over night from the scene in Indiana.
Eleven species of helminth parasites were obtained as a result of this study. Six of these had been reported previously by various authors from this host. While five species obtained in this study con stitute new host records, all eleven species are new records for the 48 state of Indiana. This report constitutes the first record of
Crepidostomum cooperi from a warm "blooded host.
The tables which follow tabulate the results of this study, indicat ing the number of hosts examined from each region of Indiana, the kinds and numbers of parasites, and the percentage of infection with each kind.
Table 1 will give the reader some idea of previous work that has been done, showing the published records of helminth parasites from this host.
Table 7 gives a comparison of the helminth parasites of four species of related birds. These were selected because they are the only four species of passeriforms which have had surveys made of their helminth parasites. These surveys were made by different workers in different parts of the country. 1*9
Table 1. Published records of helminth parasites from the Common
Grackle, Quiscalus quiscula versicolor
Phylum and Class Species Reported by
Acanthocephala Mediorhynchus grandis Van Cleave, 1916
Platyhelrainthes
Cestoda
Trematoda Brachylecithum americanum Denton, 19^5
Conspicuum icteridorum Denton, 1951
Tanaisia bragai Byrd and Denton, 1950
Aschelminthes
Nematoda Capillaria quiscali Read, I9U9
Dispharynx spiralis Walton, 1923
Splendidofilaria quiscali Odetoyinbo and Ulmer, 1959 50
Table 2. Helminth parasites obtained from the Common Grackle in Indiana
Phylum and Class Species Site of infection
Acanthocephala Mediorhynchus grandis intestine
Platyhelminthes
Cestoda Orthoskrjabinia quiscali intestine
Paricterotaenia parina intestine
Trematoda Brachylecithum americanum bile ducts
Conspicuum icteridorum gall bladder
Conspicuum macrorchis gall bladder
Echinostoma revolutum intestine
Crepidostomum cooperi gall bladder
Aschelminthes
Nematoda Dispharynx spiralis intestine
Capillaria quiscali intestine
Splendidofilaria quiscali sub-pia mater 51
Table 3 . Percentage of infection in the Common Grackle of different
species of helminth parasites in relation to
geographical regions of Indiana
Species North Central South
Mediorhynchus grandis 14.5 11.4 6.3
Orthoskrjabinia quiscali 17.2 11.4 18.3
Paricterotaenia parina 3-6 3-5 3-0
Conspicuum icteridorum 81.0 93.6 90.9
Conspicuum macrorchis 8.09 9.5 9.02
Brachylecithum americanum 6.3 6.0 12.0
Echinostoma revolutum 4.5 — -----
Crepidostomum cooperi 0.9 — ------
Dispharynx spiralis 7.2 3.8 6.0
Capillaria quiscali 4-5 4.9 15.0
Splendidofilaria quiscali 100.0 100.0 100.0 52
Table 4. Percentage of infection of male and female Common Grackles
with different classes and species of helminths
U
Class and Species Male Female Male Female
Acanthocephala - - 20.1 27.0
Mediorhynchus grandis -- 20.1 27.0
Cestoda 15.8 11.6 18.1 1^.1
Orthoskrjabinia quiscali 15.1 10.5 17.2 9.k
Paricterotaenia parina 3-3 1.1 2.6 5.8
Trematoda 91.6 93-6 88.8 87.0
Conspicuum icteridorum 91.6 93-6 88.8 87.0
Conspicuum macrorchis 10.3 10.5 8.9 9 .^
Brachylecithum americanum 7.5 8.1 6.0 k.6
Echinostoma revolutum 1.6 1.1 0.9 1.1
Crepidostomum cooperi 0.8 - --
Nematoda 100.0 100.0 100.0 100.0
Dispharynx spiralis 7-5 3 .^ 5.1 2.3
Capillaria quiscali 5.0 k.6 8.6 3.5
Splendidofilaria quiscali 100.0 100.0 .100.0 100.0 53
Table 5* Totals of male and female birds taken in the spring and autumn
in the three geographical regions of Indiana
Region Sex of host Spring Autumn Total
Male 29 36 65 North Female 22 23 ^5
Male 83 66 1^9 Central Female 57 57 ll^
Male 13 8 21 South Female 7 ^ 12
Grand totals 211 195 ^ 6 Table 6. A composite table of the number of Common Grackles infected with different species of helminth
parasites in Indiana
North Central South Total Spring Autumn Spring Autumn Spring Autumn Spring Autumn Acanthocephala
Mediorhynchus grandis - - 15 - - 30 - - 2 - - . ^7
Cestoda Orthoskrj abinia quiscali 10 9 14 16 3 3 27 28
Paricterotaenia parina 1 3 4 4 - - 1 5 8
Trematoda Brachylecithum americanum 3 4 10 6 3 1 15 11
Conspicuum icteridorum 45 45 128 118 17 13 190 176
Conspicuum macrorchis 5 4 14 11 2 1 21 16
Echinostoma revolutum 3 2 3 2
Crepidostomum cooperi l 1 «■ m m
Nematoda Disphaiynx spiralis 6 . 2 5 5 1 1 12 8
Capillaria quiscali - - 5 8 5 2 3 10 13 Splendidofilaria quiscali 20 32 57 5b 8 8 85 99 55
Table 7* A comparison of helminth parasites in four passerine birds;
including the Common Grackle
Common Blue-.. Eastern Parasites Grackle^ jay2 Crowg Starling^
Cestodes 17.2 2.1 71.1 71.0
Trematodes 91.1 31.9 7.7 1.0
Liver flukes 91.1 29.8 5-5 0.3
Intestinal flukes 1.2 - - 2.1 0.3
Cloacal flukes - - 3.2 - - 0.3
Nematodes 100.0 1*0.0 96.7 65.7
Intestinal capillaria 5.6 17.2 61*.6 60.7
Filarial worms 100.0 31.0 93-0 - -
Proventricular worms 4.9 12.8 1*2.5 k.7
Acanthocephala 11.8 - - k.9 6.0
"Soata from present paper
^Data from Boyd (1956)
^Data from Jones (i960)
^Data from Bqyd (1951) INTERPRETATION OF DATA
Hiere are many factors which work together to affect the total distribution of parasites in a host species other than the seasoned and geographical range of the host. Many of these interrelated environment al factors, which may play a role in determining the parasitic fauna of the Common Grackle, are not considered in the scope of this report. The present report does not include any of the factors of the environment other than seasoned periodicity and geographiced distribution in the state of Indiana.
This report includes results from only four hundred and six hosts, which, in a population of birds of the magnitude of the Common Grackle, is in itself a limiting factor in making any definite assertions concerning the frequency or distribution of the parasites. It will, however, be assumed that from the foregoing observations that certain inferences may be formulated intelligently.
The Common Grackle, being a migratory bird nesting in Indiana and wintering in the south, might become infected with parasites from both places. One purpose in making this study was to determine, if possible, which parasites were obtained locally. The results do not justify a statement concerning which parasites were obtained by the birds during the winter. However, it seems that at least two were obtained by the birds locally during the summer. These are Mediorhynchus grandis
(Acanthocephala) and Echinostoma revolutum (Trematoda).
Mediorhynchus grandis was the only species of Acanthocephala taken from the host in this study. It was found only in collections made in
56 57 the autumn. Of 211 birds examined during the spring months none were found to be infected with Acanthocephala. This would indicate that this parasite was not brought in with the birds on their migratory trip from the south. Unfortunately, since collections were not made during the summer, it cannot be determined at what time or at what rate this parasite was obtained by the host. However, infestations with this species of acanthocephala were rather uniform in the autumn collections for each of the four years, and for each of the three regions of the state although they were never represented during the spring. It would seem that the intermediate host or hosts must be widely distributed in the state. If the intermediate host is the grasshopper, as Moore (19^1) determined experimentally, this would'provide a widely distributed intermediate host. Examinations of gizzard contents of birds in this study revealed that grasshoppers are included in the diet of these birds.
Mediorhynchus grandis, although obtained from all three regions, had its maximum frequency in the north and its minimum in the south.
The low frequency of 6.3 percent in the south as compared with 1^.5 per cent in the north possibly is accounted for, in part, by the smaller sample of birds taken from the south. Only thirty-three birds were collected from the southern part of the state compared to 263 from the central and 110 from the northern part. Grackles were found to be extremely uncommon in Southern Indiana. Collections from Central
Indiana showed a lower incidence of Mediorhynchus grandis than those from the north, even though more birds were taken from the central region. It would make an interesting study to check the frequencies of this parasite in a certain number of birds collected from a number of graduated zones in Indiana ranging from north to south. In this way it could be determined if there was a regular decrease in the percentage of infections with acanthocephala from north to south in this state.
Collections for this study were made from three large areas as indicated on the map. The fluctuation in frequency of Mediorhynchus grandis in the Common Grackle is perhaps more closely associated with the distri bution and availability of the intermediate host than with the geography of the state. It seems that the birds undoubtedly become infected locally with this parasite, but lose it during the winter.
There was a variation in the frequency of Mediorhynchus grandis in male compared with female birds, the females having a frequency of
27.0 percent while the males had 20.1 percent. This variation in frequency, although apparent, may not be significant since the sample of birds examined was too small for this variation to be of much significance.
There were two species of cestodes obtained in this study. No cestode has been reported previously from this host. Both
Orthoskrjabinia quiscali and Paricterotaenia parina were taken from male and female birds during both the spring and autumn and from all three geographical regions of the state. Only one specimen of Paricterotaenia parina was taken from the south. However, in view of the fact that only
33 birds were examined from this area, along with the low frequency
(3 .2 ) of the species for the state, it might be. expected that the 59 frequency would vary. No other inference can he made from this study concerning this variation. If further study revealed that the variation holds true for the entire population of birds in the south, it is likely associated with the distribution and availability of the intermediate host. In the spring, Paricterotaenia parina parasitized 3*3 percent of the males and 1.1 percent of the females. Yet in the autumn it was found in 5.8 percent of the females and 2.6 percent of the males.
An examination of table 7 reveals that the frequency of infection with cestodes was much lower for the grackle than for either the crow or the starling. However, it was considerably greater for the grackle than for the Blue-jay. It is probably significant to note that insects, which serve as intermediate hosts for cestodes, form a smaller per centage of the diet of the grackle and Blue-jay than for the starling and crow. It must be understood that the comparisons made in this table are from the results of three workers in four geographical locations.
The low incidence of Orthoskrjabinia quiscali in the grackles of
1 three areas of Indiana possibly indicates that the intermediate host
is either scarce or not frequently eaten by this bird. The variations
in incidence were not sufficient to justify any discussion.
Five species of trematodes were obtained in this study. Previous
records show only three from this bird, one of which was not found in
this survey. This one, Tanaisia bragai, has been reported only once
from the Common Grackle.
Echinostoma revolutum was obtained only from birds collected in the
north or lakes region of Indiana. Even though birds collected in the 6o spring were found to harbor this trematode, inference can still be made that the birds were infected locally. Birds in general tend to return to the same nesting sites of previous years. The Common Grackle selects the same areas for nesting sites year after year. Therefore, if
Echinostoma revolutum was obtained by this bird during the winter, it seems logical that some of the birds taken in other parts of Indiana would be found infected with it. However, it is possible that the birds which nest in Northern Indiana are also somewhat segregated in their winter range and obtain the parasites there. On the other hand, since the Common Grackle does not nest in the winter months this segregation is not likely.
Echinostoma revolutum had the greatest variation in geographical distribution of any of the helminths, being limited to the northern part of the state. Infestations with this trematode appear to be associated with birds that feed from lakes and especially lakes where the snail
Physa is found. This snail is known to serve as the intermediate host for Echinostoma revolutum. Grackles were seen feeding along the shore line of the lakes. The lakes of northern Indiana contain the snail
Physa which serves as the intermediate host for Echinostoma revolutum.
This species of trematode has not been reported previously from any sub species of grackles. Moreover, its frequency in the grackle in Indiana was extremely low, which suggests that the conditions under which the grackle becomes infected are limited. It is possible that this bird may not be a natural host but an accidental one. The infection may be 6l acquired under local conditions, such as scarcity of preferred food, prevalence of the intermediate host, or accidental consumption of the intermediate host along with other food material.
Jones (i960) reported Echinostoma revolutum from the Eastern Crow in Ohio. He obtained this trematode from both juvenile and adult birds.
Juveniles would have had to become infected locally in Ohio. The environmental conditions which prevail in Northern Ohio are similar to those of Northern Indiana. It can therefore be inferred that the life history of the species can be completed in Northern Indiana. Inter mediate hosts and parasites do not observe state boundaries.
Two species of trematodes of the genus Conspicuum were obtained from the grackle. Their frequencies in the host were quite different.
Conspicuum icteridorum was found in ninety-one percent of the four hundred and six birds examined, while Conspicuum macrorchis was found in only nine percent although it always was found in the same birds along with Conspicuum icteridorum. These two species are very similar, differing primarily in the size of the testes in relation to the size of the body. There are strong indications that only one species is represented here. The variation in size of the testes may not be justifiable reason for separation into separate species. Experimental studies with other trematodes have shown that the size of the testes varies considerably with age in the same species. The original descriptions of both species were made by the same author from different hosts. This is the first report of the two being found in the same host. 62
The frequency of infection with Brachylec ithum americanum in the south was almost twice that of--either the north or central part of the state. This variation in frequency could be, in part, the result of the smaller sample of birds taken from the southern part. However, most likely the distribution of the intermediate host or hosts in the state is responsible for the variation. If it could be determined from this study that the Common Grackle became infected locally it would give more weight to the inference that the greater incidence of Brachylecithum americanum in Southern Indiana was the result of more readily accessible intermediate hosts. Other environmental factors, such as temperature, moisture, wind, etc., undoubtedly influence the development of these cestodes. These factors were not considered in this work.
Crepidostomum cooperi was obtained from only one bird out of the
^06 posted. This bird contained two living adult flukes in its gall bladder. Apparently, until the present work, Crepidostomum cooperi has not been taken from any warm blooded host although a related species,
Crepidostomum metoecus, was described by Braun (1900) from two species of bats. This latter species was described from preserved material which Braun found in a vial in the Vienna Museum, labeled as having been taken from bats.
Hopkins (193*0 obtained Crepidostomum cooperi from fish; amphibia, and reptiles. Normally it is considered to be a parasite of the intestine of fish, with amphibia and reptiles being considered accidental or occasional hosts. Brown (1927) reported that another species, Crepidostomum farionis, was found to inhabit the gall bladder 63 as well as the intestine of trout and grayling. All reports of the other species of this genus indicate that the site of infection is the intestine of a cold blooded host, which in many cases is a fish.
The bird from which the two specimens of Crepidostomum cooperi were obtained was collected in Northern Indiana on Silver Lake, near Angola,
Indiana. By using Hopkins' (1931a) key to the species of Crepidostomum, the specimens from the Common Grackle were identified as Crepidostomum ambloplitis; however, this species was placed in synonomy with
Crepidostomum cooperi by Van Cleave and Mueller (193*0. The difference between the two species, according to Hopkins, is the relative position of the ovary in relation to the acetabulum. Further study by Hopkins and Mueller revealed that size and position of ovary in Crepidostcmam cooperi are highly variable. After a study of the descriptions of
Crepidostomum cooperi, Crepidostomum ambloplitis, and Crepidostomum solidum, all of which are now in synonomy, the present author concluded that the species taken from the Common Grackle was Crepidostomum
How and why this parasite of the intestine of fresh water fish became established in the gall bladder of a bird is difficult to explain. It undoubtedly should be considered an accidental parasite of the grackle. According to Mueller (193*0 and Hopkins (I93**a) the metacercariae, which are encysted in both the nymphal and adult stages of mayflies, are precocious. Mueller reported eggs between the cyst membrane and the metacercaria within the nymph. The fluke is, therefore, sexually mature when it is eaten by the definitive host. The grackle 6k might become infected by ingesting either infected mayfly nymphs or adults. Uhder normal conditions the active metacercaria possibly are digested or destroyed in the grackle. However, if the fluke were to wander into the bile duct and reach the gall bladder, it might find a more compatible environment in which to establish itself and mature.
Such a fluke might find within the gall bladder of the- bird an environ ment more like that of a cold-blooded host.
Hopkins (193^-a) suggested that Crepidostomum metoecus, which has been considered a bat parasite, might be a variant of Crepidostomum cooperi: Because materials for study and comparison were not avail able, he was unable to justify this synonomy. If Crepidostomum metoecus is a variant of Crepidostomum cooperi, the present report, which constitutes a new record, has completed the picture as far as classes of vertebrate hosts for this species are concerned. It has been previously reported from fish, amphibia and reptiles by Hopkins (I93^a), and from bats by ^"aun (1900b), and now from a bird in the present report.
Infection of the Common Grackle with Crepidostomum cooperi was likely obtained in the state. It was found only in one of the three geographical areas of Indiana and this was in a region where the inter mediate host, mayflies, were in abundance.
Since the parasite was found only in one bird, a male from Northern
Indiana, it was not possible to make comparisons in relation to area, season, or sex of host.
Three species of nematodes were obtained in this study. Two of these were from the intestine and one from the cranial cavity.
Capillaria quiscali was the only nematode that had a noticeable variation in frequency, in the three areas of Indiana. The highest per centage of infection with Capillaria quiscali was again found in the south for each season. As has been mentioned previously, in relation to other variations in parasite frequency, this may be the result of the relatively small sample of birds collected in the south. The distri bution may be related to intermediate hosts, but Waite (1920),
Clapham (1935)* and Cram (1936) reported that Capillaria does not need an intermediate host to complete its life history.
Dispharynx spiralis, although obtained from birds in all three areas, showed the lowest incidence in the central part of the state.
Male birds were more frequently infected with this nematode than were the females. There was this same variation for both spring and autumn over the four years in which collections were made.
Splendidofilaria quiscali was the only helminth for which one hundred percent infection can be reported, there being no variations in frequency either in distribution, season, or sex. Odetoyinbo and Ulmer
(i960) working with the grackle in Iowa, found a high percentage of infection with this worm.
Jones (i960) found a high percentage of infection with filarial worms in fledgling as well as adult crows. This led him to assume that the initial infection with filarial worms took place while the birds were devoid of feathers at a time when the arthropod vector had a better opportunity to bite the bird and transmit the microfilariae. Jones also 66 found a lower percentage of filarial infection in adults than in fledglings, and a lower percentage in winter months than in spring and autumn. He associated the geographical and seasonal distribution of filarial infections with the distribution of black flies which con ceivably might serve as the vectors. From the results he obtained,
Jones concluded that all crows at some time are infected with filarial worms. Boyd (1956), working with the Blue-jay, reported that micro filariae are obtained while the birds are in the nest. He attributed the vector to a blood sucking louse. Robinson (1955) was able to demonstrate that microfilariae were obtained by the juvenile Blue-jay while in the nest, obtaining filarial worms from only a five week old bird. Later, he raised a jay, taken from the nest, which had microfilariae at the age of sixteen days. He suggested that gnats possibly served as the vector or intermediate host. If either a gnat or blackfly is the vector of the filarial worms of the Common Grackle, it might account for the paucity of ectoparasites on this bird and yet its high percentage of infection with filarial worms.
While the present investigator found one hundred percent infection with adult filarial worms, this does not include any collections from either the summer or winter months. At some future date collections should be made, locally, during the summer to check the frequency of filarial worms. If the frequency decreases during the summer, another problem is presented, namely, how do these worms escape from the brain of the bird without doing serious damage? No such study can be made locally during the winter months because of the migration of these birds. The one hundred percent infection with filarial worms is based on 184 examinations rather than 406. The first 222 birds collected were not examined for brain nematodes. However, the 184 birds included collections from all three areas and from both seasons.
It would seem that the greater frequencies of Paricterotaenia,
Brachylecithum, and Capillaria in the Common Grackle from Southern
Indiana is the result of the diet of the birds more than any other factor. Southern Indiana differs in several aspects from the rest of the state. It is unglaciated and extremely hilly, and the land is used mostly for grazing and forestry. As a result much of the cultivated grains that serve as food for the birds in other areas is sparce or lacking in the southern part. Birds would likely turn to other sources of food, including more insects in their diet. This in turn could account for the greater percentage of infection with certain types of helminth parasites in the birds in this area. In the same way that lakes and their fauna in the northern region influence the distribution of some species, the sparcity of preferred foods in the southern part of
Indiana might cause the birds to select other foods when these are available.
An effort was made to leam if any variations existed in the rela tive frequencies of helminth parasites between the sexes of the Common
Grackle. A study of Table 4 indicates that no species of helminth parasite was found in one sex that was not found in the other, with the exception, of course of Crepidostomum cooperi. which was obtained from only one bird. However, this species can not at the present be 68 considered a natural parasite of the grackle. In general, the percentage of infection of helminths was greater for the male bird than for the female except in the case of Mediorhynchus grandis which was more frequent in the female bird.
This study does not allow one to make any inference concerning the cause of the variation in frequencies of helminth parasites between the sexes. In this study it can only be stated that during the four year period covered by this research, the percentage of infection with certain species of helminth parasites varied somewhat between the sexes of the
Common Grackle. It might be conjectured that in the spring the females' diet would be different from that of the males, especially during the time of incubation of the eggs and while feeding the young.
However, this would not account for the variations of frequencies of parasites between male and female birds in the autumn. GENERAL. DISCUSSION
A survey of the literature revealed that seven species of helminth parasites had been reported previously for the Common Grackle. Of
these, only one, the trematode Tanaisia bragai, was not found and there
fore is not reported in this paper. This trematode infects the ureters
near the kidneys; however, it probably is not a common parasite of the
grackle. Boyd (1950), in an examination of the ureters of twelve
Common Grackles from Georgia, found only one bird infected with it.
Moreover, this is the only record of it from this bird. While five of
the eleven species in the present report represent new host records,
all eleven are new records for this host in the state of Indiana.
The following helminth parasites have been reported previously
from the Common Grackle: Trematoda: Conspicuum icteridorum,
Brachylecithum americanum, and Tanaisia bragai; Nematoda: Dispharynx
spiralis, Capillaria quiscali, and Splendidofilaria quiscali; and
Acanthocephala: Mediorhynchus grandis.
The five species reported from the grackle for the first time are:
Cestoda: Orthoskrjabinia quiscali and Paricterotaenia parina;
Trematoda: Conspicuum macrorchis, Echinostoma revolutum, and
Crepidostomum cooperi. The record of Crepidostomum cooperi constitutes
the first of the family Allocreadiidae in an avian host, and the first
record of this species in a warm blooded animal.
The author believes that the results obtained warrant the inference
that at least two species of helminths, vis, Mediorhynchus grandis and
69 70
Echinostoma revolutum were obtained locally by the host.
Geographical distribution of the host's parasites can best be
explained by the distribution of intermediate hosts. The environmental
factors which might regulate the development of either the intermediate
host or the parasites were not considered in this study.
Several idiosyncrasies of the grackles might account for their
helminth fauna; vis, these birds are omnivorous, they flock together at
roosting time, and they are migratory. In 1900 a report of the Division
of Biological Survey of the United States Department of Agriculture was made on a study of the contents of the stomach of this bird. It was
found that thirty percent of the contents was animal matter and seventy
percent vegetable. Insects averaged twenty-seven percent for the year,
grain forty-seven percent, fruit five percent, and weed seeds four per
cent. Forty-one percent of the grain was com. However, in July and
August wheat formed twenty-six percent of the grain found in the
stomachs, although it formed less than one percent for the remainder of
the year. These facts indicate that grackles are a serious menace to
the farmer when they occur in large numbers. On the brighter side,
insects formed twenty-seven percent of the diet for the year. In May
this went up to sixty-three percent, but it had decreased to twelve per
cent in October. Insects are selected for food when the cereal grains
are less abundant. Beetles frequently formed the major part of the
insect diet. Barrows (1912) reported that Common Grackles do a vast
amount of good by destroying the mature May-beetle, Lachnostema, and
its larva, the "grub worm." He attributed the good condition of the 71 university lawn at Lansing, Michigan, to the work of these birds. Corn weevils and grasshoppers were found in the stomachs in increasing amounts from nineteen percent in May to twenty-three percent in August.
The grackle is a migratory bird, a fact which makes it difficult to determine the exact place of infection for each species of parasite. It spends the winter in the southern and southeastern states. Reports have been made of its wintering in isolated areas of the northern states.
The present author has not seen it in Indiana later than November or earlier than February. It has not been recorded in the Christmas Bird
Census taken by the Audubon Society. The birds usually leave Indiana in
October, and do not return until March. A few males were seen by the author in late February, 1959•
Because of the migratory nature of the birds, it is evident that some overlapping of areas would occur. It would be impossible to say that birds collected in Central or Southern Indiana early in March were not on their way to Northern Indiana. Likewise, birds collected in
Central or Southern Indiana in late autumn might be migrants from the north. Once the birds became established in their nesting areas this problem was eliminated for the three geographical territories included in this report. The differences in the three geographical regions should influence the distribution of intermediate hosts and therefore influence the distribution of the parasites of the birds within the state.
Failure to obtain ectoparasites from the birds was both dis appointing and puzzling. Even though the primary purpose of this work 72 was to study the helminth parasites the author was interested in the apparent absence of ectoparasites. It is difficult to explain the paucity or absence of ectoparasites, in view of the close proximity of the birds in both feeding and roosting. It seemed only logical to expect ectoparasites. There may be some resistant factor, or factors, which limit the ectoparasites. It can not be stated that ectoparasites do not exist on the Common Grackle. However, it can be stated that ectoparasites are either totally lacking, very infrequent, or most difficult to obtain by the usual procedures. Since birds examined in this study were not collected in individual bags to be brought into the laboratory, loss of some ectoparasites might be accounted for; however, some of the birds were still alive, although injured, when brought in for examination.
Boyd (1956) stressed that she had to search diligently for ecto parasites on the Blue-jay, whereas they were "swarming" on the starling.
The starling nests two or three times a season, using the same or an abandoned nest. Nest ectoparasites undoubtedly increase with the season, when nests are re-used. Grackles, and Blue-jays, in contrast to starlings, nest only once per season. The nest is newly constructed high in a tree, another fact which might explain the absence of ecto parasites on the Common Grackle. Boyd reported a higher incidence of ectoparasites for juveniles than adults in the starling and Blue-jay, and a higher frequency in the summer than autumn or spring. All birds collected for the present study were adults and collections were not made in the summer months. Ectoparasites might appear on the grackle in the summer or on very young birds.
Certain minor variations in the prevalence of parasites were noted between the sexes of the grackle. The work done does not justify any explanation of this. The author conjectured that the diet of the female differed from that of the male during incubation of the eggs and feeding of the young in the spring. SUMMARY
Four hundred and six Common Grackles were examined for helminth parasites. Birds were taken from three general regions of Indiana, designated as the north, central, and south areas. Collections were made during the spring and autumn months of the years 1958, 1959, 19^0, and 1961.
A survey of the literature revealed only isolated reports of helminths taken from this host. No record was found of any work done on the helminth parasites of the Common Grackle in Indiana. The following eleven species were obtained in this study: Acanthocephala:
Mediorhynchus grandis; Cestoda: Orthoskrjabinia quiscali (new host record), and Paricterotaenia parina (new host record); Trematoda:
Brachylecithum americanum, Conspicuum icteridorum, Conspicuum macrorchis (new host record), Crepidostomum cooperi (new host record), and Echinostoma revolutum (new host record); and Nematoda: Capillaria quiscali, Dispharynx spiralis, and Splendidofilaria quiscali.
The finding of Crepidostomum cooperi, in the gall bladder, con stitutes the first record of this species found in a warm blooded animal. Only one other species, Crepidostomum metoecus, in the family
Allocreadidae has been obtained from a warm blooded host. With the exception of Echinostoma revolutum, all other species of helminths were obtained from both sexes of the host from all three areas of the state. Echinostoma revolutum was found only in birds collected from the northern part of the state. The acanthocephalan, Mediorhynchus
7^ 75 grandis, was found only in the autumn collections. Since none of the birds collected in the spring were infected with this species, it would seem logical to infer that the host becomes infected with this parasite locally.
Of the birds examined, 17*2 percent were infected with cestodes
(two species); percent were parasitized with trematodes (five
species); 11.8 percent harbored acanthocephala (one species); and
100 percent were infected with nematodes (three species). A higher percentage of infection with trematodes occurred in the spring than in
the autumn, while cestodes and acanthocephala were more plentiful in
the autumn than in the spring.
The omnivorous feeding habits of the common grackle probably
accounts, in part, for the diversity and high frequency of its helminth parasites. APPENDIX
Plates I - VII
76 Plate I
•*
Fig. 1. Map of Indiana showing the three geographical areas in which
collections were made (unshaded).
77 PLATE I
SOUTH SEND SILVER L. HOOBACK L. ,^2b*0AN aO LA ^ 0 & ft NORTH
FT. WAYNE
M a r Io n :
MUNCIE
CENTRAL
INOIAN APOLIS
BEDFORD
o SALEM
o PEKIN SOUTH PLATE II
Fig. 2. Mediorhynchus grandis - entire animal , -
Fig. 3* Mediorhynchus grandis - everted proboscis
Fig. b. Mediorhynchus grandis - embryonated egg
Fig. 5* Mediorhynchus grandis - basal proboscis spine
Fig. 6 . Mediorhynchus grandis - anterior proboscis spine
79 p l a t e n
c={ Plate III
Fig. 7* Paricterotaenia parina - scolex
Fig. 8 . Paricterotaenia parina - mature proglottid
Fig.' 9* Paricterotaenia parina - hook
Fig. 10. Orthoskrjahina quiscali - scolex
Fig. 11. Orthoskrjabina quiscali - mature proglottid
Fig. 12. Orthoskrjahina quiscali - gravid proglottid
81 FIG. 10 ujui I Q FIG. 7 PLATEIH FIG. II FIG. 9 82 0®e * FIG. 8 7 /° FIG. 12 Plate IV
' • Fig. 13. Brachlecithum americanum - entire animal
Fig. I1*-. Brachylecithum americanum - cirrus sac
Fig. 15- Brachylecithum americanum - egg
Fig. 16. Conspicuum icteridorum - entire animal
Fig. 17* Conspicuum icteridorum - genitalia removed, showing excretory system
83 I mm I TSL 84 PLATE PLATE FIG. 16 FIG. I mm Plate V
Pig. 18,. Conspicuum macrorchis - entire animal
Fig. 19- Conspicuum macrorchis - cirrus sac
Pig. 20. Echinostoma revolutum - cephalic spines
Fig. 21. Echinostoma revolutum - eggs
Fig. 22. Echinostoma revolutum - entire animal
85 p l a t e a:
FIG. 19
fig. ie
0.5 mm
FIG. 20
8 6 Plate VI
Fig. 23. Crepidostomum cooperi - entire worm
Fig. 2b. Crepidostomum cooperi - genitalia
Fig. 25. Crepidostomum cooperi - papillae
Fig. 26. Capillaria quiscali - cauda of male
Fig. 27- Capillaria quiscali - lateral cauda of male
Fig. 28. Capillaria quiscali - egg
Fig. 29. Capillaria quiscali - end of spicule
87 PLATE 3ZT
FIG. 23
o U1 FIG. 24 3 3
FIG. 25
FIG. 26
FIG. 28 FIG. 27
FIG. 29
8 8 Plate VII
Fig. 30* Splendidofilaria quiscali - cauda of male
Fig. 31* Splendidofilaria quiscali - mouth and esophagus
Fig. 32. Splendidofilaria quiscali - microfilaria
Fig. 33* Splendidofilaria quiscali - vulva
Fig. 3^* Dispharynx spiralis - cauda of male
Fig. 35- Dispharynx spiralis - anterior end
89 p 3 3
ujuj 10 PIG. 30 FIG. 33 PLATE PLATE 3ZIE FIG. 34 9 0 FIG. 31 FIG. 32 FIG.35 .BIBLIOGRAPHY
Achert, James E., 1919* On the life history of Davainea tetragona, a fowl tapeworm. J. Parasitol. Sept. No. 1.
Anderson, R. C., 1956* Two new filaroid nematodes from Ontario birds. Can. J. Zool.,'XXXIV, 213 - 218.
Anderson, R. C., 1961. Revision of the genusSplendidofilaria. Canadian Journal of Zoology. 39: 201 - 20ol
Anderson, R. C. and Chabaud, A. C., 1959* Remarques sur la classifica tion des Splendidofilarinae. Ann. Parasitol. 34(1-2): 53 - 63.
Ando, A. and Ozaki, Y., 1923* On four new species of trematodes of the genus Echinostoma. Japanese text. Dobuts. Zasshi. Tokyo (413), 35: 108 - 119
Baer, J. G., 1925* Quelques Cestodes d'oiseaux nouveaux et peu connus. Bull. Soc. Sc. Nat. Neuchatel. 4-9: 138 - 154.
Barker, F. D., and Beaver, C. A*, 1915* Abstract of unpublished research listed in Barker, F. D., Parasites of American Muskrat. J. Parasitol. 1915* 1: 185*
Barker, F. D., and Irvine, R. S., 1915* Extract of unpublished research cited in Barker, F. D., 1915* J* Parasitol. 1: 189.
Barrow, W. B., 1912. Michigan Bird Life. 822 pp. Michigan Agri cultural College, Lansing.
Bassett, Lucy W., 1958. Tissue response to infection with Conspicuum icteridorum in the gall bladder of the Brewer Blackbird. J. Parasitol. 44: 471 - 476.
Beal, F. E. L., and Barrows, W. B., 1900. Division of Biological Survey of the U. S. Dept, of Agriculture. Bull. 13.
Beaver, P. C., 1937* Experimental studies on Echinostoma revolutum (Froelich) a fluke from birds and mammals. Illinois Biol. Monog. 15: l-9o.
Bhalerao, G. D., 1936. Studies on helminths of India. Trematoda I, II, III, and IV. J. Helminthol. 14 (3 ) 163 - 180; 14 (4) 181 - 206; 14 (4) 207 - 228; 15 (2 ) 97 - 124.
Boughton, D. C., Byrd, E. E. and Lund, H. 0., 1938* Microfilarial periodicity in the crow. J. Parasitol. 24 (2 ): l6l - 165.
91 92
Boyd, E. M., 1951- A survey of parasitism of the starling (Stumus vulgaris L.) in N. A. J. Parasitol. 37: 56 - 84.
Boyd, E. M., 1956. Two new species of stomach worms (Nematoda: Spintroidea) from the blue jay, Cyanocitta cristata, L. Helminthol. Soc. Wash. Proc. 23: 70 - 74.
Boyd, Dininno, Nesslinges., 1956. Metazoan parasites of the blue jay. J. Parasitol. 42: p 332.
Brackett, S., 1942. Five new species of avian schistosomes from Wisconsin and Michigan with the life cycle of G. gyrauli. J. Parasitol. 28 (l): 24 - 42.
Braun, M., 1900. Die Arten der Gattung Clinostomum Leidy. Zool. Jahrb. Syst. 14 (l): 1 - 48.
Braun, M., 1900a. Die Fasciolidengattung Clinostomum Leidy. Centralbl. Bakteriol. I. 27 (l): 24 - 32.
Braun, M., 1900b. Tremetoden der Chiroptera. Ann. k. k. naturhist. Hoffm. Wien, 15: 217 - 236.
Braun, M., 1901. Zur Revision der Tremetoden der Vogel. Centralbl. Bakteriol. I. 29: 560 - 568, 895 - 897, 94l - 948.
Bridre, J., 1910. Un dispharge du pigeon. Bull. Soc. Path. Exot., 3: 38 -39-
Brooke, M. M. and Melvin, D. M., 1955* Treton X - 100 in Geimsa staining of blood parasites. Stain Tech., 30: 263.
Brown, F. L., 1927* On Crepidostomum farionis 0. F. Muller (■Stephanonhiala laureate Zeder), a distome parasite of the trout and grayling. I. The life history. Parasitol. 19: 86 - 89.
Byrd, E. E. and Denton, J. F., 1950. The helminth parasites of birds. 1. A review of the trematode genus Tanaisia Skrjabin, 1924. Am. Mid. Nat. 43 (l): 32-57-
Chandler, A. C., 1955- Introduction to Parasitology. Ninth edition. John Wiley and Sons, Inc., New York.
Chauban, B. S., 1940. Two new species of avian trematodes. Proc. Ind. Acad. Sci. 12 (3): 75 - 83.
Chabaud, A. G. and Anderson, R. C., 1959- Nouvel essai de classifica tion des filaires (Superfamilie des Filarioidea), II Ann. Parasitol. 34 (1-2 ): 64 - 67. 93
Chaubaud, A. G., Anderson, R. C. and Brygoo, E. R., 1959* Sept filaires d'oiseaux malgaches. Ann. Parasitol. 3^ (1-2): 88 - 109*
Chabaud, A. G. and Choquet, M. T., 1953* Nouvel essai de classification des nematodes Acuariidae. Ann. Parasitol. 3^: 331 - 3^9 .
Check-list of North American Birds, 1957* 5th Ed. American Omith. Union.
Chitwood, B. G. and Chitwood, M. B., 1950* -An Introduction to Nematology. 213 PP* Monumental Printing Co., Baltimore, Maryland.
Clapham, P. A., 1935* On the experimental transmission of Syngamus trachea from starlings to chickens. J. Helminthol. 13: 1 - 2.
Clapham, P. A., 1S&0. On wild birds as transmitters of helminths parasites to domestic stock. J. Helminthol. 18 (l): 19 - i|-0.
Clerc, W., 1906. Note sur les cestodes d'oiseaux de l'Oural. I. Centralbl. Bakteriol. I. Abt. h2: ^33 - J+36.
Cohn, L., 1900. Zur Kenntnis einiger Vogeltaenien. Zool. Anz. 23: 91 - 98.
Cooper, A. R., 1915* Trematodes from marine and fresh water fishes, including one species of ectoparasitic turbellarian. Trans. Roy. Soc. Canada, ser. 3*9: 185 - 205.
Cram, E. B., 1927* Bird parasites of the nematode suborders Strongylata, Ascaridata, Spirurata. U. S. Nat. Mus. Vol. ltO: pp. 1 - 465.
Cram, E. B., 1931* Developmental stages of some nematodes of the Spiruroidea parasitic in poultry and game birds. U. S. D. A. Tech. Bull. No. 227: 1 - 27.
Cram, E. B., 1932. New records of nematodes of birds. J. Parasitol. 19: 93-
Cram, E. B., 193^* Recent records of the gizzard worm, Acuaria anthuris (Rudolphi) with observations on its life history. Proc. Helminthol. Soc. Wash. 1 (l): **8 - ^9 .
Cram, E. B., 1936. Species of Capillaria parasites in the upper digestive tract of birds. U. S. D. A. Tech. Bull. 516: 1-27* Crawford, Wiley W., 19^3* Colorado Trematode Studies. I. A Further contribution to the life history of Crepidostomum farionis (Muller). J. Parasitol. 29 : 379 - W?-
Coil, W. H., 1955* Oligorchis cynocitti, sp. nov. a hymenolepid cestode: Parasitic in the Stellar Jay. Helrainthol. Soc. Wash. Proc. 22: 112 - Hk,
Daly, E. F., 1959* A study of the intestinal helminths of the Southern Crow (Corvus brachyrhynchos paulus) in Virginia. Proc. Helminthol. Soc. Wash. 2(3: 60.
Dawes, Ben., 19^6 . The Trematoda. Cambridge University Press. Cambridge. 644 pp.
Denton, J. F., 19^5* Studies on the life history of Brachylecithum americanum n. sp., a liver fluke of passerine birds. J. Parasitol. 31 (2): 131 - 1^1 ■
Denton, J. F. and Byrd. E. E., 1950. Helminth parasites of birds. Amer. Mid. Nat. 43 (l): 32 - 57*
Denton, J. F. and Byrd, E. E., 1951* The helminth parasites of birds. III. Dicrocoeliid trematodes from N. A. birds. Proc. U. S. Mus. 101: 157 - 202.
Denton, J. F. and Rausch, R., 19^9* The occurrence of Playnosomum illicience (Braun, 1901) in a N. A. Hawk, Proc. Helminthol. Soc. Wash. 16 (1): 7-9*
DeMarvel, L., 1905* Monographic des Acanthocephales d'oiseaux. Rev. Suisse Zool., 18; 195 - 386.
Dietz, E., 1909* Die Echinostomiden der Vogel. Zool. Anz. 3^ (8): 180 - 192.
Dollfus, R. Ph., 1925* Distomiens parasites de Muridae du genre. Mus. Ann. Parasitol. Hum. et comp., 3: 85 - 102, 185 - 205.
Dollfus, R. Ph., 1951* Miscellanea helrainthologica macroccana. I. Quelques trematodes, cestodes et acanthocephales. Archaeol. Inst. Pasteur Maroc. 4 (3): 104 - 229•
Dujardin, F., 1845. Historie naturelle des helminthes ou vers intestinaux. Paris.
Fallis, A., 193^* A note on some intermediate hosts of Echinostoma revolutum (Froelich). Proc. Helminthol. Soc. WashT 1: 4 - 5* 95
Farner, D. S. and Morgan, B. B., 1944. Occurrence and distribution of the trematode, Collyrichum faba (Bremser) in birds. Auk 6l: if21 - 426 . '
Faust, E. C., 1918. Studies on American Stephanophialinae. Trans. Am. Microscop. Soc. 37: 183 - 198.
Forbush, E. H., 1927• Birds of Massachusetts and other New England States. Part II. Land Birds from Bob-whites to grackles. Norwood Pres. 461 pp.
Freeman, R. S., 1957* Life cycle and morphology of Paruterina rauschi n. sp. and P. candelabraria (Goeze, 1782) (Cestoda) from owls, and significance of plerocercoids in the order Cyclophyllidea. Canad. J. Zool. 35 (3 ): 3^9 - 370.
Fuhrman, 0., 1932. Les Tenias des Oiseaux. Mem. Univ. Nauchatel 8 : 1 - 381.
Geist, R. M., 1928. Mallophaga of passerine birds in Central Ohio. Thesis M.S.B. 313* Ohio State.
Goble, F. C. and Kutz, H. I., 1945* Notes on the gapeworms (Nematoda: Syngamidea, of Galliform and Passerform birds in New York State. J. Parasitol. 31: 394 - 400.
Goble, F. C. and Kutz, H. I., 1945a. The genus Dispharynx (Nematoda: Acuariidae) in galliform and passerform birds. J. Parasitol. 31: 323 - 331.
Guberlet, J. E. On the life history of the chicken cestode, Hymenolepis carioca. J. Parasitol. Sept. 1919/ No. 1.
Hamann, 0., 1892. Das System der Acanthocephalen. Zool. Anz. 15: 195 - 197.
Hewitt, R. Geimsa preparation for staining blood films. Science 86, 1937: 5W.
Hopkins, Sewell H., 1931* Studies on Crepidostomum. I. Crepidostomum isostomum n. sp. J. Parasitol. 17: 145 - 150.
Hopkins, Sewell H., 1931a. Studies on Crepidostomum. II. The "Crepidostomum laureatum" of A. R. Cooper. J." Parasitol. 18: 79 - 91*
Hopkins, Sewell H., 1933* The morphology, life histories and relation ships of the papillose Allocreadiidae (Trematodes) Zool. Anz. 103 (3-*0: 55 - 74. 96
Hopkins, Sewell H., 193^* Studies on Crepidostomum. III. Crepidostomum brevivitellum n. sp. J. Parasitol. 20 (5): 295 - 297.
Hopkins, Sewell H., 193to. The papillose Allocreadiidae. Illn. Biol. Monogr. 13 (2): 1 - 79-
Hopkins, Sewell H., 1937* Anew type of Allocreadiid cercaria: the cercariae of Anallocreadium and Microcreadium. J. Parasitol. 23 (1): 9^ - 97.
Hughes, R. C., 19^1- A key to the species of tapeworms in Hymenolepsis. Trans. Am. Microscop. Soc. 60: 387 - toto
Hunnenin, A. V. and Hunter, G. W. III. 1933* On the species of Crepidostomum in trout. Trans. Am. Microscop. Soc. 52 (2): 150 - 157.
Hyman, L. H., 1951* The Invertebrates. III. Acanthocephala, Aschelminthes, and Entoprocta. The Pseudocoelomate Bilateria. 572 pp. McGraw-Hill Book Co. Inc., New York.
Johnson, J., 1920. The life cycle of Echinostoma revolutum (Froelich). Univ. Calif. Pub. Zool. 19: 335 - 388.
Jones, Joseph, Jr., i960. A survey of the parasites of the Eastern Crow, Corvus brachyrhynchos brachyrhynchos Brehm. 1822, in Ohio. Dissertation. The Ohio State University. Columbus.
Kaiser, J. E., 1893* Die Acanthocephael und ihre Entwicklung. Biblioth. Zool., 7: 1 - 136.
Kostylev, N., 19lto Ein Beitrag Zur Anatomie von Gigantorhynchus otidis Miesch. Centralbl. Bakteriol. Abt. I, 72: 531 - 536.
Kostylev, N., 1915* Contributions a la faune des Acanthocephales de la Russia. Ann. Mus. Zool. Acad. Imp. S., St. Petersburg. 20: 289 - 304.
Krull, W. H., 1935. A note on the life history of Echinostoma coalitum Barker and Beaver, 1915* Proc. Helminthol. Soc. Wash. 2 : 76 .
LaRue, G. R., 1957* Classification of the digentic trematodes. Esper. Parasitol. 6 : 306 - 3to.
Leidy, Joseph, 1888. On the trematodes of the muskrat. Proc. Acad. Nat. Sc. Philadelphia, to - 126. 97
Levine, P. P. and Goble, F. C., 19^2. Parasitism and disease in ruffed grouse in New York state. In "Report of New York Ruffed Grouse Investigation," edited by G. Bump. (Uhpublished M.S.)
Li, H. C., 1933* Report on a collection of parasitic nematodes, mainly from north China. Parasitol. XXV, 192 - 210.
Looss, A., 1899. Weitere Beitrage Zur Kenntnis der Trematodenfauna Aegyptens, Zugleich Versuch einer naturlichen Gliederung des Genus Distoma Retzius. Zool. Jahrb. Syst. ,12: 521 - 78k.
Looss, A., 1902. Ueber neue und bekannte Trematoden sus Seeschildlcroten Nebst Erorterungen Zur Systematik und Nomenclatur. Zool. Jahrb.,
Lopez-Neyra, C. R., 1951* Analisis critico de los generos Choanotaenia, Anomotaenia y afines con redescripcion de la Taenia porosa Rudolphi, 1810 e invalidez de g4nero Paricterotaenia (Primera parte). Rev. iber. Parasitol., 11, 337 - 36ti. —
Luhe, M., 1910. Cestoden. In A. Bauer, Die Susswasserfauna Deutschlands. Heft 18, pp. 1 - 153.
Lutz, A., 192^. Untersuchung uber die Entwicklungsgeschichte brazilianscher Trematoden. Spezieller Teil. I. Echinostomidae. Mem. Inst. Oswaldo Cruz, Rio de Janeiro, 17: 75 - 93.
Madsen, H., 19^5* The species of Capillaria (Nematoda, Trichinelloidea) parasitic in the digestive tract of ^ gallinaceous and anatine birds, with a revised list of upecies of Capillaria of birds. Danish Rev. Game..Biol. 1: 1 - 112.
Matevosian, E. M., 19^* New cestodes of birds in Russia. Collect, pap. of helm. ded. to Skrjabin, 178 - 188.
Mawson, P. M., 1956. Ascaroid nematodes from Canadian birds. Cand. J. Zool; 3^: 35 - ^7.
Mawson, P. M., 1956a. Capillarid worms from Canadian birds, ibid. 3^: 163 - 16k.
Mawson, P. M., 1956b. Three new species of spirurid nematodes from Canadian birds, ibid. 193 - 199*
Mawson, P. M., 1957* Filariid nematodes from Canadian birds. Canad. J. Zool. 35 (2): 213 - 219.
Mayhew, R. L., Studies on the avian species of the cestode family, Hymenolepidae. Illinois Biol. Monogr. 10: 1 - 126. 1925 - 26. 98
Mettrick, D. F., 1958* Helminth parasites of Hertfordshire birds. I. Trematoda. J. Helminthol. 32 (1-2): 49 - 64.
Mettrick, D. F., 1958a. Helminth parasites of Hertfordshire birds. II. Cestoda. J. Helminthol. 32 (3): 159 - 19^-
Meyer, A., 1931* Neue Acanthocephalen aus dem Berliner Museum. Begrundung eines neuen Acanthocephalensystems auf Grund einer Untersuchung der Berliner Sammlung. Zool* Jahrb., Abt. Syst. 62: 53 - 108.
Miki, T., 1923* On the secondary intermediate host of Echinostoma cinetorchis Ando and Ozaki. Aichi Ig. Kw. Z., 30: 169 - 17^*
Molin, R., 1858. Prospectus helminthum, quae in prodromo faunae helminthologicae Venetiae Continentur. Sitzungsb. Akad. Wissensch. Wien., Math. Nat. Cl. 30: 127 - 158.
Moore, D. V., 1941. Life history and development of Macracanthorhynchus ingens and Mediorhynchus grandis (Acanthocephala"]^ J. Parasitol. 27: 34 (Suppl. 6 ).
Mueller, J. F., 1934. Two new trematodes from Oneida fishes. Trans. Amer, Microscop. Soc. 53 (3 ): 231 - 236.
Mueller, J. F., 1935- A manual of drawing for science students. Farrar and Rinehart. New York. 122 pp.
Nicoll, W., 1909* Studies on the structure and classification of the digenetic trematodes. Quart. J. Microscop. Sci. LIII. 391 - ^87.
Noble, E. R., and Noble, G. A., 1961. Parasitology. Lea and Febiger. Philadelphia. 767 PP«
Odetoyinbo, J. Adeniran and Ulmer, Martin J., 1959* Studies on avian filarial worms of the subfamily Splendidofilarinae (Nematoda: Dipetalonematidae). J. Parasitol. 45: Abstract No. l4l.
Odetoyinbo, J. Adeniran and Ulmer, Martin J., i960. Microfilarial periodicity of Splendidofilaria quiscali (m. comb.) Nematoda: Onchoceridae. J. Parasitol. 46: Abstract No. 50.
Odhner, T., 1905* Die Trematoden des arktischen Gebietes. Fauna Arctica. 4; 289 - 372.
Odhner, T., 1911* Zum naturlichen System der digenen Trematoden. Zool. Anz. 37 (8-9), 181 - 191. 99
Patten, J. A., 1952. The life cycle of Conspicuum icteridorum. Denton and Byrd 1951 (Trematoda: Dicrocoeliidae). J. Parasitol. 38: 165 - 182.
Peters, Harold S., 1936. Bird-hand. Vol. 7, No. 1. 9 “ 27. A list of external parasites of birds of the eastern part of the U. S.
Petri, L. H., 19^2. Two new dicrocoeliid trematodes from birds. Trans. Amer. Microscop. Soc. 6l: 57 - 6l. 1 pi.
Piana, G. P., 1897* Osservazioni sul Dispharagus nasutus Rud. dei polli e sulle larve nematoel-mintiche delle mosche e dei porellioni. Atti. Soc. Ital. Sci. Nat., Milano. 36 : 239 - 262.
Poche, F., 1926. Das System der Platodaria. Arch. Naturg. A. 91 (2-3), ^58 PP.
Railliet, A., 1900. Trematodes hepatiques des oiseaux. Ccmpt. Rend. Soc. Biol. 52: 239 - 3^2.
Railliet, A., and Henry, A., 19°9* Les cestodes des oiseaux par 0. Fuhrmann. Rec. Med. Vet. 86 : 337 - 338.
Railliet, A., and Henry A., 1915* Sur les Nematodes du genreGoezia Zeder. Bull. Soc. Path. exot. T. 270 - 275. (Camallanus n. g. pro Cucullanus elegans. Spinitectus cristatus n. nom. pro Filaria serratia Linton, 1901 non 1892)'.
Railliet, A., Henry, A-, and Sisoff, P., 1912. Sur les affinites des disparages (Acuaria Bremser), nematodes parasites des oiseaux. Compt. Rend. Soc. Biol. 73: 622 - 62k.
Rankin, J. S., Jr., 19^*6• Helminth parasites of birds and mammals in western Massachusetts. Am. Mid. Nat. 35: 756 - 768.
Ransom, B. H., 1909* The taenoid cestodes of North American birds. U. S. Nat. Mus. Bull. 69: 1 - lkL.
Rausch, R. and Morgan, B. B., 19^7* The genus Anonchotaenia (Cestoda: Dilepididae) from North American birds, with a description of a new species. Trans. Amer. Microscop. Soc. 66 : 203 “ 211.
Read, C. P. Studies on North American helminths of the genus Capillaria Zeder, 1800 (Nematoda) iii capillarids from lower digestive tract of North American birds. J. Parasitol. Lancaster, Pa. 35: 2^0 - 2^9 . 19^9 .
Robinson, E. J., 1955* A description of attempts to infect mosquitoes with avian filarial worms. J. Parasitol. 4l: 176 - 178. 100
Rodgers, L. 0., 19^-1• A new Dilepidid tapeworm from a Cardinal. Trans. Amer. Microscop. Soc. 60: 273 - 275.
Rudolphi, C. A., 1819. Entozoorum synopsis, cui accedunt mantissa duplex et indices locupletissimi. 1 - 811. Berlin.
Rysavy, B., 1955* Parasitische WUrmer der Singvogel (Passeriformes) in der Reservation Lednice Mme. Soc. Zool. tcheosl. 19: 99 ~ H 8 .
Sandground, J. H., 1928. A new nematode parasite, Tetrameres paucis- pina, from a South American "bird, Amblyromphus holocericeus. J. Parasitol. No. 4.
Sasser, J. N., and Jenkins, W. R., i960. Nematology. The University of North Carolina Press. Chapel Hill.
Seurat, L. G., 1913* Sur un Dispharage de la Cheveche et les affinities du genre Acuaria Bremser. C. R. Soc. Biol. Paris T. 103 - 106.
Seurat, L. G., 1916. Dispharages d 1 Algerie. Compt. Rend. Soc. Biol. 79: 931*- - 938.
Singh, S. N., 1952. Cestodes of birds. Indian Helminthol. k: 1 - 72.
Skrjabin, K. I., 1916. Nematodes des oiseaux du Turkestan russe. Ann. Mus. Zool. Acad. Imper. Sc. Petrograd 20: 457 - 557.
Skrjabin, K. I., 1923* Beitrag zur kenntnis der Vogalfilarien Russlands Rev. de Microbiol, et Epedemiol. 2: 90 - 91*
Spassky, A. A-, 19^7* (On the position of the genus Echinorhynchotaenia Puhrmann, 1909, in the system of Cestodes) (Russian text) Dokl. Akad. Nank SSSR, n. s., 58 (3), 513 - 515-
Staff ord, J., 190^. Trematodes from Canadian fishes. Zool. Anz. 27 (16-17), J*8l - 495.
Stossich, M., 1891. II genere Dispharagus Dujardin. Lavoro monographico. Boll. Soc. Adriat. Sc. Nat., Trieste. 13: 8l - 108.
Stossich, M., 1898. Saggio du una fauna elmintologica di Trieste e provincie contermini. Progr. Civ. Scuola Sup. Trieste, 162 pp.
Stossich, M., 1903- Note distomologiche. Boll. Soc. Adriat. Sc. Nat. 21: 193 - 201.
Strom, J., 19*10. Notes on the classification of the Dicrocoeliinae (Trematoda). Magasin de Parasitol. de l'Instit. Zool. d L ’Acad. des Sciences U.S.S.R. 8 ; 176 - 188. 101
Travassos, L., 1917* Helminthos du colleccae do musen Paulistra Brazil Medico 31: 121 - 122.
Travassos, L., 1926. Pilaires bresiliennes. Compt. Rend. Soc. Biol. 9^: 163 - 166.
Travassos, L., I9M*. Revisao da Familia Dicrocoeliidae odhner, 1910. Monogr. Inst. Oswaldo Cruz, No. 2, 357 PP* Rio de Janeiro.
Tuhangui, M. A., 1932. Trematode parasites of Philippine Vertebrates. V. Flukes from Birds. Philippine J. Sci., Vf: 369 - 40^.
Van Cleave, H. J., 1916. Acanthocephala of the genera Centrorhynchus and Mediorhynchus (New genus) from North American birds. Trans. Amer. Microscop. Soc. 35: 221 - 233*
Van Cleave, H. J., 1918. Acanthocephala of North American birds. Trans. Amer. Microscop. Soc. 37: 19 - **8 *
Van Cleave, H. J., 1938. The recognition of a new orderin the acanthocephala. J. Parasitol. 22 (2): 202 - 206.
Van Cleave, H. J., 19^7• The Acanthocephalan genus Mediorhynchus, its history and a review of the species occurring in the Uhited States. J. Parasitol. 33: 297 - 315*
Van Cleave, H. J., 19^- Table of characters available for the recognition of "classes" and "orders" in Acanthocephala. J. Parasitol. 3^: 18-
Van Cleave, H. J., 1952. Speciation and formation of new genera in the Acanthocephala. Syst. Zool. 1: 72 - 83.
Van Cleave, H. J., and Mueller, J. F., 1932. Parasites of the Oneida Lake fishes. Part I. Descriptions of new genera and new species. Roosevelt Wild Life Annals. Vol. 3, No. 1.
Van Cleave, H. J., and Williams, R. B., 1951* Acanthocephala from passerine birds in Alaska. J. Parasitol. 37: 151 - 159*
Vevers, G., 1923* Observations on the life-histories of Rypodaerium conoideum (Bloch) and Echinostomum revolutum (Froel): Trematode parasites of the domestic duck. Ann. Appl. von Linstow, I9O 1!-. Archiv fur Naturgeschichte. 70: 297 - 309*
Waite, R. H., 1920. Earthworms— The important factor in the trans mission of gapes in chickens. Maryland State Col. Agric. Exp. Sta. Bull. 23^. 102
Wallace, F. G., 1940. Studies on two species of live flukes. J. Parasitol. 26: 37 (Suppl. 6 ).
Walton, A. C., 1923* Some new and little known nematodes. J. Parasitol. 10: 59 - 70.
Ward, H. B., 1917* On the structure and classification of North American parasitic worms. J. Parasitol. 4 (l): 1 - 11.
Ward, H. L., 1956. A new species of Centrorhynchus (Acanthocephala) from the Kite, Melvus migrans, in Egypt. J. Parasitol. 42: 39 - 41.
Wardle, R. A., and McLeod, J. A., 1952. The zoology of tapeworms. Minneapolis, Univ. Minn. Press. 780 pp.
Wehster, J. Dan, and Addis, C. J., 1945* Helminths from the Bob-white quail in Texas. J. Parasitol. 31: 286 - 287.
Webster, J. Dan, 1947* Helminths from the Bob-white in Texas, with description of two new cestodes. Trans. Amer. Microscop. Soc. 66: 339 - 343-
Wehr, E. E., 1935* A revised classification of the nematode super family filarioidea. Proc. Helminthol. Soc. Wash. 2: 83 - 88.
Wehr, E. E., 1936. Earthworms as transmitters of Capillaria annulata, the crop worm of chickens. North Am. Vet. 17: 18 - 20.
Wehr, E. E., 1952. Recent studies on the transmission of Capillaria spp. of poultry, with special reference to Capillaria contorta. J. Parasitol. 38 (4): 17 (Suppl.).
Wehr, E. E., and Hwang, J. C., 1957* Systematic position of Filaria helicina Molin 1858, from the brain cavity of the snakebird, Anhinga anhinga. J. Parasitol. 43: 649 - 655*
Williams, 0. L., 1929* A critical analysis of the specific characters of the genus Acuaria, nematodes of birds, with descriptions of new American species. Uhiv. Cal. Pub. Zool. 33: 69 - 107.
Yamaguti, S., 1958* Systema Helminthus I. The Digenetic Trematodes of Vertebrates. 1575 PP.* Interscience Publ. Ltd. New York.
Yamaguti, S., 1959* Systema Helminthus. II. The Cestoda of Vertebrates. 859 PP* Interscience Publ. Ltd. New York.
York, W. and Maplestone, P. A., 1926* The Nematode Parasites of Vertebrates. 536 pp. P. Blakistone Son and Co., Philadelphia. AUTOBIOGRAPHY
I, George William Welker, was born in Montgomery County, near
Cumberland City, Tennessee, on July 4, 1923* I received ray secondary- school education in the public school of Cumberland City, Tennessee, and my undergraduate training at Middle Tennessee State College in
Murfreesboro, Tennessee, which granted me the Bachelor of Science degree in 1 9 ^ . During my tour of duty in the armed forces I was selected to study chemistry for one term at Oxford University in
Oxford, England. I taught high-school science in New Richmond, Ohio, from 19*1-6 to 19*+9* From George Peabody College in Nashville, Tennessee,
I received the Master of Arts degree in 1950* While in residence there,
I was assistant to Professor Jesse M. Shaver, head of The Biology
Department. I became an instructor of science (Biology) at Ball State
Teachers College in Muncie, Indiana, in 1950* I enrolled in the
Graduate School of the Ohio State University in the summer of 195^* In
1955 I was promoted Assistant Professor at Ball State Teachers
College. In 1957 1 was granted a Danforth Teacher Study Grant, at which time I completed five consecutive quarters of residence at the
Ohio State University, specializing in the department of Zoology and
Entomology.
103