This dissertation has been microfilmed exactly as received
Mic 60-41OJ.
JONES, J r ., Joseph. A SURVEY OF THE PARASITES OF THE EASTERN CROW, CORVUS BRACHYRHYNCHOS BRACHY- RHYNCHOS BREHM, 1822. INQHIO.
The Ohio State University, Ph. D ., 1960 Zoology
University Microfilms, Inc., Ann Arbor, Michigan A SURVEY OF THE PARASITES OF THE EASTERN CROW,
CORVUS BRACHYRHYNCHOS BRACHYRHYNCHOS
BREHM, 1822, IN OHIO
DISSERTATION
Presented In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University
By JOSEPH JONES, JR., B.A., M.Sc.
'k'k'icfc
The Ohio State University 1960
Approved by
Department of Zoology and Entomology ACKNOWLEDGMENTS
The completion of this study was made possible
through the cooperation and assistance of many persons.
The author wishes to thank the teachers and graduate
students in the parasitology laboratory who gave many helpful suggestions during the course of this study.
The writer wishes to acknowledge the cooperation of
Mr. Max L. Mitchell, Chief Engineer of the Miami
Conservancy District, who gave permission to shoot crows in the Englewood Dam area. Mr. Dalrymple, Lorain County game protector, helped in the collection of crows in northern Ohio. Appreciation is expressed to Dr. Allen
McIntosh of the Animal Disease and Parasite Research
Division of the Agricultural Research Service for the
loan of slides from the United States National Museum
Helminthological Collection. Appreciation is expressed to
Dr. Jean Gaud of the Institute of Parasitology, Teheran
University, for his help in the identification of the feather mites obtained in this study.
The author wishes especially to express his gratitude
for the many helpful suggestions and constructive criticisms given by his adviser, Dr. Joseph N. Miller, of the Department of Zoology and Entomology.
My wife has typed and retyped the manuscript. Her efforts greatly facilitated the preparation of this report.
-ii- TABLE OF CONTENTS
Page
INTRODUCTION...... I
HISTORICAL REVIEW ...... 3
MATERIALS AND METHODS ...... 11
CLASSIFICATION, DESCRIPTION, AND DISCUSSION OF THE PARASITES ...... 18
Protozoa ...... 18
Acanthocephala ...... 29
C e s t o d a *...... 33
Trematoda ...... 49
Nematoda ...... 63
A key to the species of the genus Splendldofllarla Skr 1abln. 1923.... 103
Arachnlda ...... 108
Insecta ...... 124
OBSERVATIONS AND RESULTS ...... 134
INTERPRETATION OF DATA ...... 158
Comparison of the Intensity and percentage of Infection of nestling, fledgling, juvenile and adult crows with various species of parasites 160
Blood Sporozoa ...... 160
Intestinal Sporozoa ...... 162
Acanthocephala...... 162
Cestoda ...... 164
Trematoda ...... 165
Nematoda ...... 166 Arthropoda ...... 172 -ill- Page
GENERAL DISCUSSION...... 174
SUMMARY ...... 180
LITERATURE CITED ...... 184
EXPLANATION OF FIGURES IN PLATES ...... 204
AUTOBIOGRAPHY...... 240
-iv- List of Tables
Table Page 1 136 Published records of parasites taken from the eastern crow, Corvus brachvrhvnchos brachvrhynchos. 139 Parasites obtained from the eastern crow in this study. 141 The percentage of infection of the eastern crow with different classes of parasites in relation to age of host and season of year. 142
Range, intensity, and percentage of infection of the eastern crow with different species of parasites in relation to the age or the host and the season of the year. 146
Range, intensity, and percentage of infection of male crows with different species of parasites in relation to the age of the host. 150
Range, intensity, and percentage of infection of female crows with different species of parasites in relation to the age of the host.
154
Percentage of infection of eastern crows with different species of parasites in relation to geographical region of the state.
-v- APPENDIX ILLUSTRATIONS
Plate Page
I ...... 206
Haemoproteus danilewskii LeucocvtozoonTsakharofn Isospora corviae
II ...... 208
Mediorhvnchus grandis Orthoskr1abinia rostellata
III ...... 210
Hvmenolepis corvi Anomotaenia constrlcta
IV ...... 212
Hvmenolepis serpentulus Hvmenolepis varxabilis
V ...... *...... 214
Echinostoma revolutum Brachylecithum americanum Amphimerus speciosus Conspicuum macrorcKTs
VI ...... 216
Capillaria anatis Capiilaria contorta
VII ...... 218
Porrocaecum ensicaudatum Syngamua trachea
VIII ...... 221
Phvsocephalua sexalatus Microtetrameres helix
-vi- Plate Page I X ...... 223
Acuaria anthuris bipiotrlaena trfcuspis
X ...... 226 Splendidofilaria flexivaginalis n. sp. Sp lendidofilar la ohioensfs n. s p .
XI ...... 228
Lamlnosioptes dihvmenalis n* sp.
XII ...... 230
Svrlngophilus bipectinatus
XIII ...... 232
Analges corvlnus
XIV ...... 234 Trouessartia corvltia Gabucinla dellbata
X V ...... 236
Phllopterus corvi Mvrsiaea albiceps Briielia rotundata
XVI ...... 239
A map showing the counties and "regions" from which crows were collected during the "summer and winter seasons."
-vii INTRODUCTION
The crow is of common and widespread occurrence
throughout the United States. It may be found in Ohio
during all seasons of the year. It is an omnivore and occurs in close association with farm animals. These
facts alone tend to indicate that it may harbor a wide variety of parasites, some of which may occur also in
farm animals. Its perennial occurrence in Ohio makes it a suitable animal for a general, year-round parasitological study. No such studies of this bird have been made in America.
Several British parasitologists (Taylor, 1928;
Clapham, 1935) have investigated the role of the starling
in the transmission of gapeworms to domesticated animals.
As a result of the present investigation, inferences have been made concerning the role of the eastern crow in the dissemination of parasites. Although several workers
(Elliott, 1903; Morgan and Waller, 1941; Robinson, 1954) have found microfilariae in the blood of crows, no descriptions of the adult worms have been published. Two new species of filarial worms have been found in the present investigation, and additional information concern
ing the morphology and systematics of this group is presented.
Feather mites have been reported from relatively few
species of American birds. Only one species has been -2- found on the eastern crow (Peters, 1936). As a result of the present investigation, one new species of feather mites is being reported and four new host records are given for North America. Moreover, additional information concerning the morphology and distribution of these forms is presented.
Although various workers have reported parasites from the eastern crow, no attempt has been made to correlate the intensity and frequency of parasitism with such factors as environment, sex, and age of this host.
In this study, the frequency and intensity of parasitism is considered in relation to the age and sex of the host and to the seasonal factor. Inferences have been made concerning the relative effects of diet, migration, and roosting on the intensity and frequency of parasitism in the crow. HISTORICAL REVIEW
Although avian parasitism has been investigated by
parasitologists in many countries, only a few bird groups
have been intensively studied in order to determine the
frequency of parasitism in relation to age and other
factors. A few surveys, dealing strictly with corvid birds, have been made in Europe and North America. In most of these studies the collection of birds was
restricted to a certain season of the year. Ward (1934) examined fifty crows, taken during the fall months, in
Oklahoma. He reported that 40 per cent of the birds were infected with mites, 70 per cent with mallophaga,
60 per cent with nematodes, and 30 per cent with cestodes. A total of thirteen species of parasites was found. Morgan and Waller (1941) examined one hundred- twelve crows, shot by hunters in Wisconsin and Iowa.
Forty-five per cent of these birds harbored tapeworms,
66 per cent contained nematodes, 6 per cent were infected with protozoa, 6 per cent with flukes and 30 per cent with ectoparasites. A total of seventeen species of parasites was represented in the study. Daly (1959), after examining the intestines of one hundred-three crows from Virginia, found 54 per cent of the birds infected with cestodes, 9 per cent with nematodes, 6 per cent with trematodes, and 1 per cent with Acanthocephala. Eight -4- species of parasites were recorded. At least three studies have been made of corvid birds in Europe.
Markowski (1933) examined 139 corvid birds of seven different species in Poland, and found fifteen species of helminth parasites. Zekhnov (1947), in a study of the effects of age on the quality and frequency of endo- parasitism in the rook, Corvus frugilegus. found that certain parasites occurred only in nestling birds, while others infected adult crows only. Davies (1958) investigated the trematode parasites of the jackdaw,
Corvus monedula. in Britain. Out of sixteen birds examined, he found five species of trematodes.
Ransom (1909) made one of the earliest general surveys of the parasitic fauna of North American birds.
In his study, which was restricted to cestode parasites, over four hundred species of tapeworms were recorded from
191 species of birds. Four were taken from corvid birds.
Mayhew (1925) examined 537 birds, representing sixty species, and reported two new species of hymenolepidid tapeworms from Corvus brachvrhvnchos. Cram (1927) published a general account of the bird parasites of the nematode suborders Strongylata, Ascaridata, and Spirurata.
Her work has become a standard reference. Later (1936), she presented a report on the capillarid nematodes occurring in the upper digestive tract of birds, and gave -5
an account of the pathogenicity and life history of
Canillaria contorta. Canavan (1929; 1931) made an
extensive survey of the nematode parasites of vertebrates
in the Philadelphia Zoological Garden. He obtained
seventeen species of parasites from fifty-six species of birds, most of which were not native to North America.
Three of the species of parasites were from corvid birds.
Rankin (1946) investigated the helminth parasites of birds in western Massachusetts. In nine species of birds, he obtained ten species of parasites, one of which was taken from the crow. Boyd (1951) , in a general
study of parasitism in starlings taken from six different
states, reported a 95 per cent infection with ecto parasites and a 99 per cent infection with endoparasites,
90 per cent of which were helminthes. A total of twenty- eight species of parasites was represented, of which
fifteen were found for the first time in North American
starlings. She thought that sixteen species of these parasites were originally imported into North America by
the starling. Denton and Byrd (1945; 1951), in an investigation of seven hundred North American birds, representing thirty-two species, reported twenty-five
species of trematodes, six of which were found in corvid birds. -6-
European parasitologists were among the earliest investigators of parasitism in birds. Clerc (1903), examined sixty-seven species of birds, and found sixty- nine species of parasites, eight of which occurred in corvid birds. Fuhrmann (1932) in his "Les Tenias des
Oiseaux," gave a general account of the species of tapeworms infecting birds. His work has become a standard reference. Recently, Mettrick (1958; 1958a), in an examination of 571 birds of various species in
Hertfordshire, England, found four species of trematodes, all of which occurred in corvid birds, and sixteen species of tapeworms, give of which were from corvid birds. Soviet parasitologists have published several studies on the parasites of birds in the Soviet Union.
Bikhovskaya-Pavlovskaya (1955) examined seven hundred-six birds from Tadzhikistan and found 24.8 per cent of them infected with trematodes. Dubinins (1950) obtained forty-eight species of tapeworms from birds in southern
Tadzhikistan. Vojtechovska-Mayerova (1953) reported twenty-seven species of cestodes, thirteen species of nematodes, three species of trematodes, and two species of Acanthocephala from 735 birds examined in
Czechoslovakia. 7-
Singh (1948; 1949; 1952) and Ali (1956) have investigated the parasitic fauna of birds in India.
Singh reported thirty-one species of parasites from twenty species of birds. Ali recorded twelve nematode species, eight being reported as new. Yamaguti (1935;
1935a; 1939; 1940; 1941; 1941a) in a study of the helminth fauna of Japan, obtained fourteen species of avian trematodes, nine of which were new, and thirty- seven species of avian nematodes, twelve of which were new. The helminths of South American birds have been studied by Travassos (1916; 1926), Caballero (1948), and Lopez-Neyra (1956).
Bird malariology has been an active branch of parasitological research for the past seventy-five years.
In no other area of zoology has the work of the field zoologist been more important in the elucidation, prevention, and treatment of human malarias. Most of the early malariologists were primarily involved in the systematics and etiology of malarial parasites.
Danilewsky (1890) was the first to describe a leucocytozoan, which he obtained from the blood of an owl. Later, Sakharoff (1893) observed similar organisms
*n Corvus corax. Corvus frugilegus. and Pica pica.
Smith (1895) discovered a leucocytozoan parasite in the blood of the domestic turkey. Ziemann (1898), -8-
Berestneff (1904), and Saxnbon (1908) made additional reports of leucocytozoan and haemoproteld parasites from birds. Aragao (1908) and Adie (1924) demonstrated the stages in the schizogony and sporogony of certain haemoproteld parasites. Ross (1898) showed that mosquitos were important vectors of parasites of the genus Plasmodium. The role of insect vectors was further elucidated by the work of Reed in the Panama Canal Zone
(Matheson, 1950). Huff (1939) published an account of a survey of blood protozoa in wild birds, and later (1942), he worked cut the stages in the schizogony of
Leucocvtozoon siroondi. a parasite of the domestic duck.
This same investigator (1943; 1947) was also the first to demonstrate the presence of an exoerythrocytic stage in the life cycle of an avian Plasmodium. Manwell (1934) studied relapse and immunity phenomena in hosts infected with malarial parasites, and later (1937) conducted experiments in the cultivation of avian plasmodia.
Coatney (1936; 1937) published a check-list and host- index of the genera Haemooroteus. Leucocvtozoon. and
Plasmodium. Coatney and Roudabush (1937) examined eighty-nine birds, representing forty-four species, in
Nebraska. Twenty-six species harbored one or more genera of blood parasites. Coatney and West (1938) examined eighty-four birds, representing thirty-five species. /
-9-
Fifteen species were found to be infected with one or more genera of parasites. Five different genera of blood parasites were reported. Herman (1938) examined blood smears from 2,385 birds taken during the summer months in Cape Cod. Twenty-seven of sixty-one species of these birds were found to be infected with blood protozoa representing the genera Haemonroteus. Leucocvtozoon.
Plasmodium. Toxoplasma, and Trypanosoma.
Few investigations have been made of the feather mites of American birds. Peters (1936) listed feather mites from only thirty-six species of American birds.
Boyd (1951), who investigated parasitism in the starling, obtained three species of feather mites and a new species of nasal mites from this host.
The feather mites of birds in Indochina and Africa have been extensively investigated by Gaud and Petitot
(1948) and Gaud and Mouchet (1959). These workers described over one hundred new species, and revised the taxonomy of several groups of mites.
American entomologists have been very active in the study of avian Mallophaga. Osborn (1896) published one of the earliest general studies of the insects of domestic and wild animals, describing several new species of Mallophaga, which included two from the American crow.
Kellogg (1896) reported a series of studies on the -10-
Mallophaga of birds in Kansas and California. Peters
(1936) compiled a list of the ectoparasites of birds
in the eastern part of the United States.
British entomologists have published at least two
extensive taxonomic works on the mallophaga. Harrison
(1916) presented an outline classification and a list of
the species of Mallophaga. In a recent study, Hopkins and Clay (1932) published a checklist of the genera and
species of Mallophaga. They also revised Harrison's classification and presented a list of synonyms for the genera and species of Mallophaga. MATERIALS AND METHODS
The crows which were used in this study were collected from twenty-five counties in Ohio (Fig. 134).
They were shot with either a 12 gauge, single action
Winchester shotgun, model 37, or a 16 gauge automatic
Remington, model 11-48. Western Super-X shells containing number six shot were used in both guns. The automatic Remington provided the most effective fire power for "roost-shooting" during the fall and winter months. The Winchester was used more often during the summer months. Most of the crows taken during the fall and winter months were shot from roosts at night.
During the spring and summer months it was necessary to use decoys and a crow call. Three papier-mache crows and one papier-mache owl were used as decoys. The decoys were placed in an open field in the vicinity of wooded areas or feeding grounds. A blind was constructed of branches and twigs and placed within shotgun range of the decoys. The crows were shot as they flew in low over the decoys. During the nesting season, adult crows were shot while on the nest. Young nestling crows
(about six weeks old) were either shot from branches or removed alive from the nest.
Crows killed in the field were transported to the laboratory, either wrapped individually in paper bags,
-11- -12 tied at the top with cord, or in newspaper bounded with cellophane tape. This procedure prevented the exchange of ectoparasites between Individual birds. It also confined ectoparasites that had left the host, and therefore made it possible to collect them from the inner lining of the bag. Wounded birds were transported in a common container. Birds brought into the laboratory were either examined immediately for parasites or placed under refrigeration, at approximately 3° C., for not more than seventy-two hours before examination.
The age of each bird was designated as either nestling, fledgling, juvenile, or adult. In central
Ohio, according to Good (1952), the hatching of crows usually occurs during the latter part of April. The newly hatched birds, referred to as nestlings, remain in the nest from four to six weeks. During this time they are fed by the parent birds. Nestling crows are either naked or have only a few brownish-black feathers; their eyes are blue; the bursa Fabricii is prominent; and the throat and inside of the beak are bright pink. By the first of June, these birds, now considered fledglings, are capable of flying and also of collecting their own food. Fledgling crows are larger and possess more feathers than do the nestlings. In the fledgling crow, a bursa
Fabricii is present; the eyes are either blue or brown, -13- and the Inside of the beak, near the tip and along the edges, is dark. The crows designated as juveniles in this study were from four to eighteen months of age. In these birds, the eyes are brown; the bursa Fabricii, when present, is smaller than in fledgling crows; and, while the inner lining of the beak is dark, the throat is pink.
In adult crows, the throat and the inside of the beak are dark; the feathers are bluish-black; the eyes are brown; the bursa Fabricii is absent; and during the mating season the testes and ovary are larger than those of juveniles.
The arthropod parasites were the first to be removed from the host. Each bird was washed in a bath which contained about five grams of a detergent (Dreft) in about two quarts of water. This solution facilitated the removal of ectoparasites from the skin of the host. The feathers were then combed with a fine-tooth comb in order to dislodge any parasites adhering to them. The bath water was then strained through a double layer of cheese cloth, and the parasites, which were suspended on the cloth, were collected with the aid of a binocular dissecting microscope. The arthropods, some of which were still alive, either were placed directly on a slide in Hoyer's Mounting Medium, or were stored in 75 per cent alcohol. Mounted specimens were dried in an oven, -14- ad justed at 40° C., for several days, after which the cover glasses were ringed with 'tourrayite."
The birds were examined for helminth parasites in the following manner; the viscera were exposed by a ventral longitudinal incision, and each organ to be examined was removed from the host and placed in a physiological salt solution for warm-blood animals, e.g.,
Ringer's "warm" solution. These organs, in the order in which they were examined, were: cloaca, bursa Fabricii, intestine, gizzard, proventriculus, esophagus, gall bladder, liver, heart, lungs, trachea, brain, eyes, nasal cavity, ears, mesenteries, and oviduct.
Cestodes and trematodes were placed in Ringer's solution which facilitated the removal of extraneous materials. The parasites were then flattened under gentle pressure of a coverslip and fixed in Lavdowsky's formula of AFA fixing reagent. Whole mounts were stained in either Semichon's Carmine or Ehrlich's Hematoxylin stain. The cestodes were sectioned at ten microns and stained in Delafield's Hematoxylin and Eosin. All preparations were mounted in piccolyte.
Acanthocephalan parasites, whose probosces were deeply embedded in the intestinal mucosa of the host, were carefully teased out of the tissue with insect pins, and placed in Ringer's solution. Relaxation of the -15-
specimens was accomplished by placing them in the
refrigerator for one or two hours. The Acanthocephala were then fixed in warm AFA fixing reagent, stained in either Semichon's Carmine or Ehrlich's Hematoxylin, and mounted in piccolyte.
Nematode parasites were relaxed in warm Ringer's
solution and fixed in either warm 75 per cent alcohol or warm AFA fixing reagent. The fixed worms were then allowed to gradually clear in a petri dish containing a
10 per cent solution of glycerin-alcohol. The glycerin-alcohol was prepared by mixing 90 c.c. of
75 per cent alcohol with 10 c.c. of glycerin. The specimens were then placed in a dessicator until the alcoholic portion of the solution had evaporated, leaving the worms suspended in pure glycerin. Clear lactophenol and lactophenol containing cotton blue were also used as clearing agents. These latter solutions were especially useful in the differentiation of papillae. En face mounts were prepared by cutting off the apical end of cleared specimens with a sharp razor blade and mounting in glycerin jelly.
Microfilariae were obtained by macerating and repeatedly dousing the lungs in Ringer's solution. Some microfilariae also were removed directly from the uterus of adult female worms. The larvae were relaxed and -16- fixed In either warm AFA fixing reagent or warm 75 per cent alcohol. Living and cleared specimens were studied with the aid of a phase contrast microscope. Some of the worms were stained in either Wright1 s or Giemsa' s blood stain.
The following procedure was used in making slides of intracellular blood parasites and microfilariae. A drop of blood was removed from the heart and liver with a clean medicine dropper. The blood from each organ was placed on a separate slide. It was spread in a thin film, dried, fixed for five minutes in absolute methyl alcohol, and stained for 45 minutes in a 2 per cent aqueous solution of Giemsa's stain. Specimens stained with Wright's stain were not fixed in methyl alcohol since this fixing reagent is incorporated in the stain.
Satisfactory mounts also were prepared by staining for one minute in concentrated Wright's stain, followed by an additional two minutes of staining in a 50 per cent aqueous solution of the stain. After staining, the smears were rinsed in distilled water, dried, and mounted under a cover glass in piccolyte. Good preparations were obtained with both stains, but the
Giemsa stain gave more uniform results.
Oocysts of coccidia were obtained by removing about two ounces of feces from the lower intestine, and placing -17- it in approximately 20 c.c. of a 2.5 per cent aqueous solution of potassium dichromate. The fecal sample was mixed thoroughly with the potassium dichromate solution, and about 10 c.c. of the mixture was placed in a petri dish. Sporulation of the oocysts was accomplished by allowing this mixture to incubate at room temperature (approximately 70° F.) for two to five days, after which it was then mixed with an equal volume of water and strained through a single layer of cheese cloth. The filtrate was collected in a small beaker, mixed with an equal volume of Shearer's Sugar Flotation solution (Morgan and Hawkins, 1948), and centrifuged for ten minutes at 1500 RPM. A drop of the material was removed from the surface of the centrifuged solution and examined for oocysts under the sixteen millimeter objective of the microscope. Detailed studies of oocysts were made with the aid of a phase contrast microscope.
All drawings were made with the aid of either a camera lucida or a microprojector.
All measurements, which are given in millimeters, are based on the material collected in Ohio. The size range is placed immediately after the structure measured, followed by the mean measurement, which is placed in parenthesis. CLASSIFICATION, DESCRIPTION, AND DISCUSSION
OF THE PARASITES
Protozoa
Birds harbor at least three classes of Protozoa, viz., Sarcodina, Mastigophora, and Sporozoa. Only one class, Sporozoa, is represented in this study. Of the
three species of Sporozoa found in the crow, two are intracellular blood parasites and one is a parasite of the epithelial cells of the intestinal mucosa.
Haemoproteus danilewskii Kruse. 1890 (Figs. 2-8).
Site of Infection - Red blood cell.
Classification. according to Kudo (1954).
Order - Haemosporidia Danilewsky, 1885.
Family - Haemoproteidae Doflein, 1916.
Genus - Haemoproteus Kruse, 1890.
Synonym: Halteridium Labbe, 1894.
Generic Diagnosis (Kudo, 1954): Gametocytes in erythrocytes, with pigment granules, halter-shaped when fully formed. Schizogony in endo thelial cells of the vertebrate host.
Specific description: With the characters of the
genus Haemoproteus. Male and female gametocytes
occurring in red blood cells.
Female gametocvte. Immature and mature female
gametocytes containing an oval, compact nucleus which
stains bright pink with Giemsa's stain. -18- -19-
Nucleus 0.001-0.003 mm. In greatest diameter. Cytoplasm vacuolated; staining light to dark blue; containing pale-green, refractile granules which are ellipsoidal in shape. Granules, approximately 0.001 mm. in length.
Mature female gametocytes containing from 11-17 granules.
Mature gametocyte, completely enveloping the host-cell nucleus, 0.009-0.014 mm. (0.013) in length; 0.006-0.007 mm. (0.0065) in width.
Male gametocyte. Immature and mature male gametocytes containing a diffuse, pale pink nucleus; 0.003-0.005 mm. in maximum diameter. Cytoplasm staining pale to dark blue; containing 9-12 pale green granules, which are ellipsoidal in shape. Mature gametocyte 0.010-0.012 mm.
(0.011) in length; 0.006-0.007 mm. (0.0065) in width, located around the host-cell nucleus.
Blood smears prepared from birds which have been dead for a few hours may show rounded gametocytes which lie adjacent to the host-cell nucleus (Fig. 3). According to Wenyon (1926) , the gametocytes become rounded in a similar manner when taken into the stomach of an invertebrate host.
Discussion. The genus Haemoproteus was established by Kruse (1890), for the species H. danilewskii. from the blood of Corvus c o m i x in Europe. Haemoproteus danilewskii differs from several other avian species of -20- this genus in that the gametocyte grows completely around the host-cell nucleus. Moreover, the cytoplasm of the parasite is vacuolated and may appear to form a radial pattern around the host-cell nucleus. Because oocyst size is the main criterion used in distinguishing the different species of the genus Haemoproteus. it is probable that many species which have been described are invalid.
Haemoproteus danilewskii has been reported from
Corvus brachvrhvnchos by several workers. Coatney and
West (1938) redescribed this species from a young crow taken in Nebraska. Herman (1938) found two of four crows infected with this parasite in Cape Cod. Coatney and
Jellison (1940) obtained it from a crow in Montana.
Morgan and Waller (1941) found this species infecting one of seventeen crows collected in Wisconsin. The present report constitutes a new host record for Haemoproteus danilewskii in Ohio.
Various species of the genus Haemoproteus occur in a wide variety of birds. Coatney and West (1938) found this genus in the hummingbird, grackle, oriole, and blue jay. Herman (1938) also reported it from the black duck, mourning dove, magnolia warbler, oriole, and several species of sparrows collected in the vicinity of
Cape Cod. -21-
Life History. The complete life history of
Haemoproteus danilewskii has not been reported. Life cycle studies indicate that the species of this genus undergo schizogony in the endothelial cells of certain organs in the vertebrate host, and undergo sporogony in the tissues of an hippoboscid fly, e.g.,
Pseudolvnchia maura (Wenyon, 1926; Kudo, 1954). -22-
Leucocvtozoon sakharoffi Sambon, 1908 (Figs. 9 and 10).
Site of Infection - Red or white blood cell.
Classification. according to Kudo (1954).
Order - Haemosporidia
Family - Haemoproteidae Dofiein, 1916.
Genus - Leucocvtozoon Danilewsky, 1890.
Generic Diagnosis (Kudo, 1954): Schizogony in the endothelial cells as well as visceral cells of vertebrates. Sexual reproduction in blood sucking insects. Gametocytes in spindle shaped host cells. Cytoplasm of gametocyte without pigment granules.
Specific description: With the characters of the
genus Leucocvtozoon. Gametocytes rounded, occurring
either in immature erythrocytes or in mature leucocytes
of the vertebrate host. Host-cells rounded.
Female gametocyte. Mature gametocyte surrounded by
the cytoplasm and nucleus of the host cell. Cytoplasm of
parasite granular, vacuolated, and staining blue with
Giemsa’s stain. Nucleus compact, light pink in color,
with bright pink karyosome located at its periphery.
Nucleus 0.003 mm. in maximum diameter. Gametocyte
0.013-0.015 mm. (0.014) in length; 0.010-0.013 mm. (0.012)
in width. Host cells containing gametocytes, 0.015-0.018 mm. (0.017) in length; 0.011-0.017 mm. (0.015) in width.
Male gametocyte. Mature gametocyte surrounded by 23-
cytoplasm and nucleus of host cell. Cytoplasm of
parasite gray, non-granular, with fewer vacuoles than
female gametocyte. Mature gametocyte 0.011-0.014 mm.
(0.012) in length; 0.007-0.012 mm. (0.010) in width.
Nucleus diffuse, with brigiht pink karyosome at its
periphery. Nucleus 0.005-0.009 mm. (0.008) in length;
0.003-0.006 mm. (0.005) in width. Host cell (with
enclosed parasite), 0.013-0.016 mm. (0.015) in length;
0.010-0.013 mm. (0.012) in width.
Discussion. The genus Leucocvtozoon was established
by Danilewsky (1890) for organisms which he discovered
in the blood of the wood owl, Svrinium aluco.
Leucocvtozoon sakharoffi was originally described by
Sambon (1908) from organisms seen in the blood of Corvus
corax. Coatney and West (1938) gave a redescription of
this species from a young eastern crow in Nebraska.
Morgan and Waller (1941) found this species in three of
seventeen crows in southern Wisconsin. The present
report is a new host record for Leucocvtozoon sakharoffi
in Ohio. Leucocytozoa occur in the blood of a wide variety
of birds. Coatney and West (1938) reported species of
this genus from the yellow-billed cuckoo. Coatney and
Jellison (1940) listed L. berestneffi from a crow in
Montana. Coatney and Roudabush (1937) obtained
L* berestneffi from the magpie, Pica pica, in Montana. -24-
Al-Dabagh (1956) found L. marchouxi and L. simondi in the mourning dove and common black duck, respectively, in Ohio.
Life History. The complete life history of
Leucocvtozoon sakharoffi has not been reported. However, life history studies of L. simondi by O'Roke (1934) and
Huff (1942) indicate that the species of the genus
Leucocvtozoon undergo schizogony in the cells of certain organs of the vertebrate host, and undergo sporogony in the tissues of blackflies, such as Simulium venusturn.
.S. njgroparvum. S. occidentale. and S. ienningsi. -25-
Isospora corvlae Ray, Shivnani, Oonanen, and Bhaskaran, 1952 (Figs. 11 and 12).
Site of Infection - Intestinal mucosa.
Classification. according to Kudo (1954).
Order - Coccidia Leukart, 1879.
Family - Eimeriidae Leger, 1911.
Genus - Isosnora Schneider, 1875.
Synonym: Diplospora Labbe. 1893.
Generic Diagnosis (Kudo, 1954): The oocyst produces two spores, each containing four sporozoites.
Specific description of exogenous stages: With the
characters of the genus Isospora. Oocysts spherical to
subspherical in shape. Spherical forms 0.020 or 0.021 mm. in diameter. Subspherical forms 0.018-0.025 mm.
(0.022) in length; 0.017-0.023 mm. (0.020) in width.
Mean length-width ratio 1.10. Oocyst wall smooth.
Inner wall thicker than outer; pale-brown in color.
Micropyle absent. Oocyst with single, non-polar,
refractile granule; without residual body Sporocyst
0.014-0.015 mm. (0.0145) in length; 0.007-0.009 mm.
(0.008) in width; lemon-shaped; with residual body.
Sporulation time, three to five days, at room
temperature (approx. 70° F.). Discussion. The genus Isospora Schneider, 1881, was
described from specimens of coccidia found in a black
i -26 slug. Labbe (1893) established the genus Dlnlosnora for coccidia which he obtained from a bird. Later, Laveran and Mesnil (1902) recognized Diplospora as a synonym of
Isospora.
Isospora corviae was originally described from Corvus macrorhvnchus intermedius. the common Himalayan crow, by
Ray, Shivnani, Oommen, and Blaskaran (1952). These workers obtained only subspherical forms which were
0.015-0.023 mm. (0.020) in length and 0.014-0.0215 mm.
(0.017) in width. The present specimens found in Corvus brachvrhvnchos brachvrhvnchos. except for minor size differences, agree with those described from Corvus macrorhvnchus intermedius. Boughton (1930), in a critical study of Isospora lacazii. a coccidian parasite commonly found in the sparrow, found a wide range of variability in the mean size of the oocysts of this species. The conclusion was drawn that either two species of Isospora occur in the sparrow or that oocyst- size alone is of limited value in determining the species of this genus. Unfortunately, oocyst-size has been extensively used as the chief character in the differentiation of species of Isospora. In fact, many species have been separated only on the basis of 27- differences in the size of the oocyst. Moreover, in many cases, the size range of different species overlap.
Apparently, some species have been described which differ only in that they were taken from different hosts. Henry (1932) and Boughton (1938) have suggested that, in the absence of cross-infection experiments, Isospora lacazii may be the only valid name which can be applied to the Isospora of passerine birds. Although this view may be justified in the case of many species, it does not appear to be applicable to Isospora corviae. In this species, the oocyst is either spherical or subspherical and contains a refractile granule, but not a residual body, while in Isospora lacazii. the oocyst is subspherical, and contains a residual body but not a refractile granule. Several species of Isospora have been reported from crows. Boughton (1938) reported Isospora sp. from Corvus brachyrhvnchos in Minnesota and New Jersey and from
C . brachvrhvnchos hesoeris and C. caurinus in California and Washington, respectively. Yakimoff and Matschoulsky
(1936) described Isospora monedula from Corvus monedula in Europe. These same investigators (1938) described
Isospora rodhaini from a raven, Corvus sp., in Europe.
Yakimoff and Gousseff (1936) described Isospora rocha-limae from the European magpie, Pica pica. -28-
Re cent ly, Becker (1956) published a host-list which contained fifteen species of Isospora from passerine birds. Life History. The complete life history of X* corviae has not been determined. From previous studies, it has been shown that avian species of Isospora undergo schizogony within the epithelial cells of the host and sporogony occurs outside the body. No Intermediate host is required (Kudo, 1954). -29
Acanthocephala
Various species of Acanthocephala occur as intestinal parasites in both terrestrial and aquatic birds. While at least seven genera are known to infect birds in Europe and North America, only one species was found in the present study.
Mediorhvnchus grandis Van Cleave, 1916 (Figs. 13-18).
Synonyms: Heteronlus grandis Van Cleave, 1918.
Empodius grandis Travassos, 1924.
Site of Infection - Intestine.
Classification. according to Meyer (1931) and Chandler (1955).
Order - Archiacanthocephala Meyer, 1931.
Family - Gigantorhynchidae Hamann, 1892.
Genus - Mediorhvnchus Van Cleave, 1916.
Synonyms: Heteronlus Kostylev, 1914.
Empodius Travassos, 1916.
Micracanthorhvnchus Travassos, 1917.
Leiperacanthus Bhalerao, 1937. Generic Diagnosis (Van Cleave, 1916): Acanthocephala of medium size* Proboscis receptacle inserted near the middle of the proboscis wall. Receptacle a single walled muscular sac with inverters or 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 inverter muscles. Cement glands, usually, eight in number. Hooks on 30-
anterlor proboscis with flask shaped roots. Hooks on posterior portion of proboscis, without reflexed roots.
Specific description: With the characters of the
genus Mediorhvnchus. Body of male and female smooth,
without spines. Proboscis ending bluntly. Hooks on the
anterior part of the proboscis arranged in seven or
eight diagonal rows, each of which contains from six to
ten hooks. Female: Body length 35-37 mm.; maximum width 1.0 mm.
Proboscis length 0.814-0.820 mm.; width at apical end
0.305 mm.; width of basal portion 0.811 mm. Proboscis
sac 0.528 mm. in length. Eggs 0.040-0.043 mm. (0.042)
in length; 0.022-0.026 mm. (0.024) in width.
Male: Body length 8.5-11.0 mm.; maximum width 1.0 mm.
Proboscis 0.726 mm. in total length; 0.231 mm. in width
at apical end; width of basal part 0.429 mm. Proboscis
sac 0.528 mm. in length. Lemnisci extending to level of
anterior testis. Testes two, tandem, in posterior third
of body; 0.363 mm. in length; 0.184 mm. in width. Cement
glands containing eight cells, located posterior to
testes. Bursa muscular, 0.126 mm. in length. Discussion. Van Cleave (1948) established the phylum
Acanthocephala, with two classes and four orders. Some
writers (Chandler, 1955; Hyman, 1951) prefer Meyer's i 31
(1931) and Van Cleave's (1936) earlier classification in which the Acanthocephala were divided into three orders.
This earlier arrangement has been retained in this report.
The genus Mediorhvnchus was established by Van Cleave
(1916) from Acanthocephala collected from passerine birds in the United States. Mediorhvnchus panillosus was designated as the type species. At least four synonyms of this genus occur in the literature. According to Van Cleave (1947), the generic name Heteronlus which
Kostylev (1914) proposed, was unavailable since it had been used by Mulsant and Rey as a name for a subgenus of beetles. Travassos (1916; 1917a) proposed two new genera, Empodius and Micracanthorhvnchus‘. for species which were later recognized as members of the genus
Mediorhvnchus. Van Cleave (1947) placed Leiperacanthus
Bhalerao, 1937, in synonymy with Mediorhvnchus.
Mediorhvnchus grandis was proposed by Van Cleave
(1916) for specimens taken from the grackle, Quiscalus quiscula. in Maryland and New Jersey, and the meadowlark,
Stumella magna, in North Carolina, Ohio, and Illinois.
Later (1918), he reported this species from Corvus brachvrhvnchos in Maryland and Ohio; from Quiscalus ouiscula aeneus in Kansas, Illinois, Ohio, and Kentucky; from the rusty blackbird, Euohagus carolinus. in -32
Illinois; and from the red-wing blackbird, Agelaius phoenicius. in Ohio. Daly (1959) examined one hundred-
three crows, Corvus brachvrhvnchos paulus. in Virginia, but found this parasite only once.
Van Cleave (1916; 1947) described two additional species, Mediorhvnchus papilloaus. which he found in the
song sparrow, Melospiza melodia: and Mediorhvnchus robustus, which he obtained from the red-wing blackbird, brown thrasher, robin, and swamp sparrow, Mediorhvnchus grandis. M. panillosus and M. robustus have not been reported outside of North America.
Other species of Acanthocephala have been taken from corvid birds. Yamaguti (1935) described Mediorhvnchus garruli from Garrulus laponicus. Petrotchenko (1949) described Centrorhvnchus skriabini from Corvus corone orientalis in Asia.
Life History. The complete life history of
Mediorhvnchus grandis is not known. Some of the intermediate hosts of this parasite were experimentally determined by Moore (1941), who fed the eggs of this form to the grasshoppers, Chorotonhaga viridifasciata australior, Ornhuella pelidna. and Arnhia luteola.
Twenty-five to thirty days later infective cystacanths were recovered from each of these hosts. -33-
Cestoda
Birds harbor at least sixteen families of cestodes
(Yamaguti, 1959). Five species, representing two families, were found in this study.
Orthoskriabinia rostellata (Rodgers. 1941) Spassky. 1947 (Figs. 19-21).
Synonym: Anonchotaenia rostellata Rodgers, 1941.
Site of Infection - Intestine.
Classification, according to Yamaguti (1959).
Order - Cyclophyllidea Braun, 1900.
Family - Dilepididae Railliet and Henry, 1909.
Genus - Orthoskr1abinia Spassky, 1947.
Synonym: Skr 1 abinerina Matevosian, 1948.
Generic Diagnosis (Yamaguti, 1959): Rostellum rudimentary, unarmed. Proglottides short, much wider than long even when gravid. . . . Genital ducts ventral to excretory stems. Testes 9-12, divided into two groups separated by female gonads. . • . Genital pores irregularly alternating. Ovary simple, median. Vitelline gland small, compact, immediately postero- ventral to ovary. • . . Paruterine organ anterior to uterus, with median swelling in front.
Specific description: With the characters of the
genus Ortho skr1abinia. Small, fragile worms. Strobila 30-35 mm. in length. Rostellum rudimentary,
unarmed. Scolex 0.785-0.792 mm. in maximum width.
Suckers 0.220-0.231 mm. in greatest diameter. Mature -34- proglottid 0.429-0.435 mm. in width; 0.115-0.119 mm. in length. Gravid proglottid 0.726-0.740 mm. in width;
0.255-0.264 mm. in length; containing paruterine organ and uterus. Paruterine organ 0.135-0.140 mm. in length;
0.076-0.084 mm. in width. Uterus posterior to paruterine organ; 0.042-0.048 mm. in length; 0.133-0.138 mm. in width. Embryos globular, 0.010-0.014 nan. in greatest diameter. Excretory system consisting of dorsal and ventral canals. Ventral excretory canal
0.021 ran. in greatest diameter. Dorsal canal 0.007 mm. in greatest diameter.
Discussion. The genus Ortho skr1abinia Spassky, 1947, was established for Anonchotaenia bobica (Clerc, 1903).
The genus Skrlabinerina Matevosian, 1948, which included the species Anonchotaenia rostellata (Rodgers, 1941), A. conica (Fuhrmann, 1908), and A. oschmarini (Spassky,
1946), was considered by Yamaguti (1959) to be a synonym of Orthoskrlabinia.
0. rostellata (Rodgers, 1941) Spassky, 1947, was originally described from the cardinal, Cardinal!s cardinal!s. in Oklahoma. It has never been recorded from any other host until the present report, which therefore constitutes a new host record for this parasite.
Life History. No life history studies have been reported for any of the species in the genus -35-
Orthoskr1abinia. Freeman (1957)> in an investigation of the life history of Paruterina candelabraria. a related species in the subfamily Paruterininae, succeeded in infecting mice with the larval stages of this parasite. It is possible that a similar inter mediate host may be involved in the life cycle of
Ortho skr1abinia rostellata.
/ -36-
Hvmenolepis corvi (Mayhew. 1925) Fuhrmann, 1932 (Figs. 22-24; 28; 29). Synonyms: Weinlandia corvi Mayhew, 1925. Mavhewia corvi Yamaguti, 1956. Site of Infection - Intestine. Classification. according to Wardle and McLeod (1952).
Order - Cyclophyllidea Braun, 1900. Family - Hymenolepididae Railliet and Henry, 1909. Genus - Hvmenolepis Weinland, 1858.
Synonyms: Diplacanthus Weinland, 1858. Lepidotrias Weinland. 1858. Dicrotaenia Railliet, 1892. Weinlandia Mayhew, 1925. Fuhrmanniella Tseng Shen, 1932.
Mavhewia Yamaguti, 1956. Variolenis Yamaguti, 1959. Generic Diagnosis (Wardle and McLeod, 1952): Rostellum usually armed with a single circle of hooks. Three testes, variable in arrangement. Cirrus pouch usually large, with an external and an internal seminal vesicle and an accessory sac. A large seminal receptacle usually present.
Specific description: With the characters of the genus Hvmenolepis. Small to medium size worms. Strobila 70-110 mm. (98.0) in length. Scolex 0.238-0.251 mm. (0.245) in maximum width, bearing a single circle of hooks. Rostellum lanceolate, 0.077-0.081 mm. (0.074) in length; 0.035-0.038 mm. (0.037) in width. Hooks 10, -37-
0.033-0.034 ran. in length. Mature proglottid 0.264-0.270
mm. (0.268) in length; 0.924-0.954 mm. (0.945) in width.
Genital pores unilateral, midway between anterior and
posterior margins of proglottid. Cirrus pouch 0.231-0.429
mm. (0.349) in length; 0.145-0.198 ran. (0.185) in maximum
width. External seminal vesicle 0.198-0.231 mm. (0.220)
in length. Testes three, globular, 0.154-0.196 ran.
(0.183) in greatest diameter. Seminal receptacle
0.198-0.264 mm. (0.237) in length. Ovary bilobate,
between testes, ventral to seminal vesicle and seminal
receptacle. Vitellaria posterior to ovary; 0.098-0.133
mm. (0.121) in greatest diameter. Two pairs of
excretory canals. Dorsal canal 0.007-0.014 mm. (0.009)
in diameter. Ventral canal 0.063-0.070 mm. (0.068) in
diameter. Uterus saccular, occasionally divided into
several compartments. Embryo globular, 0.035-0.049 mm.
(0.044) in greatest diameter. Outer egg shell
0.070-0.098 mm. (0.087) in greatest diameter. Hooklets 0.021-0.022 mm. in length.
Discussion. The genus Hvmenolepis Weinland, 1858,
includes a large number of avian and mammalian species
of tapeworms. Hughes (1941) listed 328 valid species in
the genus. Wardle and McLeod (1952) listed forty-six North
American species, but did not deal with the validity of
species from other continents. All attempts to reduce -38
the size of the genus Hvmenolepis have been generally
unsuccessful. Some workers have tried to break down the
genus into several genera or subgenera each containing a
smaller number of species. Cohn (1901) proposed two
subgenera, Hvmenolepis and Drepanidotaenia. chiefly on the
basis of differences in the number of hooks. Clerc (1902)
sought to divide the genus, using the number of testes as
the main distinguishing feature. Mayhew (1925)
established two new genera on the basis of differences in
the arrangement of the testes. This scheme also was
followed by Tseng Shen (1932), who proposed the genus
Fuhrmanniella. for species having three testes arranged
at the points of an inverted triangle. In a recent publication, Yamaguti (1959) restricted the genus
Hvmenolepis to twelve species, all of which are parasites of rodents. Under the subfamily Hymenolepidinae, he placed fifty-one genera which had been established by himself and other workers, notably Spassky and Spasskaja,
and described twenty new genera. Thirty-eight of these
genera contain species which formerly were included in
the genus Hvmenolepis. Yamaguti (1959) placed
Hvmenolepis corvi and H. serpentulus in the genus Mavhewia
Yamaguti, 1956 and considered Hvmenolepis variabilis to be a species of the genus Variolepis Spassky and
Spasskaja, 1954. The descriptions given by Yamaguti for -39- the genera Mavhewia and Variolepis are essentially the same, the only apparent difference between the two genera being the shape of the hooks. The hooks in the genus Mavhewia were described as . . small, wrench- shaped," while those of the genus Variolepis were described as having a stout guard "... nearly as long as, and parallel to blade; handle usually rather short, may be slightly curved."
It is the opinion of the writer that the differences between these two genera are minor and that the characters on which they were established are too subjectively defined and probably too variable to be of taxonomic value. For these reasons, the writer prefers to consider the two genera Mavhewia and Variolepis as synonyms of Hvmenolepis.
Hvmenolepis corvi was described by Mayhew (1925) from specimens taken from Corvus brachvrhvnchos in Illinois.
Morgan and Waller (1941) found this parasite in forty- four of one hundred-twelve crows in Iowa. Fendinger
(1952) reported it from sixty-three of eighty-eigjht eastern crows which he collected in Ohio and New York.
Life History. No life history studies of this species have been reported. In related hymenolepidid species, insects serve as second intermediate hosts (Chandler,
1955). Such hosts probably are involved also in the life cycle of Hvmenolepis corvi. -40-
Hvmenoleois serpentulus (Schrank. 1788) Weinland. 1858 ------(Figs. 32-36).
Synonyms: Taenia punctata Brugiera, 1791.
Taenia serpentulus Rudolph!. 1810.
Weinlandia serpentulus Mavhew. 1925.
Dicranotaenia serpentulus Lopez-Neyra,* 1942.
Mavhewia serpentulus Yamaguti, 1959.
Site of Infection - Intestine.
Specific description: With the characters of the genus Hvmenolepis. Small to medium size worms. Stroblla
0.030-0.105 mm. (0.090) in length. Scolex 0.105-0.110 mm. (0.107) in width at level of suckers; set off from stroblla by narrow neck. Suckers 0.074-0.077 mm. (0.076) in greatest diameter. Rostellum bearing a single circle of 10 hooks. Hooks 0.025-0.029 mm. (0.028) in length.
Extended rostellum and rostellar sac 0.120 mm. in length.
Genital pores unilateral, midway between anterior and posterior margins of proglottid. Mature proglottids wider than long, 0.792-1.20 mm. (0.903) in width;
0.300-0.330 mm. (0.315) in length. Testes three, one poral and two aporal, arranged in form of a triangle;*
0.198-0.228 mm. (0.221) in greatest diameter. Ovary median, in ventral region of proglottid, composed of approximately fifteen globular follicles. Follicles approximately 0.063 mm. in diameter. Vitellaria -41- globular, 0.092-0.112 ran. (0.098) in diameter; surrounded by ovarian follicles. Cirrus pouch with anterodorsal dilation. Cirrus and cirrus pouch 0.198-
0.231 ran. (0.220) in length; 0.059-0.070 ran. (0.066) in width. External seminal vesicle 0.175-0.210 ran. (0.189) in length; 0.075-0.098 mm. (0.087) in width. Seminal receptacle 0.231-0.396 ran. (0.307) in length; 0.112-0.198 ran. (0.179) in width. Hexacanth with polar filaments;
0.046-0.056 ran. (0.053) in greatest diameter. Hooklets
0.022-0.027 ran. (0.025) in length. Outer egg shell
0.090-0.112 mm. (0.107) in maximum diameter.
Discussion. Ransom (1909) was the first to report
Hvmenolepis serpentulus from American birds. He found this parasite in the magpie, Pica pica, and in the raven,
Corvus corax. Jones (1945) reported Hvmenolepis serpentulus sturni and H. serpentulus turdi from the starling and robin, respectively. The present report constitutes a new host record for H. serpentulus.
Hvmenolepis serpentulus has been found in corvid and passerine birds in other countries. Markowski (1933), in an examination of 139 corvid birds in Poland, found a 16 per cent infection of Corvus fruaileans with this cestode, a 33 per cent infection of C. com i x , a 50 per cent infection of Pica pica and a 16 per cent infection of
Garrulus glandarius. Yamaguti (1935) found this tapeworm -42 in Corvus corone in Japan. It has been taken also from Turdus viscivorus and Fringilla montifringilla in Czechoslovakia (Rysavy, 1955).
Life History, The complete life cycle of Hvmenolepis serpentulus is not known. Jones (1934) experimentally infected grasshoppers and beetles with the cysticercoids of Hvmenolepis variabilis. a species which also parasitizes the eastern crow. The life history of H. serpentulus also may involve intermediate host similar to those of Hvmenolepis variabilis. -43-
Hvmenolepis varlabilis (Mayhew, 1925) Fuhrmann, 1932 (Figs. 37-41). Synonyms: Wardium varlabile Mayhew,” 1925. Varilepis variabilis Spassky and Spasskaja
Site of Infection - Intestine. Specific description: With the characters of the genus Hvmenolepis. Strobila 110-210 ran. (180) in length. Scolex 0.245-0.277 ran. (0.264) in maximum width; bearing a single circle of 10 hooks. Hooks 0.019-0.021 ran. (0.020) in length. Suckers 0.072-0.084 ran. (0.078) in width. Rostellum with blunt apical end, 0.095-0.098 ran. (0.096) in length; 0.077-0.080 ran. (0.079) in maximum width. Mature proglottid 0.429-0.462 mm. (0.453) in length and 0.726-0.792 mm. (0.763) in width. Gravid proglottid width 0.376-0.528 ran. (0.501); length 0.957-1.716 ran. (1.343). Mature proglottids containing two, three, or four testes (usually three). The testes are usually arranged one poral and two aporal. However, two may occur porally and one aporally, and, occasionally, they may be arranged in a transverse row near the posterior margin of the proglottid. Rarely, a proglottid may contain two or four testes. Genital pores unilateral, midway between anterior and posterior margins of proglottid. Testes 0.165-0.264 ran. (0.207) in greatest diameter. Ovary lobate, medially located in -44- anteroventral region of proglottid. Vitellaria slightly lobed, posterior to ovary. Seminal vesicle 0.119-0.168 mm. (0.135) in length; 0.042-0.056 mm. (0.048) in width.
Dorsal excretory canal 0.014-0.022 am. (0.019) in diameter. Ventral excretory canal 0.038-0*049 mm. (0.043) in diameter. Hexacanth 0.034-0.041 mm. (0.038) in greatest diameter. Outer egg shell 0.066-0.074 mm.
(0.071) in greatest diameter. Uterus saccular, completely filling gravid proglottid.
Discussion. Hvmenolepls variabilis was originally described by Mayhew (1925) from specimens which he found in Corvus brachvrhvnchos in Illinois. Morgan and Waller
(1941) examined one hundred-twelve eastern crows in Iowa and Wisconsin, but reported only one bird infected with this parasite. Daly (1959) reported a much higher incidence of infection in Corvus brachvrhvnchos paulus in which he found twenty-five of one hundred-three of these birds infected with Hvmenolepis variabilis. The present report constitutes a new host record for Ohio.
Life History. The complete life history of Hvmenolepis variabilis is unknown. Jones (1934) fed the hexacanths of this species to the beetles, -45-
Aohrodius granarius and Oxvtelus sp., and to the. .grasshopper, Melanoplus bivittatus. and eleven to
fourteen days later, recovered the cysticercoid stage of
this parasite from these hosts. She then fed these
cysticercoids to chicks and ducks, but failed to recover
the adult stage of Hvmenolepis variabilis. 46-
Anomotaenia constricta (Molin, 1858) Cohn, 1900 (Figs. 25-27; 30 and 31). Synonyms: Taenia constricta Molin, 1858.
Taenia affinis Krabbe, 1869.
Taenia nuncta von Linstow, 1872.
Anomotaenia nuncta Cohn, 1901.
Site of Infection - Intestine.
Classification. according to Yamaguti (1959).
Order - Cyclophyllidea Braun, 1900.
Family - Dilepididae Railliet and Henry, 1909.
Genus - Anomotaenia Cohn, 1900.
Synonym: Dlnlochetos von Linstow, 1906.
Generic Diagnosis (Yamaguti, 1959): With a double crown of rostellar hooks. Proglottides craspedote, sometimes campanulate, usually wider than long, but may be longer than wide occasionally. Testes variable in number, commonly posterior, some follicles may be dorsal or lateral. Genital ducts passing between excretory stems. . . . Genital pores alternating irregularly. Ovary two-winged or not, median, with vitelline gland behind. . . . Vagina posterior to cirrus pouch, seminal receptacle present.
Specific description: With the characters of the genus Anomotaenia. Medium size worms'. Strobila 110-200 mm. (170.0) in length. Scolex 0.363-0.370 mm. (0.368) in greatest diameter; with a double crown of 22 hooks.
Hooks 0.041-0.045 mm. (0.043) in length. Rostellum
0.250-0.260 mm. (0.256) in length. Suckers 0.118-0.132 -47- mm. (0.124) in greatest diameter. Immature proglottids
separated from scolex by a narrow neck. Mature * proglottids occasionally longer than wide, 1.716-1.881 mm. (1.792) in length; 0.754-0.858 mm. (0.809) in width.
Genital pores anteriorly located. Cirrus pouch extending to the median border of excretory canals,
0.084-0.112 mm. (0.102) in length; 0.049-0.060 mm.
(0.057) in width. Vas deferens highly coiled, extending dorsally above the poral lobe of ovary. Seminal vesicle absent. Ovary bilobate, occupying most of the anterior part of the proglottid. Vitellaria between posterior
lobes of ovary, 0.172-0.231 mm. (0.204) in greatest diameter. Mehlis* gland between ovary and vitellaria.
Seminal receptacle ovate, 0.091-0.126 mm. (0.111) in
length; 0.056-0.070 mm. (0.061) in width. Testes numerous, posteriorly located; 0.066-0.084 mm. (0.076) in greatest diameter. Embryos 0.043-0.056 mm. (0.052) in maximum diameter. Outer egg shell 0.060-0.070 mm. (0.067) in greatest diameter. Hooklets six, 0.021-0.023 mm.
(0.022) in length. Ventral excretory vessel 0.056-0.066 mm. (0.063) in diameter. Dorsal excretory vessel 0.003 mm. in diameter. Uterus saccular, completely filling gravid proglottid.
Discussion. The genus Anomotaenia contains about
seventy-five species which occur primarily in birds -48-
throughout the world. Anomotaenia constricta has been
reported from corvid birds in many areas of the world.
Ransom (1909) found this parasite in Corvus brachvrhvnchos. Corvus ossifragus. Corvus corax, and
Pica pica in the United States. Morgan and Waller (1941)
examined one hundred-twelve eastern crows from Iowa and
Wisconsin, but did not find this parasite. Daly (1959) examined one hundred-three crows, Corvus brachvrhvnchos paulus. in Virginia and found thirty-five infected with
Anomotaenia constricta. The present report constitutes a new host record for this species in Ohio.
Several workers have reported Anomotaenia constricta
from crows in Europe. Markowski (1933), in an examination of 139 corvid birds in Poland, found A. constricta in one of three Corvus corax. eight of forty Corvus frugilegus.
two of fifty-four Corvus c o m i x , and five of twenty-four
Coloeus monedula. Wolffhugel (1900) reported six infections in 157 Corvus corone in Switzerland. In a
recent paper, Mettrick (1958a) reported this species from
the rook, jackdaw, starling, and blackbird in England.
Life History. No complete life history studies of any of the species in this genus have been published. Mrazek
(1907) found cysticercoids, which he reported as the bladderworm stage of Anomotaenia pyrlformis. in the earthworm Lumbriculus variegatus in Czechoslovakia. -49-
Trematoda
The class Trematoda comprises a large number of species, many of which are parasites of avian hosts.
Four species, representing three families, are reported in this study. All of these were found in the digestive system.
Echinostoma revolutum (Froelich, 1802) Looss, 1899 (Figs. 42, 45, 49). Synonyms: Fasciola revoluta Froelich. 1802.
Distoma echinatum Zeder, 1803.
Echino stoma mendax Dietz, 1909.
Echino stoma paraulum Dietz, 1909.
Echinostoma coalltum Barker and Beaver. 19i5.
Echino stoma armigerum Barker and Irvine, " 1915. Echino stoma limicoli Johnson, 1920.
Echino stoma cinetorchis Ando and Ozaki. 1923.
Site of Infection - Intestine.
Classification, according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Echinostomatidae Poche, 1926.
Genus - Echinostoma Audolphi, 1809.
Generic Diagnosis (Yamaguti, 1958): Body elongate, spinose. Head collar well developed, with double, dorsally uninterrupted crown of spines. Acetabulum near anterior extremity. Testes usually in posterior half of -50
body. Cirrus pouch rather rounded anterior to acetabulum or somewhat overlapping it. Genital pore post- bifurcal. Ovary median or a little to one side of median line in front of anterior testis. Vitellaria surrounding ceca in hindbody. Uterus long, winding in intercecal field between ovary and acetabulum.
Specific description: With the characters of the genus Echino8toma. Body elongate. Body spines extending to a point approximately 0.600 mm. posterior to testes.
Head collar spines 37. Body 5.5-7.0 mm. (6.5) in length; 1.2-2.2 mm. (1.6) in width. Oral sucker sub terminal, 0.216-0.252 mm. (0.227) in length; 0.186-0.252 mm. (0.219) in width. Pharynx 0.180-0.240 mm. (0.216) in length. Esophagus 0.270-0.480 mm. (0.410) in length.
Cirrus pouch at anterodorsal margin of acetabulum,
0.360-0.420 mm. (0.385) in length; 0.212-0.240 mm. (0.235) in width. Acetabulum 0.720-0.780 mm. (0.738) in length;
0.720-0.804 mm. (0.759) in width. Ovary ovate, in middle third of body, anterior to testes; 0.300-0.420 mm. (0.373) in length; 0.225-0.240 mm. (0.237) in width. Eggs embryonated, 0.098-0.105 mm. (0.100) in length; 0.059-0.067 mm. (0.062) in width. Vitellaria lateral, originating at posterior level of acetabulum and extending to caudal end. Testes two, in tandem, in posterior third of body;
0.540-0.552 mm. (0.545) in length; 0.390-0.420 mm. (0.410) in width. 51-
Discusslon. Descriptions have been presented of at least eighteen species of flukes which possess thirty- seven cephalic spines. Two of these species occur in the genus Echinoparvphium. Beaver (1937) in a comparison of specimens of Echinostoma revolutum from a variety of avian and mammalian hosts, reported marked variations in body size, organ size, and body spination in specimens taken from different hosts. He also demonstrated that the arrangement, number, and relative size of the cephalic spines were the least variable, and therefore the most reliable, of the specific characters.
On the basis of his study, he placed eight of the species which possessed thirty-seven cephalic spines in synonymy with Echinostoma revolutum. The following species were regarded as "species inquirendae": E. sudanense Odner,
1911; E. acuticauda Nicoll,* 1914; E. callawavensis
Barker and Noll, 1915; E. erraticum. E. neelectum.
E. nephrocvtis. E. microrchis Lutz, 1924; E. echlnocephalum.
oxvcephalum Rudolphi, 1819; E. dilatatum Miram. 1840; and E. armatum Molin, 1850.
Echinostoma revolutum occurs almost universally in a wide variety of intermediate and definitive hosts.
Adults of this species have been reported from corvid birds in North America and Europe. Daly (1959) found this parasite in six of one hundred-three crows, Corvus 52-
brachvrhvnchos paulus. collected in Virginia. Morgan and
Waller (1941) and Ward (1934) in examinations of one
hundred-twelve and fifty eastern crows, respectively,
failed to find this parasite. Markowskl (1933) obtained
E. revolutum from five of fifty-four specimens of Corvus
comix and from one of twenty-four Coloeus monedula in
Poland. The present report constitutes a new host record
for this parasite.
Life History. Many life history studies have been made
of Echino8toma revolutum. Johnson (1920) recovered
cercariae from the snail, Phvsa occidentalis. which he had
experimentally infected. Krull (1935) also experimentally
infected the snail, Pseudosuccinea columella, with
cercariae of E. revolutum. Natural infections with the
cercarial stage have been reported, also, for: Lvmnaea
nervia by Tubangui (1932); Lvmnaea stagnalis by Vevers
(1923); Stagnicola nalustris by Fallis (1934); and Phvsa
rivalis by Lutz (1924). A variety of different organisms may serve as second intermediate hosts. Beaver (1937)
recovered metacercariae from experimentally infected
specimens of the snails, Fossaria modicella and
Pseudosuccinea columella, and also from the catfish
Ameirus melas. Dollfus (1925) recovered metacercariae -53- from experimentally infected tadpoles of Bufo americana.
Miki (1923) and Dollfus (1925) recovered the metacercariae from tadpoles of Rana japonica and Rana rugosa, respectively. Beaver (1937) reported metacercariae from both tadpoles and adults of Rana piniens. -54-
Amphlmerus speciosus (Stiles and Hassall, 1896) Barker, 1911 (Figs. 43 and 48).
Synonyms: Distomum longissimum corvinum Stiles and Hassall, 1894.
Qpisthorchis speciosus Stiles and Hassall,
Site of Infection - Bile duct of liver.
Classification. according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Opisthorchiidae Braun, 1901.
Genus - Amphlmerus Barker, 1911.
Generic Diagnosis (Yamaguti, 1958): Body flattened, much elongated, tapered anteriorly, spinulate. Acetabulum small, in anterior part of body. Oral sucker larger than acetabulum. Esophagus short, ceca reaching to near posterior extremity. Testes somewhat diagonal, near posterior extremity. Ovary median or submedian, a little in front of anterior testis, simple or lobed. Seminal receptacle well- developed. Vitellaria forming a series of bunches in lateral fields, divided into two distinct regions by break opposite ovary. Uterine coils anterior to ovary. . . . Excretory vesicle Y-shaped, with sigmoid stem, winding between testes.
Specific description: With the characters of the genus Amphimerus. Body elongate. Cuticle spinose.
Body 12.0 mm. in length; 1.0 mm. in width. Oral sucker, terminal, 0.343 mm. in length; 0.297 mm. in width.
Pharynx length 0.165 mm.; width 0.165 mm. Esophagus
length 0.180 mm. Acetabulum in anterior third of body;
0.251 mm. in length; 0.251 mm. in width. Oral sucker -55
.3.5-4.5 mm. from acetabulum. Ovary ovate, pretesticular,
0.561 mm. in length; 0.343 mm. in width. Testes tandem, in posterior end of body; 0.627 mm. in length; 0.409 mm. in width. Anterior testis 0.561 mm. from ovary. Mehlis' gland on left anterolateral margin of ovary. Seminal receptacle tubular, on right posterolateral margin of ovary, 0.480 mm. in length. Excretory bladder Y-shaped, between ovary and anterior testis. Excretory canal passing diagonally between testes, forming S-shaped curve. Laurer's canal prominent. Eggs light brown, with reticulate surface; operculated; embryonated; 0.028-0.033 mm. (0.031) in length; 0.017-0.020 mm. (0.019) in width.
Discussion. Stiles and Hassall (1896) described
Amphimerus speciosus from Corvus americanus and Corvus
Q8sifragus. in Maryland and Washington, D.C. No other record of this species has been presented until the present report which constitutes a new host record.
Three additional species of the genus Amphimerus have been found in birds. These are Amphimerus interruptus
Braun, 1901, from the kingfisher, Alcedo viridirufa. and the heron, Ardea virescens. in Brazil; Amphimerus anatis
Yamaguti, 1935, from the ducks» Anas platvrhvnchos.
Nvroca ferina, and Querouedula falcata, in Japan; and
Amphimerus eloneatus Gower, 1938, from swans and ducks in Michigan. -56
Life History. The complete life cycle of Amphimerus speciosus is unknown. Wallace (1940) found the cercaria of a related species, Amphimerus elongatus. in the snail,
Amnicola limosa porata. and the metacercaria in the minnow, Pimephales promelas. in Lake Francis, Minnesota.
The same investigator also recovered the adults of
Amphimerus elongatus from experimentally infected chicks and ducks.
i -57-
Brachvlecithum americanum Denton, 1945 (Figs. 44 and 46).
Site of Infection - Bile duct of liver.
Classification< according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Dicrocoeliidae Odhner, 1911.
Genus - Brachvlecithum Strom, 1940.
Generic Diagnosis (Yamaguti, 1958): Body slender. Suckers usually equal or sub equal, close to each other. Esophagus short, ceca terminating short of posterior extremity. Testes tandem or somewhat diagonal. Cirrus pouch pre- acetabular. Genital pore usually bifurcal. Ovary median, a short distance in front of midbody. Receptaculum seminis present. Uterus filling up most of hindbody. Vitellaria occupying small lateral area posterior to ovary, sometimes commencing at level of ovary.
Specific description: With the characters of the genus Brachvlecithum. Body elongate, 3.30-5.28 mm. (4.33) in length; 0.363-0.462 mm. (0.425) in width. Oral sucker subterminal, 0.224-0.264 mm. (0.251) in length; 0.198-0.251 mm. (0.231) in width. Pharynx 0.049-0.070 non. (0.059) in length; 0.046-0.059 mm. (0.056) in width. Acetabulum in anterior fourth of body; 0.264-0.495 mm. (0.369) posterior to oral sucker; 0.264-0.343 mm. (0.311) in length;
0.217-0.297 mm. (0.277) in width. Esophagus 0.132-0.244 mm. (0.186) in length. Cirrus pouch tubular, at bifurcation of esophagus; 0.217-0.330 mm. (0.251) in length; 0.084-0.132 mm. (0.106) in width. Vitellaria, -58-
0.462-0.660 mm. (0.585) in length; laterally located in middle third of body; composed of eight or nine follicles, usually eight. Seminal receptacle globular, dorsal to posterior margin of ovary; 0.091-0.132 mm.
(0.104) in length; 0.080-0.119 mm. (0.099) in width.
Uterus in posterior region of body, passing anteriorly between testes and terminating in genital atrium, which lies in the vicinity of the cecal bifurcation. Testes lobate, 0.330-0.528 mm. (0.410) in length; 0.244-0.396 mm. (0.318) in width. Ovary lobate, posterior to testes;
0.165-0.218 mm. (0.198) in length; 0.099-0.165 mm.
(0.136) in width. Eggs operculated, embryonated,
0.042-0.045 mm. (0.043) in length; 0.021-0.026 mm. (0.025) in width.
Discussion. The genus Brachvlecithum Strom, 1940, was established for a group of flukes which formerly was included in the genus Lvnerosomum Looss, 1899. Forty-nine species of the genus Brachvlecithum have been described from birds. Two species occur in mammals. Six species occur in corvid birds.
Denton (1945) described Brachvlecithum americanum from specimens obtained from the grackle, Cassldix mexicanus prosopidicola. Brachvlecithum qiwaW ga m m was found also in the grackle, Quiscalus versicolor: the meadowlark,
Stumella magna: the crow, Corvus brachvrhvnchos: the blue jay, Cvanocitta cristata: and the hawk, Buteo -59- lineatus, in Texas, Tennessee, and Georgia. Fendinger
(1952) reported this parasite from one of eighty-eight crows collected in Ohio. Other species of the genus
Brachvlecithum have been described from corvid birds outside of North America. Railliet (1900) described
Brachvle cithum lobatum from Garrulus glandarius. Corvus co m i x . C. corone. Pica pica, and Coleus monedula in
France. Brachvlecithum stunkardi was described by Pande
(1939) from specimens taken from Garrulus lanceolatus and Cvanocitta cristata in India. Brachylecithum eugenium Oschmarin, 1948, was described from material taken from Nucifraga carvocatactes and Corvus corone orientalis in Russia. Dollfus (1954) described
Brachvlecithum alfortense from Pica pica mauritanica in
Morocco. New species also have been described from cranes, in Texas (Benton and Byrd, 1951); gulls, in
Brazil (Travassos, 1917); and owls, in Japan (Yamaguti,
1939). Life History. The life history of Brachvlecithum americanum has been investigated by Denton (1945), who succeeded in infecting the snails, Polvgvra texasiana and
P. berlandieriana. recovering the cercarial stage after one hundred-six days. Metacercariae were recovered from two chrysomelid beetles, Gastroidia cvanea and Diabrotica duodecimpunctata. Apparently, certain beetles serve as the main source of infection for birds. -60
Consplcuum macrochls Denton, 1951 (Figs. 47 and 50).
Site of Infection - Gall bladder.
Classification. according to Yamaguti (1958).
Order - Digenea Van Beneden, 1858.
Family - Dicrocoeliidae Odhner, 1911.
Genus - Consplcuum Bhalerao, 1936.
Generic Diagnosis (Yamaguti, 1958): Body flattened, fusiform, with maximum width just behind middle. Cuticle distributed irregularly with conical papillae. Esophagus 3 or 4 times as long as pharynx. Acetabulum well apart from anterior end. Testes symmetrical, Immediately post- acetabular, separated from one another by uterus. Genital pore prebifurcal. Ovary submedian, posttesticular. Seminal receptacle and Laurer'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 body.
Specific description: With the characters of the genus Consplcuum. Body ovate, apical end slightly pointed;
3.46-4.0 mm. (3.67) in length; 1.7.-1.91 mm. (1.85) in width. Oral sucker subterminal, 0.363-0.528 mm. (0.477) in length; 0.363-0.495 mm. (0.440) in width. Pharynx
0.165-0.235 mm. (0.188) in length; 0.165-0.218 mm. (0.182) in width. Esophagus 0.070-0.132 mm. (0.111) in length.
Acetabulum on border between anterior and middle third of body; 0.759-0.924 mm. (0.816) in length; 0.726-0.858 mm.
(0.806) in width; 0.396-0.429 mm. (0.412) from oral sucker. Cirrus pouch between acetabulum and oral sucker; -61-
0.330-0.594 mm. (0.445) in length; 0.132-0.178 mm. (0.153)
in width. Vitellaria 1.25-1.55 mm. (1.42) in length;
composed of many small follicles. Testes two, 0.198-0.264
mm. (0.238) in length; 0.130-0.251 mm. (0.182) in width;
at postero-lateral margin of acetabulum. Ovary ovate,
median, posttesticular, 0.297-0.363 mm. (0.332) in
length; 0.182-0.198 mm. (0.190) in width. Seminal
receptacle submedian, postovarian, 0.165-0.198 mm. (0.180)
in length. Eggs non-operculated; embryonated; 0.029-0.032
mm. (0.031) in length; 0.020-0.024 mm. (0.022) in width.
Discussion. The genus Consplcuum Bhalerao, 1936, was
established for specimens taken from the robin, Turdus
amaurochalinus. in Brazil; Five additional species of
the genus have been described; four of these were from
birds and one was from a mammal. Consplcuum macrorchis
was described by Denton (1951) from four of ten crows,
Corvus brachvrhvnchos. in Texas. No additional records
have been given for this species until the present report which constitutes a new host record for this parasite in
Ohio. Another species, Consplcuum icteridorum. was
described by Denton (1951) from the grackle Quiscalus
auiscula Aeneus in Texas. Other specimens of this
species were found by the same worker in Quiscalus
quiscula: Cassidix mexicanus prosopidicola: Cassidix
mexicanus major: Euphagua carollnus; and Sturnella magna -62-
in Tennessee, Georgia, and Michigan. Conspicuum
macrorchis and C. Icterldnnm have not been £ound outside
of North America.
Life History. The complete life history of Conspicuum macrorchis has not been determined. Denton (1951) listed
the.snail, Buliimilua alternatus mariae. as the first
intermediate host of this species. Patten (1952), in
studying the life history of a related species,
Conspicuum icteridorum. succeeded in infecting the snail,
Zonitoides arboreus. from which he later recovered
cercariae. The cercariae were fed to the isopods,
Annadillidium auadrifrons and Oniscus asellus. from which he later recovered metacercariae. These were fed to grackles, and adult flukes were recovered in twelve weeks. Denton (1951) listed the snail, Deroceras laeve. as the first intermediate host of Conspicuum icteridorum.
It is possible that Conspicuum macrorchis may infect intermediate hosts that are similar to those which harbor
Conspicuum icteridorum. -63
Nematoda
Nematodes probably occur In all species of wild birds. They may be found In a great variety of tissue sites. Of the ten species reported In this study, six occurred In the alimentary canal, two were found In the mesenteries or body cavity, one occurred In the trachea, and another In the brain.
Caolllarla anatls (Schrank, 1790) Travassos, 1915 ~ (Figs. 51-54).
Synonyms: Trichocephalus anatls Schrank, 1790.
Capillaria tmnida Zeder, 1803.
Trichocephalus fiapillaris Rudolph!. 1809.
Trichosoma brevicolle Rudolph!, 1819.
Capillaria collarls von Linstow, 1873.
Capillaria resecta Dujardin, 1843.
Capillaria cadovulvata Madsen, 1945.
Site of Infection - Mucosa of intestine.
Classification. according to Chitwood (1950).
Superfamily - Trlchuroidea Railliet, 1916.
Family - Trichuridae Railliet, 1916.
Genus - Capillaria Zeder, 1800.
Synonyms: Trichosoma Rudolph!, 1819.
Trichosomum Creplin, 1829.
Lini8cus Dujardin, 1845. Thominx Dujardin, 1845.
Calodium Dujardin, 1845. 64-
Generic Diagnosis (Yorke and Maplestone, 1926): Body capillary; mouth simple; cuticle with bacillary bands, dorsal, ventral, or lateral in position; oesophagus long and gradually increasing in size posteriorly. Male: Anus terminal or subterminal, small membranous caudal alae or bursa-like structure present or absent; spicule long and slender, surrounded by sheath with or without spines on its surface. Female: Vulva near termination of oesophagus. Oviparous, eggs lemon- shaped, with the usual opercular plugs at the poles.
Specific description: With the characters of the genus Capillaria.
Female: Body 11.0-15.1 mm. (14.0) in length;
0.052-0.079 mm. (0.066) in width. Vulva 5.3-6.0 mm. (5.6) from apical end. Esophagus 5.1-6.1 mm. (5.5) in length.
Tail blunt. Anus subterminal. Eggs 0.056-0.063 mm.
(0.058) in length; 0.028-0.030 mm. (0.029) in width.
Male: Body 9.0-13.0 mm. (11.2) in length; 0.053-0.059 mm. (0.056) in width at level of posterior end of esophagus. Esophagus 5.1-5.3 mm. (5.2) in length.
Spicule sheath smooth. Spicule sword-like, 0.924-1.330 mm. (1.094) in length; 0.013 mm. in width. Spicule sheath
0.019-0.021 mm. (0.020) in width; projecting 0.147 mm. beyond body. Cloaca terminal. Tail blunt, bearing small ala-like structures.
Discussion. Capillaria anatis (Schrank, 1790),
Travassos, 1915, was originally described from specimens -65- taken from Anas guerauedula. Madsen (1952) placed
Capillaria collaris (von Linstow, 1873); C. resecta
Dujardin, 1843), and C. cadovulvata Madsen. 1945, in synonymy with Capillaria anatis. The writer has studied descriptions and drawings of these four species, and is of the opinion that they probably represent but one species. The maximum length of the specimens of
C. anatis obtained in this study is greater by one millimeter than that of specimens described by Madsen
(1945). The outermost egg shell is not undulated as
Madsen describes it.
Capillaria anatis has been found in anatine, gallinaceous, and passerine birds. Few reports have been made for corvid birds. Mawson (1956a) found females of this species, which she reported as C. collaris. in Corvus brachvrhvnchos in Canada. The present report constitutes a new host record for this parasite. Markowski (1933) in an examination of 139 corvid birds in Poland, found this parasite, which he reported as Capillaria resecta. in one of three specimens of Corvus corax. four of forty C. frugilegus. one of fifty-four £. c o m i x . and twelve of twenty-four Coloeus monedula.
Capillaria anatis has been taken from anatine and gallinaceous birds in several countries. Cram (1936) obtained it from the turkey, Meleagris gallopavo. in the -66-
United States. Madsen (1945) found it in the ducks,
Anas platvrhvnchos. Anas crecca. Anas guerauedula. and in
grouse and pheasants in Denmark.
Life History. The complete life history of this species has not been determined. Cram (1936) infected turkeys and bobwhite quail by feeding them infective eggs of a related species, Capillaria contorta. Wehr
(1936) demonstrated that Capillaria annulata. the "crop worm of chickens," could be transmitted by earthworms. The life history of Capillaria anatis may be similar to that of C. contorta or C. annulata. -67-
Capillaria contorta (Creplin, 1839) Travassos, 1915 (Figs. 55-57).
Synonyms: Trichosoma contortum Creplin. 1839.
Trichosomum contortum Dujardin, 1845.
Capillaria perforans Kotlan and Orosz, 1931.
Capillaria lophortygis Baylis, 1934.
Capillaria vanelli Yamaguti, 1935.
Site of Infection - Mucosa of esophagus.
Specific description: With the characters of the genus Capillaria.
Female: Body highly coiled, 17.0-25.0 mm. (22.0) in length; 0.112-0.132 mm. (0.128) in width, in the region of vulva. Vulva 4.5-6.3 mm. (5.5) from apical end.
Esophagus 4.3-6.0 mm. (5.4) in length; terminating
0.198-0.264 mm. (0.231) from vulva. Eggs 0.056-0.070 mm.
(0.063) in length; 0.025-0.028 mm. (0.026) in width.
Anus terminal. Tail tapering and slightly round at end.
Male: Body 11.0-17.0 mm. (13.0) in length; 0.079-0.103 mm. (0.090) in width, at the level at which the esophagus joins with the intestine. Esophagus 4.5-5.5 mm. (5.0) in length. Spicule sheath spinose, ending bluntly; extending
0.490-0.495 mm. (0.493) beyond tail. Spicule not seen.
Cloaca terminal. Tail ending bluntly.
Discussion. Capillaria contorta (Creplin, 1839),
Travassos, 1915, has been found in at least five different orders of birds. It was originally described from 68
specimens taken from the crow in Europe (Yorke and
Maplestone, 1926). It was reported from Corvus brachvrhvnchos by Canavan (1931) in Pennsylvania. Morgan and Waller (1941) found it in two of one hundred-twelve crows collected in Iowa and Wisconsin. Fendinger (1952) reported this species from thirty-seven of eighty-eight crows in Ohio and New York. In Europe, Canillaria contorta was found by Skrjabin (1916) and Korkhaus (1935) in both Corvus frueileeus and Corvus corone. Markowski
(1933) examined 139 corvid birds in Poland, but apparently failed to find this species.
Capiliaria contorta is a common parasite of ducks, turkeys, and chickens. Cram (1936) obtained specimens of this species from Bonasa umbellus. Meleagris gallopavo.
Phasianus colchicus, Perdix perdix. Anas boschas domestica. and Dafila acuta in the United States. Boyd (1951) also reported it from the starling in the United States.
Madsen (1945) took specimens of Capillaria contorta from
Anas platvrhvnchos. Anas crecca, and from pheasants in
Denmark, but failed to find it in marine diving ducks and in the black grouse.
Railliet and Lucet (1889; 1890) and Cram (1936) investigated the pathogenicity of Capillaria contorta.
These investigators reported symptoms of 'Vmiscular incoordination, leg weakness and limberneck" in heavily -69- infected ducks and chickens. Infections of twenty-five to thirty-five worms were usually fatal to ducks, but galliform birds were able to tolerate greater numbers of
this parasite. The writer examined one crow that harbored as many as forty worms apparently without ill effects.
Life History. The life history of Capillaria contorta has been investigated by Cram (1936), who succeeded in infecting six turkeys and two bobwhite quail by feeding them infective eggs of C. contorta. In one case, the eggs of the parasite remained infective even after being kept under conditions of dryness for eleven months. Wehr (1936) showed that Capillaria annulata. the "crop worm of chickens," may be transmitted by earthworms. It is possible that the earthworm may also act as an intermediate host for C. contorta. 70
Porrocaecum angleaudafcnm (Zeder, 1800) Bay 11s, 1920 (Figs. 58-64). Synonyms: Fnsarla enslcaudata Zeder, 1800.
Ascarls enslcaudata Rudolph!, 1809.
Site of Infection - Intestine.
Classification. according to Chitwood (1950).
Superfamily - Ascaridoidea Railliet and Henry, 1915*
Family - Ascarididae Blanchard, 1896.
Genus - Porrocaecum Railliet and Henry, 1912.
Synonym: Terranova Leiper and Atkinson, 1914.
Generic Diagnosis (Yorke and Maplestone, 1926): Interlabia (usually small) present in most cases; dentigerous ridges present; oesophagus with an anterior muscular portion and a posterior ventriculus of oblong shape; the latter is short in the genotype but in other species frequently long and bent at an angle so as to open into the intestine laterally; intestinal caecum present; oesophageal appendix absent. Male: Spicules equal; gubernaculum usually absent. Female: Vulva near the middle of the body. Oviporous.
Specific description: With the characters of the genus Porrocaecum. Body cylindrical, with thick, trans versely striated cuticle. Head demarcated from rest of body by slight, narrow constriction. Mouth surrounded by three lips. Dorsal lip with two papillae; ventrolateral lips with one papilla each.
Female: Body 27.0-36.0 mm. (34.0) in length in immature specimens and up to 80.0 mm. in mature worms. -71-
Vulva 52.0 mm. from apical end in mature specimens.
Vagina 4.95 mm. in length. Esophagus 4.75 mm. in length.
Esophageal ventriculus oblong in shape; 0.363-0.759 mm.
(0.583) in length; 0.396 mm. in width. Intestinal cecum rudimentary, 0.200 mm. in length. Nerve ring 0.495-1.12 mm. (0.804) from apical end* Eggs 0.070-0.080 mm. (0.077) in length; 0.056-0.065 mm. (0.063) in width. Anus subterminal, 0.462-0.693 mm. (0.602) from posterior end.
Tail tapering to rounded end.
Male: Body 30.0-40.0 mm. (38.0) in length; 0.436-0.653 mm. (0.560) in width, at level of ventriculus. Esophagus
2.50-2.83 mm. (2.75) from apical end. Ventriculus oblong,
0.383-0.396 mm. (0.390) in length; 0.196-0.231 mm. (0.215) in width. Intestinal cecum 0.120-0.132 mm. (0.126) in length. Nerve ring 0.547-0.646 mm. (0.597) from anterior end. TWo spicules, equal in size; 0.528-0.594 mm. (0.551) in length. Cloaca 0.263-0.303 mm. (0.283) from posterior end. Tail bearing fourteen pairs of preanal papillae, linearly arranged; a double pair of papillae at the level of the anus; four pairs of asymmetrically arranged, postanal papillae; and one pair of lateral phasmids.
Tail subacute.
Discussion. The genus Porrocaecum Railliet and Henry,
1912, differs from the closely related genus Contracecum by the presence of an intestinal cecum and the absence -72- of an esophageal appendix. In the genus Contracecum both of these structures are present.
Porrocaecum enslcaudatum (Zeder, 1800) Bay11s, 1920, was originally placed under the genus Fusarla. Cram
(1927), Markowskl (1933), and Mawson (1956) added to the original description. There Is some disagreement concerning the number of cephalic papillae occurring In this species. Cram (1927) reported one pair of papillae for each lip. Markowskl (1933) reported one large pair and an additional four smaller papillae on each lip.
Mawson (1956) reported two papillae on the dorsal and a single papilla on each of the ventrolateral lips. The writer's observations are In agreement with those of
Mawson (1956). Cram (1927) presented a drawing (after von Llnstow) which shows two papillae on the dorsal lip of P. enslcaudatum. Her description apparently was based on this drawing. The writer's observations on the number of caudal papillae are essentially in agreement with those of Cram (1927) . However, the papillae are arranged assymmetrically rather than symmetrically, as Cram shows them. The structures which Cram refers to as "lateral papillae" appear to be phasmids.
Porrocaecum enslcaudatum has been taken from corvid birds In Europe and North America. Canavan (1931) obtained this species from Corvus americanus in -73-
Pennsylvania. Daly (1959) found this parasite and another unidentified species of the genus Porrocaecum in nine of one hundred-three crows, Corvus brachvrhvnchos paulus. in
Virginia. Mawson (1956) obtained it from the eastern crow in Canada. The present report constitutes a new host record for Porrocaecum enslcaudatum in the United
States. In Poland, Markowskl (1933) found P. ensicaudatum in two of forty specimens of Corvus frugilegus and one of fifty-four jC. c o m i x . Porrocaecum wui Hsu, 1933, was described from Corvus macrorhvnchus intermedius in China.
Porrocaecum ensicaudatum has been obtained also from non-corvid birds. Mawson (1956) took specimens of this species from the robin, Turdus migratorjus. the grackle,
Quiscalus quiscalus. and the cowbird, Molothrus ater, in
Canada. Lopez-Neyra (1946) and Rysavy (1955) reported
Porrocaecum ensicaudatum from robins and starlings in
Spain and Czechoslovakia, respectively.
Life History. The life cycle of Porrocaecum ensicaudatum was studied by Levin (1957), who succeeded in infecting the earthworms, Lumbricus terrestris and
Octolasium lacteum by feeding them infective eggs of
P. ensicaudatum. Later, he experimentally infected the chicken, robin, starling, crow, grackle, domestic duck, -74- pigeon, and mouse by feeding them the infective larvae which were recovered from the earthworms.
Chandler and Melvin (1951), Oswald (1958) and
Pologentsev (1935) found encysted larvae of other species of the genus Porrocaecum in the shrew. -75-
Svngamus trachea (Montagu, 1811) Chapin, 1925 (Figs. 65-69).
Synonyms: Fasciola trachea Montagu, 1811.
Syngamus tracheal!s Siebold, 1836.
Syngamus primitlvus Molin. I860.
Strongylus primitlvus Hutyra and Marek, 1910.
Sclerostoma tracheale Diesing, 1851.
Syngamus gracilis Chapin, 1925.
Syngamus parvus Chapin, 1925.
Syngamus merulae Bay11s, 1926.
Syngamus tenuispiculum Manter and Pinto, 1928.
Site of Infection - Trachea.
Classification« according to Chitwood (1950).
Superfamily - Strongyloidea Weinland, 1858.
Family - Syngamidae Leiper, 1912.
Genus - Syngamus Siebold, 1836.
Synonym: Cvathostoma Blanchard, 1849* Generic Diagnosis (Cram, 1927): Sexes permanentlyjoined in copula. Buccal capsule in both sexes large, thick- walled, usually provided with 8 or 9 teeth, rarely 3, arranged about the center at the base, the teeth of 2 distinct sizes. Male bursal membrane thick; bursal rays short and thick; spicules small to very small, 150 microns to 25 microns. Vulva in anterior third of body; tip of female tail blunt or acute. Eggs of moderate size, operculated after deposition. 76
Specific description: With the characters of the genus Syngamus.
Female: Body red, cylindrical, 12.0 mm. in length;
0.854 mm. in width. Buccal cavity 0.594 mm. in diameter.
Walls of buccal cavity 0.105 mm. thick. Esophagus clavate.
Vulva 1.58 mm. from apical end. Anus subterminal. Tail tapering to subacute end. Eggs undergoing cleavage,
0.077-0.091 mm. (0.086) in length; 0.035-0.046 mm.
(0.042) in width.
Male: Body 3.6 mm. in length; 0.363 mm. in width.
Buccal cavity 0.264 mm. in diameter. Walls of buccal cavity 0.066 mm. thick. Esophagus clavate, 0.613 mm. in length; 0.178 mm. in width. Copulatory bursa 0.231 mm. in length; 0.528 mm. in width. Bursa bearing a group of four unbranches dorsal rays and a group of five lateral rays on either side. Dorsal rays 0.100-0.185 mm. in length. Lateral rays 0.179 mm. in length. Spicules of equal size, 0.085 mm. in length; left spicule curved; right spicule straight. Discussion. Syngamus trachea (Montagu. 1811) Chapin,
1925 has been found infecting at least eleven different orders of birds (Madsen, 1952). It was first taken from pheasants and partridges. Chapin (1925) described two new species: Syngamus parvus from Nucifraga carvacatactes in Europe and j>. gracilis from Corvus brachvrhynchos in -77
Pennsylvania. Baylis (1926) described £>. merulae from
Tardus merula. and Manter and Pinto (1928) described
JS. tenuispiculum from Turdus migratorius. These species were separated mainly on the basis of differences in the size and shape of the spicules, the size and number of branches in the bursal rays, and variations in body and organ sizes. Madsen (1950) demonstrated that these characters were subject to a great amount of variation, and also that variations in the size and shape of the spicules may occur in specimens taken from the same host or from different hosts. Moreover, the degree of branching of the bursal rays was found to vary in specimens taken from the same host. Cross-infection experiments by Clapham (1935) and Taylor (1928) demonstrated that gallinaceous birds could become infected with gapeworms from passerine birds. On the basis of these data, Madsen concluded that Syngamus gracilis.
S. parvus. S. merulae, S. tenuispiculum. and S. trachea were synonymous.
The writer found only one male and one female specimen of the genus Syngamus. The buccal cavity and body length of both worms were slightly larger than the size of these structures in S. gracilis. The bursal rays were similar to those,, of S. gracilis, but the spicules were 0.006 mm. longer than the longest spicule of S. gracilis. In -78-
S_. gracilis. the right spicule is curved and the left is
straight. In the male specimen obtained by the writer,
the right spicule is straight and the left is curved.
Unlike £>. gracilis both spicules were equal in length.
Syngamus trachea is cosmopolitan in distribution, and
is capable of infecting a wide variety of wild and
domestic birds. It has been taken from corvid birds in
Canada and the United States. Chapin (1925) and
Canavan (1931) reported this parasite, under the name of
Syngamus gracilis. from Corvus brachvrhvnchos in
Pennsylvania. Ward (1934) took j>. gracilis from crows in
Oklahoma. Cram (1936) found it in crows in Alaska.
Goble and Kutz (1945) found a 25 per cent infection in
juvenile crows in New York State. Morgan and Waller
(1941) did not find it in one hundred-twelve crows taken
in Iowa and Wisconsin. The present report is a new host
record for Syngamus trachea in Ohio.
Several foreign records have been reported for
Syngamus trachea. Elton and Buckland (1928) obtained
specimens of S>. trachea from crows in Great Britain. Madsen (1952) found this species in gallinaceous birds in
Denmark. Cram (1932), Campbell (1935), and Ripple (1941).
found this parasite in robins, starlings, and sparrows,
respectively.
The pathogenicity of J3. trachea has been investigated -79 by several workers. Goble and Kutz (1945) examined a pheasant and a robin which apparently had died of syngamiasis. Wehr (1937) reported nodule formation in tissue sites parasitized by gapeworms. Ripple (1941) studied the effects of gapeworm infections in game birds.
Lewis (1925), Elton and Buckland (1928), and Taylor
(1928) investigated the role of wild birds in the dissemination of gapeworms to domestic fowl. Lewis found a 37 per cent infection in the starling, and considered this bird to be an important distributor of gapeworms to domestic fowl in Great Britain. Elton and
Buckland found a high incidence of infection in the rook, Corvus frueileeus. in Great Britain. Cram (1927) stated that the turkey was probably the normal host of
_S. trachea. and considered domestic fowl to be more important than wild birds in the distribution of gapeworms.
Life History. Studies on the life history of Syngamus trachea have been reported by Waite (1920), Cram (1927), and Clapham (1935). As a result of these studies, it is now known that vertebrate hosts may become infected with
£>. trachea either by ingesting the eggs containing the -80-
inf ective larvae or by Ingesting an invertebrate host which contains infective larvae. Slugs, snails, and
earthworms may serve as intermediate hosts of this worm. These hosts become infected by ingesting infective eggs. 81-
Phvsocephalus sexalatus (Molin, 1860) Diesing, 1861 (Figs. 70-72).
Synonyms: Spirontera sexalata Molin. 1860.
Spiroptera strongylina Molin, 1860.
Filaria sexalata Perroncito, 1891.
Spiroptera strigis Seurat, 1915.
Habronema sexalata Ostertag, 1932.
Site of Infection - Encysted in mesentery and intestinal mucosa.
Classification. according to Chitwood (1950).
Order - Spirurida.
Superfamily - Spiruroidea Railliet and Henry, 1915.
Family - Spiruridae Oerley, 1885.
Genus - Phvsoceohalus Diesing, 1861.
Synonym: Leiuris Leuckart, 1850.
Generic Diagnosis (Yorke and Maplestone, 1926): Mouth with two lateral trilobed lips, each bearing three papillae externally but without teeth internally. . . . Male: Tail Inrolled. Caudal alae present. Four pairs of pedunculated preanal papillae; four pairs of very small papillae near tip of tail. Spicules very unequal. . . . Female: Vulva behind middle of body. Ovlporous, eggs subcylindrical with a thick shell, containing an embryo at deposition.
Description of third stage larva: Phvsocephalus sexalatus occurs as an encysted third stage larva in the crow. Adults are not found in this host. Cysts 0.750-
0.910 mm. (0.820) in diameter. Body 1.12-1*20 mm. (1.15) in length; 0.070-0.073 mm. (0.071) in width. Cuticle -82- transversely striated. Anterior esophagus surrounded by
several ganglia. Esophagus 0.517-0.546 mm. (0.532) in
length. Nerve ring 0.091-0.120 mm. (0.108) from apical end. Anus 0.056 mm. from posterior end. Caudal knob
0.007 man. in length; 0.005 mm. in width; bearing about
24 digitform, cuticular processes.
Discussion. Phvsocephalus sexalatus (Molin, 1860)
Diesing, 1861, was originally described from the pig.
In this host, it develops into a fully matured organism.
However, in certain hosts it fails to reach this state of development, and in such cases it occurs as a third stage larva, encysted in the mesenteries, intestinal mucosa, and skeletal muscles. Encysted larvae of P. sexalatus have been reported from bats by Alicata (1931); from the snake, Elanhe sealaris. by Chabaud (1950); and from the cotton rat, Siemodon hispidus. by Melvin and Chandler
(1950). Cram (1930) found larvae of Phvsocephalus sexalatus in the chicken; the owl, Otus asio; the hawk,
Buteo borealis: the pigeon; the bobwhite; and the turkey in Georgia and Florida. She considered birds to be important factors in the control of this worm because of its failure to develop to maturity in these hosts.
Phvsocephalus sexalatus has never been reported from
Corvus brachvrhvnchos until the.present study which constitutes a new host record for this parasite. -83-
Adult specimens of Phvsocephalus sexalatus have been found infecting pigs in Bengal (Maplestone, 1930); India,
(Baylis, 1939); and the United States (Cram, 1930).
Ezzat (1945) reported P. sexalatus from the rhinoceros,
Rhinoceros bicornis, in Egypt. Caballero and Alencaster
(1948) obtained specimens from the three toed sloth,
Bradypus griseus. in South America.
Life History. Alicata (1931) investigated the life cycle of several nematodes of swine. Two species of beetles, Ataenius coenatus and Passalus cornutus, were found to serve as intermediate hosts of Phvsocephalus sexalatus. Other beetles reported as intermediate hosts are Geotrunes douei. Gvmnopleurus sturni* Onthonhagus bedeli, Phanaeus carnifex. and Scarabaeus sacer.
Apparently, birds and swine may become infected by ingesting either infected adult or larval stages of beetles. -84”
Microtetrameres helix Cram, 1927 (Figs. 74-77).
Site of Infection - Mucosa of proventriculus.
Classification. according to Chitwood (1950).
Superfamily - Spiruroidea Railliet and Henry, 1915.
Family - Spiruridae Oerley, 1885.
Genus - Microtetrameres Travassos, 1915.
Generic Diagnosis (Cram, 1927): Long axis of female Body coiled in a simple or complicated spiral. Body usually without the four longitudinal furrows that are present in Tetrameres. If present, these furrows are also spirally coiled.
Specific description: With the characters of the genus Microtetrameres. Female: Body much larger than male; blood-red when alive. Body almost entirely filled with eggs. Cuticle finely striated and arranged about the body in a loose, sac-like manner. Body of coiled specimen 1.1-1.7 mm.
(1.5) in length. Buccal cavity 0.021-0.025 mm. (0.023) in length; 0.009-0.012 mm. (0.011) in width. Esophagus
0.891-1.584 mm. (1.188) in length. Nerve ring 0.175 mm. from apical end. Vulva 0.210-0.259 mm. (0.230) from posterior end. Tail subacute. Anus 0.140-0.189 mm.
(0.147) from caudal end. Eggs 0.040-0.051 mm. (0.046) in length; 0.031-0.039 mm. (0.034) in width.
Male: Body filiform, slightly coiled, 3.4-5.0 mm.
(4.47) in length; 0.112-0.114 mm. (0.113) in width. -85-
Buccal cavity 0.021-0.024 mm. (0.022) in length; 0.008-0.010 mm. (0.009) in width. Esophagus 0.594-0.756 mm. (0.665) in length. Nerve ring 0.175-0.195 mm. (0.190) from apical end. Cloaca 0.182-0.203 mm. (0.195) from posterior end.
Two unequal spicules; the longer 2.3-3.7 mm. (3.0) and ending posteriorly in two sharp points; the shorter
0.105-0.119 mm. (0.112) and having a blunt posterior end.
Tail subacute, ending in a small knob; bearing two pairs of preanal and two pairs of postanal papillae.
Discussion. Cram (1927) raised the subgenus
Microtetrameres Travassos, 1915, to generic rank and redescribed the genera Tetrameres Creplin, 1846, and
Microtetrameres Cram, 1927. In the genus Microtetrameres. the female body is coiled along its longitudinal axis and usually lacks the four longitudinal furrows that occur in
Tetrameres.
Microtetrameres helix was originally described by
Cram (1927) from Corvus americanus in Washington, D.C.
It has been found infecting the eastern crow in Canada (Mawson, 1956) and in Oklahoma (Ward, 1934). Morgan and
Waller (1941) obtained this parasite from forty-three of one hundred-twelve crows in Iowa and Wisconsin. The present report is a new host record for this parasite in Ohio. -86-
Several additional species of this genus have been described from corvid and other passerine birds.
0 Microtetrameres inermis (von Linstow, 1879) Cram, 1927, was described from Corvus corax. Corvus frugilegus. and
Corvus corone in Europe. Markowskl (1933) examined 139 corvid birds in Europe, but apparently failed to find this species. Mawson (1956b) described a new species,
Microtetrameres canadensis, from the owl, Nvctea scandiaca. in Canada. Schell (1953) described
Microtetrameres accipiter from Corvus corax. M. bubo from the owl and M. aquila from the golden eagle in the United
States.
Life History. The life cycle of Microtetrameres helix was investigated by Cram (1934a), who recovered the larvae of this parasite from experimentally infected grasshoppers,
Melanoplus femurrubrum and M. bivittatus. and also from the cockroach, Blatella eermanica. Later, the infective larvae were feed to the pigeon, Columba livta domestics from which the adult stage of Microtetrameres helix was recovered. Cram failed to infect the earthworm Lumbricus terrestris with the larval stages of M. helix. -87-
Acuaria anthuris (Rudolph!. 1819) Railliet. Henry, and Sissoff, 1912 (Figs. 78-82).
Synonyms: Spiroptera anthuris Rudolphi, 1819,
Dispharagus anthuris Duiardin. 1845.
Filaria anthuris Rudolphi, 1819.
Acuaria nebraskensis Williams, 1929.
Site of Infection - Beneath h o m y lining of gizzard.
Classification. according to Chitwood (1950).
Superfamily - Spiruroidea Railliet and Henry, 1915.
Family - Acuariidae Seurat, 1913.
Genus - Acuaria Bremser, 1811.
Synonyms: Spiroptera Rudolphi, 1819.
Anthuris Rudolphi. 1819.
Dispharagus Dujardin, 1845.
Generic Diagnosis (Yorke and Maplestone, 1926): Cordons directed straight backwards, not turning forwards and not anastomosing. Male: with two short slightly unequal spicules; six to eight pairs of postanal papillae.
Specific description: With the characters of the genus Acuaria. Head with two . pairs of submedian papillae and a pair of amphids. Mouth surrounded by two lips.
IVo pairs of cordons originate at either corner of the mouth and pass posteriorly over the ventral and dorsal
surfaces of the body.
Female: Body 22.0-25.0 mm. (24.0) in length;
0.390-0.435 mm. (0.422) in width, at the level of the ■88- vulva. Cuticle striated. Nerve ring 0.297-0.376 ram.
(0.350) from apical end. Vulva 11.0 mm. from apical end.
Esophagus 2.376-3.036 ran. (2.60) in length. Buccal
cavity 0.240-0.264 mm. (0.250) in length. Tail bearing one pair of papillae. Eggs embryonated, 0.040-0.053 mm.
(0.043) in length; 0.024-0.028 ran. (0.025) in width.
Male: Body 9.0-11.0 ran. (10.0) in length; 0.209-0.257 mm. (0.250) in maximum width. Buccal cavity 0.211-0.224 ran. (0.220) in length. Esophagus 2.640-3.857 mm. (3.012) in length. Nerve ring 0.285-0.317 mm. (0.310) from apical end. Left spicule 0.224-0.251 mm. (0.240) in length. Right spicule 0.178-0.224 mm. (0.211) in length.
Cloaca 0.429-0.561 mm. (0.512) from posterior end. Tail alate; bearing four pairs of preanal and eight pairs of postanal papillae, linearly arranged.
Discussion. The genera Acuaria and Cheilosoirura are very closely related and were recognized as synonyms by
Yorke and Maplestone (1926). However, Cram (1927) and
Chabaud and Fetter (1959) regarded these genera as being distinct even though they apparently differ only in the relative length of the spicules.
Acuaria anthuris has been obtained from corvid birds in many areas of the world. Acuaria nebraskensis. which
Williams (1929) described from Corvus brachvrhvnchos. was recognized by Cram (1934) as a synonym of Acuaria -89-
ant hur is . Ward (1934) took specimens of this species from
Corvus brachvrhvnchos in Oklahoma. Morgan and Waller
(1941) obtained it in nineteen of one hundred-twelve
crows in Iowa and Wisconsin. Fendinger (1952) reported
this parasite from six crows taken in New York. The present report constitutes a new host record for this
species in Ohio. Acuaria anthuris has been found also in
the eastern crow in Canada (Mawson, 1956). In Poland,
Markowskl (1933) took this parasite from one of three
specimens of Corvus corax: one specimen of C. corone. the only one examined; twenty-three of thirty-eight
C . frugilegus: and twenty-one of forty-seven C. c o m i x .
This nematode was taken, also, from Corvus corone in
Japan (Yamaguti, 1941a) and from Corvus snlendens and
Corvus macrorhvnchus in India (Singh, 1948).
Life History. Cram (1934) fed the eggs of Acuaria anthuris to the grasshopper, Melanonlus femur rub rum, and to crickets (unidentified) from all of which infective larvae of A. anthuris were recovered twenty-eight to thirty days later. She then fed the infective larvae to chickens and pigeons, but failed to recover the adult stage of this parasite from these hosts. 90
Diplotriaena tricuspis (Fedtschenko, 1874) Seurat, 1915. (Figs 83-87).
Synonym: Filaria trlcusnis Fedtschenko, 1874.
Site of Infection - Mesentery of body cavity.
Classification. according to Chabaud and Anderson (1959) and Chitwood (1950).
Superfamily - Filarioidea Weinland, 1858.
Family - Diplotriaenidae Chabaud and Anderson, 1959.
Genus - Diplotriaena Railliet and Henry, 1909.
Synonym: Triplotriaena Connal, 1912.
Generic Diagnosis (Yorke and Maplestone, 1^26): Body slightly attenuated at the two extremities; mouth simple without lips, surrounded by two lateral and four submedian very small head papillae; cuticle smooth; esophagus long, consisting of two parts, at its anterior extremity are two lateral chitinous structures, each consisting of a stalk directed forwards and a trident-like process directed backwards. Male: tail rounded, caudal alae pbsent, papillae usually absent; spicules unequal. Female: Vulva shortly behind the muscular part of the esophagus; Amphidelphys. Oviporous, eggs with thick shells and containing fully-developed embryos at deposition.
Specific description: With the characters of the genus Diplotriaena. Body surface smooth. Head with one pair of lateral and two pairs of submedian papillae.
Tridents two, laterally arranged on either side of the esophagus.
Female: Body 110-150 mm. (120.0) in length; 1.2-1.5 mm. -91-
(1.41) in width. Length of tridents 0.119 mm. Vulva
0.594-0.797 ram. (0.693) from apical end. Nerve ring
0.170 mm. from apical end. Tail ending bluntly. Anus
0.363 mm. from posterior end. Eggs globular, 0.055-0.062 mm. (0.059) in diameter.
Male: Body 50-53 mm. (52.0) in length; 0.750 mm. in width. Tail with four pairs of preanal and two pairs of postanal papillae. Spicules two, dissimilar in size and
shape. Right spicule 0.720-0.879 mm. (0.815) in length;
short and twisted. Left spicule 1.12-1.25 mm. (1.16) in
length; long and straight. Anus 0.100 mm. from caudal end. Tail ending bluntly.
Discussion. Diplotriaena tricuspis (Fedtschenko, 1874)
Seurat, 1915, was originally described from specimens taken from Corvus c o m i x in Europe. Additional descriptions of this species have been given by Seurat
(1915), Markpwski (1933), Morgan and Waller (1941), and
Singh (1949). Seurat (1915) found three pairs of cephalic papillae in Diplotriaena tricuspis. On the other hand, Markowski (1933) reported four cephalic papillae for this species. Morgan and Waller (1941) recorded four pairs of submedian cephalic papillae for this same parasite. The writer's observations on the number of cephalic papillae are in agreement with those of Seurat
(1915). Singh (1949) redescribed the male tail of -92-
Diplotriaena tricuspis on the basis of a single male and thirteen female specimens, which he took from the bay- backed shrike in India. The tail was described as having eleven preanal papillae and four pairs of postanal papillae. The writer's observations on the number of caudal papillae are in agreement with those of Seurat
(1915) and Morgan and Waller (1941).
Diplotriaena tricuspis has been reported from corvid birds in Europe and North America. Morgan and Waller
(1941) were the first to report this species from
American crows. Ward (1934) and Daly (1959) apparently failed to find this species in the eastern crow in
Oklahoma and Virginia, respectively. The present report constitutes a new host record for D. tricuspis in Ohio.
In Poland, Markowskl (1933) obtained specimens of
Diplotriaena tricuspis in two of forty Corvus fragile gus and in one of fifty-four Corvus c o m i x . Yamaguti (1935a) described Diplotriaena manipola from Garrulus glandarius in Japan. Caballero (1941) described D. roneglial from
Cassidix mexicanus in Mexico. Johnston and Mawson (1940) described Diplotriaena beta from Corvus coronoidea in
Australia.
Other species of the genus have been reported from the sparrow (Seibert, 1944), the starling (Nagaty and
Halfawi, 1942), the robin (Yamaguti, 1941a), and the -93- junco (ogren, 1950). All of these parasites were taken from either the body cavity or air sacs of the host.
Life History. Few life history studies have been reported for members of the subfamily Diplotriaeninae.
Anderson (1957) fed the eggs of a related species,
Dinlotriaenoides translucidus. to the grasshopper,
Camnula pellucida. and later recovered spirurid larvae from the haemocoel. This same investigator failed to infect ground beetles, dung beetles, field crickets,
Camel crickets, and ants.
After finding the eggs of Diplotriaena sp. and of
Diplotriaenoides translucidus in the lung and alimentary tract of grackles and other birds, Anderson suggested that the eggs probably are carried from the body cavity or air sacs to the lungs and alimentary canal, from which they later pass to the outside in the feces. He inferred that grasshoppers may serve as intermediate hosts of Diplotriaena spp. -94-
Splendidofilaria flexivagjnalis n. sp. (Figs. 88-95). Site of Infection - Encysted in mesentery near spleen.
Classification. according to Anderson and Chabaud (1959).
Superfamily - Filarioidea Weinland, 1858.
Family - Onchocercidae Chabaud and Anderson, 1959.
Subfamily - Splendidofilarinae Chabaud and Choquet, 1953.
Genus - Splendidofilaria Skrjabin, 1923. Synonyms: Chandlerella Yorke and Maplestone, 1926.
Vaerifilaria Augustine. 1937.
Parachandlere1la Caballero, 1948.
Generic Diagnosis (Skrjabin, 1923): Medium-sized worms. Cephalic end with out lips and epaulet-shaped structures. The head bears four submedian and perhaps two lateral head papillae. The esophagus is not divided into a muscular and glandular portion. Male: Posterior extremity digitiform. Caudal alae absent. Two approximately equal spicules. Female: Posterior extremity digitiform. Vulva in the anterior third of esophagus. Viviporous.
Specific description: With the characters of the
Splendidofilaria. Medium-sized worms. Cuticle smooth, without cuticular bosses. Head bearing four submedian papillae and a pair of laterally located amphids.
Female: Body 20.0-28.0 mm. (25.0) in length;
0.112-0.140 mm. (0.129) in width at the level of the vulva. Nerve ring 0.119-0.160 mm. (0.139) from the -95- apical end. Excretory pore 0.168 mm. from the apical end. Vulva triangular, 0.415-0.524 mm. (0.466) from anterior end. The vagina proceeds posteriorly from the vulva for a distance of 0.060-0.161 mm. (0.119), turns anteriorly for a distance of 0.259-0.356 mm. (0.314), forming an anterior loop, after which It bends caudally and joins the bifurcated uterus at approximately 3.0-5.9 mm. (5.1) from the apical end. Vagina filled with microfilariae. In young females the vagina may lead straight back to the uterus without forming an anterior loop or it may bend slightly forming a short anterior loop. Opisthodelphys; uteri parallel, not crossing.
Oviducts and ovaries coiled. Alimentary canal entire.
Esophagus undivided, 1.21-1.70 mm. (1.35) in length.
Anus frequently inconspicuous, 0.098-0.140 mm. (0.123) from posterior end. Tail digitiform, tapering to rounded end.
Male: Body 9.0-14.2 mm. (12.0) in length; 0.077-0.098 mm. (0.089) in width at the level of the upper end of the intestine. Nerve ring 0.126-0.140 mm. (0.136) from anterior end. Excretory pore inconspicuous, 0.175-0.189 mm. (0.182) from apical end. Esophagus 0.891-1.05 mm.
(0.982) in length. Cloaca 0.070-0.091 mm. (0.084) from caudal end. Tail digitiform, recurved, 0.042-0.053 mm.
(0.048) in width across cloaca; bearing six pairs of -96-
linearly arranged, post-cloacal papillae and a pair of
phasmids. The papillae are weakly sclerotized and some
times difficult to see. Occasionally, the two pairs of
papillae nearest the cloaca are smaller and either more mesially or laterally located than are the other caudal
papillae. Thus, they are not always visible at the
level of focus at which the other papillae are seen.
Spicules subequal. Right spicule 0.091-0.112 mm. (0.101)
in length. Left spicule 0.070-0.084 mm. (0.076) in length.
Microfilaria: Body cylindrical, unsheathed, 0.145-
0.147 mm. (0.146) in length; 0.003-0.004 mm. (0.003) in width. Head blunt, 0.004-0.006 mm. (0.005) in diameter.
Body tapering posteriorly. Tail rounded. Cuticle bearing fine transverse striations. Primodial excretory vesicle and cell at approximately 0.040 mm.
from posterior end. Three inconspicuous rectal cells at approximately 0.011, 0.009, and 0.005 mm. from caudal end.
Discussion. Skrjabin (1923) described the genus
Solendidofilaria from two specimens taken from the right side of the heart of 0 tome la phoenicuroidea in Turkestan.
Anderson and Chabaud (1959), in a study of the subfamily
Splendidofilariinae, considered the cuticular bosses as generically insignificant structures, and using the -97-
length of the tall as a taxonomic character, recognized
the genera Chandlerella Yorke and Maplestone, 1926,
Vagrifilaria Augustine, 1937, and Parachandlerella
Caballero, 1948, as synonyms of Splendidof1larla
Skrjabin, 1923. Yeh (1957), who described Chandlerella
braziliensis from a green-billed toucan, Rhamphastos
dlcolorus, in Brazil, considered the genus Chandlerella
to be valid and distinct.
Anderson and Chabaud (1959) listed ten species which
they recognized as valid members of the genus
Splendidofilaria. These were Splendidofilaria pawlowskvi
Skrjabin, 1923; J3. bosei Chandler, 1924; JS. gedoelsti
Travassos, 1926; S>. sinensis Li, 1933; IS. columbigallinae
Augustine, 1937; j>. periarterial!s Caballero, 1948; jS. verrucosa Oschmarin, 1950; JS. singhi Ali, 1956;
Si. braziliensis Yeh, 1957; and £5. gretillati Chabaud, Anderson, and Brygoo, 1959.
The species of the genus Sp lendidofilaria differ,for
the most part, in the character of certain cuticular
structures and in the number and arrangement of caudal papillae. Some species, e.g., j>. pawlowskvi. possess numerous cuticular bosses. In other species, e.g.,
£>. sinensis, the cuticle is smooth. However, the species of this genus generally possess a similar number of
cephalic papillae, viz., four, and show similarities in -98- the shape of the tail, spicules, and of the body. It is debatable whether the presence or absence of cuticular bosses constitutes a valid basis for dividing the genus.
It appears to the writer that these species are not sufficiently different to warrant generic distinction.
Splendidofilaria flexivaginalis n. sp., differs from the other species of Splendidofilaria in the presence of a pre-vulvular, vaginal loop; in the size of the spicules, body, and other organs; and in the number and arrangement of caudal papillae.
Life History. No complete life history studies have been reported for any species of the genus Splendidofilaria.
Anderson (1956) experimentally determined the life cycle of 0mithofilaria fallisensis. a related species in the family Onchocercidae, by allowing the blackflies, Simulium croxtoni. S. venusturn. _S. rugglesi. _S. eurvadminiculum. and S. latipes to become naturally infected with
Ornithofilaria fallisensis. Microfilariae were removed from Simulium venusturn, and hypodermically injected into the subcutaneous tissues of uninfected ducks and ducklings.
Thirty days later, microfilariae were found in the blood of these ducks, and adult specimens of Ornithofilaria fallisensis were taken from one of the ducks one hundred- five days after the initial date of infection. Robinson
(1955) tried to infect mosquitos of the genera Culex. -99-
Aedes. Culiseta, and Anopheles with microfilariae from the
crow, blue jay, and sparrow. The microfilariae failed to develop, except in a few Aedes aegypti that had been infected with microfilariae from a white-throated sparrow.
These apparently failed to develop into infective larvae in this mosquito. Splendidofilaria ohioensis n, sp. (Figs. 96-103).
Site of Infection - Between and within cerebral hemispheres. Specific description; With the characters of the
genus Splendidofilaria. Relatively long, slender worms.
Cuticle smooth, thin and unstriated. Head bearing four
small, submedian papillae and a pair of amphids. Tail digitiform.
Female: Body 48.0-57.0 mm. (55.0) in length;
0.084-0.126 mm. (0.110) in width across region of vulva.
Nerve ring 0.126-0.140 mm. (0.133) from apical end.
Esophagus 0.792-1.023 mm. (0.948) in length; terminating in an enlarged intestine. Vulvar opening circular,
0.429-0.759 mm. (0.637) from the anterior end. Vagina laminated, extending directly backwards from the vulva to a distance of approximately 2.5 mm. from the anterior end. Opisthodelphys. Uteri parallel, not crossing.
Oviducts and ovaries coiled. Anal opening inconspicuous,
0.237-0.264 mm. (0.250) from posterior end. Tail width across anus 0.079 mm. Male: Body length 20.0-24.0 mm. (23.0); width in the region of the upper end of the intestine, 0.112-0.132 mm.
(0.120). Nerve ring 0.133-0.158 mm. (0.143) from anterior end. Esophagus 0.720-0.910 mm. (0.891) in length. Tail recurved, bluntly rounded, 0.156-0.168 mm.
(0.161) in length; 0.042-0.053 mm. (0.048) in width -101-
across the region of the cloaca; bearing three pairs of
linearly arranged, post-cloacal papillae and a pair of
small, inconspicuous phasmids. Spicules subequal.
Right spicule 0.078-0.084 mm. (0.082) in length. Left
spicule 0.065-0.084 mm. (0.074) in length.
Microfilaria: Body cylindrical, sheathed, 0.154-0.161 mm. (0.157) in length; 0.002-0.004 mm. (0.003) in width.
Cephalic cell approximately 0.035 mm. from the anterior
end. Three inconspicuous rectal cells at approximately
0.040, 0.025, and 0.020 mm. from the posterior end.
Discussion. Splendidofilaria ohioensis n. sp., possesses the same number of caudal papillae as
Splendidofilaria sinensis. which Li (1933) described from
the trachea and lungs of Corvus sinensis and Urocissa
sinensis in China. In both parasites, the anterior end of the intestine is wider than the posterior end of the esophagus. Splendidofilaria ohioensis differs from
£5. sinensis in that the females of the former are twice as long as the latter, and the males of S. ohioensis are one and one-half times the length of those of S. sinensis.
The vulva of £>. ohioensis is at a greater distance from the apical end than that of .S. sinensis. The vagina is conspicuously laminated in £. ohioensis and the uteri do not cross as is the case in S. sinensis. Li described the esophagus of S.. sinensis as possessing both a -102- glandular and a muscular part. In SplendidofIlaria ohioensis, the esophagus is not divided into muscular and glandular portions. Moreover, the tail of male specimens of £5. ohioensis is shorter, and the right spicule is longer than are these features in S. sinensis.
Sp lendidofilaria ohioensis differs from j>. bosei, in that the body length of the former species is longer and the uterus is opisthodelphic rather than amphidelphic.
Life History. The life history of Splendidofilaria ohioensis is not known. Anderson (1956) succeeded in infecting blackflies with the larval stages of
Ornithofilaria fallisensis. a related species in the family Onchocercidae. It is possible that blackflies also may serve as intermediate hosts of Splendidofilaria ohioensis. -103-
The genus Splendidofilaria Skrjabin, 1923, comprises
twelve species of parasites, including Splendidofilaria
flexivaginalis and S.. ohioensis. described here for the
first time. In order to facilitate the identification t; of the species of this genus, a key is now presented.
A key to the species of the genus
Splendidofilaria Skrjabin, 1923.
1(7) Cuticular bosses present ...... 2
2(4) Caudal end of male bearing pre- and post-
cloacal papillae. Esophagus divided into
glandular and muscular portions ...... 3 3(2) Caudal end of male with two pairs of pre-
cloacal and six pairs of post-cloacal
papillae. Male 23 mm. in length; female
34 mm. in length; opisthodelphys. Spicules
0.063 mm. and 0.071 mm. in length...... Splendidofilaria gedoelsti Travassos, 1926.
4(2) Caudal end of male with only post-cloacal
papillae. Esophagus not divided into glandular
and muscular portions ...... 5 5(6) Posterior end of male bearing four pairs of
papillae. Male 7.4 mm. in length; female
18.9 mm. in length. Spicules 0.068 mm. and
0.077 mm. in length ......
...... S>. pawlowskvi Skrjabin, 1923. -104-
6(5) Caudal end of male with two pairs of papillae.
Male 14.0 mm. In length; female 29.0 mm. In
length. Spicules 0.045 mm. and 0.040 mm. In
length...... S. verrucosa Oschmarin, 1950.
7(1) Cuticular bosses absent ...... 8 8(16) Esophagus divided Into glandular and muscular
portions ...... 9
9(12) Caudal end of male with pre- and post-
cloacal papillae ...... 10 10(11) Posterior end of male bearing one pair of pre-cloacal and seven pairs of post-
cloacal papillae. Male 9.0 mm. in length;
female 21.0 mm. in length; amphidelphys.
Spicules 0.062 mm. and 0.077 mm. in length....
...... J3. periarterialis (Caballero, 1948)
Chabaud and Choquet, 1953. 11(10) Caudal end of male bearing four pairs of pre-cloacal and three pairs of post-cloacal
papillae. Male 27.0-33.0 mm. in length;
female 65.0-83.0 mm. in length; opisthodelphys.
Spicules 0.073-0.084 mm. and 0.091-0.094 mm.
in length......
...... S. braziliensis (Yeh, 1957) Anderson and Chabaud, 1959.
12(9) Posterior end of male with only post-cloacal
papillae ...... 13 -105-
13(14; 15) Caudal end of male with four pairs of
papillae. Male 13.0 mm. in length;
female 20.0-23.0 mm.; opisthodelphys.
Spicules 0.070-0.080 mm. in length......
S. columbieallinae (Augustine, 1937) Cnabaud and Choquet, 1953.
14(13; 15) Posterior end of male bearing three
pairs of papillae. Male 15.0-18.0 mm.
in length; female 23.0-25.0 mm. in
length; opisthodelphys. Spicules 0.050-
0.060 mm. and 0.070-0.080 mm. in length.
...... j3. sinensis (Li, 1933) Chabaud and Choquet, 1953.
15(13; 14) Caudal end of male with three pairs
of papillae. Male 18.0-20.0 mm. in
length; female 27.0-35.5 mm. in length;
opisthodelphys. Spicules 0.069-0.071
mm. and 0.060-0.062 mm. in length......
...... S . singhi (Ali, 1956) Anderson and Chabaud, 1959.
16(8) Esophagus not divided into glandular
and muscular portions. Caudal end of
male bearing only post-cloacal papillae ... 17 -106-
17(18; 19; 20) Posterior end of male bearing three
pairs of post-cloacal papillae.
Male 9.0-11.0 mm. in length; female
28.0 mm. in length; amphidelphys.
Spicules 0.070-0.080 mm. in length....
.. .£>, bosei (Chandler, 1924) Chabaud and Choquet, 1953.
18(17; 19; 20) Caudal end of male with six pairs of
post-cloacal papillae. Male 9.0-14.0
mm. in length; female 20.0-28.0 mm. in
length; opisthodelphys. Spicules 0.090-
0.112 mm. and 0.070-0,084 mm. in length.
...... £5. flexivaginalis n. sp.
19(17; 18; 20) Posterior end of male bearing three
pairs of post-cloacal papillae. Male
20.0-24.0 mm. in length; female
48.0-57.0 mm. in length; opisthodelphys.
Spicules 0.078-0.084 mm. and 0.065- 0.084 mm. in length......
...... ohioensis n. sp. -107-
20(17; 18; 19) Caudal end of male with two pairs
of post-cloacal papillae. Male 6.1
mm. in length. Spicules 0.062 mm.
and 0.078 mm. in length. Females unknown......
S. gretillati Chabaud, Anderson, and Brygoo, 1959. -108-
Arachnlda
Birds serve as host to a great many species of mites. The nasal cavities, trachea, and skin are the most common tissue sites from which mites have been
obtained. However, certain species occur inside the
quills and also beneath the skin. Of the five species of mites found in this study, four are feather mites and one is a quill mite.
Laminosioptes dihvmenalls n. sp. (Figs. 104-108).
Site of Infection - Skin and feathers.
Classification. according to Baker and Wharton (1952).
Order - Acarina.
Family - Laminosioptidae Vitzthum, 1931. Genus - Laminosioptes Megnin, 1880.
Synonym: Svmplectoptes Railllet, 1885.
The genus Laminosioptes was erected for Laminosioptes
cvstlcola. the only species heretofore included in the
genus. With the addition of Laminosioptes dihvmenalis n. sp., it is now possible to present a broader
description of this genus.
Laminosiootes: Small to medium-sized mites. Body
0.20-0.73 mm. in length, elongated and slightly
flattened. Dorsal surface smooth or finely striated.
No vertical setae on propodosoma, Body bearing a few
long setae. Gnathosoma normal, may or may not be -109- visible from above. Legs I and II short and stubby, with or without stalked caruncles. Genital suckers absent.
Hales without adanal suckers.
Specific description: With the characters of the genus Lamlnosloptes. Body elongated, white, except for forelegs and gnathosoma. Legs I and II 0.070 mm. in length, bearing caruncles and two broad, flat, curved, tarsal bristles. Pedlpalps bearing a pair of small, lateral, membranous alae. Two pairs of rostral setae, arranged In a transverse row; Inner pair shorter than outer. Propodosoma with a pair of dorso-lateral, membranous alae, originating immediately behind coxa II.
Each propodosomal ala with an apodeme at its base. A pair of lateral and a pair of dorso-lateral setae between the alae and coxa III. Y-shaped apodemes adjacent to coxae III and IV. Dorsally, a pair of relatively long setae at the level of the postero- or anteromedian margin of coxa IV. Abdomen bearing a pair of ventrolateral setae midway between coxa IV and the caudal end. Caudal end with one pair of long, stout, terminal bristles and a pair of long, stout, dorsolateral bristles. Female venter bearing a pair of small genital setae; two pairs of small, linearly arranged, submedian setae adjacent to coxae III and IV; a pair of small marginal setae between coxa III and IV; and a pair of small anal setae. Male -110- venter with four pairs of small submedian setae adjacent to the apodemes of coxae III and IV; a pair of small marginal setae posterior to coxa IV; and a pair of small anal setae. Coxa of all legs bearing a short, ventral seta. Hysterosoma with fine striations, especially adjacent to legs.
Female: Body 0.633-0.726 mm. (0.658) in length;
0.182-0.198 mn. (0.190) in width at level of coxae III and IV. Gnathosoma 0.070 mm. in length and 0.049-0.053 mm. (0.051) in width. Fedipalpal alae 0.021 mm. in length. Posterior margin of rostrum 0.161-0.168 mm.
(0.166) from apical end. Membranous propodosomal alae
0.105-0.140 mm. (0.121) in length. Genitalia medially located between coxae III and IV; 0.280 mm. from anterior end. Abdomen elongated, 0.264-0.310 mm. (0.275) in length;
0.154-0.198 mm. (0.176) in width; tapering posteriorly, caudal end truncate. Terminal bristles at caudal end of abdomen 0.330 mm. in length.
Male: Body 0.508 mm. in length; 0.175 mm. in width at level of coxae III and IV. Gnathosoma 0.067 mm. in length;
0.050 mm. in width at apical end. Pedipalpal alae 0.035 mm. in length. Posterior margin of rostrum 0.124 mm. from apical end. Membranous alae of propodosoma 0.119 mm. in length and 0.038 mm. in width at base. Abdomen slightly lobate, tapering posteriorly, caudal end truncate. Genitalia -Ill
0.084 mm. from posterior end; approximately 0.022 mm. in length and 0.023 mm. In width; Terminal bristles of abdomen 0.525 mm. in length. Dorsolateral bristles at caudal end of abdomen 0.320 mm. in length.
Discussion. The genus Laminosloptes Megnin, 1880, was established for the single species Laminosioptes cvsticola (Vizioli, 1870), Megnin, 1880, which occurs in the subcutaneous tissue of fowl in Europe and North
America. With the addition of Laminosioptes dihvmenalis n. sp,, the genus now contains two species. Laminosioptes dihvmenalis differs from L. cvsticola in that the former species is about 0.400 mm. longer than the latter, contains caruncles on tarsi I and II, and has membranous alae on the pedipalps and propodosoma. Such features are not present in L. cvsticola. Moreover, L. dihvmenalis has four caudal bristles while L. cvsticola has only two.
The two species show similarities in rostral features, the relative size of the legs, the paucity of setae, the localization of transverse striations, and, in the male, the absence of anal suckers.
Life History. The life history of Laminosioptes dihvmenalis is not known. According to Oudemans (1922), who investigated the life history of related feather mites in the suborder Sarcoptiformes, the female feather -112- mite produces eggs from which six-legged larvae are
formed. The larvae molt into eight-legged protonymphs, which later molt into deutonymphs. The latter give
rise to the adult stage. It is possible that
Laminosioptes dihvmenalis may have a similar life history. -113-
Svringpphilus bipectinatus Haller. 1880 (Figs. 109-113).
Site of Infection - Within feather shaft.
Classification. according to Baker and Wharton (1952).
Order - Acarina.
Family - Myobiidae Megnin, 1877.
Genus - Svringophilus Haller, 1880.
Generic Diagnosis (Baker et al., 1956): Svringophilus spp. may be distinguished by the elongate body with long setae, especially on the posterior margin; in having the typical cheyletid M- shaped peritremes; in having simple palpi without the thumb-claw complex; in lacking dorsal body plates; and in having short, stubby legs with tarsal claws and pulvilli.
Specific description: With the characters of the genus Svringophilus.
Female: Body 0.713-0.726 mm. (0.718) in length;
0.126-0.142 mm. (0.135) in width at the level of leg III.
Gnathosoma 0.168-0.175 mm. (0.173) in length; 0.060-0.066 mm. (0.062) in maximum width. Dorsally, the propodosoma bears three pairs of longitudinally arranged, submedian setae; a pair of lateral setae, midway between legs II and
III; and four long setae in a transverse row immediately anterior to leg III. Hysterosoma with one pair of long submarginal setae immediately posterior to leg III, and a short pair of median setae adjacent to leg IV. Abdomen elongated, bearing a pair of median setae midway between -114- leg IV and the caudal end. Posterior end with four long bristles. Abdomen 0.210-0.231 mm. (0.222) In length;
0.140-0.152 mm. (0.149) In width. Tarsi bearing pulvilli and a pair of brush-like setae immediately proximal to the tarsal claws.
Male: Males were not found in this study.
Discussion. Svringophilus bipectinatus Haller, 1880, was described from female specimens taken from the quill of fowl in Europe. It has also been reported from canaries and pigeons in Argentina and from chickens, turkeys, and pheasants in North America (Baker et al,
1956). The present report is a new host record for this species.
Another closely related species, Svringpphilus columbae. was described by Hirst (1922) from the quill of pigeons.
Life History. The complete life cycle of S. bipectinatus has not been determined. According to Baker (1956), all stages in the life cycle of this parasite apparently occur inside the feathers of the host. Ewing (1912), in an investigation of the life history of Cheyletus seminivorus. a related species in the suborder
Trombidiformes, found that the female of this species lay -115- eggs from which larvae are formed in 4.3 days. Each larva molts into a first nymphal stage which later molts forming a second nymphal stage. The latter gives rise to the adult stage. Svringophilus bipectinatus may have a life cycle which is similar to that of Cheyletus seminivorus. -116-
Analees corvinus Robin and Megnin, 1877 (Figs. 114-119).
Site of Infection - Skin and feathers.
Classification. according to Baker and Wharton (1952).
Order - Acarina.
Family - Analgesidae Trouessart, 1915.
Genus - Analges Nitzsch, 1818.
Generic Diagnosis (Gaud and Mouchet, 1959): Leg III of male "much larger than other legs. Transparent, triangular processes on the lower exterior surface of tarsi I and II. Well developed tubercles on the trochanters of anterior legs. Third pair of legs without caruncles in the male.
Specific description: With the characters of the
genus Analges.
Female: Body white except for forelegs and gnathosoma;
0.482-0.515 mm. (0.500) in length; 0.257-0.277 mm. (0.270)
in width. Gnathosoma 0.063 mm. in length; 0.052 mm. in width. Genitalia 0.150-0.165 mm. (0.160) from anterior
end. Two pairs of rostral setae, the inner shorter than
the outer. Dorsal hysterosoma bearing three pairs of
long, linearly arranged setae. Exoskeleton finely
striated. Abdomen entire, caudal end rounded and bearing
four terminal setae.
Male: Body more heavily chitinized than in the female.
Legs darker in color than rest of body. Two pairs of
rostral setae, the inner shorter than the outer. -117
Propodosoma with a pair of long, dorso-lateral setae immediately behind coxa II. Dorsal hysterosoma bearing two pairs of long, median setae, transversely arranged midway between coxae II and III. Body 0.396-0.409 mm.
(0.405) in length and 0.244-0.297 rain. (0.275) in width.
Gnathosoma 0.056 mm. in length and 0.042 mm. in width.
Abdomen entire, length 0.138-0.152 mm. (0.145); width
0.161-0.196 mm. (0.176). Abdomen bearing a terminal rectilinear, membranous expansion which is 0.014-0.017 mm. in width. Genitalia 0.154 mm. from posterior end,
0.031-0.035 mm. in length and 0.055 mm. in width. Anal suckers 0.012-0.014 mm. in diameter; 0.039-0.042 mm. from posterior end and 0.095-0.105 mm. (0.098) from genitalia. Leg III stout, ending in large claw. Coxae
III and IV with median setae. Tarsus III with rudimentary tubercle behind claw. Abdomen tapering to a rounded end which bears eight relatively long terminal setae.
Discussion. The genus Analges differs from the closely genus Megninia in that tarsus III of the latter bears a caruncle while tarsus III of the former ends in a large claw.
Few studies have been made of the feather mites of
North American birds. Peters (1936) reported feather mites from only sixty-two of 255 species of American -118- birds, including two species from the southern crow.
The genus Analges was not found on any of the birds included in his report. Ward (1934) reported Liponvssus bursa from crows in Oklahoma. Morgan and Waller (1941) found mites of the family Analgesidae on seven crows in
Iowa. The present report, therefore, is a new host record for Analges corvinus.
Gaud and Mouchet (1959) reported Analges corvinus from Corvus corone, Pvrhurus simplex. Poeoptera lugubris. and Lamprocolius splendidus in French Cameroun. Other species of Analges have been reported from the cardinal, starling, pigeon, and finch in Morocco (Gaud and Petitot,
1948).
Life History. The complete life cycle of Analges corvinus has not been determined. It may be similar to that outlined by Oudemans (1922) for mites of the families Analgidae, Sarcoptidae and Tyroglyphidae. -119-
Trouessartia corvina (Koch. 1840) Canestrini, 1899 (Figs. 120-123).
Synonym: Dermaleichus corvinus Koch, 1840.
Site of Infection - Skin -and feathers.
Classification. according to Baker and Wharton (1952).
Order - Acarina.
Family - Proctophyllodidae Megnin and Trouessart, 1883.
Genus - Trouessartia Canestrini, 1878.
Synonym: Pterocolus Haller, 1878.
Generic Diagnosis (Gaud and Petitot, 1948): Abdomen of the male attenuated behind, terminating in two narrow lobes, which are provided with transparent appendices.
Specific description: With the characters of the genus Trouessartia. Body elongated; uniformly brown in color. Dorsal and dorsolateral propodosomal shields prdsent. Posterior margin of propodosoma bearing four setae arranged in a transverse row. Vertical setae absent. Garuncles on all tarsi.
Female: Body 0.660-0.726 mm. (0.675) in length;
0.244-0.277 mm. (0.264) in width at the level of coxae
III and IV. Gnathosoma length 0.092 mm.; width 0.052 mm.
Genitalia 0*198-0.235 mm. (0.224) from apical end.
Abdomen elongated, 0.231-0.310 mm. (0.275) in length;
0.140-0.142 mm. (0.141) in width. Surface of abdomen bearing ovate markings which extend anteriorly to the -120
level of leg III. Abdomen tapering posteriorly and
ending as a bilobed structure. Two long, stout, hyaline
bristles on the lateral and terminal margins of each
lobe. One long and two short marginal setae approximate
ly 0.052 mm. In front of coxa III.
Male: Body elongated, 0.660-0.726 mm. (0.686) In
length; 0.250-0.290 mm. (0.264) In width. Gnathosoma
0.092-0.098 mm. (0.096) In length and 0.049-0.052 mm.
(0.051) In width. Posterior propodosomal margin
0.231-0.264 mm. (0.250) from apical end. Abdomen with
ovate markings, tapering posteriorly; terminal end
bilobate. Lobes leaf-shaped with serrated margins which
bear two pairs of long, stout, lateral and terminal
bristles. Anal suckers two, 0.017 mm. in diameter;
0.132-0.145 mm. (0.140) from the posterior end.
Genitalia 0.084 mm. in length; 0.171-0.198 mm. (0.190)
from posterior end. Discussion. Trouessartia corvina was reported from
the southern crow, Corvus brachvrhvnchos paulus. in
Florida (Peters, 1936). The present report constitutes
a new host record for this parasite. Gaud and Petitot
(1948) collected this species from Stumus vulgaris in
Morocco.
Other species of the genus have been taken from the -121- eastern kingbird, tree swallow, warbler, and towhee
(Peters, 1936).
Life History. The complete life history of
Trouessartia corvina has nbt been determined. Oudemans
(1922) reviewed the life cycle of Dermaleichus. a related genus in the family Proctophyllodidae. In this genus, the mites of both sexes pass through four developmental stages, viz., egg, larva, nymph and adult, but the females undergo an additional stage of development between the adult and nymphal stage. -122-
Gabucinia delibata (Robin, 1877) Oudemans, 1905. (Figs. 124-127).
Synonym: Fterolichus delibatus Robin, 1877.
Site of Infection - Skin and feathers. Classification. according to Baker and Wharton (1952).
Order - Acarina.
Family - Dermoglyphidae Megnin and Trouessart, 1915.
Genus - Gabucinia Oudemans, 1905.
Generic Diagnosis (Gaud and Mouchet, 1959): Two vertical setae present. Epimera I clear, at least in the female. Abdomen of male strongly bilobed. Genital suckers large and heavily chitinized. Legs subequal.
Specific description: With the characters of the genus Gabucinia. Uniformly medium-brown mites. Legs I and II stouter than legs III and IV. All legs with caruncles. Leg I bearing a stout spine on median margin of femur. Posterior margin of propodosoma marked by broad white transverse band.
Female: Body 0.562-0.613 mm. (0.587) in length;
0.264-0.277 mm. (0.268) in width. Gnathosoma length
0.084 mm.; width 0.073 mm. Abdomen entire, terminating in broad, rounded end which bears four long setae. Terminal end of abdomen with broad, white, U-shaped band.
Genitalia slightly anterior to coxa III; 0.073-0.084 mm.
(0.080) in length; 0.231 mm. from apical end. Male: Body 0.495-0.528 mm. (0.520) in length and
0.231-0.250 ram. (0.240) in width. Gnathosoma length
0.070 mm.; width 0.063 ram. Posterior margin of
propodosoraa marked by broad, white, band, 0.165 mm. from
apical end. Abdomen 0.175-0.204 mm. (0.198) in length;
0.204-0.217 mm. (0.211) in width. Abdomen bifurcated
posteriorly, forming two broad terminal lobes, each 0.099 mm. in length. Each lobe with a long terminal seta, a
stout median seta, a smaller lateral seta, and a long ventral seta. Anal suckers serrated, 0.028 mm. in diameter; immediately behind abdominal bifurcation.
Genitalia 0.035 mm. in length; 0.084 mm. from anal opening.
Discussion. This report is a new host record for
Gabucinia delibata. Gaud and Petitot (1948) obtained this
species from Corvus macrorhvnchus in Indochina. Gaud and
Mouchet (1959) found it infecting Corvus albus in
Cameroun. Gaud and Petitot also collected Gabucinia obtusus and G. rehberei from Diardigallus diardi and
Lobivanellus indicus. respectively, in Indochina.
Life History. The complete life history of Gabucinia delibata has not been determined. Oudemans' study (1922) indicates that feather mites pass through five develop mental stages, viz., egg, larva, protonymph, deutonymph and adult. -124-
Insecta
The Insect parasites found in this study belong to the order Mallophaga. These lice have chewing mouthparts and feed on the feathers or skin of the host. Most of the species parasitize only one or two species of birds.
Two families and three species of Mallophaga are represented in the present investigation.
Philonterus corvi (Linnaeus, 1758) Harrison, 1916 (Figs. 128; 131).
Synonyms: Lipeurus corvi Packard, 1870.
Docophorus corvi Osborn, 1896.
Site of Infection - Skin and feathers.
Classification. according to Hopkins and Clay (1952).
Order - Mallophaga.
Family - Philopteridae Burmelster, 1838.
Genus - Philopterus Nitzsch, 1818.
Synonyms: Docophorus Nitzsch, 1818.
Claviella Eichler, 1940.
Cypseloecus Conci, 1941.
Debauxoecus Conci, 1941.
Docophorulus Eichler, 1944.
Tritrabeculus Uchida, 1948.
Bitrabeculus Uchida, 1948. -125-
Generic Diagnosis (Kellogg, 1896): Body short and broad, head usually as wide across the temples as long, front broadly truncate or convex or slightly concave, rarely with a curving emargination; clypeus with distinct suture, often with a broad uncolored anterior and lateral margin; signature usually shield-shaped, with acuminate posterior angle; prominent movable trabeculae reaching to or beyond end of the first antennal segment; antennae similar in the sexes, with thick first segment, segment 2 the largest, and segments 3-5 subequal. Thorax with me so- and meta-segments completely coalesced; legs rather flattened, insertions approached; front legs smallest and usually concealed beneath the head. Abdomen usually oval, of nine segments of about equal length; last segment of male rounded, of female small and emarginated.
Specific description: With the characters of the genus Philopterus. Dark brown to black in color. Head truncated. Trabeculae prominent. Eyes large. Abdomen ovate. Dorsal abdominal segments with serrated marginal plates each with eight setae on posterior margin.
Abdominal spiracles surrounded by white linearly arranged disks, extending from the second to the seventh segments.
Female: Body 2.11-2.58 mm. (2.45) in length. Head
0.660-0.726 mm. (0.718) in length; 0.660-0.686 mm. (0.680) in width across eyes. Clypeal signature broad, brown in color. Lateral margin of each eye dark brown, bearing three posterior and two anterior setae. Trabeculae dark -126-
brown, 0.168 mm. In length. Antennae with dark brown and
white markings, 0.330 mm. in length. Thorax length 0.462
mm. Prothorax width 0.396 mm. Metathorax width 0.627
mm. Abdomen ovate, 0.990-1.22 mm. (1.15) in length;
0.910-0.924 mm. (0.915) in width; with serrated, dorsal
plates. Ninth segment without clear marginal disks.
Terminal end of abdomen bilobate. Legs with dark brown
stripes. Male: Body color similar to female. Body 1.80-1.98 mm. (1,87) in length. Head 0.627 mm. in length; 0.640-
0.693 mm. (0.680) in width across eyes. Trabeculae
0.140-0.168 mm. (0.158) in length. Antennae 0.287-0.299 mm. (0.294) in length. Lateral margin of each eye dark- brown, with five setae, three posterior and two anterior.
Thorax 0.462 mm. in length. Prothorax 0.363 mm. in width. Metathorax width 0.528 mm. Abdomen ovate,
0.858-0.924 mm. (0.900) in length; 0.792-0.858 mm. (0.825) in width, tapering to rounded, terminal end. Genitalia
0.363 mm. in length; beginning at the fifth segment and extending to the ninth.
Discussion. Philopterus corvi (Linnaeus, 1758)
Harrison, 1916, was originally described as Pediculus corvi from Corvus corax. Packard (1870) obtained specimens of this species, which he named Lineurus corvi. from Corvus americanus. Docophorus corvi which Osborn -127-
(1896) described from Corvus americanus was recognized
by Hopkins and Clay (1952) as a synonym of Pediculus
corvi Linnaeus, 1758. Philonterus corvi was collected from the eastern crow
in Ohio by Peters (1926) and Geist (1928). Peters (1936) reported this species from the eastern crow in Maryland and New York; the southern crow in South Carolina; and
the fish crow in South Carolina and Florida. Morgan and
Waller (1941) also found crows infected with it in
Wisconsin and Iowa.
Other species of the genus Philonterus were reported by
Peters (1936) from the blue jay, grackle, warbler, and
sparrow in the United States.
Life History. The complete life history of Philonterus corvi has not been determined. According to Geist (1936)
the'-eggs of bird lice are glued to the feathers of the host and develop to the nymphal stage after a variable period of incubation. The nymphs, which undergo several molts before becoming adults, are not as heavily chitinized as are the adults. -128
Mvrsidea albiceps (Piaget, 1880) Harrison, 1916 (Figs. 129; 132).
Synonyms: Menopon albiceps Piaget, 1880.
Menopon americana Kellogg, 1896.
Site of Infection - Skin and feathers.
Classification. according to Hopkins and Clay (1952).
Order - Mallophaga.
Family - Menoponidae Mjoberg, 1910.
Genus - Mvrsidea Waterston, 1915.
Synonyms: Acolpocephalum Ewing, 1927.
Allomvrsidea Conci, 1942.
Corvomenopon Conci, 1942.
Ramphasticola Carriker, 1949.
Generic Diagnosis (Waterston, 1915): Head and thorax broadand large in proportion to the abdomen. No spines on ventral surface of head. Flap across ocular emargination continuous with eye. Temples large, reclined towards the occiput. Forehead rounded anteriorly. Meso- and metathorax separated from one another. Metathorax separated from first abdominal segment by a membranous area. Sternal markings well defined. First abdominal stemite reduced; second different in size, shape or chaetotaxy (sometimes in all these respects) from the others. Pleurites well developed, but with no internal thickenings.
Specific description: With the characters of the genus Mvrsidea. Golden brown insects. Head triangular.
Clypeal signature clear. Margin of each eye bearing four long setae. Lateral margin of labium with a row of -129- short setae. Dark brown ocular band with a row of short setae along its lateral margin. Occipital band rectangular, hairy along lateral margin, and bearing a pair of longer setae on its posterior margin. Tibia with dentate seta on anterolateral margin.
Female: Body 2.11-2.37 mm. (2.25) in length. Head
0.363-0.369 mm. (0.365) in length; 0.668-0.699 mm. (0.686) in width across eyes. Antennae 0.132 mm. in length.
Thorax 0.594-0.660 mm. (0.615) in length. Metathorax
0.510-0.514 mm. (0.511) in width. Abdomen rectilinear in shape. First four dorsal plates "broken'1 on median surface. Abdomen 1.15-1.28 mm. (1.25) in length;
0.696-0.759 mm. (0.740) in width. Caudal end of abdomen bluntly rounded, bearing a terminal brush.
Male: Body 1.67-1.88 mm. (1.79) in length. Head
0.340-0.343 mm. (0.342) in length; 0.587-0.694 ram. (0.630) in width. Antennae 0.130 mm. in length. Thorax 0.528-
0.570 mm. (0.567) in length. Metathorax 0.510-0.514 mm.
(0.513) in width. Abdomen ovate, 0.850-0.990 mm. (0.957) in length; 0.567-0.627 mm. (0.583) in width. Genitalia bordering on segments four through nine; 0.567-0.600 mm.
(0.578) in length. Caudal end rounded.
Discussion. Mvrsidea albiceps (Piaget, 1880) Harrison,
1916, was originally described as Menopon albiceps from
Garrulus caledonicus. The species, Menopon americana -130-
Kellogg, 1896, from Corvus americanus- was recognized as a synonym of Mvrsidea albiceps by Hopkins and Clay (1952).
Peters (1936) reported Menopon americana from the eastern crow, southern crow, and fish crow in New York,
South Carolina, and Florida, respectively. Ward (1934) found Menopon mesoleucum infecting crows in Oklahoma.
Morgan and Waller (1941) obtained Mvrsidea albiceps and
M. interrupts from the eastern crow in both Iowa and
Wisconsin. Peters (1926) and Geist (1928) reported
Mvrsidea albiceps under the name Mvrsidea americana from
Corvus brachvrhvnchus in Ohio.
Other species of the genus Mvrsidea have been collected from the blue jay, robin, swallow, oven-bird, mockingbird, and catbird in the United States (Peters, 1936).
Life History. The complete life history of Mvrsidea albiceps is unknown. Barber's studies (1923) on lice of the family Philopteridae, indicate that the life cycle of bird lice involves an egg, nymphal, and adult stage.
Nymphal forms may molt several times before becoming adults. -131-
Briielia rotundata (Osborn, 1896) Hopkins and Clay, 1952 (Figs. 130; 133). Synonyms: Nirmus rotundatus Osborn, 1896.
Degeeriella rotundata Neumann, 1906.
Site of Infection - Skin and feathers.
Classification, according to Hopkins and Clay (1952).
Order - Mallophaga.
Family - Philopteridae Burmeister, 1838.
Genus - Briielia Keler, 1936.
Synonyms: Corvonirmus Eichler, 1944.
Meropsiella Conci, 1941.
Painiunirmus Ansari, 1947.
Traihoriella Ansari, 1947.
Guimaraesie11a Eichler, 1949.
Xobugirado Eichler, 1949.
Generic Diagnosis (Keler, 1936): Relatively large lice. Head conical and slightly truncated at apical end. Immovable trabecula in front of antennae. Tarsi with two claws. Antennae five-segmented. Terminal abdominal segment of female entire.
Specific description: With the characters of the genus Briielia. Dark brown to black insects. Head conical, tapering anteriorly, forming a slightly blunted apical end.
Female: Body 1.68-1.92 mm. (1.85) in length. Head
0.462-0.561 ,mm. (0.525) in length; 0.561-0.627 ram. (0.595) -132-
in width across eyes. Trabeculae 0.070-0.080 mm. (0.076)
in length. Antennae with light proximal segments and dark brown distal segments; 0.245-0.252 mm. (0.250) in
length. Thorax 0.396 mm. in length. Prothorax 0.277 mm. in width. Metathorax 0.627 mm. in width. Abdomen ovate, 1.02-1.32 mm. (1.22) in length; 0.712-0.858 mm.
(0.800) in width. Terminal end rounded. Dorsal plates of abdomen dark brown. Spiracles surrounded by white disks forming a linear row from segments one to seven.
Smaller, submarginal disks on segments one to eight.
Genitalia bordering on segments eight to nine.
Male; Body 1.39-1.48 mm. (1.43) in length. Head
0.368-0.462 ram. (0.420) in length; 0.495-0.594 mm. (0.540) in width across eyes; conical, slightly blunt at apical end. Trabeculae 0.079-0.084 mm. (0.081) in length.
Antennae with light basal and dark brown distal segments;
0.308-0.330 mm. (0.318) in length. Thorax 0.297-0.396 mm. (0.325) in length. Prothorax width 0.277 mm.
Metathorax width 0.508 ram. Abdomen broadly ovate,
0.712-0.825 mm. (0.805) in length; 0.627-0.660 mm. (0.635) in width. Genitalia 0.363-0.429 mm. (0.405) in length; beginning at posterior border of segment five and terminating on segment nine. Terminal end of abdomen rounded. -133
Discussion. Osborn (1896) originally described
Briielia rotundata as Nirmus rotundatus from Corvus americanus in Iowa. Ward (1934) collected specimens of this species, which he reported as Degeeriella rotundata, from the eastern crow in Oklahoma. Morgan and Waller
(1941) took Degeeriella rotundata and D. seconderia from the eastern crow in Iowa and Wisconsin. Peters (1926) and Geist (1928) collected specimens of this species from the eastern crow in Ohio.
Life History. The life history of Briielia rotundata has not been determined. Barber (1923), in a study of the life cycle of Briielia vulgata, found that the egg of this louse develops to a nymphal stage which, after undergoing several molts, becomes an adult. It is possible that Briielia rotundata may have a similar life history. OBSERVATIONS AND RESULTS
A total of 339 crows was collected from twenty-five counties in Ohio over a period of twenty months. Crows taken from the unglaciated counties are referred to under the heading "southeastern region"; those from the vicinity of Lake Erie are from the r(northeastem region"; and all others are listed tinder 'Wstern region"
(Fig. 134). Birds collected between March 15th and October 1st, are tabulated under the heading "summer season." During this period, insects, which are possible intermediate hosts of many parasites, are more active and available as food for the crow than during other seasons. Kalmbach
(1939) has shown that crows consume more insect food during the April to October period than during any other time of the year. Moreover, during the summer season, individual crows are dispersed over a wide area, and many do not roost together. At such times, a minimum number of ectoparasites would be exchanged between individual birds. Nestling, fledgling, and first year juvenile crows receive their initial protozoan and helminthic infections during the summer season.
Birds collected between October 2nd and March 14th are tabulated under '^winter season." During this season,
-134- -135- the large fall and winter flocks of crows have begun to form. At this time crows congregate in roosts, thereby making possible a greater exchange of ectoparasites.
On the other hand, insects are less active and consequently less available as food for the crow. Banding records, though somewhat inadequate, indicate that crows from more northerly located states may roost in Ohio during the "winter season" (Good, 1952). o TABLE X PUBLISHED RECORDS OF PARASITES TAKEN FROM THE EASTERN CROW, CORVUS BRACHYRHYNCHOS BRACHYRHYNCHOS.
PARASITE PUBLISHED RECORD LOCALITY
PROTOZOA
SPOROZOA
PLASMODIUM RELICIUH COATNEY (1936) Nebraska Morgan and Halccr (i94i) H isconsin Haemophoteus qanilemskii COATNEY AND HEST (l93B) N ebraska COATNEY AND JEUJSQN (1940) Montana Herman (1938) Cape Cod Morgan and Ha u e r (1941) H isconsin itUCQCYJQIflflM SAKHAROffi COATNEY AND He8T (l938) Nebraska Morgan and Ha u e r (1941) H isconsin ISOSPORA SP. BOUGHTON (1938) M innesota; Nem Jersey Morgan and Haller (1941) H isconsin
Mastigophora
lRXEAMftWHA COATNEY AND Ue ST <1938) Nebraska Morgan and Haller (1941) H isconsin
ACANTHOCEPHUA
Hediqrhynchus brandis Van Cleave (i9 i8) Maryland; Ohio; Illinois Kentucky
PUTYHELHINTHES
Cestoda
Hy h e h o l e p i s CORVI Mayhem (1925) Illinois Morgan and Haller (i94i) Iom a; H isconsin My h e n o l e p i s VARIABlLIS Mayhem (1925) ILLIN0I8 Morgan and Haller (i94i) H isconsin Am m b m b i m a n a n u c T A Ransom (1909) Nebraska UlLLlAMS (1929) Nebraska Taenia cvlindracea Hard (1934) Oklahoma lAIIRIPMH IM H Hard 0934) Oklahoma
Trenatoda
Cq m b p i c u u h m a c r o c h u Denton and Byrd (1951) Texas MACHYLECITHUH AMERICAMOM Denton (1945) Georgia Amphiheaua IPECIDMia Stiles and hassall (1896) Maryland COLLVRICLUM PABA Morgan and Haller (1941) Ioma
-136- / TABLE 1 (Contd.)
PARASITE PUBLISHED RECORD LOCALITY
NEHATHELMINTHES
Nema to d a
ACUARIA ANIHUJtlS HILLIAMS (1929) Nebraska Hard (1934) Oklahoma Cram (1934) Maryland Morgan and Ha u e r (1941) Io h a ; hisconsin Mamson (i956) Canada DlPLQTAIAENA TRLCU4PJ& Morgan an d Haller (i94i) Ioh a; H isconsin
PQRROCAECUM 1N8ICAUDATUM Mamson (1956) Canada
$.VM»URA £A£lUflfflL Hard (i934) Oklahoma
Ahioostonuh s p . Hard (1934) Oklahoma
Trichostrongylus pergracilis Hard (1934) Oklahoma
Capillaria contorta Morgan and Haller (i94i) Ioha
CAP* LLAMA COL LARI 3 Ham son (1956) Canada
i l N S A m JfeAGJULiS. Chapin 0925) Pennsylvania Canavan (i93i) Pennsylvania HARD (1934) Oklahoma Simttiia IMfiHEA. Goble an d Kutz (1945) Neh York
HlCf&TRTRAMERE?, ft&kiX, Cram 0927) Maryland Hard (1934) Oklahoha Morgan and Haller (i94i) Io h a ; H isconsin
ARTHROPODS
Mallophaga
PFtfEflHUA BOTWMA Osborn -137- TABLE 1 (Contd.) PARASITE PUBLISHED RECORD LOCALITY Acarina HAmpphraAUi.s upgm a P A L M M Ig Peters (1936) Maryland e Iaquessartia corvina Peters (1936) Maryland; No r v Carolina; New Yor k; South Carolina LlPOHVSSUS BURSA Hard (1934) Oklahoma *HAkfiMlPAE Horsan and Haller <4941) Ioha; Hisconsin Dirtera LYNCHER AMERICANA Mac Arthur (I94B) H isconsin Ornuhqica conuentA Johnson (i925) Neh Ensland States 138- TABLE 2 PARASITES OBTAINED FROM THE EASTERN CROW IN THIS STUDY (A total of thirty-one species of parasites were taken from 339 crows collected in Ohio over a twenty-month period.) PHYLUM AND C U S S SPECIES SITE OF INFECTION PROTOZOA SPORQZOA HAENOPROlEUa PANILEHSKII Erythrocyte Leucqcytqzoqh sa k h a r o f h Leucocyte or Crythrocyte ISQSPORA CQRV^At Feces ACANTHOCEPHAU Metacanthocephala Mepiorhvnchus grandis Intestine PIATYHELMINTHES Cestoda Qrthoskrjabinia ROSTELLATA Intestine AlWWTAHiiA MMaimCTA Intestine Hy he no l ep i s c o m Intestine HYNENOLEPIS 8ERPENTULUS Intestine HrMEHOkttU YAfilAfil.kU, Intestine TRENATODA Echinostqha revqlutuh Cloaca of Intestine AHPHIHERUS 8PECIQ8US Rile Duct of Liver BRAMTkKIIIWH ArttfU.CAJWM Bile Ouct of Liver C0H8P.IGUUH HASBBftSHU, Gall bladder NEMATHELMINTHES N ematoda CAP.KUMA AMAU.8 Cloaca and Intestinal Mucosa Ca p i l l a r iA co ntorta Mucosa of Esopha&us faMflfiAtcim mitAmm Intestine Sy n g a h u s tr a c h ea Trachea Ph t jw c e p h a m « m s A u i i a Encysted in mesentery and Intestinal Mucosa BisaaitiBinsatoJitLijL Mucosa of Proventriculus Ac u a r ia a n t h u r is Beneath horny lining of G izzard Diplqtriaena tricuspis Body Cavity SflLNttLBfiH.UAfUA fflUttENaia n . SP. Beneath meninge s, betheen and mithin Cerebral hemispheres of Brain Splcndipofilaria flexivaginalis n . SP. Encysted in splenic mesentery -139 TABLE 2 (Contd.) PHYLUM AND CUSS SPECIES SITE OF INFECTION ARTHROPODA Arachnida -UflLMfiMaEItl £LilXtilNAU.5 n . s p . Skin and Feathers Analges corvinus Skin and Feathers l M U 6 S 5 m . U . fiSAUM skin and Feathers isAfttfClBLiA M M f t m Skin and Feathers SYRINflttPHIWS BIPECTINATUS U ithin Feather Shaft Insecta f m i a e i m a c a m . Skin and Feathers Hyasiqea albiceps Skin and Feathers e. R U £ V U BfllUNMTA Skin and Feathers -140- TABLE 3 THE PERCENTAGE OF INFECTION OF THE EASTERN CROW WITH DIFFERENT CLASSES OF PARASITES IN RELATION TO AGE OF HOST AND SEASON OF YEAR. SUMMER SEASON (MARCH IS - OCT. l ) WINTER SEASON (O C T . 2 - March i4) YEAR-ROUND •45 Crows Examined 194 Crows Examined 323 Ck Ow A E x x m i m d : 3 3 9 Crows IMMATURE' '6 " PARASITES 4 NESTLIN6S 12 FLE06LIN6S 73 JUVENILES 56 ADULTS 70 Juveniles 124 Adults 143 Juveniles iso Adults Adult _ * * * j* * i * * » Blood Sporozqa 75.0 90.0 4.1 12.9 8.8 4.8 6.3 8.9 10.8 Intestinal --- 17.9 9.1 2 4 . 6 7.2 2 0 . 9 8.9 13.6 A porozoa Acanthociphala ~ 16.6 4.1 5.3 10.3 2.4 6.3 3.3 4.9 Cestoda --- 33.3 78.1 72.3 73.5 66.2 74.0 70.0 71.1 Trehatoda 50.0 --- 4.1 12.9 4.4 9.6 4.2 l l . l 7.7 Nematoda 25.0 83.3 95.9 90.9 94.1 99.3 95.0 94.8 96.7 Arachnida 100.0 100.0 100.0 95.0 100. 0 98.9 100.0 9 6 . 8 99.0 Insecta 100.0 100.0 58.9 7 2.9 86.1 71.7 72.8 72.3 72.6 TABLE 4 RANGE, INTENSITY, AND PERCENTAGE OF INFECTION OF CSROW WITH DIFFERENT SPECIES OF PARASITES IN RELATION TO THE AGE OF THE HOST AND THE SEASON OF THE YEAR. SUMMER SEASON (MARCH I S - D e i . i ) MINTER SEAS0H(0c T. lottARCH 15) YEAR- 145 Crows Exam NED 194 Crons Ei[AMI NED ROUND PARASITES 4 Nestlings 12 FLEDGLINGS 73 Juveniles 56 A dults 70 Juveniles 124 Adult*1 339 Crons Range A v . J t Ranse A». J t Range A v . J t Ranse A v . j t Ranse A v . Ransi A v . St s t OF NO. Inf. OF N o. Inf. OF N o . Inf. of N o . Inf. of H o . Inf. OF N o . In f . In f . Inf. Par. Inf. Par. In f . Par. In f . Par. In f . Par. In f . Par. - 142- PROTOZOA iFOROZOA ( I3.3jtY e a r l y In a c t i o n ) --- mmmm --- —. ------—- 10.6 HW W M liW MHIUBittH — --- 75.0 90.0 4.1 12.5 8.6 4.1 mm^m lEUCOCYTOZQO* SAKHAflftF.PL — --- 50.0 ------33.3 ------2.1 --- 1.8 —— mmmm ------2 . 1 ISOSFORA CQRVIAE — ------.— --- 1 6 .6 1 7 .6 mmmm 9 .1 mmmm 2 4 . 6 — mmmm 7.6 1 2 .2 ACANTHOCEPHAIA ( 4 . 9 0 Yearly Infection) MEDIORHYNCHUS GRANftlS — - .... 1-45 15.0 16.6 --- 1-3 2.0 5.3 1-4 2.1 10.01-7 3.5 2.4 4.9 TABLE 4 (Goutd.) SUMMER SEASON (March 15 - O c t . i ) WINTER SEASONtOcT. I-MARCH IS ) YEAR- 14S Crohs Exam NED 194 Croms Examined RCIIND PARASITES 4 Nestlinbs 12 PLEDBLINBS 73 Juveniles 56 Adults 70 JUVENIU8 124 Adults 339 Croms RANBE Av. * Ranbe Ay . * Ranbe Ay. * Ranbe Ay . * Ranbe Ay . * Ranbe Ay . t * OF N o. In f . OF N o. In f . of No . Inf . of No . In f . of N o . Inf . of N o. In f .Inf. Inf. Pa r . Inf. Pa r . Inf. Par . In f . Pa r . In f . Pa r . Inf. Pa r . PLATYHELHIHTHES Cestoda (7i.ij< Yearly Infection) QBltiOimABIiUA ROSTELLATA .... 2.0 8.3 ...... «... ._.... rmmri r . . . 0.3 Hyhenqlepis CORVI — —- .... 6.0 8.3 1-63 12.6 34.8 1-32 7.5 39.1 1-25 6.8 16.2 1-21 4.4 20.4 25.1 AMOMDTAENIA CONSTRICTA ----- .... 2.6 4.0 3 3 . 3 1-13 3.1 40.9 1-4 2.1 28.3 1-23 7.3 3 2 . 4 1-28 4 . 6 2 2 . 5 2 9 . 2 Hyhenqlepis serpentulus ----- ....------2 - 1 8 10.0 5.3 1-4 2 . 5 2 . 8 ...... 1.2 HYHENQLEPIS VARIABILlS .... 2-9 6.0 33.3 1-35 4.3 40.9 1-37 7.4 4 3 . 5 1-30 7.4 3 5 . 3 1-26 3.8 48.6 4 3 . 4 Trematoda ( 7 . 7 ^ Yearly Infectio n) SchiHgiigiy w m v T W i 4.6 s . o 50.0 ...... 1— 5 3 . 7 4.1 2.0 1.8 -r-r, 1.0 1.4 2-14 8.0 1.6 2.1 AMPtiJMERy# SPfCIOSUS —— ----- ...... —— .... ------.... ------— - — — .... 2 . 0 0.8 0.3 BRACHYLIC11HUM AMERICANUH —— ------.... - ---- —— 4-15 9.5 5.3 ------—.... 5-30 17.0 3.4 2.1 CONttUUWH HACRDRCHI8 ------. . . . |----- 1-5 3.0 6.5 ----- 1.0 2.8)1-2 1.3 4 .1 3.4 TABLE 4 (Contd.) SUMMER SEASON (MARCH 15 - Oct. i ) MINTER SEASON(O) it. i-March 15) YEAR- 145 Crons Exam NED 194 Crows Exi(MINED ROUND PARASITES 4 Nestlings 12 Fleoglinos 73 Juveniles 56 Adult 3 70 Juveniles 124 fDULT}) 339 Crows Range A v . ft Ranoe Ay . t Range A v . t Range A v . * Range A v . * Range A v . ft T of NO. Inf . of N o. In f , of N o . In f . o f N o . Inf . of N o. Inf . o f N o . Inf. In f . lllF. PAR. Inf. Par . Inf. Pa r .In f . Par. Inf. Par . Inf. Pa r . NEHATHElMtNTHES -144 Nematoda (96.7)5 Yearly Infection) E a m m u A . a m a h s ___ 7.0 2 5 . 0 2-42 il.I 60.0 1-113 15.4 69 . 7 1-35 6.8 34.8 1-147 17.5 4 8 . 5 1-235 13.4 31.7 4 3 . 5 CAP1LLAR1A contorta -— - 1-9 4.0 50.0 1-24 7.0 60.8 1-14 3.6 58.7 1-35 7.4 61.7 1-40 5 . 7 66.2 6 4 . 6 gflMQCAECUH 1NA1CAUDATUH ------1.0 8.3 1.0 4.1 --- .... 1-2 1.3 4.2 2.0 1.6 2.1 Syngamus v a c h e a •••• — -— —— - mmmrn ~ — --- 2 . 0 1.4 —- 0.3 tUZIfifilUIAKiUi —— —— 8.0 8.3 3 - 8 5.2 4.1 —- 2 5 . 0 1.8 1-25 8.0 7.0 1-75 2 4 . 0 3.4 4 . 0 HJCROTETRAMEBES HELIX —— 2.0 8.3 1-69 12.0 38.3 1-172 13.0 48.2 1-35 9.7 2 2 . 8 1-65 7.0 35.2 3 4 . 8 A coaria Ar t h u r u ———— 1-4 1.2 2 5 . 0 1-9 2.5 40.9 1-10 3.4 60.8 1-13 3.3 2 9 . 4 i-i e 3.4 42.2 42 . 5 £LEUtIftU£M& UL1£US£I£ —- 1.0 16.6 1-9 3 . 0 13.6 —— 2 . 0 1.8 1-4 2 .0 4.2 2-4 3 . 0 1.6 4 . 7 SPLENPIOOFILARIA FLEXIVA8INALI8 N. SP. —-—— 6-J8 9.0 41.7 5-25 12.7 62.9 1— 6 3 . 5 10.7 2-23 8.5 52.4 1—15 4.5 25.8 3 3 . 7 SPLENDIWFILARIA 9 H I9 1 N 8 IS N. ap. — - ———— ——— 13.0 8.3 1-13 10.2 2 8 . 6 —— 1.0 5.3 1-15 3.4 2 0 . 0 2-8 4 . 0 6.5 10.7 Microfilaria m — — -— ------m a— i m ------WOWS ------—• 83.3 80.4 ------80.4 — 91.2 93.6 88.2 TABLE 4 (Contd.) SUMMER SEASON (MARCH 15 - OCT. l) WINTER SEAS0N(0CT. I-MARCH 15) YEAR- 145 Crons Examined R0UN0 PARASITES 4 NeSTLIHBS 12 Pleoblinbs 73 Juveniles 56 1DOLTS 70 Juveniles 124 Adults 339 Crows Ranbe AV. * Ranbe AV. * Ranbe Av. $ Ranbe Av. t Ranbe AV. * Ranbe Av. * . * or No. Int. or No. Inf* of No. Inf. OF No. Inf. or NO. Inf. OF No. In f . In f . Inf.Par .Int. Par. Inf. PAR. lur. Par. I n f. PAR. I n f . Pa r . ARTHROPOD* Arachnida 145- (99.0jt Ye a r l y I n f e c t io n) UMINMICPTti DiHYmNAl.lt N. IF. ———— — — —— —— ———-—— —- —- m m m rn —- —- —— 0.8 0 . 6 ANALBES CORYINUS • m ...... m —■ m .... —- — — m m m m 36.6 .... 70.9 50.8 lKflV»SARTIft CMUUM, —- — —- 16.6—- 43.7 25.0 -— 97.5 — - ——— 94.5 81.8 fiAPVCIHfft PftlJATA —— 100. ——— —- 91.6 —- -— 75.3 ———- 75.0 —— —- 90.2 ——— —— 92.4 91.7 ftffllN»rmW MJltflMPIM m m m m —- ——*■ — .1.6 0 . 8 iNSECTA - ( 72. 6 0 Ye a r l y I n f e c t i o n) I h i u w h u i CORVI 3-13 8.0 100. 1— 4 2.5 2 5 . 0 1-13 4.5 31.7 1-51 6.2 35.1 2-25 8.7 36.6 2 - 7 6 9.1 57.1 42.3 My r s id e a ALB1c e p s 2-61 25.0 75.0 2-278 49.3 83.3 1-278 19.8 38.3 1-145 16.0 58.6 1-77 28.3 79.4 5-125 2 5 . 8 71.1 72.6 f i R i k U A ROTUND ATA 2-10 5.0 100. 1-47 15.0 33.3 1-47 7.8 22.7 1-24 4.1 32.6 I-4C 9.9 64.7 1-100 11.6 52.1 4 5 . 7 TABLE 5 RANGE, INTENSITY, AND PERCENTAGE OF INFECTION OF MALE CROWS WITH DIFFERENT SPECIES OF PARASITES IN RELATION TO THE AGE OF THE HOST. KALIS 175 Crows Examined PARASITES 2 Nestlings 6 Fledglings 73 Juveniles 94 Adults Range Av. Range AV. * Range A v . Range A v . 5% o r N o. Inf. o r N o . Inf. of N o . Inf. of N o . Inf. Inf. Par. Inf, Par. In f . Par. Inf. Par. PROTOZOA SPOROZOA ------50.0 100.0 —— ----- 16.6 - — --- 6.3 LEDCQCYTQZOON SAKHARQFFI — — — ■*- 50.0 ------50.0 -— 6.6 —— --- 1SQ6P0RA CORVIAC 3 3 .2 ------2 2 . 6 ------9 . 8 ACANTHOCEPHAIA Hedjqrhynchub grandis — - - ~ — — — 2 - 4 5 24.0 3 3 .2 1 -4 2 . 0 9 .1 1 - 5 2 . 3 3 .1 TABLE 5 (Contd.) HALES IT S Crow Examined 94 Adults PARASITES Range Av Range Av , of Inf, Inf or Inf, Inf Pa r , Inf, PLATYHELHINTHES I Cesioda »-» ORIHDSKRJABtNIA BOSIELLAIA I HVMENOLEMS CORVI 6.0 16.6 1-63 12.2 27.3 6.4 31.3 AnOHOTAENIA CONSTRICTA 3.5 33.2 4.6 40.9 l-ll 2.8 19.2 HYHENOLERIA SIRPEHTULUS 1.3 2.0 HYHENOLEHS VABIABILIS 7.3 27.3 4.8 47.4 Trehatoda Echinqstoha reyolutum 6.0 50.0 2 . 6 5.2 6.0 3.1 togjiiHava srKieiua BRACHVLECIIHUM AMEBICANUN 5-25 13.7 4.1 CONSPICUUH HACRORCHI£ 1.0 2 . 0 3.11-3 TABLE 5 (Contd.) HALES 175 Crows Examined PARASITES 2 NE8TLIN6S 6 Fledglings 73 Juveniles 94 Adults Range Av. jl Range Av. * Range Av . Range Av. or No. Inf. or No. In f , or No. Inf. of No. Inf. I Nr. Par. Inf. Par. Inf. Par. In f . Par. NEHATHELH1NTHES 148- Nematoda CMIUHHft AMAIU ------2-13 5.0 66.6 1-147 20.1 68.2 1-235 15.0 34.3 Ca m l l a r i a coniprta ------1-5 3.0 66.6 1-35 9.1 60.6 1-15 4.5 64.6 PORRQCAECUH EN8ICAUOATUM------■ ■■ ■ 1.0 16.6 1— 2 1.2 7.6 — - 2.0 1.0 TRAtHEA ______2 . 0 1.3 PWVSOCErHAUIS SEXALAIUS 8 . 0 1 6 .6 1 -2 5 8.0 9.1 1-75 29.2 4.1 HltfflHWAWRta HELIX „ — — ___ ------1 -6 9 7.2 30.3 1*65 7.7 39.3 Acuaria anthuris ______------1 -1 3 3.0 40.1 1-18 3.7 55.5 PlFUHRIAINA TRICUAPI1------1 .0 1 6 .6 1 -4 2 . 0 9.1 2.0 1.0 SFUEMPIDQFILARtA FLiXlVABlMAHS N. »P. ------II.0 6 6 . 6 3 - 2 3 9.4 58.6 1-11 5.1 23.7 SFLENPIPOFILARIA OHIOENSI1 N. 3P.------1 3 .0 1 6 .6 1 -1 3 5.6 17.2 1-8 3.1 10.2 M icrofilaria . ______82.1 — - 84.2 .... - 84.0 TABLE 5 (Contd.) MALES 1 7 5 Crqwa Ex a m in e d PARASITES 2 Nestlinba 6 Fl e d b l in b a 73 Juveniles 94 Adult a Ra n b e A v. ft Ra n b e A v . $ Ra n b e A v . Ranbe Av. A r achnida I I-* 4> VO UHIHOAIOPIEA PIHVHEHALI8 N. 8P. I Am a l c e a corvinus 3 5 .5 — . 7 1 .6 iRQUERAARI IA CORVIMUR 1 6 .6 8 3 .5 ------8 8 .3 CABUCINIA DILIBAT* 100.0 100.0 9 0 .3 9 8 .3 SVBtHBOPHILUS BIFICTINATUA INSECTA PHILOHERUA CORVI 10-13 12.0 100.0 1-4 2.5 33.2 1-20 7.1 42.8 1-76 9.3 44.9 Hvraidea a lbicepa —— 38.0 50.0 16-278 87.0 83.3 1-278 38.0 66.6 1-75 22.7 70.7 BrO eLIA ROTUNPAtA 3-10 6.5 100.0 4-47 19.0 50.0 1-47 8.2 51.5 1-60 I0.1 45.4 TABLE 6 RANGE, INTENSITY, AND PERCENTAGE OF INFECTION OF FEMALE CROWS WITH DIFFERENT SPECIES OF PARASITES IN RELATION TO THE AGE OF THE HOST. FEMALES 164 Croms Examined FA RASHES 2 N e stlin g s 6 Fledglings 70 Ruvebiles 86 Adults Range Av. Range Av. * Range Av. fk Range Av. PROTOZOA Sporozoa HAEMOPROTEUS PANILMSICH 100.0 8 3 .3 1 6 .6 8 .4 iJJjCQCYTQZOOW SAKtiAROFFl SO.O 16.6 2 .5 2 .4 Isospoea copviftp 2 0 .7 4 .0 ACANTHOCEPHAIA HEDIORHYMCHUS CRANPIS (.0 16.6 i*4 2 .0 6 .3 1-7 3 .6 3 .4 TABLE 6 (Contd.) FEMALES 164 Cr o w Examined PARASITES 70 Juveniles 86 Adults Range Av, Range Av, Range Av, Range Av, or I np, or I Nr, or I Nr, or Inp I Nr, Imp, (Mr. Par I Nr, PAR, PUTYHELAINTHES CE8T0DA flMHMKMABIHtA R9STELVATA 151 Ht h e n o l c p i s c o m I 1-25 9 .2 2 0 .9 l-ll 4 .5 2 0 .3 Am OHOTAEXIA CONSTRICTA 2-6 4 .0 3 3 .2 1-15 5 .6 3 2 .8 1-28 4 .4 3 0 .9 ittMEMOLEM* SERPENTULUS 4 .0 1 .3 1 8 .0 1 .2 RVMENOLEPIS VARIABILI8 2 -9 6.0 66.6 1—30 4.8 46.3 1-37 4 .5 4 7 .6 Trematoda Echinostoha revolutuh 4 .0 5 0 .0 4 .0 1 .3 Am phimerus 8PECI08U8 3 .0 1.2 toYSHYLioTmw anaicANvti 4-30 1 6 .3 3 .4 CflMSPICWH MACHQRCHIS 1 .0 1 .3 l-S 7.1 TABLE 6 (Contd.) F EMILES 164 Croms Examined PARASITES 2 Nestlinss 66 Adults Ranoe Av, Ranoe Av , nf, OF Inf,I I n f of Inf, Inf, Par Inf NEHATHEIMINTHES Nematoda 152- CAflLLARIA AMAHS 7.0 SO.O 3-42 19.0 50.0 1-1)3 53.7 7.7 30.9 CAHLLARIA CONTORT* 2-9 5.5 33.2 1-15 5.3 62.7 1-40 4.8 66.6 PflRROCAECUH EN8ICAUDATUH 2 .0 1.2 Sy m s a m u s TRACHEA PHYSCCEPHALUS SEXALATU8 3.0 1.3 3-25 14.0 2.4 H icroieiraheres helix 2 .0 16.6 1-69 15.0 25.3 1-172 9.6 40.4 A cuaria animuris 1-2 1.3 50.0 1-11 2.3 34.3 1-5 2 . 7 40.4 PlPLOTRIAEWA TRICUSPIS 1-9 3.6 9.1 2-4 3.0 2.4 SPLENDIDOFILARIA FLEXlVACINALIS » 2-25 10.0 51.9 3.5 26.0 Sf LENDIOOFILARIA OHIQENSIS H. 3P. 1-16 5.0 2.4 M icrofilaria 84.3 67.1 87.0 TABLE 6 (Contd.) TBKTB--- 164 Crows Examined 2 Nestlings 6 Fledglings 7 0 Juveniles 8 6 Adults PARASITES RAN6E A v . * Range AV. $ Range Av. * Range Av. * o r NO. Inf. of NO. Inf. of N o. Inf. of N o . Inf. Inf. Par . Inf. Pa r . Inf. PAR. In f . Pa r . 0 ARTHROPQM Arachnida 153- ttjaflENM IS n. sp. ------WM . — — ------ftHAhIK WRYtflVt ------3 8 . 6 ------— 6 0 . 4 ------lB8iiesaMm jc s m m ------5 0 .0 mm+m 8 6 .2 ------— 6 6 . 3 MfiUfiUU. fiSUSATA 2 - 8 4 . 0 1 0 0 . 0 ------— 1 0 0 . 0 ------8 6 .2 ------8 5 . 7 jttauwop.M,Mtt wucTjNAT.v.a ------— ------mm+m ------— 3 . 4 Insecta MumsitiLSflBJLL 3 - 6 4 ,.5 1 0 0 .0 ------1 -2 5 7 . 5 2 6 . 3 1 -5 1 9 .1 4 7 . 3 ttrmiwft A u i a p s 2-61 32.0 100.0 2 - 1 5 1 2 .0 3 3 .3 1 -4 5 1 9 .0 6 0 .0 1 -1 4 3 2 4 . 8 6 3 .1 toikviA KaiHMPAJUL 2 - S 3 . 5 1 0 0 .0 1 .0 1 6 .6 1 -4 0 I I . 1 3 8 . 8 1 -1 0 0 1 1 .8 4 7 . 6 TABLE 7 PERCENTAGE OF INFECTION OF EASTERN CROWS WITH DIFFERENT SPECIES OF PARASITES IN RELATION TO GEOGRAPHICAL REGION OF THE STATE. WESTERN REGION SOUTHEASTERN REGION NORTHEASTERN REGION PARASITES 238 Crows Examined 50 Crons Examined 51 Crows Examined Infection Infection Infection PROTOZOA I S p o r o z o a H* KJl 4> Ha e m p m o t e u s danilewskii 11.6 8.0 8.6 l ItVCOmOZOON 3AKHAR0FFI 2.9 2 . 0 Iw snm GflMfiAi 12.0 16.0 8.9 ACANTHOCEPHAU HIPICRHTNCHOS BRANDIS 4.9 8.0 TABLE 7 (Contd.) WESTERN REGION SOUTHEASTERN REGION NORTHEASTERN REGION PARASITES 238 Crow s Examined ______so Cr o m s Examined______Si Cr o m s Examined j> Infection______j» Infection Iiifectioh P LA TV HE LPII NT HES CESTODA QRTHOSKRJABIM A ROSTELLATA 0.4 Hy h e nq l ep i s corvi 20.9 22.0 33.6 ANQHaTAiNU CQNSTAIJTA 20.3 32.0 35.6 Ui Cn iftMENOLCFIS SERPENTULUS 1.7 l Hyhenqlepis variabilis 41.4 42.0 45.2 Trehatooa ECHiNOSTOMA BfYOLUTUM 2.0 4.0 Ahph ih e ru s speciosus 1.7 fiRMftYUSHHW AHERK»Wfl 2.1 2.0 1.9 CONSEICUUH HACRORCHIS 2.5 8.0 1.9 TABLE 7 (Contd.) WESTERN REGION SOUTHEASTERN REGION NORTHEASTERN REGION PARASITES 23S Crows Exam iie p s o Crows_Examined______S i Chows Examined # Infection j Infection______# Infection nehathelminthes Nehatqda Ca pil l a r ia a n a iis 4 0 .6 3 0 .0 6 2 .0 Ca pil l a r ia contoata 6 0 .2 70.0 6 4 .8 156- t S M M A M N H ^MaicAUDATUH 2 .1 4 .0 t m a m a tr a c h e a 0 .4 ------.... Physqcephalus sexalatus 4 .5 4 .0 3 .8 TU£M1£MAHER£S HE^tf, 3 4 .2 3 0 .0 3 8 .0 ACUARIA AWTHUHIS 4 4 .3 4 2 .0 4 0 .6 PJPLOIRIAENA TRICUSPIS 5.1 — - 7 .6 SftENPIMFILARIA FLEXIVAGINALIS N. SP. 3 4 .6 4 0 .0 2 4 .0 S plendjoo .fi laria 0HI0EN3I3 N. SP- 1 7 .0 14.0 Microfilaria 8 9 .7 8 6 .0 8 7 .5 TABLE 7 (Contd.) WESTERN RESION SOUTHEASTERN REGION NORTHEASTERN REGION PARASITES 2 3 8 Crons Examined SO Crows_Examined______S i Crows Examined ^ Infection______ ARTHROPODA Arachnida itAMIMQSIOPTES PI HYMEMALIS H. SP. 0.7 A n a l g e s CORVI HUS 52.1 46.0 53.1 I m u e s s a r t I a c o r v in u s 76.2 92.0 75.4 fiABUCINIA DEHBATA 91.2 92.0 89.0 SjtR J H6PPHILUS BIPECTHIATUS 2 . 5 ------ INSECTA tHILOPTERUS CORVI AI.3 44.0 42.3 IfrR&JPEA ALBICEPS 72.5 74.0 69.1 BRUELIA ROTUNDATA 43.5 48.0 44.0 INTERPRETATION OF DATA The species and number of parasites that may be acquired by a host are affected by a multitude of interrelated environmental factors such as temperature, humidity, light, food, and water which regulate not only the activities of the definitive host, but also the time occurrence and prevalence of the intermediate host. Since environmental factors vary with the season and locality, there may be an attendant fluctuation in the kind and number of parasites harbored by a host. The effect of seasonal variations on the prevalence of insects such as blackflies, which are known to harbor the developmental stages of filarial worms, is indicated by their widespread occurrence in certain localities during the late spring and summer months. Corresponding ly, nestling and fledgling crows, which frequently are heavily infected with filarial worms, also are more prevalent during this season. The absence of the intermediate host of a parasite during a particular i season or in a given locality may result in the absence of this parasite in the definitive host during the same season or in the same locality. The scope of the present study is such that experimental data are not available from which an evaluation can be made of the effects of specific -158- 159- environmental factors on the frequency and intensity of parasitism in the crow. Furthermore, the comparatively small size of the sample (339 crows) imposes a limitation on inferences which may be made from the data. However, despite these limitations, certain relationships may be inferred from the information obtained in this and other studies. Accordingly, the present interpretation is presented as a tentative explanation which eventually may become modified by more extensive studies. A comparison was made of the parasitic fauna of crows of different ages (Table 4). Eight species representing five classes of parasites were taken from nestling crows. Fledgling crows, which were not more than two and one-half months older than the nestlings, harbored twenty-three species representing six classes of parasites. Both nestling and fledgling groups had a relatively high frequency of infection with individual species of parasites. The difference in the number and kinds of parasites harbored by nestlings and fledglings may indicate differences in the developmental periods of the various parasites, but, also, it might be explainable as a difference in food habits of crows of these two age groups. It is interesting to note that there was a high percentage of nestlings infected with Echinostoma revolutum. This fact may be associated with the 160- relatively greater number of Insects consumed by nestling crows. The early transfer of ectoparasites from adult to nestling birds Is evidenced by the occurrence of at least four species of ectoparasites on nearly all nestling birds (Table 4). The early occurrence, in fledglings, of both adult filarial worms and microfilariae indicates that initial infections with this parasite probably occur while the young bird still is devoid of feathers. At such a time during the nestling age the bird would be more susceptible to the bite of a vector carrying the immature stages of a filarial worm. The relatively low incidence of filariasis in adult birds during the summer and winter season may Indicate the importance of feathers as a barrier to attack by blood-sucking vectors (Table 4)« Blood sporozoa. Seventy-five per cent of the nestling and 90 per cent of the fledgling crows examined during the summer season were infected with Haemoproteus danilewskii (Table 4). In most of these cases, multiple infections of individual red blood cells were observed. During the same season, juvenile crows had a lower frequency of infection than did adult crows. However, during the winter period, the percentage of infection with Haemoproteus danilewskii was higher in juveniles than in adults (Table 4). The fact that the summer incidence of haemoproteid infections was generally higher 161- than that of the winter season may be associated with the greater activity and prevalence during the summer season of hippoboscid flies, which are known vectors of haemoproteid parasites. The very high frequency of infection of nestling and fledgling crows may be related to the fact that such birds have fewer feathers than juvenile and adult birds, and therefore are more susceptible to attack by blood-sucking vectors. It is possible also that juvenile and adult crows may have acquired a degree of resistance to infection by Haemoproteus danilewskii. Gingrich (1932), in a study of plasmodial infections in pigeons, showed that some of the birds with latent and chronic infections were immune to superinfection by the same or related species of Plasmodium. It might be that, as a result of latent or chronic infections, older crows, such as juveniles and adults, are more resistant to haemoproteid infections than are younger birds. The higher incidence of infection in wintering juveniles as compared with wintering adults could be related to the migration of juveniles into Ohio from areas in which this parasite is more prevalent. Leucocytozoan infections were highest during the summer season. Nestling and fledgling crows had the highest percentage of infection, while the frequency of infection was essentially the same for both juvenile and -162- adult crows. Although leucocytozoa were never observed in the blood cells of crows taken during the winter season, it is possible that some of the birds had chronic leucocytozoan infections which were not detected. The fact that leucocytozoa were found in the blood cells of crows only during the summer season may be associated with the greater activity and prevalence during this period of blackflies, known vectors of leucocytozoan parasites. Intestinal sporozoa. The youngest crows infected with coccidia were fledglings (Table 4). Juvenile crows taken during the summer and winter seasons had a higher incidence of infection than did adults taken at the same time. The low frequency of coccidiosis in adults as compared with that of younger crows, such as fledglings and juveniles, might indicate that mature birds are more resistant to infection with coccidia than are immature birds. The highest percentage of coccidiosis occurred in juveniles taken during the winter season. It is possible that these birds may have migrated from areas in which this parasite was more prevalent. Acanthocephala. Fledglings were the youngest birds infected with Mediorhynchus grandis. One bird, a fledgling, harbored forty-five immature specimens. A 5.3 per cent infection was recorded for adult crows -163- taken during the summer season. Acanthocephala were not found in juvenile birds during this period. The frequency of infection in wintering juveniles was four times greater than that of wintering adults, but 38 per cent less than in fledglings collected during the summer season (Table 4). It is possible that this reduction in the percentage of cases from those of the summer period may be correlated with a decrease, during the winter season, in the prevalence of insects, such as grasshoppers, which are known to harbor the immature stages of Mediorhynchus grandis. While the intensity and frequency of infection were essentially the same for adult birds of both sexes, juvenile males had a higher percentage of infection than did juvenile females. Likewise, the percentage of infection in male fledglings was double that of female fledglings (Tables 5 and 6). The highest percentage of cases occurred in crows taken from the southeastern region. Acanthocephala were not found in birds taken from the northeastern region (Table 7). This regional difference in the distribution of Acanthocephala might be associated with a difference in the prevalence of the intermediate host of this parasite. The fact that Acanthocephala were not found in crows taken from the northeastern region also might be accounted for by a possible low yearly frequency of -164- acanthocephalan Infections for the period of time covered In this study, correlated with the comparatively small number of birds which were taken from this part of Ohio. Cestoda. Fledglings were the youngest crows from which cestodes were taken (Table 4). Four species were found in birds of this age group. One species, Orthoskr1ablnia rostellata, was never found in any other age group, and in fact was found in only one fledgling crow. Hymenolepid tapeworms were found in fledgling, juvenile and adult birds but were never found in nestlings. The frequency of infection with hymenolepid parasites and the mean worm burden were generally highest during the summer season (Table 4). The high percentage of infection during this period probably is associated with the greater activity and prevalence of insects, which are known to harbor the developmental stages of tapeworms. The frequency of hymenolepid infections was greatest among the mature crows, but immature crows generally harbored a higher mean number of worms than did adult crows (Table 4). It could be that crows are susceptible to infection with hymenolepid tapeworms during most of their lives and that initial infections are maintained over a period of several years, but at a lower level of intensity in mature than in immature crows. Adult males -165- were more frequently infected with Hymenolenis corvi than were adult females. However, juvenile females had a higher incidence of infection with this parasite than had juvenile males. Birds taken from the northeastern region were infected more frequently with tapeworms than were those collected in other areas of Ohio. This could be accounted for by a regional difference in the prevalence of the intermediate hosts of these parasites* The percentage of birds infected with Anomotaenia constricta was greater during the summer than during the winter season, but the mean number of A. constricta harbored by individual birds was highest during the winter season (Table 4). The high summer percentage of infection with this parasite possibly is correlated with seasonal variations in the occurrence of intermediate hosts infected with the bladderworm stage of Anomotaenia constricta. The fact that the number of worms per individual bird was highest during the winter season might be correlated with a high mean number of bladder- worms in the intermediate host during the winter period. Trematoda. At least one species of trematodes, Echinostoma revolutum. occurred in nestling crows. This was the only species of flukes taken from immature birds during the summer season. Although liver flukes were never found in immature birds, Conspicuum macrorchis. a -166- gall bladder fluke, was found in both juveniles and adults taken during the winter season. This fluke was never found in juveniles in the summer but was taken from adults during this time (Table 4). The relatively low overall percentage of infection of crows with trematodes probably would make untenable any inferences that may be made with reference to the relative effects of various factors on the frequency and intensity of infection by this group of parasites. On the other hand, the higher overall frequency of crows infected with trematodes during the summer as compared with the percentage of birds harboring these parasites in the winter definitely appear to be associated with a seasonal factor. As in the case of cestode infections, this high summer value probably is related to a more widespread occurrence of infected intermediate hosts, such as insects, during this season. Nematodes. The intensity and frequency of nematode infections probably should be considered in relation to those species of roundworms in which an intermediate host is required in the life cycle and those which probably do not require such a host. The studies of Clapham (1935), Cram (1936), and Waite (1920) indicate that Capillaria anatis, C. contorta and Syngamus trachea do not necessarily require an intermediate host in their life cycle. -167- Capillarid infections occur early in the life of the crow (Table 4). The frequency of infection was high in birds of each of the four age groups. The percentage of immature birds infected with Capillaria anatis in some instances was as much as 20 per cent higher in the summer than during the winter season (Table 4). Capillaria anatis was found more frequently in immature than in mature crows during both seasons. Moreover, the mean worm burden was highest among the immature birds (Table 4) . It would appear that immature birds might be more susceptible to infection with this nematode than are mature birds. Ackert and Herrick (1928) demonstrated that older chickens were more resistant to infection with Ascaridia lineata than were younger chickens. The phenomenon of age-resistance to helminthic infections has been shown also in dogs, cats, and a wide variety of other animals. The mean worm burden and the incidence of infection with Capillaria anatis were generally higher in immature and mature male birds than in females of the corresponding age group. Moreover, the frequency of infection was 28 per cent higher in juvenile males than in juvenile females (Tables 5 and 6). The percentage of infection with the closely related species, Capillaria contorta. was relatively high in birds of each of the four age groups during both seasons. -168- Adult birds were infected more frequently with this parasite than were immature birds. Among the immature birds the juveniles had the highest incidence of infection (Table 4). Because, in the crow, Canillaria contorta is embedded in the mucosa of the esophagus, it is possible that initial and subsequent infections with this parasite are maintained over a number of years. Therefore, adult birds might be expected to have the highest percentage of infection with this species. While both of the capillarid worms occurred in crows from all parts of the state, the highest incidence of infection with Canillaria anatis was in birds taken from the northeastern region. On the other hand, the percentage of infection with Canillaria contorta was greatest in crows collected in southeastern Ohio (Table 7). Svngamus trachea was found only once, occurring in a juvenile male bird taken from the western region. The very low frequency of "gapes11 in Ohio crows appears to indicate that this bird is not an important disseminator of gapeworms in Ohio. The percentage of Porrocaecum ensicaudatum infections was highest during the winter season. It was not found in birds taken from the northeastern region. This fact might be accounted for by the relatively low yearly incidence of infection of crows with this parasite i -169- correlated with the small number of birds taken in this region. Physocephalus sexalatus develops to maturity in pigs and similar vertebrates. In the crow, it occurs only as an encysted third stage larva. Since it is not likely that hogs become infected by eating crows, this worm undoubtedly infects other animals as intermediate hosts. While the percentage of birds infected by this parasite was somewhat lower than that of most other nematode species, the mean number of worms per bird was higher (Table 4). This relatively high mean worm burden tends to support the suggestion by Cram (1930) that birds may be important factors in the control of this nematode. Male birds showed a higher average number of worms per individual bird than did female birds (Tables 5 and 6). The highest percentage of infection with Physocephalus sexalatus occurred in male juveniles. This parasite was taken from birds in each of the three regions of the state (Table 7) . The frequency of infection with Microtetrameres helix and Acuaria anthuris, both of which have been experimentally transmitted by grasshoppers (Cram, 1934), was highest during the summer season. In general, the mean worm burden also was higher in the summer (Table 4). Since both parasites were taken from fledglings, crows -170- obviously are susceptible to infection with these parasites at a very early age. The percentage of infection was higher in adults than in juveniles, but the mean worm burden per infected bird was essentially the same in both (Table 4) . Acuaria anthuris was found more frequently in male than in female birds (Tables 5 and 6). Juvenile females harbored the highest mean number of Microtetrameres helix (Table 6). Since, in the crow, Microtetrameres helix and Acuaria anthuris are embedded in the proventriculus and gizzard, respectively, it is probable that these parasites are retained in individual birds over a number of years. Therefore, adult crows might be expected to have the highest incidence of infection with these species. Fledgling crows were the youngest birds from which Diplotriaena tricuspls was obtained (Table 4). The percentage of infection with this parasite was highest during the summer period. Anderson's studies (1957) indicate that the grasshopper may act as an intermediate host of this nematode. Kalmbach (1939) found that crows consume a large quantity of grasshoppers during the period from April to October. This fact may account for the higher percentage of crows infected with Diplotriaena tricuspls in the summer as compared with the winter period. Diplotriaena tricuspls was not found in crows collected in the southeastern region. The frequency and intensity of -171- infection with this parasite apparently were unrelated to the sex of the host (Tables 5 and 6). Juvenile birds were more frequently infected with this species than were adult birds (Table 4)* Apparently the crow becomes infected with the filarial worms, Splendidofilaria flexivaginalis and j>. ohioensis at an early age. Fledgling crows showed a very high incidence of filariasis. The frequency of microfilariasis in the crow indicates that most, and perhaps all, crows are infected with filarial worms during some stage of their lives. The very high percentage of filarial infections among immature birds indicates that most initial infections probably occur late in the spring when the nestlings, still devoid of feathers, would be more susceptible to the bite of some vector. Blackflies, which also occur in large numbers during this period, are known to harbor microfilariae. Whether such microfilariae are the developmental stages of any species of filariae found in the crow could be determined only by experimental procedures in which nestlings might be submitted to the bites of infected blackflies and other possible vectors. The mean filarial worm burden per infected bird was highest in juveniles during the summer season (Table 4). Likewise, a comparison of winter and summer percentages for Splendidofilaria flexivaginalis shows that, among juveniles, the summer value is 19 per cent higher than the -172- winter percentage. The high incidence of filariasis in adults and juveniles during the winter season may be correlated with the probable migration of crows into Ohio from areas in which these parasites are more prevalent. It may be explained also as a retention of worms that were acquired during the summer season. Snlendidofilaria flexivaginalis occurred in crows taken from all regions, but Splendidofilaria ohioensls was found only in crows collected in the western and south eastern parts of the state (Table 7). Arthropoda. Nearly all nestling crows were infected by at least four species of ectoparasites. Fledgling crows were infected by the same four and an additional species of mites (Table 4). Juveniles and nearly all adults were infected with one or more species of mites. Lice infected a somewhat smaller percentage of birds than did mites (Table 4). Nearly all species of ectoparasites were found throughout the winter season. Gabucinia delibata also occurred on a large percentage of birds during both seasons. The high percentage of infection with ectoparasites during the winter season is undoubtedly associated with the very close bodily contact of crows in winter roosts. Gabucinia delibata produced the highest percentage of mite infections during both seasons. All nestlings harbored this species. The mite burden of a few crows -173- was recorded during the summer and winter seasons. In general, G. delibata occurred In largest numbers during the latter part of the summer season. Trouessartla corvlna and Analges corvlnus were more prevalent during the early part of the winter season. Lamlnosloptes dlhvmenalls and SyrIngophllus blpectlnatus were found late In the course of this study and are reported from only a few birds. Gabucinia delibata. Trouessartla corvlna. and Analges corvlnus occurred on birds taken from each of the three regions of the state (Table 7). During the summer season, the percentage of mature crows that harbored Mallophaga was greater than that of Immature crows. On the other hand, In the winter, a greater percentage of immature birds harbored lice. The incidence of lice infection among male juveniles was greater than that among female juveniles (Table 5 and 6). The mean number of Mallophaga per bird was highest during the winter season in both juvenile and adult birds. This fact appears to indicate that the winter period is the time of peak reproduction of the lice found in this study. i GENERAL DISCUSSION It has been shown in this study that the eastern crow serves as host to a wide variety of parasitic species. Several factors may account for its diverse parasitic fauna, viz., (I) its resistance, or lack of resistance to attack and infection; (2) its omnivorous feeding habit; and (3) its migratory and roosting behavior. Since the eastern crow is a migratory bird (Kalmbach, 1939; Good, 1952), it is not possible to determine the source of its parasites. However, from the data collected in this study, as well as that of other workers (Ward, 1934; Morgan and Waller, 1941), it is evident that regional ecological conditions affect the kind and number of parasites which the crow may acquire. For example, Ward obtained five species of parasites from the eastern crow in Oklahoma, none of which were found in the present survey (Table 1). Morgan and Waller took four species from crows in Iowa and Wisconsin, none of which were obtained in the present investigation. On the other hand, fifteen species, three of which are new, are reported here for the first time. The effect of regional ecological differences on the frequency of infection of crows with certain parasites, is evidenced by the data presented in this and other studies. For instance, Morgan and Waller obtained -174- 175- Hvmenolepis corvi from 39.3 per cent of the crows which they collected in Iowa and Wisconsin, while in the present study only 25.1 per cent of the crows were infected. The data collected in this study appear to indicate that variations in the frequency and intensity of parasitism in the crow may be associated with varying seasonal factors which affect the activity of the crow and also of intermediate hosts of the parasites involved. In nearly all cases the highest incidence of parasitism and the highest mean worm burden were associated with the season when the intermediate hosts of certain parasites were most active and prevalent. This correlation was demonstrated in the case of the blood sporozoa, acanthocephalan parasites, cestodes, trematodes, and certain nematodes. The age of the host may affect the frequency and intensity of its infection with certain parasites. Immature birds, both nestlings and fledglings appeared to have been more susceptible to attack and infection with blood sporozoa than were adult birds. Coatney (1933) also found wide differences in the degree of susceptibility of pigeons to haemoproteid infections. Likewise, Gingrich (1932) found that birds with chronic or latent plasmodial infection were immune to super infection. Moreover, an infection with one species of -176- Plasmodium sometimes resulted in an immunity to infection with other species. Whether most adult crows are chronically infected with Haemonroteus danilewskii and Leucocvtozoon sakharoffi cannot be inferred from the data collected in this study. However, the difference in the percentage of infection in immature and mature crows would appear to indicate the operation of a resistance mechanism associated with the age of the host. Immature birds generally showed a higher percentage of infection with helminth parasites than did adult birds. This was especially true in the case of filarial infections. On the other hand, certain trematode, cestode, and nematode species occurred more frequently in adult birds. The phenomenon of age resistance to helminthic infections has been studied by a number of workers. Ackert and herrick (1928) demonstrated that older chickens were more resistant to infection with Ascaridia lineata than were younger chickens. Ackert (1942) showed that this resistance may be associated with an increase in the number of goblet cells in the intestinal mucosa. Africa (1931) found evidence of age resistance of rats to Nippostrongylus muris. Age resistance has been also demonstrated in dogs (Herrick, 1928; Sarles, 1929), humans (Keller, Leathers, and Ricks, 1934), and a wide variety of other animals. The data presented in the -177- present study appear to Indicate a definite association of the age factor with the incidence of parasitism. The frequency and intensity of helminthic infections may be affected by the sex of the host (Chandler, 1932). Hie evidence presented in the present study indicates that, in the crow, the degree of sex associated resistance varied with the species of parasites causing the infection. Moreover, this resistance also varied with the age of the host. For example, in the case of Hymenolepis variabilis. juvenile females were infected more often than were juvenile males, but adult males and females had the same percentage of infection for this parasite. The high percentage of birds infected with ecto parasites during the winter season appears to be associated with the close bodily contact of crows in winter roosts. The comparatively high mean burden of ectoparasites during the early part of the winter season evidently corresponds to the period of greatest reproduction of these forms. The omnivorous feeding habit of crows may partly account for the wide variety of endoparasites harbored by them. Grasshoppers constitute a large part of the summer diet of the crow (Kalmbach, 1939). Several parasites that this host harbors have been experimentally -178- transmitted by grasshoppers. These parasites are Mediorhynchus grandis, Hymenolepis variabilis, Microtetrameres helix, Acuaria anthuris, and, possibly, Diplotriaena tricuspis (Moore, 1941; Jones, 1934; Cram, 1934; Anderson, 1957). Beetles are readily eaten by the crow (Good, 1952). Brachylecithum americanum and Physocephalus sexalatus have developmental stages which occur in beetles (Denton, 1945; Alicata, 1935). Again, crows feed on earthworms (Kalmbach, 1939). These invertebrates are known to harbor the developmental stages of Syngamus trachea and Porrocaecum ensicaudatum (Waite, 1920; Levin, 1957) and possibly harbor the immature stages of Capillaria contorta and Anomotaenia constricta (Cram, 1936; Mrazek, 1907). Crows are known to feed also on aquatic animals (Kalmbach, 1939). The developmental stages of Echinostoma revolutum and various species of Amphimerus occur in aquatic animals (Beaver, 1937; Wallace, 1940). Crows also feed in pig sties and in close association with fowl (Good, 1952). At least four parasites which occur in ducks, turkeys, and pigs have been found also in crows. These parasites are Capillaria anatis, in ducks (Madsen, 1945); Capillaria contorta and Syngamus trachea, in chickens and turkeys (Cram, 1936; Clapham, 1935; Madsen, 1952); and Physocephalus sexalatus,in pigs (Alicata, 1935). The high percentage of crows infected -179- with Capillaria contorta and C . anatis strongly indicates that this bird may be an important reservoir host and disseminator of these parasites to domestic fowl. Syngamus trachea was found in only one crow. Therefore, the eastern crow probably is not an important disseminator of this parasite to domestic fowl in Ohio. SUMMARY Thirty-one species of parasites were taken from a total of 339 specimens of the eastern crow, Corvus brachvrhvnchos brachvrhvnchos. collected over a period of twenty months. These birds were taken from twenty-five counties in Ohio, representing three ecological areas which are referred to as "northeastern region," "southeastern region" and "western region," (Fig. 134). The crow harbored three species of protozoan parasites, one species of Acanthocephala, five species of cestodes, four species of trematodes, ten species of nematodes, three species of lice, and five species of mites. Fifteen species of parasites are reported for the first time from the eastern crow. These are Isospora corviae, Ortho skr1abinia rostellata. Hvmenolepis serpentulus. Amphimerus speciosus. Echinostoma revolutum. Capillaria anatis. Physocephalus sexalatus. Porrocaecum ensicaudatum. Splendido filaria flexivaginali s n . sp., Splendido filaria ohioensis n. sp., Analges corvinus, Gabucinia delibata. Laminosioptes dihymenalis n . sp., Svringophilus bipectinatus. and Trouessartia corvina. Nine species are reported for the first time from the eastern crow in Ohio. These are Haemoproteus danilewskii. Leucocvtozoon sakharoffi. Anomotaenia constricta, Hvmenolepis variabilis. Conspicuum macrorchis, Acuaria anthuris. Microtetrameres helix. Diplotriaena tricuspis. and Syngamus trachea. -180- -181- Sporozoa infected 13.6 per cent of the birds. The highest percentage of blood sporozoan infections occurred in juvenile birds during the summer season. Acanthocephala infected 4.9 per cent of the birds. Wintering juveniles had the highest percentage of acanthocephalan infections. Cestodes infected 71.1 per cent of the birds. In general, the highest incidence of cestode infections occurred during the summer season. Trematodes infected 7.7 per cent of the crows. These parasites were found more often during the summer than in the winter. Nematodes infected 96.7 per cent of the crows. The period of the highest incidence of nematode infections varied with the species of parasites involved. Lice and mites were found in 72.6 and 99.0 per cent of the birds, respectively. The peak period of infection with ectoparasites usually occurred during the winter season. Several factors may account for the wide variety of parasites which the crow harbors, viz., (1) its resistance or lack of resistance to attack and infection, (2) its omnivorous feeding habit, and (3) its migratory and roosting behavior. The frequency and intensity of infection appear to be associated, to a certain extent, with such factors as season, and sex and age of the host. Within the State of -182- Ohio, regional ecological factors do not appear to appreciably affect the species of parasites which are harbored by the crow. A comparison of winter and summer percentages indicates that variations in the frequency and intensity of infection probably are associated with varying seasonal factors which affect the activity and prevalence of the crow and also of the intermediate hosts of the parasites involved. The effect of the age of the host is evidenced by differences in the intensity and frequency of infection in immature and mature birds. The incidence and intensity of certain parasitic infections appear to be modified also by the sex of the host. The omnivorous diet of the crow probably accounts for its wide variety of helminthic parasites. Several invertebrate animals which form a normal part of the crow's diet are known to be intermediate hosts for some of its parasites. The high percentage of crows infected with Capillaria contorta and C. anatis strongly indicates that the crow may be an important reservoir host and disseminator of these parasites to domesticated animals. For the benefit of others who may be interested in -183- the eastern crow, a complete list has been compiled of the parasites which have been reported from this host by other investigators (Table 1). A key to the species of the genus Splendidofilaria Skrjabin, 1923, has been presented as an aid in the identification of the species of this genus. LITERATURE CITED Ackert, J. E. 1942. Natural resistance to helminthic infections. J. Parasitol. 28: 1-24. Ackert, J. E. and Herrick. 1928. Effects of the nematode AscarjLdia lineata (Schneider) on growing chickens. J. IParasitol. 15: 1-13. Adie, J. R. 1924. The Sporogony of Haemoproteus columbae. Bull. Soc. Path. Exot. 17: 605. Africa, C,. M. 1931. Studies on the host relations of Nippostrongylus muris, with special reference to age resistance^and acquired immunity. J. Parasitol. 18: 1-13. Al-Dabagh, M. A. 1956. Blood protozoa in wild and domestic birds in Ohio. Unpublished Master's thesis. The Ohio State Univ. Ali, S. M. 1956. Studies on the nematode parasites of fishes and birds found in Hyderabad State. Indian J. Helminth. 8: 1-83. Alicata, J. E. 1931. Infective larvae of Phvsocephalus sexalatus in bats. J. Parasitol. 18: 4/. Alicata, J. E. 1935. Early developmental stages of nematodes occurring in swine. U. S. Dept. Agric. Tech. Bull. No. 489. 96 pp. Anderson, R. C. 1954. The development of Omithofilaria fallisensis Anderson, 1954 In Simulium venustum Say. J. Parasitol. 40: 12 (Abstract 2). Anderson, R. C. 1956. Two new filaroid nematodes from Ontario birds. Canad. J. Zool. 34(4): 213-218. Anderson, R. C. 1957. Observations on the life cycle of Diplotriaenoides translucidus Anderson and members of the genus Diplotriaena. Canad. J. Zool. 35(1): 15-24. Anderson, R. C. and Chabaud, A. 6. 1959. Remarques sur la classification des Splendidofilariinae. Ann. Parasitol. 34(1-2): 53-63. -184- i -185 Aragao, H. B. 1908. Uber den Entwicklungsgang und die Ubertragung von Haemoproteus columbae. Arch. Protistehk. 12: 154. Augustine, D. L. 1937. Description of a new filariid from ground doves of St. Croix. Virgin Islands. Tr. Soc. Trop. Med. Hyg. 31: 47-54. Baker, E. W. and Wharton, G. W. 1952. An introduction to acarology. 465 pp. Macmillan Co., dew York. Baker, E. W. et al. 1956. A Manual of parasitic mites. 170 pp. Nat'l. Pest Control Assoc. Inc. Sew York. Barber, B. A. 1923. Notes on the life history and habits of Mallophaga. Papers of Michigan Acad. Sc. 1: 391-395. Baylis, H. A. 1926. A new species of the nematode genus Syngamus. Ann. and Mag. Nat. Hist. 18(Ser. 9) Baylis, H. A. 1939. The fauna of British India. including Ceylon and Burma. Nematoda. II. (£ilarioidea, DloctopKymoldea, and Trichinelloidea). 274 pp. Taylor and Francis, London. Beaver, P. C. 1937. Experimental Studies on Echinostoma revolutum (Froelich) a fluke from birds and mammals. Illinois Biol. Monogr. 15: 1-96. Becker, E. R. 1956. Catalog of Eimeriidae in genera occurring in vertebrates and not requiring intermediate hosts. Iowa State Coll. J. Sc. 31(1): 85-139. Berestneff, N. 1904. Ueber das Leucocytozoon danilewakvi. Arch. Protistenk. 3: 376. Bikhovskaya-Pavlovskaja, I. E. 1955. 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Biol. 94: 163-166. Tseng, Shen. 1932. Etude sur les cestodes d'olseaux de Chine. Ann. Parasitol. 10(2): 105-128. Tubangul, M. 1932. Observations on the life histories of Eunarvphlum murlnum Tubanqui, 1931 and Echinostoma revoiutum (Froelich. 1802). Phil. J. 3 S T 5 7 :' W ' t t l T . ----- Tubangul, M. A. and Masilungan, V. A. 1937. Nematodes in the collection of the Philippine Bureau of Science, III. Phil. J. Sc. 64: 257-268. Van Cleave, H. J. 1916. Acanthocephala of the genera Centrorhvnchus and Medlorhynchus (New genus) from N. American birds. Tr. Am. Micr. Soc. 35: 221-232. Van Cleave, H. J. 1918. Acanthocephala of North American birdB. Tr. Am. Micr. Soc. 37: 19-48. Van Cleave, H. J. 1936. The recognition of a new order in the Acanthocephala. J. Parasitol. 22(2): 202-206. Van Cleave, H. J. 1947. The acanthocephalan genus Mediorhvnchus. its history and a review of the species occurring in the United States. J. Parasitol. 33: 297-315. Van Cleave, H. J. 1948. 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Yorke, W. and Maplestone, P. A. 1926. The nematode arasites of vertebrates. 536 pp. Blakiston1s §on and Co. Philadelphia. Zekhnov, M. I. 1947. Age alterations in the infection of the rook (Corvus fragilegus L.) by endoparasites. Helminth. Abstr. 16: 181. Ziemann, H. 1898. Neue Untersuchungen uber die malaria und den Malaria-erregem Nahestehende Blutparasiten. (Abstract) Centralbl, Bakteriol. 23: 758-760. Explanation of Figures in Plate I ' c • Uninfected erythrocyte of the crow Haemoproteus danilewskii. gametocyte (early stage of development.inside erythrocyte) H. danilewskii« female gametocyte, (developing around host cell nucleus) H* danilewskii. female gametocyte, (later stage of development) H. danilewskii« mature female gametocyte, (parasite completely surrounding host cell nucleus) H. danilewskii. male gametocyte, (early stage developing around host cell nucleus) H. danilewskii. immature male gametocyte (later stage of development) H. danilewskii. mature male gametocyte, (host cell nucleus atypically located) Leucocvtozoon sakharoffi, mature female gametocyte enclosed by nucleus and cytoplasm of host cell L. sakharoffi, mature male gametocyte enclosed by nucleus and cytoplasm of host cell e Isospora corviae, unsporulated oocyst * , I. corviae. sporulated oocyst LEGEND. FOR PLATE I CH - Cytoplasm of host cell CP - Cytoplasm of the parasite G - Granule XL - Inner wall of Oocyst K - Karyosome NH - Nucleus of the host cell NP - Nucleus of the parasite NS - Nucleus of sporozoite 0 - Outer membrane of oocyst OR - Refractile granule of oocyst S - Sporocyst SB - Sporoblast SP - Sporozoite SR - Residual body of sporocyst -205- / PLATE I Explanation of Figures in Plate II Figure 13. Mediorhvnchus grand!s. adult male, ventrolateral view 14. M. grandIs. caudal end of male, ventrolateral view 15. M. grandis. hooks on basal part of proboscis, lateral view 16. M. grandis. proboscis extended, lateral view 17. M. grandis. single hook from anterior part of proboscis 18. M. grandis. egg 19. Orthoskr1abinia rostellata, scolex 20. 0. rostellata, mature proglottid 21. 0; rostellata, gravid proglottid containing uterus and paruterine organ -207- PLATE II Explanation of Figures in Plate III Figure 22. Hymenolepis corvi, scolex 23. H. corvi, mature proglottid, dorsal view 24. H. corvi, mature proglottid, cross section 25. Anomotaenia constricta. mature proglottid, cross section 26. A. constricta,. scolex 27. A. constricta, hook 28. H. corvi. hook 29. H. corvi. hexacanth 30. A. constricta, mature proglottid, ventral view 31. A. constricta, hexacanth -209- PLATE III Explanation of Figures in Plate IV Figure 32. Hymenolepis serpentulus, scolex 33. H. serpentulus. mature proglottid, dorsal view 34. H. serpentulus. hexacanth 35. H. serpentulus. mature proglottid, cross section 36. H. serpentulus, hook 37. Hymenolepis variabilis, scolex 38. H. variabilis, hook 39. H. variabilis. mature proglottid, ventral view 40. H. variabilis, hexacanth 41. H. variabilis, mature proglottid, crosssection / PLA TE IV Explanation of Figures in Plate V Figure 42. Echinostoma revolutum. ventral view 0 43. Amphimerus speciosus. ventral view 44. Brachvlecithum americanum. ventral view 45. E. revolutum. egg 46. B . americanum. egg 47. Conspicuum.macrorchis. egg 48. A. speciosus. egg 49. E. revolutum. cephalic spines, ventral view 50. C. macrorchis, ventral view -213- PLATE V Explanation of Figures in Plate VI Figure 51. Capillaria anatis. male tail showing spicule and sheath, lateral view 52. C. anatis, male tail showing spicule retracted, ventral view 53. C. anatis, vulva and esophagus, lateral view 54. C. anatis. egg 55. Capillaria contorta, egg 56. C. contorta, esophagus and vulva, lateral view 57. C. contorta, male tail showing spicule sheath, lateral view I. PLATE VI Explanation of Figures in Plate VII Figure 58. Porrocaecum ensicaudaturn. head, dorsolateral view 59. P. ensicaudatum, head, ventrolateral view 60. P. ensicaudatum, esophagus, ventrioulus, and intestinal caecum 61. P. ensicaudatum, male tail, ventral view 62. P. ensicaudatum, head, en face view 63. P. ensicaudatum, egg 64. P. ensicaudatum, vulva, lateral view 65. Syngamus trachea, egg 66. j>. trachea, spicules, ventral view 67. S. trachea, copulatory bursa and bursal rays, ventral view 68. S.. trachea, male and female in copula, male and anterior end of female 69. 6. trachea", female buccal cavity and teeth • • • \ • * • O « t * * * • * .4 * • • « ® * m t , • -217- PLATE VII Explanation of Figures in Plate VIXX Figure 70. Phvsocephalu8 sexalatus. third stage larva, lateral view 71. P. sexalatus. anterior end, lateral view 72. P. sexalatus. posterior end, showing digitiform cuticular processes, lateral view 73. Microtetrameres helix, egg 74. M. helix, female tail showing posteriorly located anus and more anteriorly located vulva, lateral view 75. M. helix, caudal end of male, lateral view 76. M. helix, female 77. M. helix, female, anterior end -219 Legend for Plate VIII C - Cephalic subdorsal ganglion CL - Cephalic lateral ganglion CS - Cephalic subventral ganglion E - Excretory sinus L - Lateral ganglion LB - Lumbar ganglion N - Nerve ring P - Excretory pore PL - Posterolateral ganglion PV - Posteroventral ganglion R - Retrovesicular ganglion RG - Rectal gland SO - Subventral ganglion -220- PLATE VIII Explanation of Figures in Plate IX Figure 78. Acuaria anthuris, male buccal cavity, ventral view 79. A. anthuris. head, en face view 80. A. anthuris« female tail showing papillae, ventral view 81. A. anthuris. egg 82. A. anthuris. vulva, lateral view 83. Diplotriaena tricuspis, anterior end of female showing vulva and tridents, lateral view 84. A. anthuris. male tail, ventral view 85. D. tricuspis. egg 86. D. tricuspis, head, en face view 87. D. tricuspis, male tail, ventral view 0 PLATE JX Explanation of Figures in Plate X Figure 88. Splendidofilaria flexivaginalis n. sp., cephalic end of young female showing vagina without anterior loop, lateral view 89. S.. flexivaginalis n . sp.. cephalic end of female showing vagina with short anterior loop, lateral view 90. j>. flexivaginalis n. sp., cephalic end of female showing vagina with long anterior loop, lateral view 91. £>. flexivaginalis n. sp*, caudal end of male, lateral view 92. S. flexivagina11s n. sp., head, en face view 93. j>. flexivaginalis n. sp., caudal end of male,, ventral view 94. J3. flexivaginalis n. sp., female tail, ventral view 95. S. flexivaginalis n. sp., microfilaria- 96. Sp lendido filaria ohioensis n. sp., spiciiles £ ., » 9.7. j3. ohioensis n. sp., male tail, lateral view 98. S; ohioensis n. sp., male tail, ventral view *• *' r j * • -224- Explanation of Figures in Plate X (Contd.) Figure 99. S . ohioensis n. sp., female tail, ventral view 100. S. ohioensis n. sp., female anterior end, lateral view 101. S. ohioensis n. sp., microfilaria 102. S>. ohioensis n. spV, head, en face view 103. S. ohioensis n. sp., female esophagus and vulva, ventral view -225- PLATE X - 226- Explanation of Figures in Plate XI Figure 104. Laminosioptes dihvmenalis n. sp., female, ventral view 105. L. dihvmenalis n. sp., tarsus I, showing caruncles and claw-like bristles 106. L. dihvmenalis n. sp., female genitalia 107. L. dihymenalis n. sp., male, dorsal view 108. L. dihvmenalis n. sp., male genitalia -227 i PLATE XI Os Explanation of Figures in Plate XII Figure 109> Svringophilus bipectinatus. female, dorsal View 110. .S. bipectinatus.■ female. ventral view 111. j3. bipectinatus. female. tarsus 112. SI. bipectinatus. female. tarsus 113. S. bipectinatus. caudal end of abdomen, female, ventral view PLATE XII • • • • W l / / no \ _. • 112 230 o Explanation of Figures in Plate XXII Figure 114. Analges corvinus, female, ventral view 115. A. corvinus, female genitalia 116. A. corvinus, male, ventral view 117.' A. corvinus. male genitalia 118. A. corvinus. female, dorsal view 119. A. corvinus, male, dorsal view PLATE XIII * 119 232 Explanation of Figures in Plate XIV Figure 120. Trouessartia corvina. female, dorsal view 121. T. corvina, female genitalia 122. T. corvina, male, dorsal view 123. T. corvina. male genitalia 124. Gabucinia delibata. female, ventral view 125. G. delibata. female genitalia 126. G. delibata. male, ventral view 127. G. delibata. male genitalia -233- s PLATE XIV 2 t 124 -234- Explanation of Figures in Plate XV Figure 128. Phllopterus corvi, female, dorsal view 129. Myrsidea albiceps, female, dorsal view 130. Briielia rotundata, female, dorsal view 131. P. corvi, male, dorsal view 132. M. albiceps. male, ventral view 133. 13. rotundata, male, dorsal view -235 i PLATE XV o Explanation of Figures in Plate XVI Figure 134; Map showing counties and "regions" from which crows were collected during "summer and winter seasons" -237- / Legend for Plate XVI S - Counties in which crows were collected during the "summer season." W - Counties from which crows were taken in the "winter season." o -238- PLATE XVI T o t t a w a CUTAH09A COSHOCTON CLANK AUTOBIOGRAPHY I, Joseph Jones, Jr., was born on June 3, 1928, in Albany, Georgia. '1 attended Madison High School, in Albany, from which I was graduated in June, 1945. I received my undergraduate training at Morris Brown College, Atlanta, Georgia, receiving the degree Bachelor of Arts in June, 1950. In September, 1950, I entered Northwestern University from which I received the Master of Science degree in June, 1952. Following graduation, I accepted a teaching position at Saint Augustine's College, Raleigh, North Carolina. My studies at The Ohio State University were begun in the summer of 1955. I was admitted to candidacy for the Ph.D. degree in March, 1959. -240-