m^ i. 1. ItU ICtbrarg A ^^ SJnrtli (Earnlina ^late SF997 .F6 NORTH CAROLINA STATE UNIVERSITY LIBRARIES S011 44495 S 12776 This book is due on the date indicated below and is subject to an overdue fine as posted at the Circulation Desk. i-&-^> -|\lo-V/) H&t 1 6 1977 FEB 2 4 Z DISEASE IN CAPTIVE WILD MAMMALS AND BIRDS INCIDENCE, DESCRIPTION, COMPARISON BY HERBERT FOX, M.D. PATHOLOGIST TO THE ZOOLOGICAL SOCIETY OF PHILADELPHIA, DIRECTOR OF THE WILLIAM PEPPER CLINICAL LABORATORY, UNIVERSITY OF PENNSYLVANIA WITH A FOREWORD BY CHARLES B. PENROSE, M.D. PRESIDENT OF THE ZOOLOGICAL SOCIETY OF PHILADELPHIA PHILADELPHIA, LONDON ^ CHICAGO J. B. LIPPINCOTT COMPANY S" COPYRIGHT, 1923, BY HERBERT FOX THIS BOOK IS DEDICATED TO CHARLES BINGHAM PENROSE, M.D., Ph.D., LL.D. IN GRATEFUL APPRECIATION OF HIS FRIENDSHIP AND OF HIS GUIDANCE AND ENCOURAGE- MENT IN ALL MATTERS PERTAINING TO THIS LABORATORY. CONTENTS PAGE Foreword — Dr. Charles B. Penrose 1 SECTION I. Introduction 17 Zoological Classification Numerical List of Autopsies Upon Which the Work is Based. II. Diseases of the Heart 48 III. Diseases of the Blood Vessels 66 IV. Diseases of the Blood and Bone Marrow 83 V. Diseases of the Lymphatic Tissues Including Spleen... 114 VI. Diseases of the Respiratory Tract 134 VII. Diseases of the Alimentary Tract. The Tube Proper 166 The Liver 222 The Pancreas 244 The Peritoneum 260 VIII. Diseases of the Urinary Tract 263 The Kidney IX. Diseases OF the Female Reproductive Organs 287 X. Diseases of the Male Reproductive Organs 313 XI, Diseases of the Ductless Glands. The Thyroid 316 The Thymus 336 The Suprarenal 336 XII. Diseases of the Skeleton and its Joints 343 XIII. Diseases of the Central Nervous System and Special Senses , 372 The Eye 402 XIV. Constitutional Diseases 410 XV. The Relation of Diet to Disease by E. P. Corson- White, M.D 415 XVI. Neoplasms 462 XVII. The Infectious Diseases. Tuberculosis 483 Mycosis 558 Streptothricosis 567 Miscellaneous Infections 596 XVIII. Animal Parasites, their incidence and significance, by F. D. Weidman, M.D 614 LIST OF TABLES PAGB 1 . The incidence of degenerative and inflammatory changes in' the heart. 5 1 2. The incidence of hypertrophy and dilatation of the heart with princi- pal associated lesions 57 3. A condensation of the important features of Tables 1 and 2 63 4. The heart-to-body-weight ratios 63 5. The incidence of arteriosclerosis 71 6. The differential percentage of the circulating leucocytes in a number of different animals 84 7. The various types of splenitis 126 8. The incidence of the various types of bronchitis and of pulmonary parasites 142 9. The incidence of the various forms of pneumonia 150 10. An analysis of cases of abscess and gangrene of the lung 156 1 1. The incidence of gastro-entero-colitis, its distribution and an analysis of the causes 182 12. An analysis of the inflammations of the alimentary tract in Ungulata . 196 13. An analysis of the inflammations of the alimentary, tract in Marsupialia 200 14. An analysis of the cases of pancreatitis, the associated pathology and the details of the anatomy of various orders 252 15. The weight of the thyroid body in relation to body weight 318 16. The incidence of the various enlargements of the thyroid body 326 17. The incidence of cases of degenerative bone disease in the various orders 357 18. The weight of the brain and the relation of this to the body weights in 196 animals 388 19. An analysis of pathological conditions in terms of diet 423 20. An analysis of the diets used in the Philadelphia Garden 426 21. The incidence of tumors 464 22. An analysis of the breeding, captivity and visceral distribution of neoplasms, of tumor-bearing animals 466 23. An analysis of the incidence, character and distribution of tuberculosis 486 24. The incidence of parasites in the zoological orders 630 25. The incidence of heterakis in pheasants 635 26. The incidence of various parasitic orders and families 636 27. The visceral distribution of parasites 639 28. The occurrence of blood parasites in zoological classes 651 vii FOREWORD BY CHARLES B. PENROSE The work on which this book is based was begun in November, 1901. From that date, systematic autopsies were made on animals dying in the Philadelphia Zoo- logical Garden. Previously autopsies had been made very rarely and only on animals of especial importance and interest. Pathological conditions were occasionally noted in animals subjected to anatomical study. The work was strictly volunteer, for there was no one on whom the Society had the right to call. Dr. Henrj'^ C. Chapman, a former Director, whose interest was in physiology and comparative anatomy, made nearly all the autopsies that were made before the beginning of the present work. In the annual reports from the foundation of the Garden in 1872, very few such examinations are recorded. In some reports there are lists of important deaths, but no record of the cause of death. This state of things was not peculiar to the Philadel- phia Garden. It existed in every zoological garden in the world. It exists in most of them today. When an animal died it had no interest or value except for its hide and bones. Rare specimens were sent to the Academy of Natural Sciences from the Philadelphia Garden. The great majority, however, were immediately destroyed, and thus during the twenty-nine years from the founda- tion of the Garden, preceding this work, there have been lost many opportunities — some of which will never recur — of increasing our knowledge of pathology. This book gives results of the study of animals that have not been subjected to experimental procedures and conditions, and though their mode of life has not been 1 2 FOREWORD that strictly natural to them, yet they have not been influ- enced by any of the artificial procedures of the laboratory which are usually followed in investigations on animals. Though the object of the work was the pursuit of knowl- edge for its own sake, yet results of practical value have followed: hygiene has been improved; disinfection has been made scientific ; epidemics have been arrested ; some diseases, notably tuberculosis in monkeys, and spiro- pteriasis in parrots, have been practically eliminated. The value of the work is recognized by the keepers. Their interest has increased and their morale has improved. The frequent deaths in a large collection of animals are discouraging, and a good animal man must have courage and optimism — traits that are stimu- lated by the knowledge that every animal that dies is studied to determine the cause of death, and to prevent its recurrence. Some results of this study are to be considered as * * raw material, ' ' while others permit cautious or tentative conclusions. The first include — visceral weights, inci- dence of certain lesions like anemia and hepatic cirrhoses, occurrence of calculi and observations on weak hind quarters, convulsions and constitutional diseases. The second include — eradication of tuberculosis and spiro- pteriasis, the response of the mammalian and avian heart to strain and disease viruses, the origin of pneumonia in birds, the genesis of bronchiectasis, the nature of osseous degeneration with relation to diet and alimentary tract, the comparative anatomy, physiology and pathology of the female genitalia, the occurrence of lesions in the thyroid comparable to those in man, the character of kangaroo disease. Great praise is due Dr. Herbert Fox and his assist- ants Drs. F. D. Weidman and E. P. Corson- White, for the splendid scientific spirit and thoroughness mth which they have done this work, with no adequate remuneration, FOREWORD 3 except the intangible reward appreciated only by the real research worker. It is a mistake to think that all animals in the -wild state are healthy. The healthy wild animals that we see are the survivors, the sick and the weaklings having died. Undoubtedly diseases are fewer than in captive animals ; but on the other hand wild animals are exposed to dangers to which captive animals are not exposed — such as lack of care in sickness and old age, starvation, and attacks of enemies that feed on them. The maximum longevities in some species are probably in captive animals. It is doubt- ful if a wild animal ever reaches the greatest age possible to it. Many kinds of parasites have been found in wild animals captured only a short time before arriving at the Garden, so that they must have existed in the wild. Thir- teen wild cats received from South Carolina within a few days after capture were infested with intestinal, muscu- lar, and pulmonary parasites. Several years ago there were received from the Island of Chincoteague, off the coast of Virginia, a number of native birds recently cap- tured, all of which had mould disease of the air sacs. This happened on several occasions, so that it became necessary to reject all birds from this locality. Among the birds were meadow larks and cardinals. Hunters not infrequently kill animals with disease, and if more hunters knew enough to recognize the disease there would be more recorded cases. Sick animals are not as Ukely to be killed by the hunter as healthy ones, as sick animals seek seclusion and do not move about. In many localities of the United States white-tailed deer are infested with the liver-fluke. Trout and other fish in remote mountain streams are found with tapeworms. Round worms and other parasites infest the white rhinoceros in the African forest. 4 FOREWORD In many instances the parasites and the host, when in health, get on very .well together ; but when the host weak- ens the parasites may give him the finishing stroke. Though some morbid conditions described in this book are peculiar to capti^dty, yet it seems reasonable to assume that many of the diseases found in captive ani- mals occur also in the wild. Some of the extinct races of animals may have been wiped out by disease. Captivity causes numerous physical and mental dis- arrangements. Unaccustomed, unnatural and unvaried food, change of climate and environment, physical and mental degeneration from disuse of muscle and brain, fear, ennui, nostalgia, lack of the exhilaration of chasing and being chased, unsatisfied sexual feeling — all react harmfully on the captive. No captive animals get their natural food ; and though some, like the carnivora, may get approximately their natural food, yet they do not get it in the natural way. They have but little variety and may miss elements impor- tant to their well-being. They get only certain cuts — muscle and bone ; they do not get blood, guts and glands. The lion 's meat is handed to him. He does not tear down his prey ; and one result is that the jaws of the captive- bred lion differ decidedly in shape from the jaws of the T\^ld-bred lion. He gets his food regularly, with awful monotony — twelve pounds of meat at 3.30 p.m. da}'^ after day; there is no alternation of feasts and fasts, with consequent change in the balance of the body reserves. Some animals, such as caribou, the moose, the black cock, the hoatzin, the koala, do not long survive captivity, because it is impracticable to obtain their natural food. Gastrointestinal disease is the commonest disease of wild animals. Twenty-five per cent, of sick humans in civilized communities also suffer from it. On the other hand, many animals become accustomed to the new diet and thrive on it. FOREWORD 5 Animals also often thrive in a climate very different from that of their natural habitat. Some animals from warai comitries, though kept outside without any artificial heat, get on very well during the severe weather of Phila- delphia winters, when the temperature often drops to zero Fahrenheit. For the past two years it has been the custom to put outdoors all monkeys — of value — that are suspected of having tuberculosis. The cheap monkeys are killed. The outdoor monkeys are kept in fair sized cages — five feet square and six feet high — usually singly, though sometimes two are together. The vervet, the grivet, the leonine macaque, the yellow baboon, the Hainan gibbon and many others, have not only survived the climatic con- ditions but have not succumbed to tuberculosis. South American monkeys do not stand cold as well as Old World monkeys, and cannot be kept out in severe winter weather. The monkeys that are kept indoors during winter have free access to the outside through swinging doors and they often go out voluntarily in the coldest weather. The same is true of other animals. A note made several years ago (February 12, 1914) states : ''Temperature last night was 1° F. At 2 P.M. outside temperature 13° F., in car- nivora house 25° F., elephant house 38° F., giraffe house 39° F., monkey house 42° F. Two Bengal tigers were voluntarily outside. Monkeys that have been out all winter are : two Barbary apes, Hainan gibbon, lion-tailed macaque, yellow baboon, grivet monkey, pig-tailed macaque, and eleven rhesus macaques. All the animals appeared comfortable." The physical condition of the animal and the kind of cold — damp or dry — have much to do with its ability to stand low temperatures. Monkeys have passed through very severe winters without damage, and subsequently in a less severe winter have had frozen fingers, toes or tail. The size of the cage or pen has not as much effect upon the well-being of the animal as might be expected. Rep- tiles, birds and mammals do as well in cages and pens of 6 FOREWORD medium size as in very large ones. A deer or antelope in a large enclosure does not use all its domain. It usually has a favorite corner near the food, water and shelter, and stays there. Nor has a large enclosure been found perceptibly to diminish mortality from cage-mates. The stronger will follow the weaker until he gets him, no matter what the enclosure. Even in the large flying cage for birds it is necessary to keep out those of a scrappy disposition. From the point of view of the public small enclosures are more satisfactory. Mental disease in captive animals offers a fasci- nating field for study — now chiefly speculative. Many conditions are present to produce it — all the con- ditions that cause prison psychoses in man. And many if not all the insane and perverted acts of animals have their counterpart in the human. Masturbation is very common in mammals. Eating of their own feces, coprophagy, is not infrequent, and is especially notable in one of the highest types — the Chim- panzee. Perhaps occasionally coprophagy may be due to some defect in diet. There is no instinctive disgust at excrement in the lower animals any more than there is in the uneducated child. Nevertheless, eating feces cannot be healthful, and probably does not occur in nature ; and occurs' only in the human with mental disease. Some of the insane acts of animals if prevalent in the wild would probably cause the extinction of the race. Such are killing of the young by the father and by the mother ; killing of the female by the male, usually during rutting time, in some instances reminding one of Sadism in man. This kind of sexual killing does not often occur in the wild. The female has a better chance to escape, and the male probably does not feel so inclined to damage her when he chooses her himself as when she is chosen for him by his gaoler. When the mother devours her young it is usually shortly after birth. I have, however, the record of a Jungle cat {Felis chaus) who ate two of her FOREWORD 7 kittens when they were seven weeks old. Some of the domestic animals devour their young ; the sow often does it, and occasionally the bitch. Sucking, gnawing and eating parts of itself or of a cage-mate is not uncommon in a variety of animals. Bears lick their paws until they are sore ; a monkey may gradu- ally gnaw away its tail from the tip to the body ; an ocelot (Felis chihigonazon) bit open his scrotum and devoured his testicle; a Tasmanian devil (Sarcophilus ur sinus) bit off one of his front feet at the wrist; a monkey may gradually gnaw away its fingers; and numerous other self-inflicted mutilations occur. Often there is a local irritating cause, as skin disease, lice, or freezing. But in many cases no local cause can be found, as no local cause can be found for thumb-sucking or nail-biting in the human. The surgery done by monkeys on their frozen fingers and toes is interesting. After the flesh has sloughed the monkey bites off the protruding phalanges, apparently without pain, so that satisfactory well-covered stumps are made. Animals often mutilate their cage-mates in an amicable way as distinguished from fighting. A bear may lick its mate's ears until the hair and skin are gone. A monkey may eat its mate's tail or patches of its skin, the victim lying placidly while the process goes on. Many animals are addicted to perverted acts on their own or their mate's sexual organs. It is probable that the phenomena just mentioned are due to confinement, idleness and ennui, and that they do not often happen in nature where an animal is kept busy seeking his food, fighting and avoiding his enemies, attending to his mate or mates, and meeting the various vicissitudes of his environment. It should be remembered, however, that the members of a wild species vary in intelligence and temperament, as humans do. There are morons and perverts among 8 FOREWORD animals in the \\'ild ; but not being coddled by the normal members of the species, they have a poorer chance than has the subnormal human of surviving and of transmit- ting their peculiarities. Fear, ennui, loneliness and nostalgia, by affecting the minds of captive animals, react on their physical condi- tion. Some animals have the fear of man bred in them. The young often show it from time of birth. This is especially common in animals that have survived for gen- erations in proximity to man. It is one reason for their survival. The mother and father may have become tame and gentle in captivity and yet the young one may be a wild thing from birth. Such fear is sometimes uncontrol- lable, an apparently slight cause making the animal dash itself against the fence of the enclosure. It is not mere speculation to discuss the physical effects of the emotions on the animal body. It has been shown that fear, anger, and grief bring about distinct measurable physical changes. Dr. Corson- White has found that the red cor- puscles are increased by over two million per cubic centimetre in the blood of a cat frightened by a dog bark- ing at it. The amount of blood sugar is also increased. Such observations are suggestive in a consideration of the changes that may occur in a captive animal sub- jected to acute and chronic fear. The monotony of a captive animal 's life is broken only by feeding, the sight and sounds of others in the same building or nearby, and by visitors. Many animals show their appreciation and pleasure when visitors approach, and some of the more intelligent animals, bears and monkeys and some birds, '' show off " apparently to keep the visitors there. When there is nothing doing, some stand swaying their heads, like a weaving horse, or pacing the cage, inanely tagging at each turn the side of the cage with the head or other part of tlie body — often so persist- ently that a sore is produced. Nearly all animals are social and suffer from loneliness when kept by themselves. FOREWORD This is true even of the lowly forms. The keeper of the reptile house reported that a giant tortoise became despondent and refused to eat when his companion, a leopard tortoise, was taken from him, and that he braced up as soon as the leopard tortoise was returned. It is not necessary that the companion be of the same species, or even of the same family. A lion or a tiger may be satis- fied with a little dog for a companion, and there was an African rhinoceros at the Philadelphia harden that was very discontented and unhappy when alone and became perfectly satisfied when she was given a domestic goat as a cage-mate; and the huge rhinoceros stood for a good deal of butting and bullying from the goat without retaliating. A sympathetic keeper may do much to relieve the loneliness of the animals in his care. Nostalgia, or homesickness, has been felt by all men. Some have died of it. The tradition among writers is that it affects young people and those who have been living nearest to a state of nature. In this country the American Indian and the negro are affected more than the whites. Much was written of it after the Franco- Prussian War and the American Civil War. It is a real condition, capable in extreme cases of causing death and of so weakening the sufferer as to make him more suscep- tible to the invasion of other diseases. At the present day we hear less of it among civilized people than formerly, perhaps because the conception of home has been broad- ened by modem methods of intercommunication. The wild animal's conception of home is narrow; he comes directly from it into an environment where he may see many other animals, but not one of his own kind. Pre- disposing causes of nostalgia are stronger with him than with the human. That home means a great deal to animals is shown by the migration of birds — the return of the carrier pigeon, and of the lost dog, and of the swallow, w^hich returns every year to the same nesting place. 2 10 FOREWORD All animals long for the things of nature — open air, earth, grass and water. They are thrilled when their feet touch the sod. Even the hippopotamus gambols when he leaves his concrete house and his feet touch earth and grass. The face and carriage of many animals cannot express feelings as in the human, though it is not unreasonable to assume that animals may indicate feelings by expression understood by their mates, though not understood or even noticed by man. When they can express it in a human way their feelings may be read. The dejection of nos- talgia is especially shown by anthropoid apes. Gorillas have been noted for it from the earliest writers. The orang is prone to it, shows it by his attitude and expres- sion, and sometimes dies of it. It is stated in A Handbook of the Management of Animals in Captivity in Lower Bengal, p. 130, that elephants have been observed to shed tears abundantly if forced to leave their old home and surroundings. How much other animals who cannot express homesickness may feel it, and how often it is a cause of unhappiness, depression and predisposition to disease, it is impossible to say. Most wild animals in captivity are sterile. The reason is not known. It shows the profound effect of captivity. It would be diflScult to determine whether the sterility of a mating is the fault of the male or the female. There is no apparent rule for sterility. Some families are always sterile in captivity, others are fertile, even with very unfavorable surroundings. The deer, horses, hippopotamuses, pigs, goats, sheep and oxen, are good breeders; while antelopes, rhinoceroses, giraffes, ele- phants, are poor breeders. Some members of a family may be good, others poor. The lion and puma breed fairly well ; the tiger, leopard and jaguar, very poorly. Bears breed well, but the mother usually destroys her young. FOREWORD 11 We cannot mate wild animals and birds simply by putting males and females together in the same cage. Domestic mammals and birds usually mate under such conditions, but wild ones often refuse. Many mammals and probably all birds that are not polyandrous or polygamous reserve the right to select their mates, and if the sexes are put together by man they may view each other with indifference or with animosity. There are many males and females of the same species of mammals at the Zoological Garden that will not consent to live together. A male monkey in a cage with several females will very often select one female for his mate and will have nothing to do with the others. Among monkeys fertility varies greatly. It is not practical to determine the ratio among the various kinds, as some kinds are much commoner in zoological collec- tions than others. I think that in general the Old World monkeys (Cercopithecidce) are better breeders in cap- tivity than those of the New World (Cebidce). The anthropoid apes are very poor breeders indeed; of the many gibbons, orangs, and chimpanzees, that for years have been captive in Europe and America, it is probable that only the chimpanzee has bred, and that very rarely. Refusal to mate, sterility, infanticide by father and mother, and sexual killing keep down reproduction in zoological gardens; and the number of young ones is a good indication of the character of a garden and of the provisions made for the happiness, comfort and health of the animals. With the birds in a zoological garden the conditions for nesting and laying are not good. Caged birds have no material for a nest, no privacy and rarely lay an egg. The outdoor water birds and the outdoor upland birds with natural surroundings, with secluded retreats, lay and hatch very well. Birds like mammals apparently are indifferent to publicity when copulating, but seek 12 FOREWORD seclusion for laying and nesting — from maternal fear for the safety of the young. The sexual instinct in indoor caged birds in a zoologi- cal garden is dormant. Very few copulate and very few lay eggs; pigeons and Mexican conures {Conurus liolo- chlorus) are exceptions to this general rule. Probably because the sexual instinct is dormant the males never fight over the females among perching birds, and veiy rarely kill each other. In some species of finches, however, as the chestnut-eared {gn. Amadina), the females fight among themselves if there are not enough males. Ovoviviparous reptiles breed more often in captivity than one would expect; and egg-laying snakes often lay eggs, which of course are only hatched artificially. Birds suffer less than mammals from the psychologi- cal effect of captivity. The mental development of a bird is much lower. With few exceptions, like the ruffed grouse, the bird accepts captivity easily and becomes tame, though he will not stand being touched. He views his keeper and visitors with indifference or friendliness. If a bird house is bright, cheerful and sunny, all the inmates thrive and appear to be happy. These conditions undoubtedly affect the health of the bird, as is evidenced by their plumage; bright colors that are lost in a dark and gloomy house are retained and developed when the house is cheerful and sunny. There are some birds, how- ever, that never retain their colors in the captivity of a zoological garden. Among them are the scarlet ibis, the American flamingo, and the roseate spoonbill. It has been suggested that the loss of color is due to the lack of something in the diet, mineral or organic, that the bird gets in its natural habitat. Tame scarlet ibises living with the chickens about the dwellings of natives in Venezuela retain their brilliant color. The material of the beautiful red color on the under surface of the wings of the touracou contains copper, yet these FOREWORD 13 birds retain this color very well in captivity, even after several moultings. The source of the copper has long been uncertain. In nature the birds are fruit-eaters, and their diet in cap- tivity consists of bone meal, zweibach, corn meal, wliite potatoes, eggs and carrots — foods that are usually assumed to contain no copper; and no copper utensils are used in the preparation of the food. Dr. John Marshall, however, writes me that all the common cereals contain minute quantities of copper; and Dr. Leon A. Ryan, University of Pennsylvania Medical Bulletin, June, 1907, states that copper may be found in animal tissues. Dr. E. P. Corson- White has found traces of copper in the bone meal used at the Philadelphia Zoological Garden. The copper in the red color of the touracou's wing therefore comes from the food. It is probable that a bright and cheerful bird house does not influence the color of birds by the direct action of light on the color as much as indirectly by improv- ing the health and spirits of the birds. Coloration in birds is a very complicated proposition. It depends upon age, sex, season, health, light, heat, moisture, mode of life, and food. No one bird house can combine all the conditions necessary for the retention of natural colors in every species. The desert species from a habitat of intense light and dryness require for their color a dif- ferent environment from the forest species, from a habitat of shade and moisture. The suppression of sexual feeling in captive birds may influence color. In nature the finest colors are attained by mating birds. It may be said that all animals — except those of noc- turnal habit — have a feeling of joy and well-being in fine -weather and bright surroundings that reacts favorably on the general health. The variability of the breeding period induced by captivity in many animals may be mentioned with sterility. It was sho\\Ti some years ago in the Philadel- 14 FOREWORD phia Garden by the European brown bear which in one year gave birth on January 16th, and in the following year on July 25th. It is another evidence of the profound effect of captivity on the captive animal. I know of no observations of the effect of captivity on the period of gestation. There is considerable mortality among captive animals from killing of cage-mates. I do not refer to sexual kilhng, already mentioned, or to fights over a female. Often males, with no females near, cannot be kept together ; probably sexual jealousy is at the bottom of it. Antelope and deer are especially inclined to scrap. Even a large enclosure will not save the weaker male ; the stronger follows him with horrible persistency, some- times for days, around and around the enclosure, often at a walk, but always on the offensive, at least during the day; until, careless from weariness, the weaker is caught unawares and finished by a horn-thrust in the side. Both birds and mammals often kill their mates when the mate is sick, or ^' down " from injury or disease. All animals hate sickness and death, and show their dislike by attacking or shunning it. Birds may get on happily together for months until one becomes sick, and as he crouches in a corner with ruffled feathers the others pick on him and finish him. The same is true of mammals, the sick one being horned or tramped to death by the mate with whom he had formerly been on most friendly terms. The keeper often reports an animal "killed by its mate," whereas the mate has only given the coup de grace. This brutality is not universal. Rarely a parrakeet will stand guard over his sick and dying mate; and we have seen a ratel — of a ferocious family — stand guard over and resist the removal of his sick companion. The diagnosis of disease in wild animals is unsatis- factory ; usually impossible ; clinical study as we know it in the human is impossible. "We know that the animal is sick, but not why. A certain group of symptoms accom- FOREWORD 15 panies all diseases — dull, rough coat or feathers, refusal to eat, weakness in the hind quarters, and finally getting down. They rarely show symptoms of pain — or at least we cannot read the symptoms. The pain of acute pan- creatitis in man is violent, yet many animals die with it and we cannot tell that they suffer. Animals do not suffer as much as the human, and they stand the ravages of disease better than the human. At autopsy we often wonder how the animal could have lived with the condi- tions that are found. A monkey may be apparently well until a few days before his death, though his lungs and abdominal organs may be a mass of tubercle. A small red howler monkey {Alonata seniciilus) was in apparent good health, playful and lively until twenty-four hours before his death from acute pancreatitis, though liis stomach and intestines contained fifty-one nematode worms, some of which were eight inches in length. As diagnosis is unsatisfactory, so is treatment. Usually all we can do is to treat symptoms ; and by the time disease has advanced to the point of becoming exter- nally noticeable, it has usually gone beyond the reach of medical treatment. It must also be remembered that drugs vary very much in their action in different families of animals. Nux vomica will not kill the gallinaceous birds of North America, and Tenant says that in Ceylon the hornbill feeds on the fruit of strychnos nux vomica. The pigeon is immune to opium. The Felidce are said to be unusually susceptible to carbolic acid ; veratrum viride is harmless to sheep and elk, but poisonous to the horse ; dogs can take with impunity large quantities of cyanide of potassium. These statements are true when the drugs are administered by mouth— the usual way of giving them to wild animals. The action may be different if the drugs are administered intravenously or subcutaneously. Variations in effect when they are administered by mouth are often due to chemical variations in the digestive secre- tions. It is probable that the action of cyanide of potash 16 FOREWORD on dogs depends on the amount of hydrochloric acid in the digestive tract. When worms or their eggs are found in the stools vermifuges are used, and with some animals especially liable to infestation by intestinal worms, periodic doses of vermifuges are given as a prophylactic. Turpentine is given to the zebra at fixed intervals for the round worm ; santonin, male fern and areca nut to the carnivora for the various worms that are so common in the intestinal tracts of these animals. The work of the Laboratory of Pathology is throwing light on the subject of diagnosis, and though from the character of the clinical material diagnosis can never be as satisfactory as in the human, yet we may fairly hope for improvement. Prophylaxis is our chief reliance, and always will be. DISEASE IN CAPTIVE WILD MAMMALS AND BIRDS SECTION I INTRODUCTION " We have also parks and enclosures of all sorts, of beasts and birds; which we use not only for view and rareness, but likewise for dissections and trials, that thereby we may take light Avhat may be wrought upon the body of man." The purpose of a menagerie under the auspices of a zoological society can scarcely be put into better words than those found in this quotation from Sir Francis Bacon's New Atlantis. Apt as this description of the mythical island's collection may be, it is but a reflection of the teachings of Plato 's original legend of a perfected community, and the practical applications of these teach- ings by Aristotle in his Anatomy and Physiology of Animals. The history of human study shows a constant investigation of lower forms of life, ever broadening in its scope, ever more satisfying in its explanation of biologic problems and ever increasing in value from an economic standpoint. If, however, all animals are to be subjected to ''dis- sections and trials" there inevitably will come under observation many specimens presenting variations from the accepted mean or standard or even from an average for their kind and therefore approaching what may be called pathological. The desire to explain the abnormal has had the effect, during the half century since Virchow defined cellular pathology and Darwin systematized the world's knowl- edge of comparative biology, of directing attention to comparative pathology and of stimulating the study of 17 18 DISEASE IN WILD MAMMALS AND BIRDS veterinary medicine. Moreover in the past twenty-five years much work has been done and many isolated pub- lications have appeared upon the diseases of wild animals, notably Bland-Sutton's work, Evolution and Disease (1895), a thoroughly scientific and most charmingly writ- ten book, but rather elementary in its approach of the subject of pathology, and Wood Hutcliinson's Diseases of A7iimals, a more or less popularly presented treatise. I am unaware, however, of any systematic monograph upon the subject wherein we may find data showing the char- acter of pathology in the various animal groups or the incidence of the various lesions. The reports of some zoological gardens contain the result of medical and pathological data collected for the report period. The publication of greatest merit and value is that from the Zoological Society of London, whose huge collection studied by a large official personnel makes it possible to present valuable data. The New York Zoological Park uses its material in a similar manner and has been able to explain some of the knotty problems so frequently met in wild animal collections. Here at Philadelphia it has been our practice now for twenty years to perform an autopsy upon every mammal and bird that dies, and upon all of the large or important reptiles. There is no aquarium connected with the Garden. The office of the society keeps a record of the arrival and a general description of every animal so that a brief history of the specimen is usually available. The keepers are required to observe their charges regularly and closely and to report any abnormalities to the officials of the Garden. Somewhat detailed discussions upon signs of sickness will be given at appropriate places, where also a few remarks upon treatment will be included, but as this work is not intended to be a treatise on therapy and since this subject does not differ from that referable to domes- ticated animals, little space mil be devoted thereto. Upon death a complete autopsy is performed and the INTRODUCTION 19 findings are recorded upon a printed form from which, when the histological, bacteriological and parasitological studies have been completed, a set of cross index catalogue cards are typed; these are divided into the principal diagnoses and determinations. The report of the Zoo- logical Society, appearing at the end of their fiscal year, February 28th, contains a resume of the observations for the year, together with notes of interesting cases and experimental work. There have accumulated the records of nearly six thousand autopsies and upon them as a basis has been founded the following report of the incidence and nature of pathological manifestations in the various animal groups, using also as additional data, published reports from other gardens. The book might be described as a collection of our studies, parts of which have appeared as separate articles, but most of which are entirely new, put together with as much connection as the subject matter will permit. The zoological and pathological literature has been consulted very extensively, but except for the reports of zoological societies and the publications of special students, it usually rep- resents isolated notes by travellers and veterinarians so that many articles may have been overlooked. Therefore no claim of perfect completeness of reference is made, the statements resting chiefly upon our own records. The subject will be approached from the stand- point of description and incidence, but it is inevitable that comparisons and contrasts must be noted. Into the realm of evolution (1) I shall not venture because I appreciate a lack of adequate preparation for such an attempt, and because, even if such were not the case, the material at hand is lacking in data upon fishes, many kinds of reptiles and invertebrates. ( 1 ) Those who are interested in the subject of disease in its effect on evolution are referred to Morley Roberts, Proceedings, Zoological Society, London, 1918, p. 247. 20 DISEASE IN WILD MAMMALS AND BIRDS A direct and practicable application of these data will be in the direction of explaining some of the pathological states in domestic animals and man. There are indeed many disease entities or syndromes in these groups for which no useful hypothesis has been advanced, while for others a partial explanation has been offered, usually, however, inadequate wherewith to form the basis of rational prevention or therapy. Thus, for example, essential emphysema seems to be limited to the animals of civilization ; on the other hand, the anatomical basis of exophthalmic goitre may be seen throughout nearly the whole animal kingdom yet the clinical phenomena belong characteristically to man, and are occasionally seen atypically in the dog. While it may be impossible to give a complete comparative anatomy aiid physiology for each of the pathological states, the attempt will be made to treat all subjects analogically through the zoo- logical orders. The experimental pathologist may find the records of the Garden useful in his work. For example, he can know that rodents are not prominent among the orders shomng spontaneous arteriosclerosis, but that nephritis occurs among them in about a quarter of natural deaths ; or he may learn that the Primates have a good cardiac reserve while the Marsupialia have not. Too often experimental work is not based upon facts including natural probabilities. A collection of pathological data such as is presented in the following pages may be of assistance to vet- erinarians and managers of zoological gardens in the diagnosis of sickness in animals, both mid and domesti- cated. We do not presimie to offer a system of veterinary medicine, but it is possible to introduce certain objective findings of practical hygienic and therapeutic value. Such observations are, however, limited and in our experience at the Philadelphia Garden the diagnosis of disease in a wdld animal, excepting of course those which are per- INTRODUCTION 21 fectly self-evident, is more often speculation and conjecture than at all well grounded. It is not uncommon for animals to come to autopsy presenting a perfect galaxy of abnormalities, yet the closest antemortem observation failed to reveal unusual conduct or appear- ance. On the other hand specimens are frequently opened whose organs fail to contain any lesions discoverable even by careful study. Dr. Henry Chapman, sometime prosector to the Society, once made a remark in this con- nection — * ' Why do they die or how can they live so long. ' ' Space is given to this phase of the observation of wild animals in order to emphasize the difficulties of interpret- ing their conditions, but of course it should be understood that certain data of value may be gained by close atten- tion to the details of their normal behavior and to changes which occur indicating that something is wrong. The naturalist and the trained animal keeper are, in our opinion, better judges of a wild animal's condition than is the veterinarian, miless he be at the same time a zoologist and have long experience with a menagerie. My own observation of dogs and horses leads me to think that more acumen is needed to interpret the actions of wild animals since they seem to have greater natural reserve, and of course in regard to them there are many more variables since we see fewer specimens of each species than we know familiarly among domestic varie- ties. The principal objects for observation are, as in veterinary medicine, the eyes, the hair and skin, the mucous surfaces, the droppings, the condition of the abdomen, the appetite and the desire for water. Physical examination is limited to tractable beasts and those which can be caught and handled mthout danger to the per- sonnel or unusual fright and damage to themselves. In the interpretations of physical signs in tractable animals, such as many ungulates and some monkeys, the experi- ence of the trained veterinarian is of the greatest value, but this fails amongst carnivores and birds. It might be 22 DISEASE IN WILD MAMMALS AND BIRDS said that anesthetics could be used for a thorough exam- ination, but this would be undesirable for a seriously sick animal and it is, in our experience, none too safe a pro- cedure although often perfectly practicable. Animals do not like to be molested much as they may seem to enjoy attention, and when it is possible it is our practice to avoid handling them. It might be contended that observations upon diseased states in captive animals would not represent natural developments, in other words, not that which occurs in the wild. Such indeed may be true in regard to the infec- tious diseases, but since we are imperfectly informed as to the pathology of the wild state, we are obliged to accept and use the best substitute at hand. Moreover it seems perfectly fair to consider as characteristic for an animal or group, the physical and even physiological expressions of morbid agencies as we know them, even though the animals be at the time under conditions not natural to them. It would be perhaps incorrect to say that cirrhosis of the liver occurs in .6 per cent, of animals in the wild as is the case for our autopsies, since incorrect food and infections are potent in its causation ; on the other hand, our experience and some few data from naturalists and pathologists make it conceivable that tumors occur to this number in native states. The incidence of tumors in mid rodents is quite well known. Degenerations and fibroses, the result of parasitism, are known to exist throughout the entire animal kingdom. Further to illustrate how pathology is distributed in wild life, Plimmer's experience (2) with 500 rats {M. decumanus) might be cited. He found the following: Tubercle 3 times, tape worm cysts 10, Tryp. lewisi 49, empyema 2, tumor of jaw from old injury 1, pleuritis and hydrothorax 1. Bacteria were found in 71 rats, 40 times in the lungs, 31 times in the spleen; saccharomyces were found 16 times in the lungs. Dr. W. L. Abbott reports to us personally that he (2) Proceedings, Zoological Society, London, 1911. INTRODUCTION 23 has repeatedly found coiled exproventricular worms in the wild specimens he has collected. Not only are we informed of some isolated and individual pathological states but the existence of epizootics of communicable disease among wild life is well authenticated. The simple citation of the extermination of deer in one section of Colorado by pleuropneumonia will suffice to illustrate this point. Other examples are, however, interesting. The occurrence of changes in the jaw bone ahnost certainly those of actinomycosis is reported by Blair, the specimens being shot in the mid and believed never to have been near civilization. The white-tailed deer of the Swan River Valley in Montana, are knomi to be constant carriers of liver-flukes. It would seem therefore that it is not unfair to use material gathered from animals under somewhat unnatural conditions as representing the reaction of the zoological orders to pathogenic agencies. Such conclu- sions must however be made very guardedly, for it is probable that not over ten per cent, of the total number of mammalian and avian species are to be observed in captivity. Because of the number of orders and the great variety of genera included in the present study it is probable nevertheless that the lesions are fairly representative of the whole animal kingdom. However, the nimibers and percentages given should be read to indicate the probabilities and should not be interpreted as implying the mortality relationships smce different varieties have differing powers of resistance to the same pathological state. The margin of safety in any given group for one or several dif- ferent disease entities cannot at present be stated mth any degree of precision but this factor is doubtless very great. The work of physiologists suggests that there is a reserve power in the human lung sufficient to sustain life until five-sixths of the functionating organ is useless, and 24 DISEASE IN WILD MAMMALS AND BIRDS I shall quote a case of an opossum wherein only one-tenth of the respirable surface seemed to have remained; we have repeatedly seen both lungs of a monkey apparently entirely solid. Such physical vital incompatibilities might be exemplified by many other cases, but when one reviews the physiological margin of safety, inexplicable and contradictory instances are equally numerous. 1 have seen a male deer run a doe against the fence and butt her, without result, whereas in an apparently similar occurrence the animal would be dead in a short time. Numerous instances of slight enteritis of a short stretch of duodenum or ileum have killed, with almost nothing to be found microscopically, and on many occasions we have been chagrined in being unable to discover the cause of death. The capacity of self-healing is a variable one, but seems in direct proportion to the quietness and seclusion possible for the animal and inversely to the chance of bacterial infection. The effect of captivity has been the subject of much speculation. For the preservation of health it would seem that animals require periods of rest and activity, thor- ough elimination, possibly a moderate exercise of their procreative functions, but most of all, appropriate food obtained by the physical effort we term chase. All but the very last condition is supplied in a measure in well managed collections. The degenerating effect of the absence of chase must be admitted. An interesting and suggestive example of this was noted by Mr. Jones at the London Zoological Gardens. He observed the skull of a lion that had been in captivity thirteen years, in which the canine area of the face and the part of the skull acting as the insertion for the seizing and holding muscles had undergone atrophy while the chewing muscles with their bony bases had remained normal. Numerous examples of disease atrophy are on record and those of a physical nature must have counterparts in the realm of physiology. The size to which an animal will attain INTRODUCTION 25 cannot be estimated by the examples seen in menageries. Judging by the accounts of collectors and hunters and upon the more reliable of the moving picture displays of wild animals in their native haunts, it would seem prob- able that under normal conditions of habitat the average size of wild beasts is considerably in excess of that in park specimens. The effect of captivity may also be felt in the direction of reduced resistance to infectious diseases. Brooks, of the New York Park, expresses the view that captivity increases susceptibility to bacteria and causes paren- chymatous degenerations. In the latter direction it is interesting to learn that Seligman of London claims to have seen sudden deaths in wading and struthious birds from myocardial disease, without valvular or other lesions, for which he holds the enervating effects of cap- tivity responsible. It is well recognized that a species may be unusually susceptible to a disease that it has not encountered in its phylogenic development. Man illus- trates this peculiarity very clearly. Europeans were found exceedingly susceptible to sleeping sickness when they went first to the part of Africa inhabited by the tsetse fly, and the American Indians died in hordes when they met the tubercle bacillus for the first time. Judging by the ravages of tuberculosis in captive monkeys a similar susceptibility probably explains the matter for there are no entirely satisfactory records of this disease among them in the wild state. In so far as general susceptibility to infection is con- cerned, it may be in part due to one of the artificial conditions of captivity, that of inbreeding. This influence is undoubtedly very great, both by chance in families, and by intention on the part of dealers as well as the mating which occurs in menageries. However, it is not known how far inbreeding may go in the wild state so that one must be very careful about drawing conclusions in this particular. Several years ago, at the time we reported 3 26 DISEASE IN WILD MAMMALS AND BIRDS the neoplasms found at the Garden, discussion arose as to the effect of inbreeding, and thereafter some observa- tions were made in this direction. With the exception of the hyperplasias of the thyroid, not certainly of neo- plastic nature, in a much mixed-up family of wolves, we could find no evidence that inbreeding was responsible for tumors. Plimmer and Murray of London, seem to imply that some of their inbred animals are likely to have tumors ; reference to this matter will be made later in this book. In so far as diseases of the organic systems are concerning those of the bones seem to be the only ones in which inbreeding is significant. The individual resistance will be reduced of course by the unsanitary surroundings incident to trapping, ship- ping and storage, but this need not affect the figures or pathological tendencies of classes or orders. The effect of captivity is felt in another way. A very large percentage of wild life perishes during the first weeks or months after its capture, and in gardens the heaviest mortality occurs among the recent arrivals. The London Garden figures that from thirty-three per cent, to fifty per cent, of their total mortality is in animals that have not been in the garden six months and that die because they are not yet accustomed to their new sur- roundings. It seems to us, both from an academic and a practical standpoint, that this is a long time and should afford ample opportunity for the garden to study the specimen and for the specimen to become acclimated. These early deaths are perhaps to be ascribed in large part to failure of acclimatiza- tion but many are doubtless the result of infection acquired in the wild, in transit while in the hold of ves- sels, at quarantine, or in trains, or at the establishments of dealers. We have seen a few deaths which have fol- lowed behavior that might be likened to homesickness. Perhaps the age at arrival has an influence upon the morbidity and mortality of wild animals, for it is easily INTRODUCTION 27 conceivable that the young and the very old might adapt themselves to new surroundings with much less readiness than the sturdy middle-aged adult. The age of animals upon arrival is verj^ rarely knowm, and can only be recorded as "young," "fully developed," and "old." This will have an effect upon statistics and when possible is noted in the text, but this is not practicable to the extent we desire. The meaning of "young," "adult," and "old" is not the same throughout the animal orders nor even \vithin orders. Mitchell (3) has attempted to gain concrete ideas of the expectancy of life among animals by analyz- ing the records of the London Gardens. This gentleman bases his figures upon kno^vn ages and the length of time in captivity, from a combination of which data the specific viability and the potential longevity may be estimated. Such results, he admits, can only be approximate and they show within classes and orders, a decided lack of uniformity. The terms "spe- cific" and "potential" longevity, coined by Sir Ray Lankester, apply, for the first, to the average length of life as it is affected by external conditions and those incident to procreation, while, if an animal be under ideal conditions it will attain the potential longevity which is longer than the former. These considerations have a biological and economic importance, while a knowledge of the pathology shown by the various groups may help to explain these durations of life. Contrariwise figures of the expected longevity may assist us in evaluating youth and senility in the causes of death but can hardly affect the comparative nature of the lesion. A resume of Mitchell's studies indicates that the higher apes have a potential longevity and a hardihood much less than man but still upwards of thirty years. As one investigates lower in monkeys, life periods become shorter, while in the next order, Lemures, the length of (3) Proceedings, Zoological Society, London, 1911, p. 425. 28 DISEASE IN WILD MAMMALS AND BIRDS life rises. Carnivora have a reasonably good vitality, their potential periods varying from ten years in the foxes to thirty-three years in bears. Insect eating animals are short lived, three years being a maximum. The Bat family shows great variations, the greatest life being not over seventeen years. The Rodentia have long lives compared to their sizes — twenty years in porcupines, fifteen years in squirrels, thirteen in marmots, nine in agoutis and capybaras, and three in dormice (which is also about the maximum for the rat). Hy races live four years on the average. Proboscidea, although reputed to live to great age, probably rarely live a half century and may be said to have an expectancy of twenty to thirty years. Perissodactyla (horses, tapirs and rhinoceroses) while they may live half a century, have an average life of between fifteen and thirty years. The closely related Artiodactyla fall into two groups, a first comprising antelopes, sheep, goats and deer which rarely exceed seventeen years, and a second consisting of cattle, camels and giraffes, which vary in expectancy from eighteen to thirty years. The smaller members of the Ungulata have in relation to size a relatively greater viability, the ruminants, however, having on the whole a low viability. Marsupials vary from a maximum of seven years in the opossum to eighteen in the wombats, but none of this group has a good viability. The Aves as a class or if compared according to dietary requirements, have longer potential ages and better viability than mammals. Pas- serine birds average twenty years and many live to sixty, while the Picariae approach the former figure but do not have such good viability. Psittaci and Striges may live a half century but the resistance of the latter is much reduced by any unfavorable surroundings. The rapta- tory birds live fifty years, but their viability is variable. Herodiones have a maximum expectancy of thirty years and good resistance, while their relatives, Steganopodes, may live fifty years, and Odontoglossae have a good INTRODUCTION 29 viability, up to twenty years. Anserine birds may live to be fifty, and, unless conditions are quite unfavorable, have a good resistance. Columbae may under good con- ditions live to be fifty. Gallinaceous birds may only be expected to survive twenty years, a figure also given for Fulicariae. Alectorides may live up to fifty years. Limicolas, though they do not thrive in captivity, may live thirty years. Impennes live poorly under artificial con- ditions, the greatest record being twelve years, a figure also holding for Crypturi. Struthiones, if the conditions be right, may live fifty years. Because of the variable specific longevities, it is fre- quently difi&cult to decide when an animal is senile. Man is said to be as old as his arteries, and his span of life nowadays is in the neighborhood of half a century. Par- rots exhibit lesions of the vascular system comparable to the arteriocapillary fibrosis of human beings, and their expected longevity is about the same or a little greater. From a study of our cases of this lesion in parrots it can be said to appear quite early in life and not to lead to organic disease as it is alleged to do in man. It is, how- ever, interesting to note that in those animals which are supposed to have the longest specific lives — elephants, snakes, anserine and raptatory birds, parrots — there is relatively low mortality and fewer infectious diseases are encountered. The last part of this statement should be qualified by stating that anserine birds and parrots are quite susceptible to mycoses, in all probability from musty food, which raises their death rate, but as this is accidental and artificial, it can be excluded from, consideration. In a rough way there is a direct relationship between the size of an animal and its longevity, but this is not close enough to be a reliable guide ; whales and elephants live a long time, but so do snakes and parrots. Within orders this relation of size and expected longevity is more easily seen but is not absolute. I cannot state, according to my 30 DISEASE IN WILD MAMMALS AND BIRDS present studies, that there is an unqualified relationship between the size and expected longevity of an animal and its pathological lesions. The immediate surroundings and the management of captive animals have a very direct and important bearing upon the mortality and perhaps upon the incidence of morbid processes but probably not upon the character of the latter. A full knowledge on the part of the personnel of a zoological society concerning the habits and habitat of eveiy animal in their keeping is essential, to wliich must be added a group of interested keepers. In engaging the last, it should not be forgotten that certain men have ''a way" with animals and that others cannot manage themselves. The enormous literature at the disposal of the naturalists permits executive officers to formulate a plan of housing and feeding with fair accuracy for each kind of animal, but of course it is rarely possible to obtain in sufficient quantity the natural food {e.g., ants for ant- eaters). In so far as food is concerned it seems that with a few exceptions like the one just mentioned, the substitu- tions made at the zoological gardens are nearly satisfactory. The elements in which the captive diet is poor are the inorganic salts and vitamins since Dr. Corson- A\Tiite, some of whose work is included in a later chapter, has shown that for those animals which our statistics indicate as most prone to have rickets and osteo- malacia, the available phosphorus and calcium are low, and one vitamin was also below the desired quantity. In this regard, however, I am not at all con\inced that diet alone mil suffice to explain these degenerative osseous diseases ; I shall take this up more fully later. Careful inspection of all food should be made and cleanliness (sifting of cereals, protection of meat from flies, etc.), is indispensable. The mortality among our camivora has materially decreased since the horse meat after butcher- ing was placed in covered galvanized iron pans. There INTRODUCTION 31 are many problems of feeding, too numerous to be cov- ered in a survey of this sort, which must be solved, and it is a credit to superintendents that this they have studied carefully. There are two problems in the management of animals upon which much difference of opinion exists, namely the heating of houses and the material of which cages are made. It seems to be the practice in many gardens to keep animals very warm. Dr. Chalmers Mitchell states unqualifiedly that adult animals do not have to be kept warm, and that even an equable temperature is not demanded, variations in temperature having a distinctly stimulating effect. However he main- tains that they should be kept dry and must be supplied with a shelter. This is in accord with the experience at the Philadelphia Garden, since for many years we have allowed access to the open air all winter to every animal that could stand it. A large group of macaques has now lived entirely in an open ''band stand" cage for nine years with a lower mortality than in the rest of the monkey collection, which is permitted to go indoors some of the time. Occasionally one in poor health is frozen to death, and healthy ones may lose fingers, toes, or a part of the tail, but the general condition is so much improved that they present an attractive exhibit to visitors. Unless a storm be of great severity, wild animals are usually indifferent to it although they may seek their shelter. Snow apparently is no source of fear to them, and many enjoy playing in it. The general principles of the enclosure should be proper lighting, free access of air, dryness and shelter in time of storm, the last so arranged that the sleeping place is well protected. Appropriate arrange- ments should be made for nocturnal animals, regardless of their visibility to visitors, if their preservation is of importance. The hygiene of communicable disease has influenced everyone to use concrete and metal for cages. These sub- 32 DISEASE IN WILD MAMMALS AND BIRDS stances are without doubt most simply kept clean, but they are heat-conducting and remain cold or damp longer than wood or the ground. It may be claimed that the latter two cannot be disinfected so well, but this need not militate against their use. Wood can be disinfected by sunlight or by mechanical cleaning plus disinfectants, by a blast lamp and by paint. The ground will disinfect itself if allowed to lie fallow for a time, or it may be turned over after sprinkling with lime. It is fair to note that the New York Zoological Garden reduced their mor- tality, especially from verminous pneumonia, by changing some deer herds to concrete paved enclosures; if that were the only change made the result would be very significant, but it should not be forgotten that another clean ground range might have served as well to a herd from which the infected ones had died. My own observa- tions with guinea-pigs, rabbits, mice and dogs lead me to believe that they thrive and breed better on wooden floors than on metal or stone. I have tried to work out figures to show that more animals die when housed in enclosures of stone and metal than when upon the earth or on wood, but the attempt has been unsuccessful chiefly because of the presence of epi- demics and parasites, principally among the birds. The attempt was further embarrassed because some members of an order are housed on both floorings. However, there was no great advantage for the metal and concrete floors even after the epidemic had been discounted. This Gar- den does not have a great number of pneumonias, a disease said to be favored by dampness and cold, but those that occur are chiefly among the small mammals, on wooden floors and in the large bird house in cages of con- crete and metal. However, the construction of both these houses permits the visitors to approach very close to the cage, a factor that doubtless explains the dispropor- tionate incidence of inflammation of the lungs. In so far as outdoor fowl and ungulate ranges are concerned, they INTRODUCTION 33 should be changed frequently under the best conditions since occasionally one will find groups doing badly until moved. Moreover the ground becomes contaminated with parasites such as esophagostomum and heterakis, infestation with which while not very serious in itself, may lead to fatal infection with bacteria. The effect of animal parasites upon the morbidity and mortality of wild beasts and birds in captivity is by no means clear, and Doctor Weidman aiid I are inclined to be sceptical, with certain reservations of course, of their great importance in the death rate. Doctor Weidman has kindly agreed to contribute a chapter upon the general distribution of protozoal and metazoal parasites with a summary of their probable pathogenic importance. The groups known to have a decided pathological power might be divided into the toxic, the tumor formers and the mechanically obstructive ; certain parasites have properties placing them in two of these classes. The first group comprises the hemosporidia and hemogregarines, the uncinaria and some of the cestodes, forms which pro- duce hemolysis and hemorrhages with varying grades of anemia. The importance of this group is shown chiefly among the Aves, in which high grades of anemia are occasionally met from malarial infections, but cats and dogs or even herbivores also frequently suffer from hookworm. The tumor-producers are chiefly echinococ- cus worms, the cysts of which may grow large enough to occupy nearly the entire abdomen. A certain grade of anemia and general ill health accompany this hydatid disease, partly the result of a toxin and partly by damage to important viscera. Those parasites which obstruct mechanically do so by their own bulk or by an accom- panying inflammation, incited by them as foreign bodies or by bacteria which have gained entrance at the irritated point. This is exemplified by the enormous collection of nematodes sometimes found in reptiles (a pailful was removed from a python) and by the tightly coiled or 34 DISEASE IN WILD MAMMALS AND BIRDS tangled thread and tape worms frequently found in birds. The effect of swelling by the mucous membrane under the influence of worms is illustrated by the infestation of the proventricle in parrots. Here spiroptera penetrate into and under the glandular layer which sweUs and pours out mucus, the total mass of nematodes, mucus and tissue obstructing the passage. Very many animals show parasitic infestation at post- mortem, but the percentage in which they can be said to be principal causes of death is quite small, while that in which they play a role as activator of the terminal condi- tion is also small but indeterminate. The latter group comprise, together with the anemias mentioned above, certain forms of pneumonia, of hepatic and vascular lesions. Inflammations of the lungs from ascaris and paragonimus are fairly well knowai ; fortunately we have been troubled less with this than have most gardens, pos- sibly because we do not have such large herds of herbivora susceptible to it. Hepatic diseases from flukes, from coc- cidia and from amoebae we have always with us in small numbers, but they are unimportant excepting enterohep- atitis, a condition which appears in nearly all orders. This last disease, be it purely amoebic as in dysentery of man and monkeys, or like blackhead of turkeys and chick- ens or in the forms of quail disease, arrests the attention at once and evokes a desire to explain the association of large intestines and liver. Parasitic vascular lesions are relatively unimportant. Taking parasitic infestations by and large, there are close similarities throughout the entire animal kingdom, and the effects produced by a given genus will be repeated almost exactly in several others. The pathological pic- tures of anemia, of hepatic degeneration, of cystic degeneration, of colonic ulceration or of fibroses are simi- lar in different hosts, only slight variations in the type of inflammation being noted, for instance in reptiles and birds as against the mammals. We have made rather close INTRODUCTION 35 observations upon the effect of parasites in the produc- tion of neoplasms, incited by Fibiger's discovery of nematodes in the rat 's stomach cancer, but, with the pos- sible exception of a papillomatous growth in the stomach of opossums from the action (?) of physaloptera, we have been unable to establish such an etiological relationship. A decision of the importance of parasites in any given case is not without its difficulty, and we are inclined to reserve judgment pending further analysis unless the effect of the invaders is unequivocal. Leiper (4) does not seem to credit animal parasites with a great effect on the mortality after a specimen has been in the collection six months since all the intestinal varieties he studied came from animals dying in that period. On the other hand the forms which invaded the internal organs and tissues were, in his series, from specimens resident several years in the garden. He seems to think the conditions of life at the garden favor the expulsion of intestinal worms. To what extent some intestinal worms may be commensal remains as uncertain as the value of certain bacteria in the gut tract. In man considerable importance has been ascribed to certain fermentative and putrefactive germs in the maintenance of a reaction unfavorable to strict pathogens and some observers have looked at them as possessing a digestive power. In the digestive tract of the animals eating large quantities of carbohydrate as cellulose, nature provides for its use by rumination and by supplying a large hind gut, by which means secondary mastication and bacterial decomposition of the cellulose capsule insures its fuU use. Possibly a similar usefuhiess may be finally ascribed to some animal microbes or even larger metozoa. The role of vegetable parasites in the causation of dis- ease among wild animals seems as undoubted as it is in the human being and the pathologic results are usually as (4) Proceedings, Zoological Society, London. 1911, p. 620. 36 DISEASE IN WILD MAMMALS AND BIRDS clear, at least for the entities of which we have exact data, based upon comparisons with man and domestic animals. There seems to be no essential difference among mammals between the pathological pictures of infectious septicemias, the mucous and serous membrane inflam- mations and tuberculosis for example. They are characterized by fibrinous, purulent or infiltrative inflammations which may go on to necrosis or repair, by fever, by leucocytosis and by evidences of resistance — all of these things occurring in a similar way throughout the class. Of course not all animals are receptive to all infections since specific racial and generic immunities exist, but the basic response in terms of pathology is similar. There are no normal means of judging the sus- ceptibility of wild animals on their native heath to the important pathogens of civilization, pneumococci, strep- tococci, staphylococci, cholera bacilli, the typhocolon group, the Friedlander group and others, but it is inter- esting to note that in captive conditions they evince some receptivity to these germs or their congeners. The pneu- mococcus takes a fairly heavy toll in zoological collections every year and the Friedlander bacillus, not a very com- mon cause of human pneumonitis, has been seen here and at London. Among the birds, however, quite distinct differences in some pathological processes occur, not only from the mammals but also within the class. ' As a whole birds do not produce pus as we know it in man, probably because of the absence from their leucocytes of a protein-splitting ferment; their leucoc>i;e-producing organs do not seem to respond as readily to a virus, the place of purulent exudate being taken by a coagu- lum or necrosis. The former varies from a clear gelatin-like material seen upon serous surfaces to a thick mat or mass of coarse but short fibrinous strands. Necrosis may succeed upon the latter or occur so promptly as to appear like the original form of INTRODUCTION 37 degeneration. It is usually rapid, accompanied by a circumferential congestion but not associated with active phagocytosis. Giant ceU production is variable, but when developed the appearance is like that of large syncytia. Hemolysis is not marked in the simple infections but a hyperplasia of the mononuclear nodes of the liver is the rule. The function of this nodal increase is not quite clear. It has been always thought that the scanty bone marrow would supply the necessary erythrocytes, but we have seen these mononuclear areas full of pale red cells fitted with round nuclei and without pigment. The fibrin mentioned above does not have the delicate interweaving that we know in a fibrinous exudate in man. This is inter- esting when we consider the composition of the blood and its coagulation in the Aves. The cell upon which human coagulation seems to depend, the platelet, is rep- resented in birds by the thrombocyte, which appears only up to about 50,000 per cubic millimetre. Coagulation time is relatively short and the resulting clot is firm and irregular. Perhaps this may have something to do with the nature of an inflammatory exudate. The response to infection on the part of birds may to some extent depend upon differences in anatomy, which are quite distinct, not only from the mammals within which class the anatomy is more uniform, but also from one avian order to another. These differences among the birds may be exemplified by the large foramina between lungs and air sacs in the water birds, a passage which permits infection, notably mycosis, to spread from the first to the second. Again the close apposition of the pancreas to the duodenum over a long stretch permits easy infection of the former from the latter. Still again the large renal-portal vein in the gallinaceous birds explains some of the infections of the liver sec- ondary to intestinal disease. The position of the lungs, deep in the thorax and fitted into recesses made by the sharp anterior border of the ribs and overlaid anteriorly 38 DISEASE IN WILD MAMMALS AND BIRDS by a rather firm air sac wall, makes it difficult for these organs to expand and therefore renders even a simple congestion a dangerous thing. The position of the ovary subjects the shell-less egg to much danger from the intestinal area. These and many other peculiarities of anatomy affect the pathological picture in birds. To be sure there are also noteworthy differences among the Mam- malia, notably in the intestinal and genital tracts, but the pathologic response is not so varied as in the birds. When due allowance is made for the kind of stomach and absorptive area, apparent differences can be reconciled. For example, there is little confusion experienced in comparing acute erosive gastritis or the follicular enteritis of an omnivorous intestinal tract (man or pig), of a sacculated stomach and absorptive tract (the marsupial), of a carnivorous gut (cat) or a herbivorous compound stomach with its long digestive and water- absorbing surface (cow or camel) and an expansive muscular organ with a very extensive digestive area (seal). The type of lesion seems the same, in that inflam- mation, pus, necrosis, granulation tissue and cicatrices are comparable throughout the series. The size of the hind gut has been taken by Metchnikoff as an indicator of the possibility of intoxication by degradation products of digestion. He believes that the capacious colon of herbiv- ora and the short small one in carnivora explain the relatively greater life in the latter, because here less stagnation and absorption can take place. A reference to the expected lengths of life given before hardly substan- tiates this, and in our later chapters there will be found no strong indication that animals with large colons suffer with degenerative visceral changes more than those with small ones ; nay even the reverse may be found ti-ue. In regard to epizootics the behavior of man and lower animals is similar except perhaps that during an outbreak a smaller percentage of the latter give evidence of indi- INTRODUCTION 39 vidual immunity and whole groups are apt to be carried off. Occasionally hygienic measures stay the ravages, at other times nothing seems to avail. Fortunately it is sometimes possible to sacrifice infective specimens and remove contagion. We have had few serious outbreaks, unless one might call our former heavy infection with tuberculosis in monkeys an epizootic. The principal ones were an unexplained water fowl disease which carried off one hundred and forty-six birds, an imported epi- zootic of quail disease which killed about the same number, a few cases of blackhead among wild turkeys, and a small group of cases of amoebic dysentery in monkeys and of thrush in passerine birds, and a small number of tuberculous pneumonias in snakes. Pathology may be difficult upon an anatomical basis, but when we engage to explain functional physiological defects we are surely embarked, with a poor compass and weak rudder, upon an uncharted sea. One knows, of course, that all animals require the same amount of food elements per kilo of body weight, that man eliminates his nitrogen as urea and uric acid, that monkeys do the same, that most other mammals destroy uric acid and excrete allantoin, that birds and reptiles form uric acid but chiefly urates, that there is an adaptation of alimentary tract and diet, that herbivores have a high threshold for carbohydrates, that there is a variable quantity of enz\Tne present in different organs and in different ani- mals, that vitamins, whatever they may be, are necessary for the growth of young animals, that hormones exist whereby correlations of parts are kept normal — ^but these things, rather than being learned thoroughly from ani- mals, have merely been substantiated by comparisons with man. Constitutional diseases so-called, from which the necessity to investigate much of this physiology origi- nated, are little known in the wild animal. Many cases of so-called gout have been encountered and we have seen an instance of diabetes in a fox, but more extensive 40 DISEASE IN WILD MAMMALS AND BIRDS experience is needed for definite practical comparisons. This applies to thyroid and pituitary disorders and to the vague conditions we have at times been obliged to call marasmus or inanition. Some attention has been given to the study of diets for the ^vild specimens of our Garden, but no systematic observations have been made or records kept upon purely physiological subjects. Reference will be made at appro- priate places to accepted comparative physiological facts, but our statistics permit additions to such knowledge only in a limited manner and in isolated instances. Doctor Corson- White has very ably summarized the diet, ali- mentary tract and physiology of the zoological groups with the pathology as found in our records. A word might be added here as to the destruction of animals by injury from fighting and harassment by others in the cage. Fighting doubtless causes death, especially when males are together, but it is our experi- ence that in cases of traumatic death search should always be made to see if the resistance of the dead animal had not been reduced by some disease. This is well illus- trated in birds. Very frequently a specimen will come to autopsy with its head feathers plucked out, or with a bill- thrust in the wing or pelvic region. Such birds are not infrequently suffering from malaria, or heavy intestinal parasitism or from organic disease whereby the resist- ance and self-preservatory power has been decreased. The foregoing survey of the approach to our subject reveals the multiplicity of factors wliich affect the study of comparative pathology. No one of them can be entirely omitted, no one is without some effect upon the origin and expression of disease, and no one is fully understood. Yet it is to be hoped that a study of our material, accumu- lated under routine conditions and uninfluenced by any experimental procedures, will demonstrate the natural response of various zoological groups to morbific agencies. Perhaps reactively some of the modifying INTRODUCTION 41 conditions may thus be understood. It is also not unreasonable to expect that alterations observed as natural responses in a large number of specimens in nearly normal surroundings would serve as more reliable guides to investigative speculation than would changes in a few animals under artificial technical experimenta- tion. We hope that the few facts we have been able to record may afford someone a basis for further biological studies. It is also to be hoped that something has been learned which in the end will afford an explanation of the diseases of man. Too great optimism in this direction should be guarded against because the human being is indeed an animal sui generis and, from the standpoint of normal conditions of nature, a wild animal. The zoological classification found on pages 43-46 was compiled in 1903 by Dr. A. E. Brown on the basis of the British System. With a few exceptions the computa- tions in the text are made on the basis of zoological orders since the number of specimens in families is often too small and the complications of so many different figures would be confusing. The tables will be found to correspond to the sequence of the classification. Dr. Corson- White has, however, used for her analysis the dietary groupings. A carnivore in her chapter implies strictly a meat-eater, in the rest of the book one of the zoological group Carnivora. The Laboratory of Comparative Pathology at this Garden speaks for the earnest desire on the part of the Directors to use the material to its fullest extent, and I, acting for myself and my associates, wish to record our appreciation of the facilities offered to us for study, and for the broad-minded, scientific cooperation the Board has always displayed. The President, Charles B. Penrose, M.D., Ph.D,, LL.D., was the active originator of the plan whereby this department was started, and he has given to it continuously the support of his rich experience. I wish to express for myself the deepest appreciation of his 4 42 DISEASE IN WILD MAMMALS AND BIRDS personal interest in my studies, and assistance which has been constructive and stimulating. Whether or not this present work prove useful to the extent that is hoped, the results from the Laboratory are such as to make the scientific world debtor to this gentleman. It is a duty, and a pleasant one, to record, though unfortunately in memoriam, my association with Arthur Erwin Brown, A.M., Sc.D., Ph.D., C.M.L.Z.S., for many years the Secretary of the Society and Executive Officer of the Garden. Doctor Brown as teacher was ever ready to help in the broad subject of biology, and I am proud to recall that he guided me also as a friend. The first director of the Laboratory was Courtland Y. AVhite, A.M., M.D., who served from 1901 to 1906, retiring then to accept a position in the City Laboratory. The foundation of the recording system is still in use essen- tially unchanged from his plan, and is a credit to his fore- sight. Our clerk and technician. Miss Harriet M. Phelps, has served the Garden faithfully and well since 1906. The condition of the museum is very much due to her interest and watchfulness. Thanks and appreciation for her work are felt by every one, the author most of all. Dr. F. D, Weidman has been our first assistant since 1911, and his work on parasitology has been of the greatest value, practically and scientifically. It is to be hoped that we shall be able to retain him indefinitely. Dr. E. P. Corson- White has in recent years taken an assistant position with us, armed for the work with a thorough knowledge of applied organic chemistry and immunology, and has already obtained useful results. ZOOLOGICAL CLASSIFICATION MAMMALIA PLACENTALIA Primates Simichdce Anthropoid apes. Cercopithecidce Old World monkeys (macaques, baboons) . CebidcB New World monkeys (capucins, howlers, spiders). Hapalidce New World monkeys (marmosets) . Lemubes Lemuridce Lemurs, Loris, Galagos. Carnivoba Felidce Cats. YiverridoB Civets, Genets, Paradoxures, Ichneumons. HycenidcB Hyena. Canidce . . .Dogs, Wolves, Foxes, Jackalls, Etc. Mustelidce Marten, Skunk, Weasel, Otter, Badger, Etc. Procyonidce Raccoon, Bassaris, Coati, Kinkajou. Ursidce Bear. These are grouped sepa- Otariidce Eared Seal, Sea Lion. J lately as sub- PhocidcB Common Seal, Walrus. \ pET)I A illus- trating water carnivores. Insectivora Tenrecidce Tenrec. Solenodontidce Solenodon. TalpidcB Moles, Shrews. Erinaceidce Hedgehog. Chiroptera I'teropodidcB Fruit Bats, " flying foxes." VespertilionidcB Common Bats. Emballonuridw .... Snouty Bats, Free-tailed Bats. RODENTIA Sciuridce Squirrels, Spermophiles, Marmots. Castoridce Beaver. Muridce Rats, Mice. GeomyidcB Pouched Rats, " Gophers." DipodidcB Jumping Mice, Jerboas. H eteromyidce Kangaroo Rats. Octodontidce Capromys, Coypu. Hystricidw Porcupines. CMnchillidce .*. Viscacha, Chinchilla. DasyproctidcB Agouti, Spotted Cavy. Caviidw Guinea-pig, Capybara. Leporidce Rabbits, Hare. Proboscidea Elephant. Hyracoidea Cape Hyrax. 44 DISEASE IN WILD MAMMALS AND BIRDS Ungulata Perissodactyla (odd toed) Rhinocerotidce Rhinoceros. ' Tapiridce Tapir. Equidce Horse, Ass. Artiodactyla (even toed) Bovidce Oxen, Antelopes, Sheep, Goats. Cervidce Deer, Moose, Elk. Antilocapridce Prong -horned Antelope. Girafjldce Giraffe. Tragulidce Chevrotains, Muis Deer. Camelidce Camels, Llama. Hippopotamidce Hippopotamus. BuidcB Swine, VVarthogs. Tayassuidcc Peccaries. Sirenia Sea-oow, Manatee, Durong. Cetacea AVliales, Porpoises. Edentata Bradypodidce Sloths. DasypodidcB Armadillo. Myrmecophagidoe .... Ant-eaters. MARSUPIALIA Marsupialia Didelphyidce Opossums. Dasyuridce Dasyures, Tasmanian " Devils." Feramelidcv Bandicoots. Phascolomyidce Wombat. Phalangeridce Phalangers. Macropodidm Kangaroo, Wallabies. MONOTREMATA Monotremata Echidnidce Echidna, Ornithorhynchus. AVES ' '-^^'^'''' Passeres Turdidcp Thrushes, Robins, Etc. Sylviidce Warblers, Kinglets. ParidcB Titmouse. Troglodytidce Wrens, Mockingbirds, Catbird, Etc. Pycnonoiidw Bulbul. Crateropodidce Babblers, Jay -thrushes. Oriolidw Griole. MotacilUdo} Wagtails. Dicruridw Drongos. Mniotiltidce Chats, Warblers, " Woodwarblers," Etc. Ccrrebidce Sugarbirds. Vireonidce Vireos. Laniidce Shrikes. Ampelidce Waxwing. Hirundinidce Martins, Swallows. Meliphagidce Honeyeaters. Tanagridce Tanagers. Ploceidce Weavers, Whvdah birds, Waxbills, Finches, Etc. INTRODUCTION 45 Fringillidce Finches, Sparrows, Buntings, Grosbeaks, Etc. Icteridce Hangnests, Troupials, Grackles, " Black- bird," Etc. Sturmdw Starlings, Mynahs. Corvidce Crows, Jays, Magpies, Jackdaws. AlaudidcB Larks. Tyrannidce Tyrans. Cotingidce Bellbird, Cock-of-the-rock, Etc. PiCARLE TJpwpcB Hoopse. Trochilidce Hummingbirds. Cypselidce Swifts, " Chimney Swallow." CaprimulgidcB Night hawk. Whip-poor-will. CoraciidcB f-. Roller. HALCYONES Alcedinidce Kingfisher. MomotidcB Motmots. BUCEROTES Bucerotidce Hornbill. TROGONES Trogonidce Trogons. SCANSORES Picidw Woodpeckers. Rhamphastidw Toucans. Capitonidce Barbets. COCCYGES Citculidce Cuckoos. Musophagidce Touracous. PSITTACI Loriidce Lories, Lorikeets. Cacatuidce Cockatoos. Psittacidce Macaws, Conures, Amazons, Parrots, Parrakeets. Striges StrigidcB Barn-owl. Btibonid(B All other owls. ACCIPITRES FalconidcB Buzzards, Hawks, Falcons, Eagles, Etc. Serpentaridce Secretary Vulture. Catharidce Vultures. COLUMB^ Treronidce Fruit pigeons. Columhidte All other pigeons and doves. Pterocletes Pteroclidce Sand grouse. Gali.i Tetraonidw Grouse, Ptarmigans. Phasianidce Pheasants, Fowls, Turkeys, Quail, Etc. Cracidce Curassows, Guans, Etc. MegapodidcB Brush turkey. 46 DISEASE IN WILD MAMMALS AND BIRDS Hemipodu TumicidcB Hemipodes. FULICABI^ Rallidce Rails, Porphyries, Gallinules, Coots, Etc- Alectobides Aramidce Courlan. Eurypygidoe Sun bittern. OruidcB Cranes. CariamidcB Cariama " Crane." PsophiidcB Trumpeters. LlMICOL^ CEdicnomidce Thicknees. CharadriidcB Plovers, Sandpipers, Curlews, Woodcocks, Etc. Chionidce Sheathbills. Gavle Lariidce Gulls, Terns. StercorariidoB Jaeger Gull. Pygopodes Colymhidce Loons, Grebs. Alcidce Auks, Murrs, Puffins. Impennes 8pheniscidce Penguins. Steganopodes SulidcB Gannets. PelicanidoB Pelicans. PhaldcrocoracidcB . . . .Cormorants. AnMngidce Darter " Water turkeys." Tubinares Procellariidce Petrels, Fulmars. Hebodiones Ardeidce Herons, Bitterns, Egrets. CicomidcB Storks, Ibises. Plataleiidce Spoonbills. Odontogloss^ Phoenicopteridce Flamingoes. Palamede.e Palamedeidce Screamers. Anseres Anatidce Swans, Geese, Ducks. Stbuthiones ApterygidcB Kiwis, Apteryx. Casuariidce Cassowaries. Struthionidce Ostriches. Rheidce Rheas. Cbtptubi Tinamidce Tinamous. INTRODUCTION 47 List of Animals subjected to Autopsy giving the number of each. These Figures are used to obtain the percentages quoted in the Tables and Text. MAMMALIA Primates 498 Lemures 86 Carnivora 481 Pinnipedia 20 Insectivora 6 Chiroptera 5 Rodentia 198 Ungulata 365 Proboscidea 3 Hyracoidea 7 Edentata 16 Marsupialia 175 Monotremata 1860 AVES Passeres 1355 Picarias 87 Striges 133 Psittaci 689 Aceipitres 196 Columbffi 157 Pterocletes Galli 299 Hemipodii 2 Fulicariffi 35 Alectorides 37 Limicolas 6 Gavipe 20 Pygopodes Impennes 5 Steganopodes 21 Tubinares Herodiones 98 Odontoglossae 6 Palamedes 5 Anseres 317 Struthiones 32 Crypturi 5 3505 5365 SECTION II DISEASES OF THE HEART The heart is an organ whose duty, throughout the two classes considered in this study, remains entirely identic, purely a physical one in driving the blood through the corresponding vascular system. The physics involved naturally differs between mammals and birds, but energy is derived from the automatic power lodged in the cardiac musculature. Whether this be neuromyogenic, as seems to be the case in all mammals, or purely myogenic, as is probably the case for the birds in which MacKenzie and Robertson (1) say there is no atrioventricular bundle, the result is the same, since in both classes there is some continuity of muscle fibres from auricle to ventricle. The gross anatomy varies little if any more than the physi- ology, albeit there is proportionately greater auricular capacity in the mammals than in the birds, and indeed there are differences within the classes which cannot now be readily explained ; certain minor variations of valvu- lar arrangement exist, such as the absence of the membranous right tricuspid in Aves. When, however, one considers the cardiac power available for various animals, the subject becomes one of greater breadth and complexity, for no consistency obtains even within families, since the demand for cardiac strength will vary more with habits than with zoological relationships. Thus for example the domestic rabbit has a small heart volume while the wild hare has a great one. Although, of course, the size of an organ may not be an absolute measure of its efficiency (a flea's leg muscle has relatively greater power than a man's), yet size is the only physical gauge one has for estimating nature 's preparation for expected demand. Perhaps this (1) Br. Med. Jour., 1910, 2, 1161. 48 DISEASES OF THE HEART 49 will be shown later when after discussing the pathological anatomy of the heart in the wild mammals and birds, we can study these changes in light of statistics upon the relative size of the heart. Expression of Cardiac Disease. The diseases of this organ are known only by their physical effects, chiefly by causing physical or functional defects in other organs and to a minor degree by purely physiological irregularities in the heart itself (tachy- cardia, arrhythmia). All the latter and most of the former are subjects discovered by observation during life and unfortunately cannot be included in the study at hand. Both states are well known to the veterinarian who diagnoses them with reasonable ease in animals that can be handled ; I saw one case of arrhythmia in a monkey for which no adequate gross morbid explanation was found postmortem. Some of these functional abnor- malities are certainly caused by myocardial disease and cardiac failure has occurred among many orders. An interesting observation was made by Plimmer (2) on sev- eral large birds (ostriches, storks, cassowaries) which apparently died from this condition ; at autopsy he found myocardial degeneration, or epicardial edema or only a flabby heart. Lack of exercise was held responsible by this observer. Such cardiac deaths have probably been encountered at this Garden but we have accounted them to shock, or gastrointestinal disease; this matter will be discussed on a later page. Coronary Artery Disease. If a degenerative sclerosis of coronary vessels be the cause of angina pectoris then perhaps paroxysms of this kind occur,,for we have seen such anatomical changes in the heart of three widely separated varieties of animals, a Nylghaie, a Hamadryas Baboon, a Macaque, and a Brown Pelican; The history of these animals does not (2) Proc. London Zool. Soc, 1907. 50 DISEASE IN WILD MAMMALS AND BIRDS register anything resembling the clinical picture of angina pectoris in man, and they did not come to their death from the arterial changes in the heart alone since sufficient other pathology also existed. Kinds or Pathological Change. As an introduction to the strict pathology of the heart it might be well to outline the headings of the scheme upon which it seems desirable to study the subject. It is hardly profitable to take up seriatim the ordinary general pathological processes as discussed in systems of pathol- ogy for it is our purpose to show the distribution of basic aberrations from thenormalintermsof zoologicalposition. To this end one must consider the response of the heart (a) to damaging influences and (b) to a demand for increased work. In the first group come degenerations and inflammations, upon which may succeed an incom- petency in the form of dilatation. The response of the normal heart to any physical demand greater than customary has usually been thought to lie in the direction of hypertrophy, but on occasion it has seemed to be in the form of dilatation, especially if the strain has been sud- den and severe. Starling thinks that the primary and normal reaction of the heart to physical strain is always dilatation. The idea of hypertrophy must not be con- fused with an understanding of the relatively large hearts in animals whose habits demand great cardiac power, for then it is their norm and might be called ''physiological cardiac hyperplasia." I shall use the terms ''increased muscle bulk" and "increased chamber space ' ' as preferable to hypertrophy and dilatation ; this also focuses attention upon the two features of an enlarged heart. Degenerations and Inflammations. Degenerative changes in disease are recorded in our system as amyloid, hyaline, fatty metamorphosis, gran- ular and cloudy degeneration. While there is perhaps DISEASES OF THE HEART Table 1. 51 Table Showing Incidence, in the Orders, of Degenerations and Inflammations, or in Other Words the Response to Infectious and Toxic Agencies. Heart of 1 ■■S Q .2 .2 i 1 CD i 1 * 1 21 .t iJ Primates. . . 8 1 19 2 5 3 2 12 4 2 io 2 2 .',4 2 16 2 12 14 "9 3 5 19 10 17 3 18 li "6 "8 1 6 9 3 1 7 8 13 1 1 '. '. '. "1 11 5 io ;; 2 12 2 2 6 1 38 1 54 3 16 "1 20 7 33 19 6 1 24 36 3 28 "3 2 '2 "1 4 "4 50 11 '5 1 1 7.6 Lemures 1.2 Carnivora 11. Pinnipedia 16. Rodentia 8. Insectivora Chiroptera Proboscidea 33. Hyracoidea Ungulata 5.4 Edentata U- Marsupialia 19. Monotremata Passeres 1.4 Picariae 6.7 Striges .8 Psittaci 3.4 Accipitres. . . . 18.3 Columbae .... 2. Pterocletes. . Galli 9.3 Hemipodii FiilicariEB 8.6 Alectorides. . 5.4 Limicolae. . . Gaviffi 10. Pygopodes Impennes 16. Steganopodes 20. Herodiones 4.1 Palamedes 15.7 Struthiones . . . . 34. Crypturi Total... 98 118 112 44 367 8 * Total is the number of individual animals showing lesions, not the aum of the listed changes. In this and subsequent tables, figures in italics are for small groups of animals coming to autopsy, usually less than one hundred, and from which percentages may be misleading. The number of autopsies upon such groups may be found by consulting the list given on page 47. 52 DISEASE IN WILD MAMMALS AND BIRDS between some of these conditions and true myocarditis a matter only of degree, the records have been analyzed as filed and perhaps some lessons can be learned from the responses of the various zoological orders. In the accompanying Table 1 will be found the distribution of degenerative and inflammatory lesions through the zoo- logical orders. The percentages speak for themselves but deserve as well some consideration from the standpoint of normal heart value ; this will be taken up later after the other lesions have been discussed. Endocaeditis, Myocarditis. Romberg said in his classical work on the heart that there is always some form of myocardial disease with endocarditis. The 44 cases of valvular disease detected at this Garden are 15 of chronic nature, 29 of acute or sub- acute character. In the former, the chronic, 9 showed some grade of muscular involvement, while 21 of the 29 acute cases of valvular inflammation were accompanied by myocardial damage ; the percentages are 60 for chronic and 72 for acute, a relation that would be expected if one credit the theory that many valvular inflammations start at the root of the valves, but, at all events, they indicate that after the acute stages have passed the myocardial damage may be repaired. The large number of cases of endocarditis among the Camivora, Accipitres, Anseres and Marsupialia is note- worthy and can hardly be explained by other argument than a special vulnerability of this organ in these groups. However, the unusual number of cases in our only native marsupial, the opossum, seems worthy of a special note since ten of the twelve instances in the order Marsupialia affected this particular animal. When seen these ten cases were acute in five instances, subacute in three and chronic in two. The Streptococcus pyogenes was isolated in three of the five acute cases; bacteriology of the others was negative or not done. All of the acute and one of the sub- Fig. 1. — VEGETATIVE AND ULCERATIVE ENDOCARDITIS OF AORTIC VALVE. OSTRICH (STRl'THIO AISTRALIS). ORGANISMS ISOLATED CORRESPONDED CLOSELY TO BAC. AFANASIEFFI- CHESTER. DISEASES OF THE HEART 53 acute cases were combined with some evidence of general septicemia. The type of lesion was in no way peculiar, unless the facts that all were vegetative when acute and markedly deforming when chronic, he noteworthy. In two chronic aortic cases the valvular orifice was almost closed, yet the left ventricle could not be considered as greatly hypertrophied and no dilatation existed. In one acute aortic and mitral case, general dilatation existed. The mitral was involved nine times, three times alone, four times with the aortic, once each with the tricuspid and pulmonary. Once the vegetations were hmited to the mural endocardium. Nine of these animals came from one enclosure over a period of two and a half years, dur- ing which time other opossums died from similar bacterial infections (pneumonia) despite repeated cleansing of the place. No unusual number of cases of this or similar kinds occurred elsewhere in the Garden at this time, but it would seem that we had in this cage a continued bac- terial infection. Unusual Pericardial Changes. The appearance of a stiff gelatinous exudate in the pericardial sac has attracted our attention on ten occasions (8 birds, 2 mammals). The substance seems quite homogeneous and almost entirely acellular. One specimen became solid on heating and another became turbid when put into Kaiserhng 's fluid. In two cases bac- terial cultures were made ; nothing grew. It has not been associated with tuberculosis or tumors nor has any one pathological lesion more than another appeared to accompany it. A peculiar lesion occasionally seen in birds is ''uratic pericarditis ' ' a process not infectious at all, according to Plimmer, but due to renal disease. It has been seen here in association with retention of urates in the kidney, with gout of birds, and apparently quite independent of any renal or constitutional disease. Both layers of the sac 54 DISEASE IN WILD MAMMALS AND BIRDS are pearl gray or irregularly salted with a whitish granu- lar material so that they are entirely opaque ; occasionally the distribution is spotty. The deposit does not seem to penetrate the myocardium. There is at times some involvement of other serosse, but this is usually much less marked than around the heart. It does not seem that this of itself should be fatal, but it has been the most decided pathological factor in some of the autopsies. Aside from pericarditic exudates, twenty-one instances of pericardial effusion have been encountered. They offer little that is peculiar in etiology, chemistry or cytology, but as there has been some question of the position of the accumulation of the fluid in human beings, it might be well to note the position in our material. It is recognized in veterinary medicine that the cardiac dullness is increased especially to the right, and that most of the fluid will be on that side and posteriorly. At our autopsies on mammals this is the position usuall}^ occu- pied by the fluid, the apex being covered by pericardium, unless the quantity be great enough to make the sac taut, and this position is retained whether the animal be laid upon the one side or the other ; nor does the fluid all leave the base of the heart when the body is placed prone. The crown of the heart is nearly always well covered. In birds, on the other hand, the fluid occupies the apical part of the sac, probably due to the fact that this membrane is attached by its tip to the transverse air sac wall which takes the place of a diaphragm, so that the tip of the heart is always free and the pericardium of the base fairly closely applied to the epicardium. This holds good even for the birds whose cardiac apex is normally attached to the pericardium by a fibrous band. ', Hypertrophy and Dilatation. The response of the heart to a continued demand upon its working capacity will, as already indicated, lead to increased muscle bulk or to larger chamber size. Whether DISEASES OF THE HEART 55 hypertrophy be purely the building of a bigger engine or be accompanied by, or due to, muscular disease as had been suggested by certain authors, was a question to which an answer was hoped, but it would seem that the solution is no nearer than can be obtained in human pathology. Fifteen of the 34 cases showed some degree of myocardial damage. Some of the other cases may have been instances of so-called essential hypertrophy, enlargements due to hard work or to low-grade hidden infection. Aubertin (3) ascribes such cardiac muscle increase to overwork under the stimulus of intoxi- cation from intestinal sources or from irregular constitutional functions. For information concerning this and pathological enlargements one may consult the Table (2) of Hypertrophies and Dilatations; in advance the method of charting must be known. Since it is usually impossible to decide what may be the single important factor in the cardiac disease, all of the accredited factors have been listed with the hope that the resulting figures would be significant. Thus an animal may have recorded pericarditis, nephritis and arterio- sclerosis — ^who shall say which was primary or most potent in the cardiac change. Essential hypertrophy is limited to those cases for which there was no concomitant pathology that might have been responsible for the overgrowth. There was one in a carnivore (fox) and one in a raptatory bird (buz- zard). Idiopathic dilatations on the other hand are much more common, but they stiU bear a relation to the apparent vulnerability of the heart. Their distribition is as follows : Primates 1, Carnivora 1, Pinnipedia 1, Ungu- lata 1, Marsupialia 4, Anseres 1. These may be cases such as Plimmer described, of cardiac failure, indicated by dilatation, the result of inactivity. Besides these special instances and the ones accounted for in the list, there were three acute dilatations apparently due to shock, (3) Comptes Rendus Soc. Biol, T, 62-206. 56 DISEASE IN WILD MAMMALS AND BIRDS two ungulates and one marsupial, probably incidental to fright when being caught by the keepers. The association of secondary dilatation with hyper- trophy is only evident in three instances. One case and perhaps the most interesting, is that in which the princi- pal antecedent pathology was thyroid hyperplasia and nephritis; the dilatation was perhaps agonal or shortly before the last struggles. It would seem that all of the dilatations occurred shortly before death because long standing passive congestions and dropsies of cardiac origin are exceedingly rare; only one certain case is recorded (carnivore). Let us now examine the Table (2) according to orders and then as to causation. The Primates' heart is appar- ently well able to increase in size in response to increased work, a demand most often made by pulmonary, pleural and pericardial diseases. Two of these cases occurred in animals suffering with pulmonary tuberculosis suffi- ciently extensive to impede cardiac action while in another case the tuberculous lesion was mild but a peri- carditis existed. When the right hand columns are inspected it would seem that on occasion dilatation may occur; one of the tuberculous pulmonary cases had a dilated heart. The slothful lemurs apparently have no call upon their cardiac mechanism. Carnivora with their large organ, which, it would seem, should be prepared for excess work either simply as a reserve or as an inherent ability to grow, present in about equal numbers, hypertrophy and dilata- tion. It is admitted that there are within this order, genera of differing habits, but analysis of the canida?, felidae and ursida? for examples, in the first place, offer too few specimens for conclusions and, secondly, have upon trial actually shown nothing definite, so that we are forced to use the larger group, the order. It is interest- ing to note that long continued infection is in this order the most potent factor in enlargements of the heart. Four DISEASES OF THE HEART Table 2. 57 Table Showing Incidence -per Order of Hypertrophy and Dilatation, and the Principal Associated Lesions Believed to Have Etiological Importance. Hypertrophy Primates Lemures Carnivora Pinnipedia Rodentia Insectivora Chiroptera Proboscidea. . . . Hyracoidea .... Ungulata Edentata Marsupialia. . . . Monotremata . . Total Mammals Passeres Picariae Striges Psittaci Accipitres Columbse Pterocletes Galli Hemipodii Fulicariae Alectorides Limicolae Gaviae Pygopodes Impennes Steganopodes . . Tubinares Herodiones Odonotoglossae . Palamedes Anseres Struthiones .... Crypturi Total Birds .... Total For meaning of italics see foot note Table 1 5 34 2 7 7 3 13 10 6 3 15 54 3 10 9 14 8 3. 12.5 4.5 1.5 58 DISEASE IN WILD MAMMALS AND BIRDS of the ten cases show myocarditis. Nephritis does not seem very important in relation to cardiac muscular increase, but occurs with great frequency in association with dilatation. Three of the cases of hypertrophy were associated with thyroid disease and two of these showed dilatation as well. The general causes of chamber distention are more diverse, and we see associations that do not appear with hypertrophy, namely arterio- sclerosis and diseases of the chest. The rodents seem to have no power to increase muscle bulk, but a sufficient number of cases of dilatation occur to make one conclude that this is their method of response to unusual strain. Pulmonary disease, mostly of infec- tious nature, and myocardial degenerations are the principal causes. The next order to show cardiac enlargement is the Ungulata where nephritis is the most frequent associa- tion with hypertrophy and disease of the pleura and peri- cardium with dilatation, or the reverse of the factor value in the Camivora. These animals, fairly well prepared for flight, with moderately large hearts, seem more often to show dilatation than hypertrophy. Two Edentata (armadillo) showed dilatation but no hypertrophy. Marsupials behave somewhat like rodents in that the heart does not seem to increase muscle bulk, but our records do not explain this clearly. As already men- tioned four cases had no sufficient internal reason for dilatation, but as one was probably the result of shock three only remain to be accounted for. Nephritis seemed to exist in all three, but two of them had kangaroo-mycosis of the jaw and a general chronic infection. If now our attention be given to the Aves we find the highly specialized Passeres and Striges not represented and their closely related well-organized orders Picarife and Psittaci with only an isolated single case. This is the more interesting since the last order suffers reasonably DISEASES OF THE HEART 59 often with arteriosclerosis. Accipitres, the birds of pug- nacious habit and carnivorous diet, seem well able to increase their muscle upon demand, but do not often suffer dilatation. Vascular and renal diseases stand out most prominently in the etiology, and one-half of them show myocardial change. The Galli, which includes both ground and flying birds, are represented but fail to exhibit any unusual accompanying disease. Anserine birds apparently have a low power to increase the size of the heart, but most often allow it to dilate. Struthi- ones, large stalking and rapidly travelling birds, apparently have a good margin of safety in their cardiac mechanism. ) Summary of Lesions Associated with Hypertrophy and Dilatation. Analysis of the associated pathology will reveal that among the mammals, renal disease, chronic infections and diseases of the thoracic serosa are most often respon- sible for hypertrophy, and that something over one-third of the hearts showed myocardial damage. Among the Aves arteriosclerosis and renal disease are most impor- tant in enlarging the heart; half of the cases had myocarditis. In so far as dilatation in mammals is concerned, renal disease and acute infections are decid- edly more important than other influences, even than the next in order — chronic infections and pulmonary dis- eases ; only one-fifth of the cases had myocardial disease. Acute infectious disease is the most potent cause of dilata- tion in birds ; only two of the eight cases had degeneration of the heart muscle. Comparison of Mammalia and Aves. If a comparison of the incidence of increased muscle bulk in the two classes be made (4) it will be found to occur two and one-half times more often in mammals, (4) This is done by determining the percentages of hypertrophy and dilatation for the total number of each class examined at autopsy. 60 DISEASE IN WILD MAMMALS AND BIRDS while dilatation occurs nearly ten times more often among the mammals than among the birds. Hypertrophy is accompanied by myocardial change in 44 per cent, of the cases, whereas muscular degeneration was only seen in 24 per cent, of the dilatations ; this change is conspicu- ously lacking in the Primates, Ungulates and Marsupials. The usual teaching has been that dilatation, which means enlargement of chambers and thinning of walls or at least no thickening thereof, imphed an inability on the part of the heart to keep up with increased demand — a decom- pensation. If Starling be correct that dilatation is not a degeneration of pump value but merely one of adapta- tions to increased demand, then this method is more characteristic of mammals than of birds. There is, however, the reserve power to increase the muscle bulk inherent in the mammalian, not possessed or needed by the avian heart. The large-hearted class Aves certainly dilate their blood pump less frequently than mammals and indeed have less cardiac disease. An analysis of the incidence of hypertrophy versus dilatation shows that hypertrophying power resides in the Primates, Accipitres and Struthiones, their hearts relatively seldom dilatating. Lack of such power and consequent dilatation resides in Rodentia, Ungulata, Marsupialia and Anseres. Hypertrophying power lies therefore chiefly in the heart of average size for its class, dilatation occurring in the small heart. (See page 63.) Avian Hypertrophy. There is little to be learned from the nature and anatomy of the hypertrophies and dilatations except per- haps their character among the birds, in which the physics of the circulation is somewhat peculiar. In this class both the hypertrophy and distention are predomi- natingly left-sided, a state probably explained by the pressure against which the pump must work in flight because then the lungs and the viscera are somewhat DISEASES OF THE HEART 61 compressed by the pressure of an excess of air in the pneumatic sacs. At all events while concentric hyper- trophy was mentioned once, it is difficult to estimate the degree of increase in the right chambers because they are not uncommonly well filled when diastole occurs at death. Grober (5) asserts that the normally large heart (or what I have called "physiological hyperplasia") shows a * 'hypertrophy" of the right ventricle because of the extra work entailed in flying. This is certainly not the case in the material we have seen under pathological conditions. Right-sided increase might be expected if pulmonary or serous membrane affections were prominent, but left- sided increase, following arteriosclerosis and nephritis is the actual finding. The best examples of concentric hypertrophy are in the dogs with thyroid disease and the best examples of concentric dilatation in ungulates suffering shock. , Summary. The foregoing pathological data can now be sum- marized by grouping the facts under the headings of absolute and relative vulnerability of the heart. By the former is meant the actual number and quality of lesions in the various orders, but here at once one comes upon the irregularity of examples of zoological and pathological character, and if one trust entirely to the percentages, fallacious conclusions might be reached. Basing judg- ment upon the incidence of pathological lesions in mammals and birds, it is evident that the former has greater vulnerabiUty, as 13 is to 6.2. This is noteworthy as we shall learn that the bird has a larger and appar- ently better prepared heart than the mammal. Attempts to discover the order or kind of animal having the great- est or lowest vulnerability are difficult for the reason given above. Thus, for instance, Pinnipedia, Probos- cidea, Edentata, Gaviae, Impennes, Steganopodes, and (5) Arch. fur. Ges. Physiologic, 1908, 125, 507 62 DISEASE IN WILD MAMMALS AND BIRDS Strutliiones present the highest percentages of cardiac lesions, but the total specimens examined are so few that these figures may well be misleading. (See Tables 1 and 2.) If, however, figures mean anything in such small groups, these are the animals which have the greatest cardiac vulnerability. They have little in conmaon in regard to zoological relationships and habits ; four of the seven orders are rather slothful and three are active. It is much better to limit our observations to those orders from which sufficient examples have been subjected to autopsy and upon which we have some standards for comparison in the heart-body weight ratio. It so happens that in the above seven orders I was unable to obtain any reliable figures of heart weight. Table 3 is a combination of data from Tables 1 and 2 for the principal orders from which we have enough material (at least one hun- dred autopsies) and for which it is possible to obtain as comparative standards figures indicating the weight of the normal heart in kilograms of body weight; Table 4 gives these ratios for normal hearts. The information about the weights was obtained from some of our own figures and the references given in the footnote. (6) There are no extensive data upon weights and measures in exact terms, such as body weight, so that we are limited to the numbers quoted in parentheses besides the orders in the table. The ratios might be modified slightly by a greater number of examples, but they show certain things by comparison of the classes ; in a rough manner the heart ratios correspond to the pathology. (6) dal Piaz: Papers from the Department of Anatomy, University of California, 1912. Bergmann: Dissertation, Munich, 1884. Loer: Arch. f. die gesamte Physiologic, 1911, V. 140-293. Grober: Arch. f. die gesamte Physiologic, 1908, V. 125-507. Grober: Deutsch Archiv f. Klin. Med., 1907, V. 91, 502. Welcher and Brandt: Arch, fiir Anthropologie, 1903, V. 28. Vierordt: Tahellen, 1906. Parrot: Zoologischer Jahresbericht, 1893. Hasenfeld and Romberg: Arch. f. Exp. Path, und Pharmacol, 1897, V. 39-333. Joseph: Jour. Exp. Med., 1908, V. 10-521. DISEASES OF THE HEART 63 Table 3. Table Containing a Condensation of the Two Foregoing Tables and Showing Figures for Degenerations, Hypertrophy and Dilatations for Orders Having the Largest Number of Autopsies. Degenerations, &c. Hypertrophy Dilatation 7.6 11. 8. 5.4 19. 1.4 6.7 .8 3.4 18.3 2. 9.3 4.1 15.7 .8 2.1 0. 1.2 .6 0. 1.1 0. 0. 4.3 0. .7 0. .6 .8 2.2 4.2 3. 5. 0. 0. Striges 0. Psittaci Accipitres Columbae GalU .1 .5 0. .7 Herodiones Anseres 0. 1.5 Table 4. Table Showing Weight of Normal Heart in Relation to Body Weight. Number of Specimens used to Determine Weight Quoted in Parenthesis. Average Heart of Grams per Kilogram of Body.* Man ( 4) 5.67 Primates ( 4) 6.56 Carnivora ( 6) 6.78 Rodentia ( 5) 5. Ungulata (10) 5.8 Marsupialia ( 3) 5.1 Average 5.82 Passeres (43) 19.8 Picarise ( 9) 21.3 Striges ( 4) 7.33 Psittaci ( 6) 8.89 Accipitres ( 7) 12.32 Columbae ( 4) 14.47 GalU (16) 11.08 Fulicarise ( 3) 23.82 Limicolse ( 2) 8.78 Anseres (14) 11.8 Struthiones ( 1) 12.7 Average 13.84 * These values are obtained in part from the literature, in part from our own specimens. At this laboratory the hearts of animals killed or dying from a short illness, organs showing no pathological change, were removed by cutting at the base of the vessels, washed free of blood and weighed. The weight of the whole animal was obtained after death. For meaning of italics see foot note Table 1. However, there are many reasons why great caution should be used in evaluating the relative size of the heart. Welcher showed in his work that the proportion is 64 DISEASE IN WILD MAMMALS AND BIRDS greater in small and young animals than in large and adult ones. All the authors quoted agree that in birds and to less degree but still clearly in mammals, there is a direct relationship between the bodily activity of an animal and its cardiac bulk. This is fairly well shown in the list of avian heart ratios, but not so clearly in the mammals. What shall be considered the most active mammals — the monkey, perhaps, with his tendency to be occupied constantly, yet we find the greatest heart bulk among the Carnivora, animals prepared for travel and struggle, and the smallest among the Rodentia, quiet and timid animals. The avian order showing the greatest cardiac ratio, the Fulicariae, shore birds, is made up of some quiet hiding varieties, and of some capable of very prolonged flight; the most constantly active fliers (Pas- seres) also have a high cardiac weight proportion. The inactive owls have the smallest heart bulk. The contrast between the average heart-to-body weights of mammals and birds is striking, the latter hav- ing two and one-half times as much as the former, 5.8 vs. 13.8. Since this is the most prominent and best supported statement in the table of weights, it may be used to compare mth the incidence of the pathology as seen in the two classes. Degenerations and inflammations occur in mammals and birds as 9.5 is to 5.5.(7) Hj^pertrophies occur in mammals and birds as 10.3 is to 4.3.(7) Dilatations occur in mammals and birds as 2.4 is to ,28.(7) ' In other words, mammals are much more susceptible than birds to degenerative and inflammatory processes, show an ability to increase the muscle bulk two and a half times as great and are liable to chamber dis- tention nearly ten times as often. It might also be put / (7) These lipures are obtained by determining mathematically the percentage of each feature in each class and then reducing the numbers to their lowest value. DISEASES OF THE HEART 65 that birds cannot or do not need to increase their muscle, and that the chamber and muscle balance is more perfectly arranged. While in the preceding pages hypertrophy has been discussed rather from the standpoint of its value as a compensating and reserve capacity, and dilatation as a degenerative or decompensatory process on the part of the cardiac mechanism, it may be that dilatation of the mammalian heart is the usual method employed by the class in response to increased demand. It seems certain, however, that the originally and normally larger heart, both mammalian and avian, more often uses an increase of its muscle to this purpose. Hypertrophy was accompanied by myocardial disease in 44 per cent, of the cases, while dilatation showed this change in only 24 per cent. This supports the theory that dilatation is a normal response of the myocardium under strain and the belief held in many quarters that the muscle increases its bulk because some of it is damaged. The differences between classes are not so conspicuous between orders. However, the large heart of the carni- vores increases both its muscle and chambers, while the small heart of the rodents and marsupials more often dilates. Analysis of the avian orders is inconclusive and somewhat contradictory. Let it suffice to say that the birds which fly most, with exception of the ducks, have a relatively low vulnerability, and the soaring carnivorous Accipitres and the largest birds, Struthiones, apparently have a high susceptibility to damaging influences and enlarge their muscle bulk in response to increased work, i Aneurysms of the heart are quite rare ; only two have been seen. They were both located at the apex of the left ventricle in birds ; they did not rupture. Myocardial damage is evident in both cases but the cause is not clear ; parasites could not be demonstrated. Plimmer reports a case of cardiac aneurysm at the apex from infestation of the heart muscle by sarcosporidia. SECTION III DISEASES OF THE BLOOD VESSELS The gross anatomy of the blood vascular system is constructed upon the same general scheme throughout mammals and upon a comparable basis in birds. Micro- scopically there is little variation throughout the orders unless it be in the relative proportion of muscular and connective tissues. The origin of the great vessels at their cardiac base and their distribution to the pul- monary and to the greater circulations in no way differ in these two classes in that it always consists of an efferent pathway to the lung and a root vessel above the aortic orifice. The foiTner has usually quite a distinct origin on the right side, but in some birds the posterior wall of the pulmonary artery may overlie the entire aortic base ; this, however, is not the rule for birds. The aorta in most mammals remains a separate and distinct vessel for some distance, after which it gives off the innominate and sub- clavians. In the birds on the other hand, the stretch immediately above the aortic valve is usually ballooned out somewhat, into a sort of sac or ampulla from which the subclavians and descending aorta arise. This forms a structure of rather trident shape, the lateral prongs being the subclavians, the middle and posterior being the aorta proper. In some birds the aorta may have the length of a centimetre or more then dividing into the left subclavian and right aorta from which the right sub- clavian comes off. There is definitely more support to the heart and vascular roots in mammals than in birds, in the latter class these structures lying quite free between the lateral air sacs and well in front of the lungs. Nor is there the richness of mediastinal areolar and fatty tissue in the winged creatures. DISEASES OF THE BLOOD VESSELS 67 The vessels of mammalia retain a considerable wall throughout nearly their entire length. At first the wall is thin compared to the calibre of the vessel while the arteries smaller in calibre, have a heavy wall. In birds the arterial stalk at the heart is supplied with very heavy walls, but after the second branching the relation of wall to calibre seems to continue about the same. In this class the stalk vessels have wall to calibre relation of 1 to 3 (measurements in 2 Passeres, 1 Psittaci, 1 Accipitres) whereas in mammals the relation varies from 1 to 5 to 1 to 7 (observations on 2 carnivores, 1 ungulate, 2 rodents). In mammals the consistency^ of a normal artery wall remains much the same, a firm, resilient, yellow-white tissue, quite opaque and standing open upon cross section. In birds this description covers the main stalk, the aorta in the abdomen and the first part of the carotid and iliacs. When these characters are lost, the arteries become semitranslucent bluish strands so that they are difficult to follow in the muscles of the neck and extremi- ties. This is particularly true in the Passeres, Picariae, Galli and Columbae while in the Psittaci, Accipitres, and Anseres the arteries are distinctly whiter than the veins but yet quite soft. In the Herodiones and Struthiones, thick walled vessels may be followed as far as the second joint in both extremities. These differences depend in part upon the grosser construction of the central arteries in Aves and in part upon the larger amount of elastic tissue in them than in the secondaries and smaller vessels, and than in comparable mammalian vessels. It is impracticable to go into the minutiae of histology in the different orders, which indeed varies but little, although attention might be directed to the facts that in all central vessels the relative amount of elastica is greater than in smaller ones and that muscular tissue seems to exceed in the latter. Considerable work has been done upon the amount and arrangement of muscle bands in isolated genera, but no comprehensive data are at hand 68 DISEASE IN WILD MAMMALS AND BIRDS upon orders. The strands of muscle do not seem arranged so regularly as in mammals; the pulmonary artery of the cat, for example, has a muscle arranged like an oblique band in waves or festoons along the length. The mammals as a class seem more richly supplied with arteries and veins than do the birds, and the square area of the vascular system is likewise larger. This is dis- tinctly different from the amount of heart bulk as given in the discussion of kilogram-heart ratios so that one might say that the birds are * ' overhearted and undervesseled. ' ' In so far as the physiology of the two classes is con- cerned'it is obvious that a different regulatory system is necessary because, aside from the variations of pressure incidental to pulmonary, muscular and visceral work, there remains the altering pressure within the air sacs of Aves, a force different under states of rest, of running, of deep water swimming and of flying with or against the wind. Part of the internal air pressure variation is cared for by the ability a bird has to respire the air in its sacs and bones, but in prolonged exposure to the pressure under water or during protracted flight some compen- satory mechanism doubtless exists. This seems to reside in part in the heavy elastic quality of the arterial stalk and the very rich venous supply of the abdomen, includ- ing the renal-portal system and the distensible pelvic veins. Just where the governing power for this mech- anism resides is as much a matter of debate as in the case of the human being, but certain researches would place it in the caudate lobe and pituitary body.y Having discussed these general comparative data we can now pass to a consideration of the pathology seen at this Garden. The subject will be studied from the stand- point of the vessels as a system and the changes peculiar to it. Naturally the most important lesions affect the great stalks and the principal trunks, from which the processes may continue into the smaller vessels. The DISEASES OF THE BLOOD VESSELS 69 essential alterations are inflammatory and degenerative, of which the latter are by all odds the more important. The former are either involvements of the vessel walls by frankly infectious processes, or less easily proved to be bacterial in origin, as is the case with periarteritis nodosa. Acute arteritis and phlebitis are constantly encountered and present nothing unusual. General nodal periarteritis has been seen in the lower animals, Lupke having reported (1) before the German Pathological Society a big outbreak in cows, but it is less common than among men ; we have not discovered it here. Thromboses. Thrombosis is practically always a parasitic or an infectious process although at times considerable difl&- culty is encountered in explaining the source of the worms or bacteria.j Thus, for example, the iliac or femoral thromboses which are at the bottom of intermittent claudication, are frequently quite vague in origin. We have had one such case in a deer in which a partly occlud- ing thrombangeitis existed in both femoral arteries and veins. Mesenteric thrombosis, a serious condition in cattle and horses from infestation with sclerostomum or strongylus, has not been proven at the Garden, but we have seen one case of numerous thromboses of the venous radicals in the jejunal wall apparently due to some nema- tode larvae; the specimens were so soft by decomposition that determination was not attempted. There occurred a thrombosis of the cava and aorta originating from a necrotizing cloacitis, apparently streptococcal in nature, in a Demoiselle crane. The clot, while not totally occlu- sive, extended nearly as far as the heart in the vein and the abdominal aorta. There is also on record a throm- bosis of the vena cava and right pulmonary vein in an American beaver, harboring Hepaticola hepatica in the liver, with a fibrosing pneumonia due to this parasite. (1) Verh. deutsch Path. Gesel, 1906, X, 149. 70 DISEASE IN WILD MAMMALS AND BIRDS Another case in which parasites seemed to take a hand concerned a common raccoon with tapeworms (sp. ?) in the small intestine and microscopically discoverable parasitic parts in the lungs ; these organs were the seat of extensive congestion and venous thrombosis, the latter containing really enormous numbers of diplococci. The parasites probably paved the way for bacterial invasion. A frank case of septic thrombotic aortitis was noted in a Rice Grackle, the infectious focus apparently being a vegetative * tricuspid " valvuUtis.) Arteritis. In addition to these cases, productive inflammatory changes were discovered five times affecting vessels in or near frank inflammatory processes. The animals affected with this productive process were three birds, a rodent and an elephant. In the case of two birds and the rodent the process was associated with chronic intestinal lesions, while in the elephant it was found as an endarteritis oblit- erans in large vessels of the lung of chronic pulmonary tuberculosis occurring in this animal. These instances serve as examples of the truly productive inflammatory processes affecting vessels and illustrate the distribution through the animal kingdom. Pathogenetically there are no essential differences, and histologically they corre- spond to the forms seen in man. Had every piece of tissue been subjected to microscopy wherein such lesions might have existed, more examples might have been discovered, but these processes excite no peculiar secondary effects so that attention is not dra^vn to them directly. The only noteworthy difference between mam- mals and birds is the fragile character of the clots in the latter class. This is peculiar because the principal response of this class to an infectious irritant is coagula- tion necrosis, liquefying enzymes apparently being absent or smaU in quantity. DISEASES OF THE BLOOD VESSELS 71 Fatty deposits in the aortic intima are by no means uncommon in the human subject and are encountered at all ages, even in youth at a time when progressive arteriosclerosis does not accompany them. There is a belief in many quarters that this fat may be laid down Table 5. Tabh Showing the Incidence of Degenerative Arterial Disease, the Percentage in Animals Subjected to Autopsy and the Principal Associated Pathology. "o 1 F ■3 o CD is 6^ ^S Ulu s ■« S m H II 2 r §1 > 0.T3 1^ ^ •< S > rt o^ 3 16 13 .6 3.3 3.5 5 7 3 1 1 1 ? 1 3 1 5 2 3 1.8 Total 35 3 2 1.8 .22 2.2 7 7 2 5 1 2 4 8 Picariae Psittaci 13 3 1.8 2.2 1 1 6 ?, 3 2 Striges... . . . Accipitres. 13 6.6 1 4 2 5 ? 4 GaUi* 5 1.6 I 4 2 1 2 Steganopodes* 6 25. 1 3 Herodiones 1 1. Palamedes 1 20. 1 1 Anseres 11 3.4 2 4 1 4 1 Struthiones 7 22. 1 2 3 Alectorides 2 5.A 1 1 Total 66 1.8 1 6 15 4 25 9 n Grand Total 101 1.8 8 13 20 5 27 13 19 *1 with Coronary sclerosis only. **2 with Coronary sclerosis only. For meaning of italics see foot note Table 1. and then removed. Such deposits are exceedingly rare in wild animals ; when they occur it is in small indefinite patches and not the bands or rows as found in man. Degenerative Arteritis or Arteriosclerosis. Whether or not it be exact to speak of the more pro- tracted forms of vascular disease usually called arterio- sclerosis or atheroma as degenerative, such changes fonn the most pronounced features of the lesions, and 72 DISEASE IN WILD MAMMALS AND BIRDS we have made such a separation at this laboratory. Here is not the place to engage in the academic discussion of the nature of the process, but I A\ish to state that col- lectively the changes as seen in such lesions in the lower animals are more degenerative than productive, and that we have never seen true ulcerative atheroma as it not uncommonly appears at the autopsy table in any large hospital. This disease of the vascular walls has long been attributed to alcohol, gout, syphilis and other such prolonged intoxications to which we might apply the light term of ''toxins of civilizations." Too little credit, or dis- credit has been given to chronic intestinal disorders, overeating, and overdrinking of ordinary fluids, to entirely incorrect diets, and to chronic bacterial diseases. Even though the exact counterpart of the disease in man does not occur in lower animals, we shall see the probable association with food and with habits, in a manner discordant with former teaching of the causation of the disease. The group to which the name degenerative arteritis has been applied is, as has already been indicated, more productive than the analogues seen in the human being, but indeed it is questionable whether the lesions even in the lower animals are not more degenerative than pro- ductive." Since, however, chronic arteritis is always asso- ciated with damage to the elastic and muscular fibres of the media as well as with fatty change and overgrowth of the intima, all the deforming and degenerative cases will be classed together. The general picture in mammals is one of diffuse rather than of plaque-like thickening, but well outlined raised or depressed areas are encountered. In the aorta and larger branches one may find irregular streaking and loss of elasticity with fairly clear, pale yellow or gray, flat sections of distinct opacity. Rarely these may con- tain calcareous matter, a change most often seen in the carnivores. The lesions are very largely limited to the KiG. 2.— ARTERIOSCLEROSIS AND ATHEROMA. THORACIC AORTA. JACKAL (CANTS AUREUS). THIS WAS CONTINUED TO THE MESENTERIC AND ILIAC VESSELS. DISEASES OF THE BLOOD VESSELS 73 aorta ; 26 or 76 per cent, of the 35 cases had this distri- bution alone. The arch seemed never to be affected alone, and indeed it is rather commoner to find opaque patches stretching along the thoracic or even abdominal portion ; this is especially true of the Ungulata. There have been also in mammals five cases of mesial change which have given rise to the picture described by Monckeberg and usually entitled by his name. However, the noteworthy differences between the wild animal and the human cases are the absence of advanced calcification in the media under the concavities and the prominence of the changes in the aorta near the heart to be found in the former. These few cases do not permit an associa- tion of the arterial disease with any particular pathology in other parts. Considered minutely, the outstanding lesion in the class Mammalia is the separation of the elastic fibres by fluid and debris, apparently derived from the degener- ated muscle fibres, associated with a decrease of round and elliptical nuclei. Globules and hyaline pink-staining material are often collected between split-up elastic strands, which fibres in some cases seem quite numerous, in others reduced. In the intima heaping-up of cells and fibres is very moderate in degree while usually one finds only subendothelial edema. When the process has advanced far, the microscopy is like that of well developed human lesions. Arterial degeneration due to parasites gives a different picture in that medial degen- eration is far advanced and some fibrinocellular activity is seen upon the intima when this tissue remains. When, however, the infestation has proceeded to weaken the wall sufficient for it to give way into an aneurysm, little or no vestige of the true arterial wall is left. In the Aves the distribution and anatomy of this process present some differences. The aorta is as usual most conspicuously the seat of change, but it is note- worthy that the dilatation or ampulla immediately above 6 74 DISEASE IN WILD MAMMALS AND BIRDS the aortic valves and from which the main vessels spring, is practically always free of lesions which are on the other hand most marked in the thoracic and abdominal sections. One's attention is usually attracted to the aortic surface by its roughness although visibly there may be no plaques, but upon close inspection a mottled opacity may be detected. This all seems due in the few cases subjected to tissue section, to hyperplasia of endothelia, with or with- out fibre increase. The media may show muscular granu- larity or no change at all. At the stage when plaques are formed, fairly well outlined, firm but rather brittle, raised areas are detected, seated upon a distinctly opaque gray wall. The remainder of the vessel may be smooth and elastic but sometimes, in the Accipitres for instance, a general resistance to pressure and tension is found. Microscopically such a vessel will show a media the seat of ruptured muscle fibres, split-up or broken elastica and some debris, while the intima is covered with active and distinct fibrocellular exudate. I have for comparison divided the cases into those in which the superficial productive character was prominent and those seemingly entirely a degeneration of the media, that is with inactive intima. In mammals 77 per cent, of the cases were of the degenerative type while in birds 50 per cent, were of this kind^ The exact importance of this difference is not easy to evaluate, but with the facts that the bird has a greater elastic supply for its large vessels and a greater wall-to-calibre ratio, it is interesting. The aorta alone was affected in half of the birds, the remainder showing lesions in the carotids and f emorals. , The tendeiicy for the media to degenerate would lay the basis for concavities on the intimal surfaces after the type seen in Monckeberg 's sclerosis. A number of cases of this variety have been encountered, but instead of being better developed in the vessels of extremities as in man they have presented more definite pictures in the Fig. 3.— arteriosclerosis. ATHEROMA AND ANEURYSMAL DILATATIONS IN THORACIC AORTA. WILD CAT (KELIS RUFKUS). DISEASES OF THE BLOOD VESSELS 75 aorta and pulmonary vessels. Examples will be quoted under the discussion of the various orders. Primates are not often affected with degenerative arterial disease, two of the instances observed showing this change confined to the coronary vessels. As might be expected the myocardium in both was affected, and in one animal had a definite concentric hypertrophy. A very interesting case was encountered in a Lion-tailed Macaque {Macacus silensis ). His heart showed distinct fibrosis of the conducting pathways from auricle to ventricle and of the papillary muscles. No atheroma was present in the aorta, but in the pulmonary distinct sacculations of the Monckeberg type were found. Mesial degeneration was apparently responsible, but no calcifica- tion had occurred. Death was due to acute gastritis. Carnivora present about half the cases seen in mammals and 16 per cent, of the total. Five of the six- teen cases owe their origin to parasitic arteritis and were combined with aneurysms. As will be noted by consulta- tion of the list there is no outstanding accompanying pathology, a fact which makes parasites more important. One case of mesial degeneration, resembling the Monckeberg type was observed in the thoracic aorta of a bear. Ungulata are generously represented, thirteen cases being recorded distributed rather unevenly between the odd-toed (1 or 8.5 per cent.) and even-toed groups (12 or 2.9 per cent.). The single case in the first group occurred in a Zebra {Equus burchelli) wherein was found about the middle of the thoracic aorta a diffuse thickening of media and intima in a circular plaque approximately two centimetres in diameter; it was by no means so well developed as similar lesions in the even-toed ungulates. This recalls the expression of doubt by ZinserUng as to the occurrence in the horse of arteriosclerosis similar to that in human beings. 76 DISEASE IN WILD MAMMALS AND BIRDS The lesions in the Artiodactyla are both in plaques and diffuse, the aorta and its branches sometimes being quite wrinkled but beset with firm elevations with and without calcification. These changes are fairly definite and, although they never attain the development seen in man, resemble the stage of wrinkling and roughness in the preulcerative stage of the human analogue. Two of the cases were associated with dilatations of the mesen- teric vessels and with periarteritis, a picture strongly suggesting parasitic infestation; in one case ineffectual search was made, in the other no record is made of the parasites. Renal disease occurred only twice and myo- cardial damage only once. Chronic pulmonary disease, present thrice, took the form of tuberculosis twice and pleural adhesions with atelectasis once. Marsupialia present three quite interesting cases, a Tasmanian Devil (Sarcophilus ur sinus) and two Kanga- roos (Macropus). The first showed distinct sacculations in the ascending aorta, suggestive of Monckeberg's sclerosis but equally resembling several small or incom- plete aneurysms. The underlying vessel was opaque and stiff, continuing so to the middle of the thoracic portion. The other marsupials showed distinct mesial damage with early calcification and a roughened intima ; once the pulmonary artery was involved. It will be noticed that the orders Rodentia and Lemures are missing from those showing arterial disease although we have a reasonable number of autopsies upon them. Passeres are hardly good exponents of vascular disease, an interesting thing in view of their large heart, heavy vessels, and flying habits. It is, however, striking that two of the three birds of this group had aneurysms of the aorta, one of which ruptured just above the origin, partly into and partly outside the pericardium. Despite several microscopical sections we were unable to find the tiny ends of the breach and any evidence of parasites. In Fig. 4.— arteriosclerosis IN AORTA. OTTER (I.LTRA CANADENSIS). THERE IS RELATIVELY LITTLE INTIMAL CHANGE, THE MEDIA BEING FIBROTIC AND ALLOWING THE INTIMA TO BE DEPRESSED IN SMALL CONCAVITIES. DISEASES OF THE BLOOD VESSELS 77 the other case a vegetative growth occurred on the intima near the dilatation. The PicariaB were represented by a Hornbill and a Toucan. The former presented roughened yellowish ele- vations for two centimetres above the aortic valve ; this seemed the only involvement. The Toucan had a few small scattered but deep yellow plaques in the same location. Psittaci, although failing to be accredited with a high percentage of arterial disease, nevertheless present some striking and interesting changes. In the first place, the central vessels are not so conspicuously the principal seat of atheroma as is the case in many other orders, and the lesions are not so productive. It is usual to find flat areas of opacity, perceptible as easily by section through the wall as by holding up the opened vessel to the light, the seat of the density being in the media. This can be followed into the wing arteries and, upon microscopic section, these smaller vessels will show mesial degener- ation, thus being comparable to arteriocapillary fibrosis of man. However frequent this picture may be, there are also instances of overgrowth in the intima, prominences over opacities or raised plaques at the points of branching, lesions which correspond to the activity of the inner coat. It is interesting that renal disease, chronic pulmonary and general infections occur in a goodly proportion of these birds. Striges is an order of little importance. The lesions in the two cases consisted of rather prominent plaques in aorta and large branches. Accipitres stand out as giving the greatest percentage of any order of which we have had a fair number upon which to make comparisons. Their arterial lesions are frequently accompanied by renal, myocardial and valvu- lar disease. Mesial and intimal alterations are about equal in degree, irregular patches going on to softening without ulceration, and early calcification being quite prominent. The sickle at the branching of the renals is a 78 DISEASE IN WILD MAMMALS AND BIRDS favorite site of deposit. Like the parrots their arterial damage is not confined to the aorta, but may be found in the carotids, femorals or small ^\dng arteries. The most common situation is, however, in the lower thoracic and renal regions. Gain, represented by five specimens, seem to have their vascular disease accompanied very often by myo- cardial and infectious disease. Their arterial pathology consists of raised gray patches in the aorta only. Steganopodes give the highest percentage among avian orders, but this must be held sub judice because of the small number of specimens subjected to autopsy. Renal disease occurred in three cases. Their lesions are in the form of yellow opaque streakings in the aorta and its branches, in one case following the carotid half way up the neck. Plaques are not common, and when they occur are streaky and illy outlined. Herodiones, represented only by a heron, are neg- ligible. This bird showed a diffuse thickening with early thrombotic deposits attached to the intima. ■ Palamedes are represented by a Screamer which showed around the orifices of the renal arteries an early fatty deposit and mesial opacity. Anseres present a considerable number of cases which are accompanied by cardiac, renal and general pathology. The character of the lesions is like that of the Accipitres and the distribution differs in only one particular. In four of these eleven cases the elevations or opaque areas were limited entirely to the stretch of aorta which might be compared to the arch in mammalia, that is the part with which the right subclavian is in closest contact and which bends almost directly backward to become the thoracic aorta. This excludes the ampulla just above the aortic valves. There seems no real reason for this and it may be accidental. Struthiones, with seven specimens having arterial disease in the great vessels alone, seem to have no 4 ' ■> KIG. 5.— AORTIC ARTERIOSCLEROSIS. SARUS CRANE (GRUS ANTIGONE). THE LESIONS ARE ELEVATED AND IRREGULAR. DISEASES OF THE BLOOD VESSELS 79 especial characters unless these be in the heavy furrow- ing and stiffness of the wall, with opaque, elevated, indefinite patches, seen mostly in the descending aorta; once a long tough and partly brittle stretch was found in the carotid. Alectorides, with a relatively high percentage, present irregularly outlined fatty and finely granular patches in the intima of the lower aorta and abdominal vessels, and, in two cases, as far as the vessels of the lower extremities could be followed. The orders Columbae and Fulicariae are missing from the list of Aves having arterial disease, yet a reasonable number of autopsy records are at hand. Summary on Arterioscleeosis. Having discussed the orders separately, a review of the whole situation is desirable. Chronic arteritis, or as it is usually called arteriosclerosis, is common to very many zoological orders, and its principal lesions are comparable throughout the two classes under considera- tion. A statement as to its incidence would best be made by adopting an arbitrary number of observations as the desired minimum upon which to draw conclusions, and I shall adopt one hundred as such a figure. Accepting this as reasonable, a review of the table indicates that the order of percentage incidence is : Accipitres 6.6, Ungulata 3.5, Anseres 3.4, Carnivora 3.3, Striges 2.2, Psittaci 1.8, Mar- supialia 1.8, Galli 1.6, Primates 0.6, Passeres 0.22; the other orders have less than one hundred specimens each. There is no doubt that carnivorous birds have the highest incidence of chronic arterial disease. Next in order come three varieties with nearly equal incidence, the ungulates, anserine birds and carnivorous mammals. These orders have little in common unless it be that in nature they are often engaged in prolonged or strenuous effort, as in fight or flight. We possess no measurement of their vascular supply but by consultation of the table giving 80 DISEASE IN WILD MAMMALS AND BIRDS heart weights (page 63) it will be found that three of them have values below that of the class in which they belong ; the Carnivora alone have a greater heart-to-body ratio than the average for its class Mammalia. Nor do these orders have any direct dietetic relationship. The expected longevity of these groups does not permit one to discover any reason for arterial changes except per- haps that they have a reasonably good viability under park conditions, and therefore many have a longer oppor- tunity to develop vascular disease. It so happens, however, that the first four groups are the most likely to suffer from gastrointestinal inflammation, of dietetic or bacterial origin. It is interesting, but not easily explicable that the orders of great activity. Primates and Passeres, are at the end of the list ; their food is very largely carbohydrate in character. Just why Lemures, Rodentia and ColumbaB should be missing is not quite clear, because orders of comparative habits and food are included. A review of the concomitant pathology reveals the fact that nephritis more often accompanies these proc- esses than any other single condition. Among the chronic infectious disease in the table is included chronic enter- itis ; this group falls well behind the renal diseases. The relatively small number of cases of valvulitis speaks rather against an active infectious origin of the vascu- lar lesions. Aneurysms. Aneurysms have been observed all over the world and in nearly all the larger orders. The London Garden has had an unusually large number to report, the most striking being that described by Seligman in the 1906 Report of the Society, in a tiger thirteen years in cap- tivity which had fourteen sacculations from pea to plum size scattered along the aorta. Even with the number of cases on record and those collected here it would be Fig, 6. — PARASITIC ANEURYSM IN THORACIC AORTA PARADOXURE (PARADOXURUS LEU- COMYSTAX). PARTLY SACCULAR, PARTLY DISSECTING ANEURYSM WITH OPEN THROMBOSIS AS INDICATED BY THE CLASS ROD. PIECES OF WORM FOUND IN WALL. COULD NOT OBTAIN WHOLE SPECIMEN, SPECIES UNDETERMINED. DISEASES OF THE BLOOD VESSELS 81 unwise to draw deductions as to their incidence or as to the possibilities of vascular dilatation in any given order. Horses have aneurysms occasionally, cows and dogs rarely, according to Rievel. I can, however, state that there has not occurred in our experience a large growing pulsating aneurysm in the aortic arch region comparable to the condition so well known in man. The literature to which we have had access gives a definite impression that parasitism of vessel walls is the most important factor in the causation of ectasia, and that simple non-parasitic arteriosclerosis is relatively unimportant. Two of our seven cases seem to have been free of parasites but the notes cannot entirely assure one of this. The distribution of cases at this Garden is found in Table 5. A dilatation of the first part of the arch in a seal to a size which might be described as an aneurysm caused us to make such a diagnosis, correctly enough from the size and shape of the vessel but possibly worthy of reconsid- eration in light of the fact that no damage to the wall was found. At the heart and in the descending arch the diameter measured 4-5 cm., while the first part of the aorta measured 7.5 cm. This great irregularity in width could not be found in other seals albeit this section of the arch is usually a trifle larger than its origin and descend- ing portion. The cava in seals is also large, but in this particular animal it measured 6 cm. across at the fiver where there is a normal dilatation. These two spaces are looked upon as normal reservoirs for blood during diving, but the case in question seemed to have excessive ''aneurysmoid" enlargements without mural disease. London reports an aneurysm of the aorta in a seal. (2) Aneurysms are not so common in birds, the incidence being in comparison with mammals as 1 to 2.2 Two seats are prominent for their development : the sinuses above the aortic valves and the first part of the subclavian ves- sels ; two of the six cases occupied the first position, two (2) Proc. London Zool. Soc, 1916. 82 DISEASE IN WILD MAMMALS AND BIRDS the second, while one other lay in the arch of the aorta, the last in its descending thoracic portion. Those developing- over the valves seem to arise from simple degenerative arteritis; those that occupy the other loca- tions are apt to be surrounded by plaques on the intima. Aneurysms in birds reveal by microscopy some trace of all the vascular coats and seem not to construct an adven- titia from surrounding areolar tissue. The veins have presented no peculiar pathology, except in tumors which will be taken up later. A Derby 's Tyran showed a phle- bitis and periphlebitis of the left subclavian vein, of mycotic nature, which led to death by rupture and hemorrhage. SECTION IV DISEASES OF THE BLOOD AND BONE MARROW The production and physiology of the circulating* blood seem closely similar in the two classes under considera- tion, although the anatomy is not the same in birds and mammals, variations also occurring within the latter group. Pathological responses follow comparable lines in that hemolyzing agencies, be they hemosporidia, absorp- tions from metazoan parasites or bacterial toxins, produce a reaction in erythropoietic centres, and positively chemo- tactic viruses call forth increases in the colorless elements. We have also observed a decrease of leuco- cytes in an Orang Utan suffering from influenza, a finding analogous to that in the human attack. There is, how- ever, a much less ready response on the part of birds to any leucocyte-stimulating influence, in this class the mononuclears seeming to bear much of the burden assumed by the myeloid cells of Mammalia or at least appearing on the stage very quickly so that any increase of the latter is overshadowed by them. Perhaps this apparent difference may be further explained by the greater number of colorless blood cells, structures which might be called the principal secondary defences of the body and constantly at the disposal of the organism, normally present in the birds' blood; they amount to 25,000 per cubic millimetre in birds, while in the mammals very few varieties have half this number. On the accom- panying Table (6) will be found a few differential leuco- cyte counts now known to us. It would seem, from a general observation of simple and infected wounds and from a few blood counts, that the response of leucocytes in the lower animals is greater than in monkeys and man. The ease with which animals 83 84 DISEASE IN WILD MAMMALS AND BIRDS I I. -11 || 1 ^1 ^1 •sajiqdouieoa •juao J3J •ajBapnu "inao -oqduiA'j ii^mg ■JU80 J3J •saBapnu O 00 10 c^ CO .-H CO 5B 8 00 JO t2 CO ^ 10 (N > 05 >o 00 § § s LO 10 10 ^ o — a gill ,0^ g-T &I X > §32=1 '3 C O « |fa-|ll «:b=l . fl 5 o S£^ bb ;^ ^ ,g O i::: fe: r^ C .^ ^ rt rt M Q H ^ O hJ o DISEASES OF BLOOD AND BONE MARROW 85 •" a « S-^ II i l^^l «;! S ^ « 2 llsfli c o o o ^ a ^ ^ SS 2 . g & S3 s:-5 "o a> o bo 02 '^ 73 O o o aj K^ o -S e.S -=5 1.2 •s "^ ^ c a) 0) 03 3 > 1^ -3 fl C bD Cm J ^ o.S fl CO C^ t> »o 05 CO 00 «5 o o M ■* 05 (M o CO ■* Oi CO T}H ■* to ■* ■* CO lO CO CO ^ o (N ■* ^ CO ■* »o o o Oi O 1^ iO (N rt^ 00 o CO TfH CO rf< »o Is c3 -S O it! ^ « 3 3 > JH 3 ^ C 2s "^ O 3 ^ Roo 3 o3 85 o 1^ ; o o'- -.^i^ -:iiSi a;oiitc-:T»OrtOoSo gCL, ACQ cph gon >Tnphatic hyperplasia is an exceedingly rare entity in wild animals. There has been no case resembling Hodgkin's disease of man. California Hair Seal {Zalophns calif ornianus) . Young 6 four months old. Appeared to be in good health, no loss of flesh or activity. Diagnosis. — Lymphomatosis. Fatty degeneration of kidneys. Em- physema. Chronic follicular and catarrhal enteritis. Nematodes in intes- tines. External appearance good. Both lungs distended and are tense; it seems almost like a spastic dilatation — a simple emphysema — beneath pleural surface are many minute petechias. Lung mottled red and gray. This color is present on section. Lobules clearly outlined and separable with fingers. Bronchi and vessels widely open, the latter containing small amount of fluid blood. Lymph nodes — Mediastinal, tracheal and those visible in neck are variously but definitely enlarged, firm, irregular DISEASES OF THE LYMPHATIC TISSUES 119 and roughly nodular, both in appearance and to touch. On section they are red and gray, follicular border not preserved and merging with medulla; connective tissue visible; gray juice expressed. Peri- cardium normal. Epicardium, glistening, transparent and pale. Heart position, size and interior normal. Abdomen contained no adhesions or fluid. Pale brown red, firm liver with smooth surface and sharp edges; markings not clear; section surface smooth; watery blood expressed from section. Gall-bladder normal, contents limpid, brown. Common bile duct patulous. Somewhat enlarged spleen, soft, yet resilient; capsule smooth ; section surface pulp homogeneous brown-purple ; folli- cles gray, slightly enlarged, clear, sharply outlined; trabecul^e faintly visible, more prominent where they are near follicles. Klidney normal in size, shape and location, with smooth, pale brown surface; capsule smooth, strips easily, section surface glistening, consistency soft yet resilient ; thickness of cortex slightly wide, of medulla normal ; individual lobules clear yet no increase of connective tissue between; pyramids quite pale; top of medulla a little darker; cortices, pale, striae quite faint and seem irregular ; glomeruli not visible. R. Adrenal — In upper pole are two cysts about 1.5 and .8 em. across ; no scar at this point. L. Adrenal — Wide pale purple, regular homogeneous cortex under very dense capsule; medulla reddish brown, homogeneous, quite bloody. In left adrenal is a pale gray, fairly well outlined area between medulla and cortex at upper pole about 3 mm. across. Stomach. — Contains glass and stones; mucosa, soft, smooth, flat, yellow and pink, translucent rugs ; cardia about normal ; pylorus, valve prominent but probably not hypertrophied. Duodenum — Beginning at pylorus and extending through to ileum where follicles commence, intes- tine contains small amount white, creamy material, serosa negative, mucosa smooth flat, pale pink, translucent. There are small nematodes, probably uncinaria, some of which are attached quite firmly. Here and there throughout the gut are bits of thickened mucosa or submueosa, 2-5 mm. across. Some of these have a tiny opening from which clear fluid can be expressed. Ileum shows smooth flat translucent mucous mem- brane ; empty ; individual follicles faint but discernible. Colon reddened mucous membrane, nematodes and little mucus. Pancreas, soft, normal in size and position, color pink gray. The lymphatics of peritoneal cavity are all enlarged; most of retroperitoneal are also. Regional lymph glands are also enlarged. The description given for mediastinal answers here. The glands of mesenteric stalk present chain of sausage-like masses. Glands in mesentery are but slightly affected. Glands within and without peritoneum at kidney are especially enlarged. Large gland behind stomach and pancreas and in front of upper end of spleen is much enlarged, thick and roughly circular; it shows much fibrosis. The regional glands are distinctly enlarged but proportionately not so much as the internal ones. Smear of juice expressed from peritoneal glands shows large and small lymph cells in about proper proportions. There are a few plasma cells. No granular cells. 120 DISEASE IN WILD MAMMALS AND BIRDS Smear from bone marrow shows enormous number of premyeloeytes and mast cells. Polynuclears in their early stages are not numerous. Small gi-oups of lymphocytes, lightly packed together, noted here and there. Blood smears unsatisfactory. JMicroscopical. — Lung shows distended vesicles with considerable congestion of septa and in some places red blood cells in alveoli. Kidney. — Glomeruli contracted, relatively anemic ; capsule negative ; epithelium throughout in state of granular degeneration probably fatty ; connective tissue not increased. Intestines. — Show slight hyaline change in muscle fibres and nodular degeneration and disappearance of protoplasm, leaving bare nuclei and outline of the cells; submucosa loose; mucosa shows infiltration of round and plasma cells, diapedesis, degeneration and desquamation of epithelium. Section of ileum shows a hyperplasia of lymph follicles with active centre in which large endothelial cells pre- dominate. Small swellings in submucosa consist of chronic granulation tissue, but there is no foreign body and no evidence of preexisteuee of a lymph follicle. Lymph nodes loosely arranged but in places solidly small lymphoid ; chords and sinuses contain chiefly small lymph cells in which are some mast and plasma cells; follicles small and loose, and centres contain chiefly large lymph and plasma cells; connective tissue not increased yet there are some fibroblasts in follicles. Spleen. — ShoAvs almost entirely hemoiThagic pulp ; blood destruc- tion not active; follicles large and made up of about equal mixture of small lymph, plasma and large lymph cells ; connective tissue promi- nent probably because of excess of blood around it. Local Hyperplasias. In this group and touching the purely hyperplastic, come the hypertrophies of the lymphadenoid tissue of the nasopharynx ; this can be disposed of by stating that no true chronic hypertrophies have been seen. With most cases of gastroenteritis in ungulates, and many in car- nivores, there is a swelling and redness of the tonsillar region but an exudative or necrotizing process does not occur. Thymus. The thymus is rarely visible in our specimens but when found has usually been normal. An increase in size and a decrease of consistency of this organ is noted in marasmus (inanition) from failure of care of the young DISEASES OF THE LYMPHATIC TISSUES 121 by the parent, but so far as can be determined this has not been adequate to cause tracheal compression. In a few cases of rachitis in the canines, the organ is large and pale along with the rest of the lymphatic system. In one of the tumors of the mediastinum to be discussed, the suspicion arose that the growth originated in the thjTiius ; adenomata and sarcomata have been described in the lower animals. Tuberculosis. Tuberculosis of the superficial lymph glands is rare as an independent lesion in the lower animals. Dr. C Y. White was fortunate in seeing a monkey with a chain of fibrocaseous nodes in the cervical region, upon which before death he offered the suggestion that it was of tuberculous nature. In Primates almost all drainage glands exhibit some miliary or caseous process. One monkey rejected upon the tuberculin test had what was apparently a primary lesion in the glands at the tracheal bifurcation. In the Ungulata, lymphatic tuberculosis assumes two forms, the caseous and the cellular. The fonner is generally understood while the latter is more uncommon. It is occasionally seen in the ''fungous tubercle" of cows, but we have seen it in deer and in another order, Carnivora. In the latter, tuberculosis being uncommon, examples in the Ijmiph nodes were noted but twice, once caseous and once solid ; this latter was made up of firm, homogeneous yellow pink masses of glands in the mediastinum, shomng under the microscope solidly packed epithelioid and giant cells. Lymphatic tuberculosis in the birds is rare ; only one case is recalled (unfortunately record cannot be found) as small yellow, discrete firm nodules in the mediastinum and neck. The minute picture was of a solid arrangement of large vacuolated mono- and polynucleated cells which were so packed with bacilli that the preparation could not be decolorized. 122 DISEASE IN WILD MAMMALS AND BIRDS TUMOES. The neoplastic enlargements are represented by a IjTnphosarcoma of the mediastinum in a Dorcas Goat; there were secondary growths in the liver, the kidney and several isolated lymph glands. Secondary growths from original tumors not in the lymphatic system are curiously rare, certainly much less frequent than is found in the human being. In ninety-two tumors which are known to give metastases, only three involved the lymphatic glands and only one of these could be considered as involving nodes not in the ordinary drainage pathway. The Spleen. The spleen is an organ whose exact position in the scheme of things remains mystifying through the whole series of animals. Its functions have been arrived at largely by exclusion, somewhat by conjecture and specu- lation, while the acceptably proven duties are indeed few. Although this is not the place to enter into a discussion of all the points at issue concerning the anatomy and func- tion of the organ, a few facts might be mentioned of comparative and perhaps pathological value. While the spleen has been looked upon always as the origin of blood cells in the embryo, cases in man are on record where no spleen was discovered, a condition suggesting that its absence is not incompatible with life ; that such is the case is obvious for experimental or therapeutic removal of the organ is well known. It was suggested by Virchow that at times the spleen assumes the duty of the bone marrow and the swelling of the viscus in certain anemias seems to support this state- ment. However, there is no physical relationship between the size of the animal and that of the spleen, nor between the available quantity of bone marrow, the obvious rich- ness of circulating blood and amount of lymphatic tissue. This is well shown in the water mammals which have a great deal of blood in vessels and viscera yet their spleen DISEASES OF THE LYMPHATIC TISSUES 123 is relatively smaller than birds of somewhat comparable size (the ostrich, for example). The Carnivora and Eodentia among the mammals, have quite large spleens proportionately, while comparable birds, Accipitres and Striges, have relatively small ones, and an absolutely small amount of marrow. Another anomalous example of the function of this tissue is found in the hematopoiesis of birds. While there seems to be some evidence that in the spleen and liver red blood cells may be formed, there is perhaps at most times sufficient bone marrow to keep the blood cells at a proper number. There is no inverse relation between the amount of bone marrow and that of splenic bulk ; that is in diving birds with their hollow bones the latter is no greater than in gallinaceous birds which have abundant marrow in all the skeleton. It has been suggested that the spleen supplies an activator for pancreatic ferments. This receives a sort of support from the greater size of the organ in carnivores and smaller relative size in ungulates, since in the former concentrations of digestive enzymes are more often needed. Because of the greater excretion of iron in splenectomized animals or those being starved, Fischer has thought that the spleen has some power to metabolize this element ; this receives some support from the state- ments concerning the size of this organ in the carnivorous orders. From experimental studies it appears that some relation exists between the lymphocytes and resistance to implanted tumors. If one apply this idea to the amount of lymphatic tissue and the incidence of spontaneous new growths, it is found that no definite relationship is discov- erable either in terms of size of spleen or richness in lymphatic nodes. Carnivores, rodents and marsupials show a high tumor incidence and have a good lymphatic supply. Aves, on the other hand, with a poor nodal appa- ratus and a variable splenic bulk, show many fewer 124 DISEASE IN WILD MAMMALS AND BIRDS tumors than do the mammals. The mononuclears of avian blood rise normally as high as 60 per cent., of which 40-45 per cent, are of the small size. The size of the spleen is subject to great variation not only within orders but actually within genera ; it even seems that one sometimes sees two or three members of a species kept in the same enclosure, maybe with the same disease, maybe without any obvious disease, yet with definite variation in the size of the organ. This irregu- larity is recognized by veterinarians (Hutyra and Marek) who ascribe it to some unknown disease, past or present, and to normal variation. The lack of uniformity is seen more clearly in birds than mammals. In the former it might be due at times to low grade or inactive parasitism or, conjecturally, to toxins from incorrect diet ; we shall see later that infection and intoxication seem to have a different effect. Some writers have mentioned the possi- bility of an idiopathic splenomegaly (?), a condition associated in youthful human beings, with anemia and lymphadenopathy, and occasionally going over into a sort of leucemia. The existence of such a condition is difficult to admit or deny; we have met nothing which could not be aligned in some fairly well defined group. Birds have a relatively larger spleen than mammals and in addition the organ seems to respond more actively in infections or intoxications, since it may reach, under active stimulation, a size ten times that of the nor- mal organ. Enlargements of Spleen. Acute enlargements of the spleen, be they of con- gestive or ''inflammatory" nature are quite common among the lower animals, a fact that is recognized in veterinary medicine. Congestions of the spleen are most often seen during acute infections and diseases of the heart and liver. The diagnosis has been made among DISEASES OF THE LYMPHATIC TISSUES 125 mammals proportionately more than in birds, 27 or 1.5 per cent, to 35 to 1 per cent. Eeason foe Congestive Enlakgements. Perhaps this is partly due to the circulatory anatomy since in the former the arterial and venous supply is usually by one large vessel of each kind, whereas in birds the splenic branch of the coeliac axis breaks up into several small arteries and the venous return is accomplished by numerous venules some of which reenter the posterior cava almost directly, others joining with the mesenteric to form the portal; by this arrangement a more elastic system is assured. The same condition is found when analyzing the records of chronic passive congestion; in eighteen recorded cases, fourteen were mammals and four birds. In these cases the principal associated pathologi- cal lesions were pleural and pulmonary in seven, cardiac in three, renal in three and hepatic in four. While the anatomy of the splenic blood supply may help to explain the small number of congestions in Aves, it will not answer for the inflammations which occur in large numbers in this class; this will be discussed in the following pages. Hemorrhage and infarction of the spleen are not very common, there being eleven of the former and ten of the latter and all occurring with the same indication or history of infectious disease; in one case, an opossum {Didelphys virginiana) an injury probably caused a mas- sive hemorrhage shaped like an infarct. It is, however, curious that of the twenty-one cases only five occurred in birds,, of which only one hemorrhage was in the shape to which the term infarction is best applied. There have been, as one might expect, a few cases of infected infarc- tion, with abscess. It is perhaps worth noting that no case of hemorrhagic cyst or inspissated coagulum has been seen. Only one case of rupture is recorded second- ary to acute splenitis during an acute septicemia. 126 DISEASE IN WILD MAMMALS AND BIRDS Inflammations. It has been the practice at this laboratory to classify splenitis in three ways, (1) acute diffuse splenitis — general congestion with perhaps small hemorrhages, swelling but retention of general architecture and mth no undue prominence of the follicles; (2) follicular hyper- plasia — where this is the prominent gross and minute finding, the pulp being less pronouncedly involved; (3) acute splenic tumor — where the organ is greatly enlarged but with loss of the usual markings. Perhaps this separation is not warranted upon a strict etiological basis, and yet as we shall see it holds true fairly well in the toxic and infectious lesions.. Moreover, from the fol- lowing list it is evident how the various orders call upon the splenic tissue in disease. Table 7. This Shows the Percentage of Various Forms of Splenitis in the Animals Coming to Autopsy. Acute Diffuse Splenitis. Per cent. Follicular Splenitis. Per cent. Acute Splenic Tumor. Per cent. 7. 2.5 2.4 2.6 9.2 SS. S3. 12.5 3. 6.5 5.2 1. 2.6 10. 7.5 4. 6.8 3. .7 3. 4.9 .3 4.7 3. 3. 2.3 7.2 6. 1.5 10. 3.7 3.4 6. 10. .7 1.3 2. 1.2 1. Marsupialia Pinnipedia 3.7 Proboscidea. Edentata 6.2 Passeres 3.8 Psittaci 4.1 5.2 2. Galli 4.7 TT Accipitres 1.6 GaviaB 5. Picarise 1.3 Striges FulicarisG Stecanopodes For the meaning of italics see foot note Table 1. ' The points to draw from this chart are first the greater variety of lesions seen in the birds, which appar- DISEASES OF THE LYMPHATIC TISSUES 127 ently make greater use of the organ in defence, and the preponderance of acute diffuse hyperplasias in mammals. It is noteworthy that the Primates and Marsupialia show more of the acute hypertrophy of the spleen going under the name of tumor. It would be interesting and valuable to be able to discover exactly what determines and con- stitutes the conditions usually termed hyperplasia and inflammation of the spleen and an attempt was made in this direction by tabulating the data from Table 7 in terms of each diagnosis, classifying these latter also as infectious and toxic. The results of this effort are not conclusive, and while they permit of some discussion of the lesions, do not allow finished conclusions. The figures obtained by study are not illuminating. One can state, however, that in infections, either specific or not specific, more elements of the spleen were engaged in the process than when the condition did not resemble a communicable disease, but might be called toxic. Under the former condition the diagnosis of splenitis or acute tumor pre- dominated, while under the latter follicular hyperplasia is more often recorded. Anatomical alterations in these two groups are described in the definition given in a previous paragraph and deserve no special discussion except in so far as they concern the changes in the avian splenic tumor when under magnification. Microscopically there is a total loss of the relations in the acute splenic tumor of birds, the chords and follicles being replaced by a rather regularly arranged mass of small and large mononuclears, granular cells of the circu- lating types and red blood cells ; pigment always seems increased. Endothelial cells do not take part in the general mass of the organ but along lymph and blood vessels their bulk and number are increased so that if the packing be not too dense one may find double lines of these cells passing through the hyperplastic tissue. :There is, however, a group of seven cases (six birds) to which the term endothelial hyperplasia in the spleen has been 128 DISEASE IN WILD MAMMALS AND BIRDS applied. All of them show an unusual prominence of the vascular linings and of the follicular centres and peri- vascular areas ; besides this there is a moderate general hypei*plasia. An examination of the historj^ and autopsy results in the birds indicated that they had all suf- fered with some rather protracted intoxication and showed a moderate anemia (four of the spleens were well pigmented). The more chronic changes of this organ, be they moderate or of a grade to which one must apply the term fibrosis, are rather uncommon when one considers the number of animals with prolonged infection, anemia, skeletal diseases and hepatic cirfhoses ; these are the con- ditions that take a prominent place in the associated pathology and history. There is no essential difference in the organ throughout the animal kingdom, and one may find as far down as the struthious birds analogies to the processes of domestic and human animals. Special. Subjects — Amyloid. Amyloid changes in the spleen have occurred in the follomng orders: Primates, Carnivora, Rodentia, Marsupialia, Passeres, Galli, Gaviae, and Anseres, fourteen cases in all. This infiltration is usually ascribed to long continued suppuration or chronic infection,! every one admitting, however, that once in a great while a case is seen wherein no cause can be dis- covered. In this laboratory we have a high percentage of cases without adequate accompanying pathology so that we have called six of the above cases primary or idiopathic. The gross appearance of the spleen is in all these cases that of an enlarged, firm, homogeneous body without the sago spots usually described for this change. \ Microscopically the infiltra- tion occurs first in the vessel walls, thence spreading to the sinus walls and reticulum. Joest in discussing this DISEASES OF THE LYMPHATIC TISSUES 129 infiltration (1), states that it is comparatively rare among the domestic animals, where it may appear mider the usual conditions or as a primary affection; it seems at times to follow unsuitable feeding. According to this author, the ''sago" type is more common than the ''bacony;" this has not been our experience. The most conspicuous example is given here. European Badger {Meles meles) Adult 9. Gradual failure for two weeks. Diagnosis. — Amyloidosis. Chronic parenchymatous nephritis. Emphysema of lungs. Muscles atrophic; fat scanty. Heart is dilated and muscle is red brown. Aorta is jaundiced and there is a small patch of atheroma near the anterior leaflet of aortic valve and about 5 mm. above it. Abdomen contains a slight amount of clear fluid, no adhesions. Liver is normal in size, smooth surface, sharp edges, firm consistency, yellow color. The section surface is glistening, smooth and moist. The spleen is very hard, greatly enlarged (25 x 6 x 2 cm.) and lies across the abdo- men over the intestines. It is pale red and very bacony. In iodin the tissue stains a mahogany brown. The kidney is normal in size. (5^2 x 3 cm.). The capsule is smooth and strips easily leaving a smooth, yellow surface. Organ is firm. The section surface is glistening, has a relatively narrowed cortex and relatively wide medulla. The glomeruli are barely visible but stain a mahogany brown in iodin. The adrenals are 12 xlO x 4 mm. The cortex is wide, dull yellow and regular. The zone beneath fades into the cortex although rather abruptly. The centre of both glands is occupied by an irregular pale yellow, sharply outlined nodule suggestive of solid medulla, tubercle or tumor. The zone between cortex and this is gray, irregular and firm. This is practically a normal figura- tion. The stomach is empty, mucosa apparently normal. Intestines were not opened but serosa seems normal ; when opened after Kaiserling fixation they seem normal. The pancreas is normal in size, firm and pale. Lymphatics of the mesentery are slightly enlarged, soft, homo- geneous pale yellow. Histological Notes. — Lung seems somewhat atrophic and there are some vesicular ruptures. One small patch of amyloid found in blood vessel wall. Liver shows slight capillary congestion and granularity of parenchyma. There is a marked blood vessel amyloid deposit not only in interlobular spaces but in intralobular capillaries. Spleen, no trace of splenic tissue recognizable in section. It is composed of more or less eosin-staining material surrounding single, or small collections of round or plasma cells. Kidney shows marked amyloidosis of glomeruli and slight deposit in blood vessel walls and in the increased connective tissue. There is a general moderate fibrosis; irregular tubules; low (1) Lubarsch-Ostertag, Ergeb. aus der Allg. Path., 1908. 130 DISEASE IN WILD MAMMALS AND BIRDS epithelium and hyaline casts. Heart muscle fibres are small and stain deeply. No amyloid in section. No pigmentation although section is suggestive of brown atrophy. Adrenal is practically negative. There seems to be slightly more connective tissue than noimal but parenchyma may be considered normal. At one place in the cortex there are some structures of deep layer included in vesicular layer. This seems like a structural malformation. Necroses. Focal necroses of the spleen affecting chiefly follicular centres but also chords, are not at all uncommon in avian spleens, especially where parasitism occurs, not only with hemic protozoa and embryos, but also ^\ith intestinal or visceral nematodes and trematodes. Spleen in Anemia. The spleen in the anemias shows much less definite change than one would expect. In the secondar^^ form of anemia among mammals one finds a slight excess of pigmentation and an occasional fibrosis but often the size of the organ is recorded as normal. When the impover- ished blood seems secondary to skeletal degenerations there is a diffuse or follicular enlargement. In the birds, on the other hand, there is nearly always some grade of enlargement which is due in the well studied examples to a richness of blood cells. There may be a slight increase in follicles, but these bodies are usually small and solid. In two instances a prominence of large endothelial cells was discovered. Pigment is seldom increased, but it may be very excessive. In the primar}^ anemias little more than the above is to be found. Fibrosis is more evident and perhaps pig- mentation less so, but the variations are more of degree than kind. The spleen in leucemia can only be discussed upon the case already reported {q. v.). Spleen in Hepatic Fibroses. The fibroses of the liver are not infrequently associ- ated with some enlargement of the organ under discus- sion. At this laboratory hepatic cirrhoses are divided Fig. 7. — EUROPEAN BADCl.R IM1.I.1.> Ml.l.L.i). PRIMARY AMYLOIDOSIS. THE ENOR- MOUSLY ENLARGED SPLEEN IS SEEN LYING ACROSS THE STOMACH BELOW THE LIVER, WHICH ORGAN IS EXCEEDED IN SIZE BY IT. DISEASES OF THE LYMPHATIC TISSUES 131 into Portal, Biliary, Fatty, Perilobular and Vascular. By a study of the spleens in these cases a few facts have been obtained. In mammalian portal cirrhoses there is usually a very moderate but definite increase in the size of the spleen due to connective tissue increase in the trabeculae and reticulum, with very small compact fol- licles. Among the birds the organ seldom shows more that a moderate congestion and diffuse hyperplasia. In the mammalian biliary cirrhoses there is almost without exception a definite enlargement due to fibrosis and fol- licular hyperplasia. Among the birds the process is very far from uniform, there being just as often no change, as a congestion and hyperplasia, or as a mild fibrosis ; it is notable, however, that pigmentation is commonly met in this class. No noteworthy change is met \vith in the spleen of fatty hepatic cirrhoses. We have no record of cyanotic induration of the spleen accompanying a similar condition in the liver. When the condition of perilobular hepatic fibrosis has been met a distinct increase of the splenic pigment is usually found. In reviewing the facts in this paragraph one is forced to the conclusion that, with the exception of the frank infective cases, there is no definite relation of the splenic changes to those in hepatic cirrhosis, a fact made perhaps the more significant in view of the idea held in some quarters that the primary change in this pathological process occurs in the spleen. Perisplenitis. Perisplenitis of an acute or fibrosing variety as a part of peritonitis or as the result of an injury to the splenic region, has been encountered on numerous occasions and offers nothing worthy of comment, but the so-called ''sugar-icing" spleen, associated with perihepatitis and general peritoneal thickening, and with an obscure rela- tion to tuberculosis has not been seen. As we shall learn later peritoneal or massive intestinal tuberculosis is not 132 DISEASE IN WILD MAMMALS AND BIRDS common in the lower animals. A diffuse fibrous thick- ening of a pearl gray glistening appearance, has occur- red in five monkeys harboring the Filaria gracilis in the peritoneum. Tuberculosis. In tuberculosis of this organ the capsule is almost invariably thickened over the nodules, but not over the whole organ and rarely in the diffuse variety. There is almost always, however, in this prolonged infectious dis- ease some reaction on the part of the spleen, particularly when several other viscera are involved. In such cases a low grade of general hyperplasia and fibrous tissue increase is found. This is especially true in the Primates and Carnivora in long continued pulmonary tuberculosis, and it is in this form that one does not often see tubercu- lous lesions proper in the spleen itself. Tuberculosis of the spleen is not very common in adult human beings and domesticated animals, while in the young it is seen in a fairly high percentage of cases. In the wild animals this organ seems much more susceptible to the settling of tubercle bacilli as will be attested by the facts to be cited, and yet there are some conspicuous exceptions to this statement. Just why certain groups, or orders should show splenic tuberculosis very frequently while others fail to do so is far from clear. The appearance of the lesion is fairly comparable throughout the animal kingdom, variations among the Aves being chiefly in the more frequent occurrence of the diffuse tuberculous splenitis. Miliary, caseous and conglomerate masses occur separately or together and without very distinct relation to other pathologi- cal involvements. The relative incidence of tuberculosis in the spleen is set forth in the section devoted to this infection, but may be mentioned here in a general way. The organ is about equally susceptible to the disease in mammals and birds DISEASES OF THE LYMPHATIC TISSUES 133 according to the figures, but the high percentage for the former is due to the frequency with which tuberculosis is met in the Primates. With this order deducted there is no doubt that the avian spleen has a greater vulnerability for the tubercle bacillus than has the mammalian organ. Rodentia as a mammalian order stand next to the Primates, whereas the Camivora and Ungulata relatively seldom show splenic tuberculosis. Among the Aves one finds that Columbae, Galli and Accipitres have the highest splenic susceptibility, but beyond this one hardly dare venture because of the irregularity in the number of specimens seen at autopsy. Suffice it to say that a greater number of cases of tuberculosis are seen in the spleen of the class for which the intestinal origin of tuberculosis appears most important. In the cases of actinomycosis and its congeners, such as the Kangaroo maxillarj^ mycosis, which we have seen at the Garden, no specific changes have been seen in the spleen, unless a single or double nodule of the same variety as the original focus may be considered specific. Such was found in two cases, one a tapir Avith low grade inactive actinomycosis, the other a kangaroo with maxillary strep- tothricosis. Two deer came to autopsy with a gross picture suggestive of anthrax; no bacilli could be found in the semifluid spleen, but a member of the hemorrhagic septicemia group w^as found. The latter group of infec- tions has been fairly well represented, although not in epizootic form, and the almost invariable splenic change has been that already discussed as diffuse splenitis. Three secondary tumors are recorded, two cancers and one renal adenoma. SECTION VI THE RESPIRATORY SYSTEM AND ITS RELATED STRUCTURES It is customary to divide the descriptions of normal and diseased conditions of the upper entrance to the body into respiratory and alimentary parts, the nose, naso- pharynx and larynx belonging to the former, the mouth, buccal cavity and phar^mx to the latter. As a matter of fact they can for most purposes be considered as the structures contained in the anterior head and furthermore their pathological states are more often followed by exten- sions into or implications of the respiratory organs proper than of the alimentary tract. From a comparative standpoint the incidence of specific infectious diseases and of the involvement of accessory nasal sinuses present the most interesting subjects. There are several infections, believed to be specific, observed among domestic mammals and birds but their actual indi\dduality has hardly been unexceptionally proved. This refers to the communicable rhinitis of cows, pigs, rabbits, and birds, especially parrots, the follicular catarrh of horses, and croupous nasopharyngitis, all of which have been ascribed to a par- ticular virus, without finished evidence in many instances. Some of these diagnoses doubtless cover or are confused wdth the early sj^mptoms and signs of the disease of pro- tean manifestations, distemper, and indeed the Bact. septicus and relatives of the bird cholera organisms are reported as being responsible for them. No intention of excluding well recognized entities like bird diphtheria, foot and mouth disease or influenza, exists. I shall refer below to small groups of epizootics which do not conform strictly with word pictures drawn by Hutyra and Marek, Moore, or Ward and Gallagher. 134 THE RESPIRATORY SYSTEM 135 Rhinitis, Sinusitis. The nature of inflammations of the nasopharynx sug- gests at once that there may be some anatomical reason for their distribution and character. A general review of the anatomy of the mammalian and avian nasopharynx reveals the relatively greater space in the former, espe- cially in the passage from the nose to the pharynx, and emphasizes the exposure of the opening of the upper larynx in the bird, lying as it does in the posterior part of the tongue and surrounded by the constrictor es glottidis. A dissection of the accessory nasal sinuses exposes the relatively large size of these spaces in the lower mammals, and the capacious openings into the nasal cavities. (1) In the Primates and Lemures the anatomy more closely resembles that of man, the sinuses being relatively smaller and the communicating passages narrower. In the bird on the other hand, while the sinuses may be extensive in some they are usually small, yet in all the communication with the turbinate area is by a narrower slit or tortuous canal, frequently, as in Galli, running from below upward into the maxillary sinuses. The extent of the turbinate and the richness in mucosa is probably greater in all mammals than in birds ; certainly this seems true of Carnivora and Ungulata versus Accipitres and Galli. If the seriousness of a rhinitis be dependent upon the extent of involvement of the sinuses and the blocking up of their outlets it would be expected that the variety of animal having the smallest drainage channels would show the greatest evidence of these diseases. Our records would indicate that 32 birds (.96 per cent, of the autopsies upon Aves) had rhinitis whereas only 7 mammals (.39 per cent, of autopsies on this class) presented the con- dition. Extension to the sinuses occurred in only one- third of each of these figures, a complication which in turn produced generalized infection more often in (1) Vide Sisson's Veterinary Anatomy and Owen's Anntomy of the Vertebrates. 136 DISEASE IN WILD MAMMALS AND BIRDS maminals than in birds as 4 is to 3. These figures are perhaps too small for conclusions but it would seem that rhinitis occurs more often in birds with their small sinuses and channels wliile sinusitis and general infection occur more often in mammals with their large sinuses and exten- sive turbinate apparatus. The most conspicuous orders represented are, in line of numbers Anseres 12, Psittaci 7, Accipitres 5, Carnivora 3. Nine of the twelve water- fowl were part of an epizootic which mil be discussed under specific diseases. Baeteriologically the mammalian cases that have been worked out were due to Streptococcus pyogenes in several instances, including the generalized cases, and to a mixture of streptococci, golden staphylococci and mem- bers of the colon-aerogenes group. In one case in a tapir a member of the B. septicus group was found. Moulds were discovered in three avian cases and filaria in one. No pentastomum or oestrus has been discovered. In thirteen instances the lungs have been involved, appar- ently secondary to the nasopharyngeal disease. There have been two small outbreaks of an acute non- specific infection — that is not suggestive of cholera, psit- tacosis or infectious enteritis — among the parrots in which during a short time 4 and 6 parrots died mth naso- pharyngosinusitis as the prominent lesion. One outbreak was studied baeteriologically without definite result. There was no uniform internal pathology unless, in one outbreak, congestion of the cerebellum may be mentioned. Fowl diphtheria and its associated condition from which a satisfactory separation has not been accomplished, epithelioma contagiosum, has happily given us little con- cern, so that it is not possible to record any instructive facts upon its cause or differential diagnosis. There was recorded in the 1911 Report of the Society the occurrence of two fatal cases in cassowaries from which it was pos- sible to isolate the'B. columbarum and one bird with the same clinical appearances whose recovery seemed to be THE RESPIRATORY SYSTEM 137 due to the use of human diphtheria antitoxin. In light of more inf onnation and consideration of the accepted varia- bility of this disease, it is possible that tliis bird may have recovered without the injections or with the use of normal serum. Nowadays it is possible to obtain antiroup serum which is stated by Blair of New York to be efficacious. The disease has been observed in a wild turkey and an Abyssinian Ground Hornbill, beside the two struthious birds mentioned above. Mycosis. Mycotic disease of the nasopharynx seldom restricts itself to this cavity, usually extending by continuity to the esophagus, or by inspiration to the lungs whence it spreads to the air sacs. This condition of the upper passages has occurred here only in Psittaci and Accipitres although it is reported by veterinarians as occurring in Anseres and Struthiones. In the first order four birds were affected, two showing extension to the esophagus and a like number having pulmonary and serous mem- brane involvement. These cases were all due to asper- gillus whereas those next to be mentioned were caused by an oidium close to the ' ' albicans ' ' variety. In four Mississippi kites the prominent changes were found in the pharynx and esophagus down as far as the proventricle with only a few rather trifling lesions in the nasal area. Infiltrative and necrotizing processes characterized the action of the oidium wliile that exerted by the aspergillus was more superficial and extensive. In one case of a parrot the whole nasal ca\ity was completely filled with a yellow gray exudate whereas the esophageal wall of the kites was thoroughly infiltrated by a gray brown, friable, necrotic mass. Attempts at treatment were made in the case of the latter, using potassium chlorate and saline solution on cotton swabs. The result was entirely nega- tive and the applications seemed to have no effect upon the course of the infection. 10 138 DISEASE IN WILD MAMMALS AND BIRDS There are on our records in addition to the above, several cases of necrotizing processes about the head seeming to emanate from wounds to the mucosa by foreign bodies, by decomposition of pieces of food in cre^dces or by damage by masses too large to be swal- lowed. In the few instances where we have tried bacteri- ology, no definite result has been obtained unless the frequent occurrence of organisms bearing a resemblance to Bact. necrophorus be important. Tliis organism how- ever may be found in many necrotic processes in animals ; I do not look upon it as specific in the locations just cited. Mammals as a class do not present many inflammatory conditions around the anterior head, aside from the specific diseases like distemper (?), actinomycosis. Kan- garoo disease and the like. Monkeys occasionally have acute coryza, which may indeed seem transmissible to others but it seldom leads to any serious consequences and is untreated, except by segregation. Tuberculous lesions are not recorded. There has been no glossitis aside from lesions involving the pharynx. The tonsils have been discussed under the lymphatic apparatus and it only need be repeated here that inflammation and hypertrophy of these organs are exceedingly rare. Specific or individual diseases of the salivary glands are also rare although these organs may be involved by extension. Tliis gen- eral region is not often affected with tumor, unless the jaw be included which bone is the seat of several tumors in antelopes and opossums. Aside from these we have seen an epithelioma of the tongue in a black bear {TJrsus americanus) . Larynx. The larynx is an organ of fairly uniform construction through the mammalian orders but is conspicuously dif- ferent in the Aves where it is double. The upper end of the trachea, in the latter class is surmounted by a cartilag- inous box lying beneath the root of the tongue through THE RESPIRATORY SYSTEM 139 which an anteroposterior slit-like opening forms the glottis ; there is no epiglottis. This is only an air passage, the voice being made in the syrinx or lower box which lies at the bifurcation of the trachea. The structure of the upper box is quite simple with its lateral plates controlled by the glossal muscles and two external retractors but the syrinx is very complicated and variable in the dif- ferent orders and even in the same family. It possesses an internal and external set of muscles and in some birds can be opened at one point to permit air to pass to the cervical or thoracic air sacs. Detailed discussion of its anatomy is hardly profitable since there is nothing peculiar about its diseases. On one occasion only have we seen distinct pathological change — what was probably an extension of mould disease from it to the cervical air sac. It is involved in true tracheitis and bronchitis but even these are rare in birds. The larynx on the other hand is constantly reddened in cases of pharyngitis and may be the seat of mould colonies. Edema of this structure is, however, not very common, it being recorded but twice in birds in associa- tion with nearby inflammation and five times in mammals ; in the latter cases three were of acute infectious nature, one was a tumor and the other osteomalacia. It is common to find the laryngeal and tracheal mucosa swollen and wet in chronic bone degenerations without the con- dition being severe enough to call it edema. Laryngitis. Acute laryngitis of active catarrhal or purulent nature has been met five times in mammals and twice in birds while more chronic lesions have occurred only in the former, four times. Tuberculous laryngitis has been observed in a cockatoo and a lemur; they are interesting enough to cite. There are no cases recorded among monkeys despite the large number dying from the disease ; this implies of course that no suspicion of its existence 140 DISEASE IN WILD MAMMALS AND BIRDS was had at postmortem but perhaps some would have been detected had every larynx been subjected to micro- scopic section. A citron-crested cockatoo was found when posted to have general miliary tuberculosis. The bright red rim of the glottis attracted attention and upon slitting open the organ, pinhead size, sharply outlined yellow tubercles were found on both sides. A black and white lemur was killed because of a positive tuberculin test. He was in good condition and exhibited as his only lesions retropharyngeal lymph nodes with precaseous miliary nodules and small miliary tubercles on the epi- glottis, true and false vocal chords and in the mucosa of the main ventricle, each lesion being surrounded by a narrow sharply injected zone. This seems like a recent double implantation since the retropharyngeal glands probably do not drain toward or from the larynx. The larynx has been the seat of only one tumor, a squamous cell cancer in an Azara's agouti. The tumor caused ulceration and edema of the whole mucosa sufficient to produce fatal asphyxia. The trachea is of relatively little comparative or pathological interest aside from its inflammations which however are so closely associated with bronchitis that they will be included under that heading. Perhaps the most important condition of this tube is its infestation with Syngamus tracliealis since this leads to inflam- mations not only of the related mucosa but predisposes to pulmonary infection. The occurrence in the Galli is well kno^\Ti but perhaps it is not so well recog-nized that this worm occurs also in crows (Passeres) and swans and geese (Anseres). For the diagnosis of this condition it is customarily stated that a frothy mucus in the mouth is very suggestive; this is true in the cases seen here but in addition a mucopurulent stomatitis is exceedingly common and when the two are combined the picture is almost confirmatory. Although worms are credited with considerable weight in the production of pneumonia in THE RESPIRATORY SYSTEM 141 Ungulata, they have only been seen once within the tracheal tube. The Beonchi. The bronchi will be discussed as a separate part of the respiratory system in so far as possible since they present a very decided difference in anatomy between mammals and birds and because the incidence of their disease is other than will be found for the lungs. However, distinc- tion has been made between changes in the grosser tubes and those in the finer bronchioles, especially because capillary bronchitis so-called is really a pneumonitis in which the mucosa of the larger passages need not par- ticipate. The mammalian tubes are not really greatly dif- ferent in their construction, passing through ever smaller branchings which give an increasing square area of tube capacity and more extensive mucous surface. The avian main bronchus breaks up very shortly after entering the lung into a varying number of spaces lined with low epithelium lying upon a fibrous support and without cartilage. These spaces then open into secondary air spaces of a size visible to the naked eye which are in turn surrounded by microscopic alveoli. The largest spaces, first mentioned, continue to grow smaller toward the lower part of the lung where they usually communicate with one or other of the various ostia of air sacs. Bron- chial diseases in birds must therefore be limited at the place where the bronchi lose the cartilaginous rings since below this the surface functionates as pulmonary tissue. Beonciiitis. The accompanying list. Table 8, will show the distri- bution of bronchitis not accompanying pneumonia or due to mycosis. It is striking that carnivorous animals are more prone to bronchitis than any other order (the struthious birds are too few to be important). There is a very decided preponderance of mammalian cases over a\4an, there being not only more cases but proportion- 142 DISEASE IN WILD MAMMALS AND BIRDS ately more orders affected. The character of lesions in the mammals is nearly always catarrhopurulent or freely purulent while ulcerative changes are not uncommon. Peribronchial infiltrates are seldom found without some Table 8. Showing the Percentage Incidence of Bronchitis and of Parasites in the Autopsies upon the Various Orders. Orders Simple Bronchitis Verminous Bronchitis and Pneumonitis Inactive Parasites of Lung (Encysted) &c. Cases Per cent. Cases Per cent. Cases Per cent. Primates 7 14 5 3 4 4 1 1 2 1 1.4 2.9 1.3 1.7 .29 .58 .5 1. .67 3.3 3 8 2 2 2 7 1 3 .6 1.6 1. .5 1.1 .5 .3 1. 5 1 8 10 2 3 2 1 1. Lemures Carnivora Insectivora Chiroptera Rodentia Ungulata Proboscidea Hyracoidea Edentata 1.2 1.6 2.7 12.5 Marsupialia Monotremata Passeres . 1.7 .16 Picariae Psittaci Accipitres .GaUi Hemipodii Fulicariae. Alectorides Limicolse Gaviae Steganopodes Herodiones Odontoglossse .... Palamedes Anseres 1. Struthiones For meaning of italics see footnote Table 1. evidence of pneumonia ; nor is it common to meet the pale lines extending from bronchi between the lobules, such as are seen in human streptococcal disease. Avian bron- chitis is usually hemorrhagic or catarrhal and with exceeding rarity becoming purulent ; when this occurs the THE RESPIRATORY SYSTEM 143 cause is frequently found to be tuberculosis or mycosis. Inflammation of the larger passages is nearly always accompanied by pulmonary congestion, , a serious con- dition in birds as will be seen later. ' A few of these cases have been studied bacteriologically with no definite result, nor have these cases occurred in such groups that an epizootic was suggested. Bact. avisepticum, Bact. cani- septicum, Bact. coli, Bact. aerogenes mucosum, and Ps. pyocyaneus, Streptococcus hemolyticus and non-hemo- lyticus and staphylococci have been found. A review of the active verminous lesions of the bronchi and the pneumonitis to which parasites lead, shows again the liighest number among the Carnivora, with negligible percentages among the Aves. The forms concerned are, when determined, ascarides, strongylus, hepaticola, fas- ciolopsis, cytoleichus, pneumonyssus and paragonimus; these will be discussed later. The lesions in the bronchi are mucocatarrhal or hypertrophic; occasionally actual ulcerations are seen. What is more important however is the peribroncliitis leading to interstitial pneumonitis and to bronchiectasis of the smaller bronchi, or to areas of atelectasis by total occlusion of some small air passage. This pathology is fairly well recognized among veterinary pathologists but there are two points which seem worthy of special emphasis, namely, the relative mildness of the changes in the larger bronchi and the importance of the worms as causes of pneumonia. In regard to the first it can be pointed out that the trachea and its branches need not be altered at all while the middle sized bronchi present a mottling of small recent congestion with pig- mentations from old hemorrhages, together with slight unevennesses of the surface. Small bronchi on the other hand are the seat of ulcerative internal processes and quite marked peribronchitis, as indicated by round and connected tissue nuclei or perhaps polynuclear increase under active acute inflammation; it is in the latter case that acute pneumonitis is present. 144 DISEASE IN WILD MAMMALS AND BIRDS To what extent do the parasites predispose to pneu- monia? Compare for this purpose the two columns of Table 8 showing active and inactive pulmonary parasi- tism. In Rodentia and Galli alone do we note that active parasitism is effective, there being no passive cases. In all the other orders, animal invaders of the lung are more often encountered as quiescent or encapsulated bodies, therefore as findings incidental to the autopsy and per- haps not concerned in the cause of death. In Ungulata the inactive parasitisms are five times as frequent as the active inflammatory lesions. It might be added that the list is made up of cases wherein we saw parasites whether determined or not, an explanation of the apparently small number of cases; there were many more in which such invaders were suspected but not found and there- fore excluded. Beonchiectasis. Bronchiectasis affecting the smallest tubes, or bron- chiolectasis, is not at all uncommon in verminous pneu- monitis and is explained as due to the degeneration of the wall, the surrounding progressive ulceration, to accumu- lation of inspired air and its retention by the obstruction. There is described a generalized bronchiolectasis, chiefly in young human beings, due to a destructive bronchio- litis ; this has not been seen. Non- verminous bronchiectasis of the middle sized bronchi such as is seen in human chronic bronchitis, simple or tuberculous, is quite uncommon. Widening of the bronchial lumen may be divided, as I see its path- ogenesis, into (a) that due to congenital weakness of the walls, (b) that due to obstruction permitting air to pass into but not out of a bronchus because of a ball-valve obstruction or weakness of expiratory power, (c) that due to external pressure by tumors or distortion by fibrous tissue either witliin the lung or pleura and (d) that due to inflammatory weakening of walls, augmented by loss of THE RESPIRATORY SYSTEM 145 supporting pulmonary tension, accummulation of secre- tion and the dilating etfect of inspiration preparatory to and incidental to coughing. How important the last three auxiliary factors may be in the cases explicable under a, b, c, can easily be speculated upon and may vary in different cases. Bronchiectasis is reasonably common with pulmonary diseases of man, particularly of chronic character, but is certainly not frequent among animals. Under the first group (a) we can record one case, a Siberian tiger which died of enteritis and its complications to which were added a mild inactive bronchitis and a broncliiectasis of diffuse distribution. The lungs were irregular in shape, dull, gray red in color and gave a variable boggy and vesicular sensation to the fingers. On opening the lung, dilatations of the bronchi were found, affecting chiefly the larger secondaries but apparently not the broncliioles. Para- sites were not found nor were inflammatory reactions apparently adequate to explain the distentions, so that we looked upon this case as congenital. Cases coming under the headings b and c are not recorded. Inflammation almost certainly represents the most important single factor in the pathogenesis of this lesion and could be demonstrated in two cases, a Clouded Leopard {Felis nehulosa) and a Red River Hog {Potmnockcerus porcus). While I feel that parasites probably laid the foundation for the dilatations in these cases, none were found after, in one case at least, a very thorough search, although in the second animal a single cyst of Cysticercus tenuicollis was found in the peritoneum. In both animals there was a low grade interstitial pneumonitis and peribronchitis mth dilatations of the middle sized and end bronchi, these being supplied mth thick walls but containing very scanty secretion. We have on record chronic ulcerative pulmonary tuberculosis in six primates, two carnivores and nine ungulates. It is highly probable that among this number 146 DISEASE IN WILD MAMMALS AND BIRDS some cases of ulcerative bronchiectasis occurred but if so they were not conspicuous enough to mention in the diag- nosis and in only two protocols do I find a discussion thereof, once in a monkey and once in a carnivore. All the cases of our records were diffuse ectasias, fusiform, or irregular and none of the distinct saccular variety. From the foregoing facts it would seem that in human cases more weight, should be laid to the effect of the dilating power of coughing and its preparatory exertions. While I am aware that the comparative incidence of human and lower animal' bronchiectasis cannot be based upon the meager figaires at our command, these dilata- tions certainly can be expected in a general autopsy ser\dce more in man than in lower animals. Chronic bronchitis is relatively rare, aside from the verminous varieties. I have seen little retained exudate in the bronchi, probably because quadrupeds seem with ease to raise and swallow the secretions. Nor do animals give vent to paroxysms of coughing such as the human being feels forced to do. Suggestive deductions from these points are that inflammation is the principal factor in acquired bronchiectasis and that the retention of secretion with violent inspiratory efforts are potent in man for the dilatation of the tubes. The Lungs. The essential respiratory organ of the animal body, the lung, is all through this kingdom a structure intended to expose the blood to free or combined atmosphere in order to permit gaseous interchange, therefore being arranged so that there is a close apposition of the two factors, separated only by such cells and membranes as may be necessary to protect the circulation ; perhaps these anatomical elements possess at the same time some vital force to further the exchange of useful and useless matter. In the two classes under discussion there is no difference whatsoever in the primary factors of respiration albeit THE RESPIRATORY SYSTEM 147 some variations exist in reference to oxygen and carbon dioxide interchange, moisture of the air, and the physics of inspiration and expiration. The chemical variables have in our limited knowledge of comparative physiology apparently little effect upon morbid anatomy but it is probable that some pathology may be in part explained on physical grounds. The mammalian respiratory box is a relatively elastic affair, but collapsed at the end of expiration which is largely a passive or recoil process, f The avian thorax is believed to be normally a tensely distended space from which air is expelled by pressure of the pectoral contrac- tion upon the broad sternum driving the latter back upon the air sacs which in turn drives it from the lungs. Also by this means, air is distributed through the bones and- air spaces, a measure necessary in flight, especially in a head wind when tracheal inspiration is said to be sus- pended at times.. The communications of the lungs, air sacs and bones make it possible for birds to breathe internally when the trachea is closed and externally as well if a bone be opened. The balance of air pressure in the lungs and related spaces is dependent upon the patency of the ostia communicating between the bronchial ends and the air sacs, from wliich the bony cavities obtain their supply. Should all these be closed there is first a standstill of current and a limitation of the respiration of the lungs. Fortunately it is extremely rare that this occurs for it is obvious that it is incompatible with flight, and with life indeed. The principal effect upon the lungs of obstruction to the passages seems to be expressed in congestion but in how far this is due actually to the closure of foramina and how far to the cause of obstruc- tion is sometimes difficult to evaluate. ' It should be remembered that the air sacs are usually looked upon as mucous surfaces continuous with the bronchial wall, there being a deep layer to each membrane possibly continuous with the serous membranes. In mould disease of the 148 DISEASE IN WILD MAMMALS AND BIRDS lungs there is very commonly a colony lying in the ostium supplying the anterior, lateral and posterolateral ca\"ities.^| 1 The lungs in birds are not free as in mammals, being fitted into the troughs made by the anterior ridges of the ribs, to the serous covering of which thej^ are lightly attached by delicate fibres running between the two. This more or less definite fixation, together with the pressure of the air in the sacs give the free play of the lungs a limited excursion. They are naturally very elastic by reason of a good supply of elastic fibers and large air spaces, a con- dition aided by their attachments to the supports of the diaphragm and to the insertions of the air sac walls. Not- withstanding this elasticity and the great capacity of the organ for blood, it seems as if congestion of the lungs is a very serious matter, since from the foregoing re^dew of anatomy, accommodation of excess blood and any con- solidation must be difficult. As a matter of fact the mere excess of blood known as active congestion seems able to kill small varieties.J Congestion- op Lungs. Birds of flight seem to have little resistance to this con- dition and often it is the only diagnosis one can make at autopsy. The causes of this condition include exposure, dust, gorging (?), indigestion, enteritis and infection in birds while in mammals acute gastrointestinal disease stands out as the most prominent accompaniment. To what extent dust and exposure operate I do not see, although they are frequently mentioned as causes. The overfilling of the crop, esophagus and proventricle, the turgescence incident to gastric indigestion or the pressure of foreign bodies in large amount are supposed to operate by exerting pressure on the anterolateral air sacs mth closure of their ostia and also by right lateral torsion of the heart with twisting of the very delicate pulmo- nary veins. , THE RESPIRATORY SYSTEM 149 I have sought to show that protozoa or embryo nematodes in the blood might embarrass the lungs to a state of congestion, a thought suggested by some findings in the London Gardens, but only about ten per cent, of our cases of hemic parasitism are accompanied by it. The incidence of congestion of the lungs not due to stasis as from cardiac diseases, is 2.4 per cent, in mam- malian autopsies, in only 7 per cent, of which figure did it represent the principal morbid anatomy, whereas in birds it occurred to the extent of 7.6 per cent, of postmortems, in 17 per cent, of which it was the sole or principal cause of death. This seems to bear out the feature of delicacy of the pulmonary vascular mechanism in these latter animals. This condition seems to be indicated by simple dyspnoea in birds, relief for which has occasionally been afforded by removal from the exhibition cages and pro- tection separately in a warm dry room; this is partly hypothetical of course and congestion is to be looked upon as serious, particularly in passerine birds. Pneumonia. Pneumonia as a clinical disease is a relatively un- common, although quite serious sporadic condition in animals. However accompanying the specific, more or less epizootic diseases such as influenza, distemper and the choleras it may be a frequent and quite pronounced complicating feature of the case. Pneumonia per se has exacted a reasonable toll in this Garden but unfortunately recognition being impracticable, diagnosis and treatment have not progressed. Nor has it been practicable to group our cases pathologically because of the lack of history and the difificulty of making bacteriological observations at many autopsies. Fortunately we have had practically no epizootic pneumonias, an experience shared with other gardens judging by their published reports. Etiologi- cally, and of course this applies to non-verminous, non- mycotic and non-tuberculous cases, the pneumococcus has 150 DISEASE IN WILD MAMMALS AND BIRDS stood out prominently as a cause with a few additional cases due to the streptococcus and the Bact. aerogenes mucosum group ; London reports four cases in monkeys due to the Friedlander bacillus. Some time ago Doctor Table 9. Showing the Number of Cases of the Various Forms of Pneumonia Found in Each of the Orders. Primates Lemures Carnivora. . . . Insectivora. . . Chiroptera . . . Rodentia Ungulata . . . . Proboscidea . . Hyracoidea . . Edentata . . . . Marsupialia. . Monotremata Passeres Picariae Striges Psittaci Accipitres. . . . Columbaj . . . . GalU Hemopodii. . . Fulicariae . . . . Alectorides. . . Limicolse Gaviae Impennes. . . . Steganopodes. Herodiones. . . Odontoglossse. Palamedes . . . Anseres Struthiones . . 3 "3 ■^ '> •11 i5 b O < a o 4 22 o 1 1 o 1 3 1 34 6 4 3 ' 14 5 1 1 1 1 13 1 1 1 59 3 2 2 1 15 1 2 2 1 1 1 1 Weidman subjected our pneumonias to an analysis and was able to show that there is no parallelism between the seasonal incidence of pneumonia in man and animals, rather indeed that the Garden is more apt to have a greater number of cases in the summer, a sort of *' closed THE RESPIRATORY SYSTEM 151 season ' ' for man. This I am inclined to interpret as con- nected with the larger number of visitors during that season. Doctor Weidman was further able to show that the only real examples of lobar fibrinous pneumonia strictly comparable to the human infection occurred in the Primates. I have uncovered one in a lemur and one in a carnivore. The pneumococcus has been far and away the greatest producer of our pneumonias, in two typed cases being of the IV variety. There mil be given below a summary of the pathological types of pneumonia en- countered, to be followed by some notes upon the prin- cipal gross and minute anatomy in special orders. Table 9 mil show the numerical distribution of types among the orders. All the principal mammalian orders are represented while the birds seem relatively less sus- ceptible to the disease and, except the Passeres, show a trifling incidence. Primates present a definite group of variations from the other orders, notably in having four clear cases of lobar fibrinous pneumonia, and in certain histological findings. In reference to the lobar cases, a re\aew of their history does not indicate that any might have been surely diagnosed by their symptoms, and only possibly by signs in one case during the stage of red hepatization; unfortu- nately no temperature records are at hand. In one case it was possible to see a group of alveoli wdth the fibrin collected in a strand which, according to classical descrip- tion, passes through the septum to the adjoining alveolus. There were two cases, a Chimpanzee (Pan niger) and a Galago (Gal ago maholi) with a microscopical picture suggestive of those we met in the influenza epidemic, and indeed the lung of the former resembles grossly the lung of influenza pneumonia. The spotty areas of watery purple color correspond under magnification to cellulo- edematous semisolid sections shomng a sanguineous exudate, fewpolynuclear cells and many swollen epithelia. The microscopic picture of the bronchocatarrhal pneu- 152 DISEASE IN WILD MAMMALS AND BIRDS monias shows conspicuously thickened septa decidedly ^\dder than one is accustomed to see in human cases and apparently due more to round cell infiltration than to con- gestion or polynuclear increase. Bronchopneumonia or capillary broncliitis with zones of cellular edema in the vicinity is a rather usual picture in the deaths from degenerative bone disease. It cannot be said that there is anything very peculiar about it, although a frequent note met in the autopsies describes spotty areas of hemorrhage and nearby atelectasis. The case of lobar pneumonia found in a ring-tailed lemur {Lemur catta) showed very delicate fibrinous retic- ulum and relatively few cells in the exudate, a picture apparently due in part to beginning resolution since the whole upper left lobe was in a stage of gray hepatization. The peculiarity of the Carnivora seems to lie in the reaction of the epithelia, these cells being quite large, swollen and occasionally much vacuolated. Such a picture was most pronounced in the terminal bronchitic pneu- monias in cases which might be called distemper. Many instances of pseudolobar catarrhal or bronchopneumonia are recorded but we also observed the fibrinous lobar form at the stage of red hepatization in a Texas skunk {Mephitis mesomelas) . Concerning the orders Rodentia and Edentata no especial notes seem necessary for their inflammatory reactions are essentially like the others in that epithelial cells are much swollen and prominent. Pneumonias of Ungulata are well known to pathology and offer in causation and microscopy little that is peculiar. It might be emphasized however that the gross appearance of the bronchocatarrhal variety closely simu- lates that of lobar pneumonia, therefore to be called a pseudolobar form, in that extension to various parts of a lobe seems to occur. Moreover in the bronchitic varieties associated with enteritis, with or without infec- tious foci in the pharynx or larynx, there may be two or even three stages of the pneumonitic process in one lung hl( 8— \()R\I\I WIW IKI\l\k"l \\i) -UONDXk'i \l \ 1 Ol I THi M n AI PROLONG M lON-i IHAI 1 (H M) IHl J'RI\IAR\ \1 \ 1 i Fig. 9. — EARLY HRONCHOPNKUMONIA OF SUPERFICIAL ORIGIN. NOTE SOME LITTLE EXUDATE IN SECONDARY ALVEOLUS. WIDE SWOLLEN SEPTA AND BOTH ALVEOLI REDUCED IN SIZE. THE RESPIRATORY SYSTEM 153 or lobe. It seems that this pseudolobar appearance occurs definitely more often in ungulates than in the other orders. Marsupialia offer two rather easily grouped classes of bronchopneumonia — one associated with enteritis and one secondary to " Kangaroo disease " of the jaw; they differ in microscopy correspondingly. The simple bron- chitic and peribrOnchitic infiltrate and superficial exudate occurring with enteritis or with a general infection is relatively diffuse, giving in some instances the impression of an interstitial process and showing notably swollen septa; there may be fibrin but this is exceptional and scanty. Wlien mycosis of the jaw has been the origin or occasion of the infection the picture is that of frank aspi- ration pneumonia, therefore more like a septic infarct. However the amount of fibrin is sometimes very great and whole alveoli will be filled with it, perhaps accompanied by red cells, polynuclears and epithelia. Epithelial cells however play a small part in the minute anatomy. Hemor- rhage and edema are prominent but true abscess forma- tion and gangrene are not. Possibly the animals die too soon for the latter to develop. Pneumonia in Aves aside from that due to moulds is apparently much less coromon than among the Mammalia, one order only, the Passeres, showing an incidence com- parable to the important orders of the latter class. The other orders, and this applies particularly to those of which we have an adequate number, are quite insus- ceptible to simple pneumonia, none of them showing over two per cent. / There are listed for Aves three instances of lobar fibrinous pneumonia. These cases can be described together since in all the findings were about the same. A whole lung or goodly portion thereof was uni- formly involved in a red or gray consolidation of rather fine granular character which on section study seemed to be made up of the same lesion all over, with fibrin a promi- nent part of the exudate. The coagula were largely 11 154 DISEASE IN WILD MAMMALS AND BIRDS within the secondary alveoli but the primaries also contained it. The microscopic section may not have rep- resented the process at all places, and since the arrange- ment of fibrin is similar in definitely catarrhal lesions, these may of course have been instances of pseudo- lobar pneumonia. Our data are too few to draw any conclusions as to the behavior of the various orders but one note may be per- mitted. The passerine birds have a great tendency to dense cellular infiltrates while parrots show more coagu- lative or fluid exudates. Production of Insular Pneumonia in Birds. Insular consolidations in which catarrhal and infiltra- tive processes are prominent, the bronchopneumonias, seem to arise in two ways. One course of events appar- ently follows infection via the bronchial mucosa, the other via the blood stream and a study of the resulting lesions may help toward an understanding of the development of pneumonia in man. When infection unquestionably has been superficial, that is via the bronchus, the first thing to happen is a swelling of the septal prolongations dividing the primary alveoli and an extension of their ends farther into the secondary alveoli with the result that the inlet to the primary air sacs is narrowed and the space in the second- aries is reduced. Upon the surfaces there then develops the usual catarrhal exudate while in the deeper parts marked congestion makes its appearance. Fibrin may develop and be mixed with the cells both in the larger and smaller alveoli but it is more e\ddent in the former. (Figs. 8, 9, 10.) The other process by which insular pneumonia de- velops seems to begin in the septa of the smaller alveoli and in the perivascular areas. This has been looked upon as hematogenic or pleurogenic. The first change occurs in the surroundings of the primary alveoli where there ) Fig. 10. — LATER BRONCHOPNEUMONIA OK SUPERFICIAL ORK;iN. I GREATER SWELLING OF SEPTA. PRIMARY ALVEOLI PRACTICALLY ALL HAS BECOME CONSOLIDATED. Fir.. 11. — INSL'I.AR PNiaMUMA, Bl.t,lNMNC, A> Cl.l.lA LAK. IMILIRAIION OF DEHl'ER PARTS OF SEPTA AND OF INTERSTITIAL TISSLE. FOUR AREAS OF DENSE AIRLESS CONSOI.ID.A- TION. ALL SECONDARY AND .MANY PRIMARY ALVEOLI WIDELY OPEN. THE RESPIRATORY SYSTEM 155 appears a richness of nuclei, of round, moderately well stained character, among which one may see a few gran- ular and red blood cells. Soon the epithelia of adjacent alveoli increase in number and a fibrinocellular exudate appears, at first probably in the smaller sacs. However when the lesion is intensive the course of events must be rapid for the identity of a group of primary alveoli is soon lost and the exudate may extend to the larger air space. (Fig. 11) In severe or late cases a decision as to the course of origin is often impossible. The most instructive point of this part of the study is the closing of primary alveoli by the sw^elling of their septal ends and the early occlusion of the secondary alveolus by a catarrho- fibrinous or even pus-like material. It is quite possible that a similar course of events transpires in the path- ogenesis of human pneumonia, the superficial avian form being comparable to the aspiration form, the interstitial form comparable to the septicemic variety.] Abscess and Gangrene of Lung. Abscess and gangrene of the lung are degenerative processes dependent upon embolism, or inspiration of infective matter and it is usually assumed that gangrene succeeds upon abscess when the blood or air supply of a part of the pulmonary tissue has been obstructed mechanically or by inflammation. A review of our material adds little to the etiology or pathogenesis of these two lesions, well recognized as they are by veterina- rians. As opposed to human beings, lower animals prob- ably suffer more from them, for an explanation of which one can probably look to the B. necrosis or necrophorus, an organism quite common in feed, and acknowledged to be of great importance as a secondary invader during specific infectious diseases. It has been found in embolic abscesses and in the organs in calf diphtheria and similar other conditions. It has been cultivated here twice, once from a lung abscess, once from Kangaroo disease. It 156 DISEASE IN WILD MAMMALS AND BIRDS 0.0 ■h^ 1.-=? a §■2 si l5 gs ■si Ei = & o « [|j ■ — 0) 3 ■ • B • I II i, --i ---i ■■ 3 i. ^ .a 3 • o ■ XI J3 53 2 Oi: -a c « ?i'2J^T33 'O 0-" -s s ;?;= E^ o a 8 " ft* 'Sm THE RESPIRATORY SYSTEM 157 ^ s u S ^ 5 >> 5 •>, £ o. _2 2 2 1 11 a ! m m PL| m CO (i< 1 ill 1-. !i III o >, ■1 |§ 1 03 ■|| H! II gg 1! IS 1. 1 ^ - 0) ll g>. a 2 J3 - -Is Ill if ill =11 1- lbs 1 il 1 E 1 c3 lli ill Ii II j_ « 03 ill 02 < hj ■< ^ >J m ^ ►3 ■« "eS ~A "ei S S. » 3, 1 Ji « 1 •S-^ 1 £ — •^ 2 — _2 c3 M C3 Su- Cli M M "a "m "m a o« S 3 c S (5 s s s 1 S s m CO hi i 1 i iJ i s « s i i % S S M o c to o o S ^«S ^ J -D .s ^ J 1 1 1 1 ■< te -s: < << < <:: < <3 <: *t; ■ 5.2 E E Q. J 3 .2 2 . B E II to 7' 1* 1 1 1 i 1 > ■s i 1 > 1 en 83 o 1 1 >> 1 >> ^ 1 £ E C n > •^ •s > ■J S ^ S a S C3 S s 1 > ■■S 3 £ .- m C3 C9 M.S M.^ o! "o a C3 ■w 03 M U CO c^ t! §1 u a V IB (U ca"3 (So (U C •< IS < ;z Z W M 'Z •Z :z; IS -J" « 3 3 i 3 3 1 o 1 a fl u 3 5 1 ^ 1 S § 3 2 s 1 .2 1 1 "3 "2 .s M 1 a *« O "o .2 ■c C3 1 i § § 3 3 ^ 1 ll II ■\ i .1 1 1 o a £2 II II II I a 2 1 1 1^ o c E >> O 1 2 < 03 < ^ < ^ c ti ^ u. ^ c CI c 158 DISEASE IN WILD MAMMALS AND BIRDS doubtless occurs in human necrotizing processes but is seldom emphasized or even heard about; possibly none is due to it or its congeners. The distribution of abscess and gangrene in the lungs in terms of the antecedent disease, therefore its causation, may however be of interest. The lower animals move more in the horizontal position, they seldom cough, they are subject to several different diseases A\T[th principal lesions in the anterior head (diphtheria, actinomycosis, etc.) but not to chronic lymphatic infection, they push their snouts into all kinds of filth thereby probably taking into the nose and throat many objects which can find their way to the bronchi, and finally they are not subjected to various instrumental operative procedures when they chance to have a focus of pathology in the nasopharynx. For these reasons the position of abscess and its sequels may be instructive. It has recently been stated that ab- scesses of the lung in human beings following anesthesia for infections of the upper respiratory tract, occur most often in the upper parts of the lung whereas those follow- ing pneumonia develop more in the lower lobes. For the animals of this series, these facts are not borne out. In the accompanying list will be found our acceptable cases of abscess and gangrene. Little can be said about incidence save the fact that the highest percentage and number occur in the Carnivora. It will be found that the right lung is affected nine times predominatingly while the left lung is affected seven times. The lobe most often singled out for an isolated lesion is the lower left, the right middle being the next most often affected. These figures concern the mammals alone, the three birds being considered too few to discuss. It cannot be said from these figures that there is in animals any definite distri- bution of pulmonary abscess and gangrene. Nor do these figures correspond to those appearing in literature of human pathology. In man inspiration of foreign bodies, including bacteria-laden mucus, usually THE RESPIRATORY SYSTEM 159 carries them into the right lung because of the larger and more directly vertical bronchus to that side. Emboli go as a rule also to the right lung more than to the left because of the greater size of the pulmonary artery and more direct blood current to that side. In these animals right side lesions are more numerous but the left lower lobe is the principal segment of the lung to be affected. The cases are too few and the anatomy too variable to permit any deductions. There is in the affected animals no uniform anatomical peculiarity which would explain the predominance of the right lung as a whole or the left lower lobe as a unit. Pneumonokoniosis. Pneumonokoniosis, because of its importance in indus- trial diseases, has been subjected during recent years to considerable intensive study in human medicine, from which activity some interesting and useful information has been obtained as to its genesis and effect upon the function of the lung. This condition is of course a purely environmental one, the degree and particular kind of '' dusting " being dependent upon the duration and nature of exposure of the particular individual. This Garden is situated beside an active railroad trunk line so that the opportunity for coal dust inhalation is con- tinuous. The degree of anthracotic pigmentation of the lungs and related serous membranes is really negligible and with one questionable exception, we have not seen fibrosis due to this cause in any animal. The one excep- tion, an amazon presented and living in the Garden but three months, at necropsy showed an interstitial chronic bronchitis and pneumonitis stretching out from the hilum, all of the affected area being deeply pigmented. The picture was comparable to what might be expected from a second degree anthracotic fibrosis of Landis and Pan- coast. Many specimens come to autopsy with some grade 160 DISEASE IN WILD MAMMALS AND BIRDS of pigmentation, but none, except the one above, vnth. resultant fibrosis. The degree of anthracosis is usually so slight that it has been considered important enough to include in the diagnosis but eighteen times and curiously enough seventeen of these were observed in birds. Were there more cases it might be profitable to plot their exhi- bition period but the use of this small number might lead to error ; the average leng-th of exhibition of the birds was about a year. It is common to observ^e some black speck- lings of the air sacs, as if pepper were dusted on them as has been said before, but even this is rarely marked. It is most often seen in the Anseres, Psittaci and Struthiones but a goodly number of cases occur in the long- lived Passeres. ■ The distribution of the pigment is essentially the same throughout Mammalia — peribronchial, submucous and in the Ij^mph nodes at the root of the Imig. In the birds it is first seen in the subepithelial spaces of the septa of the small alveoli where they project into the secondaries, later accumulating in the connective tissue of the main septa. Collections under the pleura and at the root of the lung are rare, the dust usually spreading out along the air passages into the air sacs. Other forms of pneumonokoniosis are unkno^vn. Although animals must inspire much dust from dry feed and from floors it must be caught early and removed by snorting or by the lymphatic drainage. It seems fairly well accepted that dusts are dangerous to the degree that they contain inorganic substance and as these animals are not exposed to concentrated mineral or metallic dusts, no effects are seen. Infarction of Lung. Infarctions of the lung, while not at all common, are interesting because of their incidence in the Carnivora and in the distribution. The figures concern the mammals THE RESPIRATORY SYSTEM 161 only since the decision for or against infarct in the birds is very difficult because of the frequency in this class of hemorrhage A\4th pulmonary congestion. There were eleven single or double non-septic infarcts, of which seven occurred in Camivora, one in an ungulate, two in Pri- mates and one in a rodent. The existence of parasites was excluded in most of the cases but could not be entirely in all. Eight of these infarcts were on the left side, five of these being in the lower lobe. Emphysema. Emphysema of the atrophic and chronic vesicular types with the soft, fluffy, pigmented or pale pink organ has not occurred in the animals under observation. Acute vesicular emphysema, such as is seen in chronic bronchial and cardiac diseases, has been encountered several times. Cardiac lesions were found four times, nephritis eleven times, acute enteric conditions seven times, hepatic dis- eases seven times. Two cases of wide spread amyloid disease showed a deposit of this substance in the alveolar walls. It is quite common to find some grade of emphysema in monkeys dying from osteomalacia and rickets. The process is then most prominent in the upper lobes and along the free anterior margins. The incidence in the orders is Primates 5, Lemures 2, Camivora 2, Pinnipedia (drowning) 1, Rodentia 1, Ungulata 4, Mar- supialia 4. The best example was found in a Skunk {Mephitis mesomelas) having a general infection, ema- nating from the cranial sinuses, and cardiac dilatation. Emphysema does not seem to occur in birds for only one was seen which seemed to present this condition. This was a Bald Eagle {Ealiceetus leucocephalus) with chronic renal and enteric disease and cardiac hypertrophy. The lungs were tensely distended under their serous covering and showed a few small bull« anteriorly. Unfortunately a histological preparation is not at hand. 162 DISEASE IN WILD MAMMALS AND BIRDS Tumors. Tumors of the lung are moderately common, both of primary and secondary origin. Thus we have seen three primary and six metastatic growths in mammals and one of each kind in birds. The primaries were : carcinomata in a civet {Viverra tangalunga), a bandicoot {Thylacomys lagotis), a kangaroo {Macropus rufus), and a lorikeet {Glossopsittacus concimius) . The secondaries were : car- cinomata in a black bear {Ursus americanus), a polar bear (Ursus maritimus), a lion (Felis leo), and a dasyure {Dasyurus maculatus) ; sarcomata in a prairie wolf {Canis latrans), and a raccoon-like dog {Canis pro- cyonoides) ; adenocarcinoma in a chestnut-eared finch {Amadina castanotis). The histological character of the primary cancers would place them in group of the nodular and infiltrative types of Kauffman. They all seem to have taken their origin from the smaller bronchi, the usual starting point. The growths were small in the civet and bandicoot and strongly suggest that the tumors arose in bronchi occu- pied by parasites ; such bodies could not be demonstrated. It is the usual tiling to find in cases of parasitism of the bronchi that if there be no ulcerative destruction of tissue the epithelium undergoes some form of hyperplasia, and even structural metaplasia in, the air tubes supplied with cuboidal or cylindrical cells. Epithelia many layers deep have been encountered, usually arranged in orderly fashion but frequently '' papillomatoid, " suggesting the epidermal layers yet not so far as to show protoplasmic bridges. Distention of various degrees, affected by the contents of the tube and the surrounding inflammation, are common. Such a picture naturally resembles epi- thelioma and indeed growths of this nature are reported as due to verminous pneumonitis. There are, especially in cats and dogs, small scattered adenomatoid groAvths(2) under the pleura and in the (2) Ball, Jour. Vet., 1907. THE RESPIRATORY SYSTEM 163 pulmonary substance, thought to originate in the alveolar epithelium and occasionally growing to large size; the case in the kangaroo may have had this origin. It ivas the only primary tumor to give metastasis (to the spleen and gastric wall), the secondaries being decidedly ade- nomatous in character. Metastatic groAvths come from the following originals : two from the thyroid, well known to give pulmonary embolism in dogs; one each from the breast, uterus, adrenal, intestine and kidney. The form assumed is a gray and red mass lying under the pleura or an isolated nodule in the substance. Sarcomatosis, the form appar- ently spreading out from the hilum and growdng in isolated grayish tubercular masses, has not been seen. The Pleuea. The pleura is a tissue apparently quite susceptible to infection in mammals and so closely associated with the air sacs in birds as to be a part of the same membrane, therefore the two being affected together. Throughout the higher class all orders give copious examples of the involvement of the pleura, principally of course as an accompaniment or a sequel to pneumonitic or bronchitic processes but also as a part of acute infectious diseases, such as hemorrhagic septicemia, pleuropneumonia and the like. However two orders present such a number of instances of pleuritis that they deserve notice. The seals, Pinnipedia, of w^hicli we have twenty autopsy records, showed inflammation of this membrane four times, three of which were dependent upon pulmonary infection and one apparently due to general septicemia with trifling damage to the lung proper. One of the first cases had gone on to empyema of the classical type, a shrivelled dry almost carnified lung with a thick fibrinopurulent cover- ing. The lung of the seal is well divided into lobules, the external surface being generously supplied with lym- phatic channels under the pleura, an arrangement which 164 DISEASE IN WILD MAMMALS AND BIRDS should carry away infection one would think. Perhaps this high percentage of pleurisy in our Pinnipedia is but accidental. The marsupials, while having a notable per- centage of pleurisy both among all the cases and in rela- tion to the number of postmortems, are not so striking from the etiological standpoint since practically all of these have suffered with Kangaroo mycosis or pneumonia. In over half the cases of tliis infectious disease some grade of pleuritic exudate has been observed, only one, however, going to the stage of empyema. ; One cannot speak so definitely of pleuritis in birds since this tissue merely represents in them the covering of the lung and is firmly attached posteriorly to the ribs and anteriorly to the air sacs. Exudates show as col- lections upon the air sac side of the combined membrane, pleuritis proper in birds being an infiltrative affair com- ing through the pulmonary tissue and therefore being a part of pneumonitis.; I notice a tendency in a few articles to write of pleuritis when the process is confined to the thorax but this gives the impression that the disease is peculiar. There seems no difference in the gross and minute appearance between thoracic serositis and pan- serositis. The course of procedure seems to be from the anterior or mesial pulmonary ostia into respectively the cervical and thoracic air sacs and this seems to hold good whether the infection be mycosis or fowl cholera or fowl pest. There are records of 104 cases of serositis in birds of which 45 were among parrots, the remainder being well distributed among the various orders ; only two each occurred in Galli and Anseres, orders prominently affected under domestication. This high percentage of pleuroperitonitis among parrots and their congeners can only be explained upon the ground of a continued infec- tion of our stock by the virus of fowl cholera and by mould. One case of undoubted fowl cholera occurred recently and as the records are reviewed a few are dis- covered where the organism was found. The virus must HA t. •o o ^ -c •«* ■ill p o S « ■« e CIS Oj g 535 =i ^l§iiil •S g I I s ^ s to e 5.2? s. -S^ to "^ ^.8 I e § S s s P>.?^ O ■:!='§ •2 g-g.'^e-?' glBuiiuv iB^ox noioQ CO Baui^eajni o « pauirajajapurj n siaafqo iB0|8.!SBJBj |Buituv eaisdo^ny JO -(jngo jaj «5 00 CD O GO CO »o CO r-l^ -rfi CD coc4 do rt S S a ^''S'P w o oj p g rt c 5o a 5 S cs £ i:-a So-t; h S b THE ALIMENTARY TRACT 183 iCi Tj< ^ (M CO O 05 CO CO CO ^ |oro^ C<)COOTt< ^ g^ in ^(M ^^^ 00 COO^ COOOkO - T}<^ ■* ^(M ^ CO Tti io (N -^ncoy-ic^ oot^>ot^ t>I rJH^IOCOCO .-H COIM CO^ --I 00 t>;t-; CD r-i 'eo eooi »o eot^»occico >0 cot- 00 CO £ 8^ a5 c^ .t; -a 2 G C "S • s 2.2 o tp SR--^ 184 DISEASE IN WILD MAMMALS AND BIRDS ous. From the observations of Kitt, Strassberger and some others, the normal flora of domesticated ani- mals is subject to wide variations so that our observations must receive confirmation before they are finally acceptable. We have on several occasions isolated from carnivorous intestines Bact. paracoli, Bact. suipestifer and other members of this group. We have no reliable cultural data upon the herbivorous intestine and can only quote the Gram pictures as mentioned above. On two occasions, an eland and an elk, a very large number of forms corresponding to necrosis bacillus were seen; to this organism Kitt gives considerable power in the production of necrotizing processes. The Table. Our records have been analyzed from the standpoint of diagnosis and the most probable cause. The first will be taken up in discussing each of the orders. The prob- able causes are divided into food, bacteria, animal parasites, physical objects and undecided, in other words a classification based upon the most prominent or definite evidences as seen at postmortem combined when possible with antemortem observations^ When findings were inconclusive or contradictory, cases were called unde- cided, naturally a very large group. Fermentative processes in the presence of obviously undigestible material, are classified with food as a cause. When evi- dences of septicemia existed in absence of the other factors, it is held that bacteria were responsible. Cases were grouped under animal parasites when these were the most definite findings. Physical objects are relatively unimportant and self-explanatory. To the etiological chart there are appended columns intended to show the percentage or case incidence of the disease of the grosser subdivisions of the gastrointestinal tract which indicate in a general manner what part of the tube in the various orders is most susceptible to disease. While of course THE ALIMENTARY TRACT 185 conclusions must be drawn with great caution, there can be little doubt, for example, that carnivores and mar- supials have more gastric disease than any other order, and that the high place for the colon is held by the galli- naceous birds. This charting was suggested by the work of Dr. Raymond Pearl upon statistics, wherein he takes as a basis of classification the part of the body which succumbs to disease-producing organisms or from which a disease starts. It caimot be stated that there is a clear cut relationship between enteritis and the expectancy of life. Mammalia. The Primates as an order have their share of in- flammations of the gastrointestinal tract and present points of interest. Acute digestive disorders succeeded by acute dilatation of the stomach, or in less fermentative cases by acute catarrh of the intestine, are not at all un- common. The reason for this is not discovered by review- ing the diet and manner of feeding. The buccal pouches, distensible esophagus, the freely movable stomach, and relatively elastic gastric wall would seem to permit of very considerable dilatation to accommodate the large quanti- ties which the monkey sometimes crams into himself. Nine fairly acceptable records of gastric overfilling exist and two of them seem to have been followed by tympan- ites sufficient to embarrass respiration, in one case there occurring an acute cardiac dilatation with myocardial degeneration. The animals give no symptoms of this condition and in the last case cited the beast, while old, ate well and was not distended the evening before death. When acute gastritis exists (twenty cases) the animal seems uneasy but does not vomit. On one occasion I was called to see a monkey which was retching and seemed in pain. Lime juice was offered and taken, followed by gentian and cardamon, which seemed to give some bene- 13 186 DISEASE IN WILD MAMMALS AND BIRDS fit. Somewhat later this was repeated in another case, but observations where this might be useful are rare. The pathology of gastric conditions offers little to contrast with that of man. The enormous distensibility of the fundal pouch often suggests to the observer the rumen of ungulates. Acute gastritis of one kind or another and acute catarrhal enteritis are the most com- mon lesions noted in the Primates. Involvement of the intestine or colon need not carry with it an increase of signs of illness, although at times one will see an evi- dently sick animal with diarrhcEa. Anatomically the lesions are commonly restricted to the stretches of gut above the ileum, it being rather rare that this division or the colon is affected. Pathologically the lesions are catarrhal with definite involvement of the follicles in about one-third of the cases. In this order toxic and pseudomembranous forms are quite rare and ulcerative lesions uncommon. Colonic disease as a sequel to inflam- mation higher up is sometimes seen in the follicular varieties, but takes a minor place compared to amoebic dysentery of which we have had several cases ; this will be discussed under a separate heading. Degenerative disease of the skeleton is almost always accompanied by a low grade of enteritis but not necessarily gastritis or colitis. The pallor of the mucosa, while at times striking, may be relieved by follicular spots and petechia or pig- mentation. Often, however, animals suffering from osteomalacia and rickets come to their end by an acute inflammation of the gut tract. The bacteriology at our disposal allows no conclusions. Aside from a case probably due to Ps. fluorescens and one with colon bacillus abscesses in the liver, no reliable data are at hand. Reference to Table 11 reveals the fact that among orders with sufficient autopsies to permit percentages, the alimentary tract in monkeys is in the group of low figures, that the intestinal section is relatively more often THE ALIMENTARY TRACT 187 affected, and that the colon is more often diseased than in other mammalian orders, and is exceeded only by the gallinaceous birds. The Lemures, of which we have eighty-six autop- sies, do not differ much in anatomy from the Primates, however greatly they disagree in habits and outward appearance ; their diet is the same. Clinically the sloth- ful behavior of a normal lemur probably obscures symptoms and signs of illness, for our antemortem notes with the exception of a few observations of loose stools, fail to offer a lead as to diagnosis. This order has a large incidence (twenty-three cases) of gastroenteric condi- tions as shown in Table 11, but some explanation of the figures is deserved. In the first place, only one case of acute gastritis occurred, and this was apparently a part of a general infection, and if induced at all by food this was only secondary. Indeed as one reviews the records it does not seem that the lemurs are easily disturbed in their gastric digestion. Acute and subacute inflamma- tions from bacterial action seem definitely more promi- nent since they take the catarrhal, follicular and deep submucous form and are frequently associated with gen- eralized infectious processes. One amoebic case was observed and there was another in which a heavy cestode and nematode infestation seemed to have paved the way for bacteria. Carnivora. The food of this order is received into the fundal part of the stomach, the distensible but normally capa- cious left and superior two-thirds of the organ. The general shape of the viscus, that of a gourd, permits a fairly sharp separation of the fundal and pyloric sections, so definite indeed that the pathology of the two parts was studied. The intestines vary in length, but in the land carnivores are relatively short, narrow in lumen and rich in waU. A cecum, or at least a blind end of the 188 DISEASE IN WILD MAMMALS AND BIRDS large gut made by the insertion of the small intestine above the tip of the colon, is suggested in all families, although, as in the bears, it may be quite insignificant or rudimentary. Theoretically no stasis should occur at this point. The colon is short in all carnivores and, like the small gut, with a heavy wall. The comparative simphcity of the carnivorous gut tract, the ability of many of these animals to disgorge, the suggestion of high resistance of the upper end of the tract to infection and the ease with which diarrhoea can clear out the tube, would seem to warrant the expectation that inflammation would not be serious. Such, however, is not the fact for, on the con- trary, they have shown a higher incidence than any other order for which we have adequate comparison. Anatomi- cally considered their stomach occupies the second place in vulnerability, next to the marsupials, and their intestines the highest place ; this indicates of course that combined gastric and intestinal disease has often occur- red. Involvement of the colon occupies the second place, in ordinate susceptibility, being exceeded only by the monkeys, due to heavy parasitic infestation, but would occupy the first place were the eleven amoebic dysenteries in monkeys subtracted from their total, a subtraction which might be allowed since it represented an epi- zootic outbreak. Etiologically considered, it would seem as if the influence of incorrect feeding were of little importance, and from one standpoint this is probably the case. Acute fermentative or irritative processes are not common at all, while more inflammatory pictures, catarrhal, erosive or ulcerative, are the rule. There is another phase to the term incorrect food, that is incorrect in its cleanness. During 1912-15 there was an increasing mortality among the cats and dogs fed upon horse meat, mutton and fowl heads. Early in 1916 the butcher shop was reconstructed and thoroughly cleaned and covered galvanized pans supplied in which to transport the food ; these pans were THE ALIMENTARY TRACT 189 scrubbed and scalded after use. Since that time, infec- tious inflammations of the stomach and intestines have shown an ever increasing downward incidence, which result, there having been no material changes in other directions, I do not hesitate to ascribe to the improve- ment of butchering and dispensing engineered by Dr. W. B. Cadwalader. Helminths seem to be of importance in this order both by reason of the percentage of autopsies in which they presented the most probable or at least most suggestive cause and because uncinaria and strongylus have been seen attached to the wall and a large bulk of known irri- tative cestodes have occupied the lumen. Physical objects, stones, bones, wire, may cause irritation enough to activate bacterial action or may actually penetrate the wall; the latter action is well known. In so far as prac- tical application of this is concerned, it teaches to feed whole, unsplintered or ground bone. The distribution and character of pathological lesions according to the region of the stomach is what might be expected from the shape and physiology of its parts. True inflammatory processes are best, and in some cases only seen in the pyloric half of the viscus, while the changes in those few cases believed to be fermentative or irritative in nature were largely confined to the fundus. Dilatation of the latter part may be understood because there the muscular coats are about equal to the mucous in thickness and one-half the width of those at the pylorus, but why inflammatory processes should not be so developed in the fundus is not clear unless the greater availability of mucus protects the secreting wall. Not only does acute inflammation reach its most definite form in the second part of the stomach, but the irregular pigmentation, mammillated overgrowth and atrophy or ulceration of chronic disease are likewise best seen in this part. 190 DISEASE IN WILD MAMMALS AND BIRDS Acute enteritis, of all varieties, is seen more beauti- fully in carnivores than in any other order of mammals, and nowhere can it be studied better. Its gross appear- ance is that of the text-book and its minute character even more instructive. I have used a slide of acute catarrhal enteritis in a lion for the illustration of this lesion for the Text-hook of Pathology by Doctor Stengel and myself. (However, as is known to all who have paid any attention to enteritis, the postmortem findings are usually much less definite than clinical observations would warrant one to expect. The Carnivora not uncommonly show intestinal congestion, mucous membrane swelling without edema or opacity, congestion of the spleen, cloudy swelling of the liver and kidneys and perhaps mesenteric lymph node edema. This picture we have viewed as a toxic affair of some sort or a bacterial infection not yet far enough advanced to produce catarrhal or ulcerative enteritis and septicemia. In such cases the carnivorous intestinal mucosa offers instruction. The epithelium is vacuolated or fringed on the free edge or may be missing altogether. In the depths mucus formation is very active, and where it is going on, round cells seem attracted, collecting in groups in the villus or in the subjacent submucosa. Peri- vascular round cell increase may be noted. Plasma cells and granular eosinophiles are common, but I cannot state how important the latter are in the general picture because of the frequency of parasites in carnivores. The central vessel of the villus and the arterioles of the sub- mucosa are injected. Lymph follicles may or may not be enlarged, but if so usually fail to show a germ centre. Colitis alone is not common in this order, but as an extension process or involvement at the same time as the upper levels it occurs occasionally. The only fact I wish to record and one which I would emphasize because of having seen it recently in a human case of chronic colitis, and since it does not appear important to systematic writers, is superficial blood supply. The capillary net- THE ALIMENTARY TRACT 191 work of the colonic villi, while rich, is in the form of a fine plexus just under the epithelium. In the cases studied these vessels become quite distinct and possess much more definite walls, often bordered by mononuclears, while connective tissue is more evident at the bases of the villi and deeper. This may help in deciding the existence of a colitis. Bacteriologically the most instructive experience to report is the discovery that a small outbreak of enteritis among small Carnivora, chiefly cats, fed upon fowl heads was due to Bact. paracoli, or at least this organism was found in the intestinal mucosa, spleen, and heart's blood of three cases. The type of enteritis was hemorrhagic and follicular. There was also a case of septicemia appar- ently emanating from enteritis due to Bact. suipestif er in a lion {Felis leo). These facts bring strongly to attention the modem teaching that meat poisonings of the Gartner type are to be considered as infectious and not of the so-called ptomaine group. PiNNiPEDiA, while related closely to the Carnivora, are grouped in a suborder in our classification and because of their restricted diet are treated here in a separate paragraph. The tract is peculiar in the strong tubular stomach sharply bent upon itself, the great length of the small gut (upwards of a hundred feet in some genera), and the practical absence of a cecum. Pathologically speaking, the most striking lesion of these animals is ulcerative gastritis, a process usually most marked along the posterior-superior surface, but not confined thereto. Upon inspection the gastric mucosa, normally supplied with low regular rugae, is much distorted by swellings upon the top of which are irregular ragged ulcers with rounded elevated but not frayed margins. The density of the edges indicates much infiltration of the deep mucosa and submucosa; this can be confirmed by micro- scopical examination. One attempt to study this gastritis bacteriologically was fruitless. Sections of one case 192 DISEASE IN WILD MAMMALS AND BIRDS showed streptothrix-like masses while in another case bacterial colonies and yeast-like bodies were found in adjacent lymph nodes. The genesis of this condition might lie in injury by fish fins or by foreign bodies, of which large numbers are found at times (a pint and a half of stones, marbles, and sticks were found in one stomach). fGastritis has been the starting point of septicemia on two occasions, and three times an acute exacerbation or new implantation of infection occurred, ^vith extension into the intestine. It is interesting that all the deaths of Pinnipedia with gastroenteric conditions occurred in the winter months. Insectivora are represented by two common Euro- pean Hedgehogs. In one there were three shallow but shelving ulcers in the stomach which had bled suffi- ciently to weaken the animal; free blood was found in the intestine. The other specimen was diagnosed at post- mortem as having catarrhal enteritis involving nearly the whole small gut, but histological section did not con- firm this. As one descends the zoological scale the first gastro- intestinal tract prepared for the nutritional care of bulky food is to be found in the Rodentia. This order pre- sents a great variety of shapes and arrangements of the stomach, but the outstanding feature, with very few exceptions (cf. spermophiles), is the development of the cardiac and fundal divisions ostensibly for the reception of a large bulk of coarse food to be digested at leisure. Some genera like the hamster {Cricetus) have a stomach closely resembling the ruminants, while that of the spermophile suggests the equine stomach. The pyloric end, variable in many ways, greatly resembles the abomasum. So too the duodenum is large, loose and dis- tensible while the copious small gut ends in a very large cecum, shaped at times in a manner which has led to the term ''colonic stomach." The colon is variable and not always supplied with longitudinal bands and sacculations. THE ALIMENTARY TRACT 193 In regard to incidence of gastrointestinal disease, rodents occupy a middle position in the table. The stomach seems a vulnerable section of the tract. Dilata- tion of the left hand section is common, due, to all appear- ances, to fermentative processes which have as a result the softening of the mucosa so that even immediately- after death it will separate almost entirely. In these cases the pyloric part need not participate but may remain flat, smooth, soft and pink. This condition is slightly more common in the compound than in the simple stomachs. In some of this order, especially rats and cavies, there is a fermentative gastroenteritis expressed by injection and edema of the pylorus and duodenum, and much frothy mucus. It was at first thought that some relation might exist between this condition and the absence of the gall-bladder, but it occurs in varieties possessing this structure. The reaction of the intestine to irritation in this order is peculiar in two ways, the occurrence of mucus and the activity of the lymphatics. In all the inflammations from and including the stomach to the cecum, mucus is conspicuous. At times it is thin or loose and mixed with contents, while at others it forms a relatively close covering for the mucosa almost like a false membrane. Rodentia are peculiar in the prompt- ness and clearness with which the follicles of the intestinal wall and mesentery enlarge in inflammation. They appear as pale, well outlined or diffuse opacities in the wall or as distinct plaques prominent on the surface. The Proboscidea are represented by one Elephant {Elephas indicus), in which a mild catarrhal change was seen in the middle stretches of the small intestine. This was of little importance as a cause of death, there being several other diagnoses, and was probably a termi- nal affair. Hyraces, of which we have a total of seven ex- amples, present two mild involvements of the intestine but none of the stomach. It would appear from the records 194 DISEASE IN WILD MAMMALS AND BIRDS that the intestinal condition had little to do with the death of the animals, and unfortunately no microscopic slides were made. Because of the curious formation of the large gut, notes of both ceca were made in one case, and can be condensed as follows: "The upper or anterior cecum presented a shaggy pearl gray mucous covering, closely attached to the mucosa. This cavity and the posterior ceca were packed with dry crumbling feces. Small thin- walled cysts were seen in the tips of the lower ceca. Duo- denum was congested and mucosa swollen. Stomach contained dry, poorly digested food. No parasites were found. ' ' The Edentata are represented by an Armadillo {Tatu novemcinctiis) and an Ant-eater {Myrmecophaga tridactyla.) The former had a prolapse of the rectum accompanied by colitis but it is impossible to decide the priority of the two conditions since the former is known to have existed long enough to have permitted the latter to develop. The Ant-eater had a distinct mucocatarrhal enteritis in which bacteria played a part since involve- ment of the liver, spleen, kidney and lymph nodes also existed. The beast was in good condition upon arrival, but did not become accustomed to the proffered diet, and was distinctly anemic at death. The Ungulata, so-called for their hoofed and horned character, are also associated anatomically by the construction of their gastrointestinal tract. However, the order of the list as given on page 44 does not represent their historical development nor does it accord with anatomical arrangement of the tube under discussion. The last three families of Artiodactyla are perhaps the simplest in the construction of this tube, or at least take an equal place mth the Peris sodactyla, while the remain- ing families of the former suborder have a complex tract of generally similar architecture. Tliis whole order has, however, an alimentary tract anatomically suited for the consumption of bulky vegetable stuffs and shows an THE ALIMENTARY TRACT 195 attempt at adjustment between the food, tlie methods of mastication, the area of digestive surface, and the bulk necessary for nutrition. The number of factors opened up by the many variations between this complex tract and that of the carnivorous simple tract is so great that I shall attempt only to contrast the anatomy and pathology of the simpler ungulate tracts and the complex rumi- nant apparatus. The simpler gastrointestinal tract is possessed by the Perissodactyla, and by the follomng families of Artio- dactyla, the Phachocoeridae, Suidas and TayassuidaB. This consists of a stomach with a single cavity (some Peccaries have partitions but no true septa with strict histological differences) divided into esophageal, cardiac, fundal and pyloric areas, dependent upon the nature of the lining epithelia and the absence or presence of glands, as well as the nature of the tubules. The duodenal section is ample and may be sacculated while the intestines are small in calibre and rather sturdy in wall. The cecum is rela- tively very large, well supplied by longitudinal bands and sacculations; the colon is relatively short but quite capacious and sacculated. The ruminants and other remaining members of the Artiodactyla have a compound stomach suited to the separation of coarse and fluid foods and the retention of water, and so arranged that boluses of different densities are distributed a& needed. These divisions are histo- logically as well as grossly different. The first three, comparable to the esophageal section of the simpler stomach, are reservoirs or channels, while the fourth or true digestive section, is divided into areas corresponding to fundus and pylorus, possessing the appropriate type of gland. The duodenum in this group is narrow, as is the rest of the small gut, and has delicate walls. The cecum proper is short and of variable width, but never as great as in the group first discussed, while the colon, an intricately wound tube, is narrow and very long. Certain 196 DISEASE IN WILD MAMMALS AND BIRDS of the first group (Peccaries )have a colon of this type, but it is not so complicated as in time ruminants. These complications seem designed to permit of a long retention of coarse food of low nutritive value per bulk for a time sufficient for full digestion ; reverse adaptation of large size of stomach and colon may be explained on the same basis. Let us now examine these two groups to dis- cover their pathological reactions and the nature of the lesions. Ungulates as a class stand in a position equivalent to the Primates in the incidence of gastrointestinal diseases. They show a conspicuous percentage of cases apparently due to incorrect food and, coupled with this, a high per- centage of gastric involvement, being exceeded in this respect by the marsupials only (there is an adequate reason for this — see below). Cases ascribed to bacterial Table 12. Showing the Percentage Incidence of Various Forms of Inflammation in the Alimentary Tract of Ungulata. Lesion Group A Periasodactyla and Swine, Wart Hogs, Peccaries Per cent. Group R Ruminants and Relatives Per cent. Acute fermentative gastritis All other forms of gastritis Acute toxic or fermentative enter- itis 18. 16.3 1.6 1.6 0. 0. 2.1 5. 2.3 All other forms of acute enteritis . . . Chronic enteritis 8.4 1.9 Cohtis and typhlitis 3.1 or parasitic agents are not numerous. Analysis of the records of the two groups discussed above brings out some interesting facts. Consultation of Table 12 shows percentage comparisons, based on the number of autopsies, of lesions in the various sections of the tract. Group A, that with the simple stomach and the short capacious colon, is represented by forty-four specimens, while Group B embraces 321 animals. Perhaps the use of these two widely differing figures for comparison is open THE ALIMENTARY TRACT 197 to some objection which might be final and decisive were not the figures so definite. At a glance one can see that Group A has involvement more marked high in the tract while Group B has more disease in the intestine and colon. Certainly gastritis is more common (five times) in the A than in the B group, while enteritis is more common in B (over three times). No case of chronic enteritis or of involvement of the colon is recorded in animals with a simple stomach and a wide hind-gut. This may be read either in terms of vulnerability of the stomach or in the degree of resistance of the respective groups. Consideration of the local factors of the stomach brings to light at once the fact that incorrect food enter- ing the simple stomach could attack the softer, less resistant glandular section of the fundal and pyloric areas whereas the. rumen and psalter of the compound organ, with their stratified epithelium devoid of glands, act as barriers or as places where detoxication of irri- tants might take place. In both groups bulky food is packed to the left, the esophageal and cardiac section in the simple form, the rumen in the compound. Soft or liquid food may pass into the psalter and abomasum of the ruminant stomach almost directly since it has not the force or bulk to push aside the valve-like fold of wall at the junction of esophagus, rumen and reticulum. For this reason, if for no other, the character of soft food supplied to this order must be unexceptionable. It has not been possible to follow out the layering of diets as Scheunert did when showing the course of various foods before they are mixed at the beginning of the pyloric compartments. We have seen two cases in ruminants which seem to indicate that soft food had passed into the right side cavities of the stomach, there to cause irritation, while the rumen remained quite normal. It seems, however, accepted by veterinarians that exces- sive soft food may be followed by trouble in the digestive stomachs, while excessive dry food may cause distention 198 DISEASE IN WILD MAMMALS AND BIRDS of the left hand sections. So far as I know, the relative incidence of gastric disorders in the above outlined groups has not been pointed out before. The pathological types of gastroenteritis do not pre- sent many unusual features. Simple injection of vessels during digestion seems more evident in this than any other order, while mucus production seems less marked. The mucosa of the reservoir portions may, in simple over- filling and fermentative distention (gastric tympanites), be quite normal or dull red ; when active fermentation has occurred it may be digested and peel off. More or less true inflammation as seen in the digestive stomachs is definitely better expressed in the compound ruminant organ than in the simple equine variety where congestion and edematous swelling with hemorrhage form the usual picture. True catarrhal changes both gross and minute, are often well seen and ulcerative lesions are not uncommon; ulcers are unusual in the simple stomach. These statements hold good also for the small intestine. Enteritis affects the duodenum and jejunum more in the Perissodactyla and swine than in the ruminants. Ungulata do not react with hyperplasia of the mural IjTuphatics as do many other orders, but the swelling of the mesenteric nodes is often noteworthy. As might be expected this is more definite mth catarrhal changes and therefore best seen in the ruminants. In the colon, how- ever, the solitary follicles are often quite prominent in simple inflammation. Histologically the greatest changes seen in this order are superficial degenerations with mononuclear increase in the deep submucosa, mostly arranged in perivascular fashion. The villi do not con- tain the large number of cells seen in the Carnivora. Maksupialia. Consultation of the figures for this order in Table 11 arrests attention at once because of the peculiar percentages found under the detailed factors. THE ALIMENTARY TRACT 199 Thus food is of no moment as a disturber of the alimen- tary tract, while bacteria and parasites are high in importance. The gastric segment is more often attacked than any other portion and slightly more often than in the nearest order, Camivora. These unusual figures deserve explanation, to which purpose it will be necessary to con- sider the anatomy of the organ and to discuss why bacteria and parasites stand so high in etiology. Marsupials are divided into six families in our classi- fication (see page 44) which in regard to their diet, range from largely carnivorous (the first two) through those choosing mixed insects, fruits and vegetables (the second two) to those eating vegetables and grain (the last two). The stomach of these animals does not vary exactly according to their diet, the first four, opossums, dasyures, bandicoots and wombats, possessing an organ closely similar in outline and construction and resembling the carnivorous variety, while the phalangers and kangaroos have a stomach entirely different from the first four although somewhat similar to one another. The first group has a round or irregularly elliptical organ with the esophagus and pylorus close together along the lesser curvature. The wombats have a bank of glands sur- rounded by a capsule, near the cardia. The stomach of the first four animals is divided into cardiac, fundal and pyloric parts by the construction of their mucosa, the first mentioned division being a high, rounded pouch, rumen-like, well to the left. The phalanger 's stomach is more elongated, the two openings well separated and a fissure is found in the right end of the lesser curvature which serves to separate the pyloric part from the rest. The Macropodidffi all have a stomach resembling the human colon in being elongated, with longitudinal bands gathering it into sacculations. There is a distinct eso- phageal section to the left with a blind sac suggesting an ungulate rumen, a long tubular fundal, and a sacculate pyloric division. 200 DISEASE IN WILD MAMMALS AND BIRDS The small intestine of the order starts with the opos- sums as a stout muscular and mucous tube fitted for meat eating, but as one proceeds to study the families do^vIl- ward in the list this tube becomes more delicate and longer. In the first two families the cecum is rudimentary and the colon very short as in cats, but the length and capacity of these parts increase through the bandicoots and wombats until in the strict fruit, vegetable and grain eaters, phalangers and kangaroos, the cecum is Table 13. Showing the Incidence of Gastroenteric Disease in the Two Forms of Marsupial Intestinal Tracts. Group A Carnivorous Stomach and Intestines 103 Specimens Per cent. Group B Herbivorous Stomach and Intestines 73 Specimens Per cent. Acute gastritis Chronic gastritis. . . . Acute enteritis Colitis incl. typhlitis Bacteria 0. 9.6 2.9 1.9 Verminous 13.6 15. Bacteria Verminous 5.4 0. 1.5 4.3 long and capacious and the colon relatively long and roomy. Table 13 contains an analysis of the forms of gastro- enteritis as they were described in the two groups of tracts, that resembling the carnivorous, that similar to the herbivorous intestinal construction, and according to the factor believed to be responsible. In the first group gastritis of an acute nature occurred in 9.6 per cent, of the 103 specimens. They were chiefly catarrhal in charac- ter and seem for the most part secondary to verminous infestation ; at least six of the ten cases were associated T\dth parasites. The process microscopically is catarrhal and deeply infiltrative. Group B has a high incidence of gastritis and here the evidence of bacteria or toxins is quite plain. Several of the cases were in animals showing also Kangaroo disease of the jaw Avith pneumonia or septicemia; the gastric lesion of streptothricosis will be described under that heading. The character of gastritis THE ALIMENTARY TRACT 201 without jaAV disease is somewhat different from that with it. Pathologically the process is a congestive and super- ficially necrotizing affair, forming upon the tips of the folds, small gray erosions or flat shallow irregular ulcers, which upon histological study consist of loss of tissue of the mucosa and some deep congestion with round cell groups but no reaction deep in mucosa or submucosa. True catarrhal inflammation has occurred, but not like in the opossums. Chronic gastritis in the simple stomachs is almost exclusively in opossums harboring Physaloptera turgida, a worm which fastens itself more or less firmly in the mucosa and probably, with the assistance of bacteria, causes sufficient irritation to produce a hypertrophic change in the deeper layers and a destruction of the glands where it holds and a distortion of those nearby. One is reminded that Fibiger found spiroptera to be responsible for adenocarcinoma in rats ; no tumor forma- tion has been found in these animals, although one opossum with such a stomach had an adenocarcinoma mammae. Small hemorrhagic spots may occur in the deeper layers, possibly where the worms have bitten. The rugae are irregular or interrupted by knobs and papillae. Group B, stomachs showing chronic change, were all kangaroos. The three cases resembled the infiltrating necrotizing lesions as discussed under ulcers (page 175). The process showed an infiltration of the subsurface tis- sues with a gray slough over the densest part. The mucosa as a whole was irregularly rugous and spotted with red gray areas. Altogether one gets the impression that in the simpler stomach, reactive inflammation is most prominent, while in the colonoid stomach degeneration is greater than reaction. Intestinal lesions in marsupials are not common and not peculiar except in that they carry out the pathological 14 202 DISEASE IN WILD MAMMALS AND BIRDS reaction as seen in the stomach. The carnivorous intes- tine shows frank catarrhal changes, the herbivorous presents congestions and superficial necroses. These two groups then follow the descriptions as already given for carnivores and ruminants. The colon presented in the first group, simple catarrhal or follicular change. In the second division the lesions resembled those in the stomach; they were only once of the three instances associated with streptothricosis. The class Aves has been treated in the chart precisely as have the mammals, but it is not possible to make the accurate diagnoses or to separate groups of lesions ac- cording to anatomy as was done for the latter class. Upon the average there is more gastroenteritis among birds than manamals, but when looking for an explanation of this, it was unfortunately necessary to enter in the ''unde- cided" column of Table 11 a very great number of cases. This column would be greater relatively were it not for the parasites in parrots and gallinaceous birds, the entries for which are high. Besides this fact and that the owl stands at the top, the pigeons at the bottom of the list numeri- cally, no further general statements are allowable. There are several orders with high values among those of which insufficient autopsies are at hand to cast percentages. The avian alimentary tract seems to have developed according to the character of food the different varieties of birds consume if one may judge by the construction of the bill, the gastric musculature and the intestinal waU. Zoological classification has only secondarily considered this point, it being made incidental to the habits and habitats of birds. For this reason the orders as outlined on pages 44-46, placed as they are in historical evolution- ary position, represent with few exceptions groups which have differing diets and, by the same token, differing gas- THE ALIMENTARY TRACT 203 trointestinal tracts. Classifications based upon habits of life (Raptores, Cantores, Natores), prove likewise too broad or too heterogeneous, while systems making character of food the chief criterion though apparently correct in reasoning and helpful in certain orders, (Accipitres, Galli) are found to present copious excep- tions ; moreover we are imperfectly informed of the exact diet that many families require or resort to in absence of their preferred food. I shall therefore discuss the chief diseases and distributions according to our classification, preceding the discussion by a brief resume of the ana- tomical peculiarities of the avian alimentary tube. The first digestive burden falls upon the proventricle where the principal juices are secreted while the muscu- lar stomach or gizzard assumes the duty of gastric mastication. The lateral muscular bellies of its heavy wall grind the food and mix well the gastric juices. Its mucosa probably supplies only lubricant. In birds whose food is hard, com and the Hke, this grinder is supplied with a dry horny internal layer, while a thick, moist, soft, epithelial surface is sufficient for carnivorous birds. All kinds of gradations exist between these extremes. The mucosa of the proventricle is always soft, but quite deep to permit the placement of compound tubular glands.;, The relation of size of these two parts is subject to many variations. (1) The proventricle is larger propor- tionately in meat eating, fish eating and fruit eating birds, the gizzard having the greater size in granivora and insectivora. In certain birds the mucosa of the two is separated by a very soft thin zone, an important fact in Psittaci since at this place spiroptera seem to penetrate to the glandular layer of both organs. The duodenum begins in practically all birds, from a spherical cavity at the pyloric end of the gizzard, to be accredited anatomically to both sections. It passes down- (1) See Magnan, Compt. Rendus d. I' Acad, de Science, 1910 and 1911, Vol. 150, 151, 152. 204 DISEASE IN WILD MAMMALS AND BIRDS ward, forms a long loop enclosing the pancreas, its distal end lying under the liver and near the gall-bladder. Near its end it receives the major bile and pancreatic ducts; smaller ducts from the liver and pancreas may enter near the pylorus or elsewhere along the loop. The small intestine is usually simple in its coils, but in the birds that eat grain, grass and greens, may be long and compli- cated. So too the colon, usually a very short segment, may be increased in the just mentioned group while the ceca are only of any considerable length in herbivorous birds. The length of the ceca is, according to Owen, related to the availability of food and the need the bird may have for exhausting the nutritive value of it. In carnivorous birds as in similar manomals, the whole gut, but especially the hind-gut, is very short and the ceca small or absent. But so they are in picarian birds which are chiefly herbivorous, but may eat meat. Gasteitis. The double-muscle stomach, that with the two lateral plates and tough epidermal internal coating, is seldom the seat of disease. An excess of greens in the diet some- times seems to soften or macerate the hning, while an excess of pebbles may cause erosions. Upon severe irri- tation this internal layer assumes the appearance of tanned leather and may crack. The proventricle of such a gizzard seems rather resistant to disease, particularly one would say, to infective processes, for catarrhal or ulcerative inflammation is uncommon. The saccular stomach with uniforai muscular walls continuous with those of the proventricle, such as is seen in raptatory birds and parrots, offers a somewhat different picture. The internal membranes of these organs are definitely softer, seeming to swell with great ease, and the glands themselves are smaller both at the fundus and outlet, a construction which may favor their closure by swelling from simple congestion and edema. Catarrhal and ulcer- THE ALIMENTARY TRACT 205 ative processes are definitely more common in sucli organs than in the first type or true gizzard. Enteritis. Enteritis of all orders is most outspoken in the duo- denal loop, but a determination of the lesion must be made with some care. All the signs of intestinal inflamma- tion — congestion, swelhng and opacity, excess mucus or mucopurulent covering — must be present to justify a gross diagnosis of enteritis and even when these exist one fails at times to confirm the finding by microscopical sec- tion. These changes may be simulated by digestive activity so that it is but reasonable to demand them all in a clear cut fashion before applying the term enteritis. However, it is well known that cases in both human and veterinary medicine giving a satisfactory clinical picture of this disease may fail to show to the naked eye and under magnification the changes expected. In the hmnan being, the carnivore and the ruminant, the ileum presents the most definite picture of enteritis. In the bird, the duodenum shows the prominent lesions, and with the exception of specific diseases like blackhead, is always involved when the smaller coil of small intestine is affected. This is true whether the enteritis be non- specific or be associated with cholera of chickens, ducks or parrots. The colon presents peculiar lesions in but few birds. Ulcerative and necrotizing processes have been encoun- tered in three orders, Psittaci, Anseres and Herodiones, suggestively like the specific forms seen in the ceca. Microscopy and one unsuccessful culture failed to reveal a mould or protozoon. The condition appears as a gray white plaque in the cloacal wall or it maj^ spread up to the colon and around the urethral orifices. At times it is superficial upon the mucosa and may be covered by a pseudomembrane. 206 DISEASE IN WILD MAMMALS AND BIRDS The chief interest in the ceca centres about entero- hepatitis either of heterakis and amoebic origin or that supposed to be due to coccidia or Bac. scoticus^ This specific form has been encountered only in Galli (three of the four families). It has been so well described by Hadley, Smith, Morse and Cushman that it is unneces- sary to discuss it since we have nothing to add to its pathogenesis or pathology. At a later time some atten- tion will be given to our experience with Quail disease. (What is more interesting from a comparative stand- point, besides having a bearing upon blackhead, is the discovery of heterakis in the ceca, and hemorrhage and fatty change in the liver without amoebas or coccidia in either place (unfortunately no bacteriology was done), in a Sebastopol Goose {Anser domesticus), a bird which has ceca not unlike the gallinaceous varieties. This is a single observation and must be treated expectantly. Microscopically the avian digestive tract in its various inflammatory states presents a few noteworthy features. The primary reaction, sometimes the only one, to irrita- tion is injection of the vessels in the villi or deeper mucosa. To this, however, is nearly always added a granularity of the epithelium, without much evident mucus (goblet cell) formation. When the epithelial degenera- tion is marked there appears a round cell increase in the deep mucosa shortly followed by a similar infiltration into the villi. True catarrhal enteritis as described for the cats is not as common as some combination of the changes just detailed, but when it occurs is best developed in the carnivorous avian tract. The most striking cellu- lar finding is the round cell of the infiltrate. It is of the middle lymphoid size with clear protoplasm, or, when late in the disease, may be small and so-called adult. Poly- nuclears, unless eosinophilic, and endothelioid cells are rare. The foregoing are general remarks concerning the pathology of the avian tract, and we are now ready to THE ALIMENTARY TRACT 207 discuss briefly the reactions of the orders. I shall, how- ever, omit mention of those in the right half of Table 11. Passerine birds are represented better than any other order because they are more numerous in nature and therefore exhibited more generously in collections. This order is not especially vulnerable as to its alimentary tract, but this tube is often affected by tiny stones, bits of rust from cages and by acute general non-specific infec- tions to which these birds are quite susceptible. Upon many occasions intense congestion with and without tiny hemorrhages in the duodenum are all that can be found in the little birds and the diagnosis of enteritis is put down. Only about one-tenth of these birds have shown more or less definite catarrhal changes. Many birds have heavy infestation with worms which may activate bacteria. The Striges and the Accipitres will be discussed together because of the similarity of their tracts and their diets. The reaction of their gastric complex has already been mentioned and what was written there can be extended to the intestine. The type of lesion is catarrhal and seems to be "meat bred" although this cannot be proven. They never have given positive heart's blood cultures so that the disease seemed not to be septicemic. Liver and spleen have harbored colon and paracolon bacilli. It will be noticed that they have the highest incidence of gastric disease. Psittaci, birds with a tract similar to the last two but with a captive diet of seeds, fruit and vegetables although they may eat small animals and insects in the wild, present figures under bacteria and parasites which explain the involvement of the alimentary organs. We have had two acute outbreaks of what seemed to be fowl cholera, judging by the pathology and the isolation of Bad. gallinarum and we have frequent deaths with the same gross anatomy from which bacterial isolation has not been tried or was unsuccessful. At all events the enteritis of parrots is often a definitely infectious affair. 208 DISEASE IN WILD MAMMALS AND BIRDS We have also isolated Bad. psittacosis on two occasions. Separation of the birds and cleansing of the exhibition spaces seemed to stop the disease. The lesions are hemorrhagic and superficially necrotizing in this group whether or not they are septicemic. Some have also sho^vn a follicular appearance both grossly and minutely, one of which cases was associated with Bad. psittacosis. The effect of animal parasites is chiefly exerted, for this group, in the proventricle where the Spiroptera incerta occupies the lumen and penetrates the mucosa or burrows under the inner lining of the gizzard. Enteritis is not especially associated with this infestation, death resulting more from inanition than infection; some few cases have had enteritis, others pulmonary disease. Columbai are not susceptible to disease in the parts under consideration. This order seems to have some tissue resistance, for their lesions are quite frankly catarrhal, more so than in most grain-eating birds. The figures set against the Galli are swollen by the number of cases of enterohepatitis of one sort or another. Extracting these from the total leaves the order among the lowest. Their lesions are congestive and hemorrhagic, although they may show catarrhal cases. They seem to be able to summon mucus more readily than many other birds. Anserine birds, though not very high in figures, pre- sent three conditions worthy of note. In the first place, acute simple gastritis occurs often, sometimes associated with foul green food, sometimes without any apparent cause. From the number of times that foreign bodies are present it seems probable that they contribute in some Avay. Excessive stones and sand, bits of glass, collar buttons and the like are sometimes found. Then the form of acute enteritis has always a hemorrhagic tendency, at least in the submucosa, while the mucosa may be swollen, opaque and covered with mucus. Upon histological study these intestines show intense swelling by cellular infil- THE ALIMENTARY TRACT 209 trate and disappearance of the tips of the villi. The third observation concerns what is apparently a subacute or chronic process although this is not supported by micros- copy. Certain birds will have a cast of mucus and epithelial detritus rather closely adherent to the wall. Under the microscope there may be slight evidence of chronic inflammation or there may be little amiss. These birds have usually been large ones, and several have come from the separate goose pens, not from the open lake where many birds are kept. The struthious birds deserve a word. They have had a great deal of enteritis and mostly of infectious nature. Two instances have arisen from bird diphtheria, one from cholera and six from what later seemed to have been anthrax but was not diagnosed at the time. The character of the lesions in the struthious intestine tends to be hypertrophic and superficially erosive if not ulcerative. The changes are found with greatest clearness in the lower duodenum and small coil. Constipation. (Having discussed the inflammatory conditions of the gastrointestinal tract we now come to the more or less definitely mechanical abnormalities, whether or not they depend upon preexisting inflammation, and the subject of constipation will claim first attention. In the human Jbeing this condition is the result of bad habits more than any other one thing or all things together, I think it will be admitted. In the lower animals perhaps no such thing as habit of defecation exists so that one can with more certainty hold incorrect food, chronic catarrhs or physical obstruction as responsible. Veterinarians look upon excess of dry food and irregularity of work and food periods as the principal causes of constipation. These factors do not hold in zoological collections. As a matter of fact constipation is of minor importance in this menagerie, but a certain few cases are worthy of note. It 210 DISEASE IN WILD MAMMALS AND BIRDS has been mentioned in the diagnoses in only a little over 1 per cent, of the total, and of these the records indicate its importance only ten times (.2 per cent.) ; a few notes of these cases are appended. The first place of incidence is taken by marsupials (six kangaroos and one opossum), the second by ungulates (largely ruminants) and the third by Primates. It will be noted that with exception of the opossum, herbivorous mammals occupy the first places of incidence, carnivores falling well behind the orders named. This condition is quite infrequent in birds and is usually associated with the presence of seeds or parasites or with impaction in the ceca. Primates, almost exclusively feeders upon carbo- hydrate and soft protein food, have shown as causes of constipation two outstanding conditions. A low grade of colonic catarrh with excessive pouchings of this tube has had constipation associated with it three times. One of these cases had small coproliths in the diverticula, one other a fecal concrement in the cecum. Another group of these cases with evidence of delayed passage of feces shows chronic peritonitis with adhesions, one of which seems certainly due to filaria in the peritoneal sac. The seat of constipation in monkeys is practically always the colon. The carnivores while occasionally showing hard fecal masses packed into the colon, more often exhibit a constipation in the ileum. One case pre- sented a nearly empty colon with a long scybalum just above the cecal valve. There is no peculiar associated pathology in the notes at my command. Ungulata, showing next to the highest incidence, has its stoppage chiefly in the colon, but the lowest stretch of the ileum may contain balls of feces. In nearly every case one finds some grade of colonic catarrh. In two instances, there being a proctitis, it seemed as if the animal volun- tarily restrained from defecating because of pain. The caput coli is the seat of stoppage in the Rodentia. THE ALIMENTARY TRACT 211 Marsupials give such a high relative incidence that especial search of their records was made, Avithout, how- ever, very definite result. In three of the seven cases an acute general infection existed, in one an acute peritonitis which seemed to emanate from a small ulcer in the ileum, in one an injury to the anal region was found while in the remaining two the notes would suggest that the lower intestine was atonic, judging by its distention, trans- lucency and pallor. In five the stoppage took place in the large bowel alone, in the others both divisions being affected. It is often difficult to establish a diagnosis of consti- pation in birds because many varieties form a long rather dry mass in the lower small intestine, to be moistened in the cloaca for discharge. Still again the groups with capacious ceca are apt to have them filled normally with firm casts. Diagnoses of fecal inspissation and stoppage in the smaller tube have been made seldom, but one must consider also the obstruction offered by excessive urate collections either in the cloaca or lower ileum which will amount to a constipation if the cloaca be over-dilated and dried urates mixed with dirt or feathers cover the anal opening. The causes of this condition in birds are usually mechanical, inflammation being found in a small minority of cases. In the small passerine birds, seeds, sand, or parasites form the commonest findings. This is also true of parrots, while excessive urate collections are noted for both these groups. The gallinaceous birds present two reasons for fecal stoppage — disease of the ceca (see pages 205-6) and cloacitis probably secondary to anal closure by excessive urate collection. Uratic stones, varying from one to five millimetres in diameter, have been found in the cloaca in several orders. In only one case, a pheasant, did they cause ulceration and cloacitis. Sand, rust, grains and the like are found frequently, and sometimes in groups of birds, indicating that the speci- 212 DISEASE IN WILD MAMMALS AND BIRDS mens had not been put upon the correct flooring or caging. Unbroken seeds may obstruct the lumen.y Mechanical, Obstruction. Although the following is not constipation it is well to cite at this place an experience which amounted to mechanical intestinal obstruction. A number of finches were subjected to postmortem and found to have whole white millet seeds in their intestines, this being the only discoverable cause of death. Investigation revealed that during the night mice ate the canary seed in the pans, leaving only the millet, w^hich the hungiy birds consumed w^hole. Small birds can take a few millet and crack them when eating leisurely, but apparently not when hungry. When the food was removed at night the trouble ceased. Obstruction by sand is well illustrated by a peculiar form of pica, in a goose, which is worth citing, and calls to mind the sand disease of horses : Canada Goose 6 {Branta canadensis canadensis). Diagnosis. — Masses of sand in entire intestinal tract. The general condition externally and internally is good. The crop is distended like a sausage, quite firm and the overfilling is obviously due to sand in which veiy few stones, Avhieh could be called pebbles, are found. This mass continues into the esophagus making the whole tract impassable for food. The mucosa is a little pink and dirt-stained in places but is not visibly inflamed. The gizzard is contracted over a mass of sand but no food. Sand in more or less definitely packed condition is found all along the gut tract, in one place in the small coil it being quite as tight as in the crop and no lumen remaining. Sand and bits of shale are found in ceca. The organs are apparently healthy, slightly pale perhaps, but certainly not distinctly anemic. No infection exists. The aorta, just above renals, has a 15 ram. x 2 mm. pale opacity of same consistency as the rest of the vessel, just perceptibly higher than sur- rounding surface. *'Sand disease" has occurred in a Persian Wild Ass (Equus onager) causing in this case ulceration, perfora- tion and peritonitis, a Common deer {Mazama virginiana) and a Chapman's zebra {Equus hurchelU cliapmani). The collection of sand is always greatest in the caput coli, but may coat the large bowel to the anus. THE ALIMENTARY TRACT 213 Larger and more definitely obstructive physical objects are found in both mammals and birds. We have on record a lion (Felis leo) and a tiger {Felis tigris), which swallowed pieces of bone large enough to be stuck in the small intestine and completely occlude it. Smaller objects like buttons have been found even in the passerine tract. Worm masses may occupy such a large part of the lumen of the tube as to constitute a physical obstruction. This is definitely less important in mammals than in birds, especially in the passerine order of the latter class. Dilatation of the intestine aside from that occurring in connection with fermentation, constipation or ileus, in other words chronic atonic dilatation, has not been encountered. Acute dilatation has been found in several orders under the picture known for domesticated animals. Its pathology and incidence have already been discussed. Ileus. (^ Ileus or acute intestinal obstruction may be divided for our purposes into intussusception, volvulus, strangu- lation and paralysis from interruption of mesenteric circulation. Examples of all these varieties have been encountered and illustrative cases will be cited. In so far as incidence is concerned, the Ungulata and Carnivora greatly outnumber all other orders, showing seven cases each ; the sum total in all other orders is but eight. Upon re-reading some of the protocols I have, however, excluded three invaginations in the carnivores, one each in the ungulates and rodents, as probably being post- mortem or shortly antemortem occurrences ; two had very early peritonitis but other things, sufficient to account for death, were present. These deductions bring the total cases of ileus in mammals to seventeen. Five cases in birds will be discussed briefly. Primates present one case of volvulus, one of intus- susception and one of internal strangulation. The first displayed the entrance of four inches of ileum into the 214 DISEASE IN WILD MAMMALS AND BIRDS colon with such swelling of the waU as to prevent reduc- tion. The exciting cause seemed to be an enteritis, the cause of death a peritonitis. A white-collared mangabey {Cercocebus collaris) was the victim of volvulus probably favored by an anomalous position of the transverse and descending colon which lay to the right, the latter traversing the abdomen obliquely from right to left to reach the pelvis. The volvulus occurred in the ileum just above the cecum, the tmsted part being found adherent by the peritonitis. The third case is a strangulation due to peritonitis from filaria and adliesions between stomach and colon from a colitis and pericolitis due to cestodes, one of which was found deeply implanted in the colonic wall. Two cases of intussusception are noted (after deduc- tions above) for the Carnivora. They both occurred in the ileum, one restricted thereto, the other extending into the colon. In both a vague history of being '' off their feed " or giving evidence of intestinal trouble could be obtained from the keeper. The three excluded cases had invaginations in the middle and lower small intestines but not at the cecal valve. Volvulus did not occur in the Carnivora. Aparadoxure {Paradoxurus hermophroditus) died as the result of a strangulation of a six-inch knuckle of gut which had passed through a hole in the omentum. The animal had not been eating well for a month but gave no signs by which this ileus could have been diagnosed. Per- haps it had existed for sometime but only shortly before death had swollen sufficiently to cause obstruction. Having excluded a doubtful invagination in a small rodent there remains an interesting though somewhat obscure case in a porcupine {Erethizon dorsatus dor- satus). This animal suffered with an acute hemorrhagic and catarrhal enteritis while the colon seemed free of change until the rectum was reached. Here was a stretch of a foot with the purple, lusterless but translucent THE ALIMENTARY TRACT 215 appearance of a strangulated intestine although no invo- lution or twisting remained. This was looked upon as a volvulus which had untwisted a few hours before death. Intussusception was seen only once in the Ungulata, a tapir (Tapirus terrestris) with chronic enteritis. Here the ileum had passed into the colon for a distance of nine inches, it being much swollen and congested but not gangrenous. Its condition warranted the idea that the process was antemortem but a peritonitis had not arisen, death having occurred from the slight extra shock in an animal suffering with chronic malnutrition. Volvulus was encountered three times, two deer and a zebra. The last was the animal already described that carried such a heavy load of sand in the gut tract, a factor in the produc- tion of the twist probably although this might have been aided by a fibromyoma of the colonic wall. The location of the volvulus in this order was twice in the dilated descending colon, the third in the jejunal area. This last was a twist which resembled an internal strangulation because of the intricate knot-like windings of the smaU bowel. The marsupials present two interesting cases. A rock kangaroo {Petrogale pencillata) had chronic gastric ulcerations with local peritoneal adhesions which appar- ently obstructed nearby coils of intestine so that they became inflated and twisted over. An opossum had a vol- vulus of the stomach which performed one and a half turns from left to right ; its protocol follows. Common Opossum 6 (Didelphys virginiana). Ileus. One and one- half complete volvulus turns of stomach on duodenum. General condi- tion fairly good. Abdomen quite prominent, a condition found to be due to great dilatation of the stomach which occupied the whole anterior part of the abdominal cavity. The organ is blue and the vessels stand out. Postmortem changes are occurring everywhere favored by the obstruction to the circulation. The dilated stomach has undergone a volvulus upon the third part of the duodenum making one and a half turns. The spleen lies upon the right side well below the liver; it is swollen, soft and deep purple. The duodenum in its upper half takes part in the dilatation and beginning gangrene. The pedicle of the 216 DISEASE IN WILD MAMMALS AND BIRDS twist is made of the duodenum, esophagus, edge of the mesentery and the middle part of the pancreas; the end of the tail of the last is gan- grenous. There is no apparent obstruction lower down to explain the twist of the stomach. Among the Aves the following cases only are worthy of report. A parrot {Melopsittacus undulatus) was found to have a tightly packed mass of worms in the end of the duodenum above wliich the bowel was distended, enlon- gated, doubled on itself and of a deep red color ; below this the small intestine was empty. A closely siioilar con- dition was found in a Screech Owl {Otus asio asio) the obstruction occurring just above the end of the small gut. A Sparrow Hawk {Falco sparverius) had an invagination two cm. in length, a short distance above the end of the small intestine. No peritonitis existed but the presence of an acute enteritis helps to explain the intussusception. Hernia. Hernia is not a common occurrence among the lower animals but our experience is instructive in two partic- ulars, to wit, its absence in the orders preceding the Eodentia and the frequency of the traumatic variety. There being no general remarks to be made upon the sub- ject, it seems well to give a summary of the findings in each of the seven cases. A Western Fox Squirrel (Sciurus rufiventer) showed a diaphragmatic defect on the right side, a, rounded open- ing with smooth edges, through which a loop of intestine had passed, entering behind the liver and reaching into the pleura as high as the pulmonary apex where it was adherent ; this was probably of long standing. Two more loops were found wedged in the diaphragmatic hole, one of which was gangrenous. An Indian Antelope {Black Buck) {Antilope cervi- capra) presented an irreducible incarcerated but not strangulated umbilical hernia. The peritoneum was fused THE ALIMENTARY TRACT 217 with the aponeurosis at the ring but the gut was not adherent at this point while it was attached within the sac outside the muscle, thus forming the incarceration. Apparently the sac had dissected between the muscular layers for it could be followed for several centimetres in some directions. A Hog Deer {Cervus porcinus) had apparently suffered an injury in the flank for at one point the muscles were irregularly cicatrized and a rent was present through which several loops of intestine and a band of omentum had escaped, being adherent to fascia. No injury to the skin was apparent. Another Indian Antelope showed a clean traumatic rupture of the muscle mid peritoneum in the right inguinal region ivithout penetration of skin. An acute hernia had occurred which was lightly adherent to fascia and an acute peritonitis was beginning. The bowel was however not strangulated. An aoudad {Ovis tragelaphus) seems to have suffered an injury hy a pointed object (horn?) just to the right of the ensiform cartilage for at this position there is a circular hole, with smooth healed edges, in the aponeuro- sis, permitting the emersion of a peritoneal sac contain- ing omentum. All parts were adherent but no acute inflammation existed. ( What may have been a hernia or a relaxation of the transversus perinei was observed in an Undulated Grass Parrakeet {Melopsittacus undulatus). A bulge about the size of the finger end was seen externally, beside and behind the anus. This proved to contain several loops of bowel and a mass of fat. A lateral abdominal hernia was seen in a Barbary Turtle Dove (Turtur risorius). It consisted of a per- itoneal sac and two loops of intestine. This protrusion, while firmly fixed in its unnatural position, was in no way constricted. 15 218 DISEASE IN WILD MAMMALS AND BIRDS Rectal, Prolapse. Prolapse of the rectum may in a sense be looked upon as a hernia or at least as a relaxation of the anal and perineal muscles with protrusion of parts normally situ- ated intracorporeally. Although not frequent it has been incurable in the animal, as it frequently is in man without operation, a measure we have not adopted. Just what determines weakness in the pelvic outlet is entirely obscure for indeed we have seen here wounds and inflam- mations of the perineal area \vithout prolapse of the rectum and in none of the cases of prolapse did the pelvic floor seem injured or diseased. It is but speculation to blame the annular muscles of the anus. Tenesmus, or at least reasons for this straining action, have been sought, with the result that in our cases lesions of the egg-laying apparatus in birds and enteritis in mammals have stood out most prominently. In no case have hemorrhoids been encountered nor has a tumor pendant from the colonic mucosa, drawn the bowel toward the anal opening. It might be added parenthetically here that hemorrhoids are practically unlaiown for quadrupeds, Hutyra and Marek failing to mention them independently and only one refer- ence being found in the Jahresbericht fur Veterindr Medizin (Schmidt 1914-169); this case is more like angioma than hemorrhoids. If tenesmus be active in the production of rectal prolapse then it would have to be assumed that this straining effort can be induced by enter- itis since eversion of the rectum has occurred with this disease in the absence of colitis, the condition usually expected in the presence of tenesmus. The thirteen cases have been seen in Mammalia, 8, (Carnivora, 2, Rodentia, 1, Ungulata, 3, Edentata, 1, Marsupialia, 1) and Aves, 5, (Passeres, Picari.T', Striges, Psittaci and Galli each one). Three mammals had enteritis, one had foreign bodies in the bowel and one had many ascarids; three had no demonstrable or suggestive causes. Two of the five birds THE ALIMENTARY TRACT 219 had enteritis high in the tract, one had uratic calculi in the cloaca, and three had trouble in the egg laying apparatus : one too large an egg, one a broken egg and one a salpingitis.; Diverticula. It is almost certain that in a human pathological service of fifty-five hundred autopsies, one or more diverticula of the Meckel variety would be encountered and perhaps several of other kinds. In our material only pouchings or false diverticula of the colonic wall are recorded, and our personnel has often spoken of the absence of these gross abnormalities of the alimentary tract. The two cases, notes of which are given, are instances of hernial pouchings of the colonic mucosa and serosa, a condition which is well known in human medi- cine. C,It may be said to occur in two varities, one in which the pouchings have heavy walls formed by a thickened mucosa, muscularis and peritoneum and on© in which the bulgings have delicate walls, then being small herniae of the inner coats through rifts in the outer. Such a division is probably unnecessary or misleading since the latter may be only a forerunner of the former. However the clinical evidence of the simple variety is scanty and may be little more than constipation while the peritonitic variety gives a clinical picture of pain, constipation and a mass in the left abdominal area, then kno^vn as diverticulitis or pericolitis sinistra. In these cases the colon is much dis- torted by the irregularity of its mucosa and by inflamma- tory thickening of the muscularis and serosa. Diverticula arise from defects of the muscular coat, or second- arily after inflammation or prolonged constipation, by weakness of muscle, or as hernial protrusions around the entrance of blood vessels where the muscle is thin. Such sacculations permit feces to collect and continue the infla mm ation, thus further weakening the gut and pro- ducing constipation, the whole vicious cycle being favor- 220 DISEASE IN WILD MAMMALS AND BIRDS able to the formation of more sacculations; coproliths may form in the diverticula. The two monkeys now reported seem to have varying grades of the same con- dition, a long standing colitis \\ith diverticula, constipa- tion and the collection of inspissated feces in the sacculations. These animals did not have hemorrhoids. Black Ape 9 {Cynopithecus niger). Coprostasis. Coproliths in diverticulum. Chronic colitis. Cor bifida. The large intestine is of the same calibre as the small intestine should be when not distended. The sacculations as seen before opening the organ are salient, forming distinct pouches. In one or two cases they are so pronounced as to constitute diverticula 7 cm. long. In two instances the serosa at the fundi of these diverticula is markedly hyperemie and very thin. In many cases the sacculations contain coproliths. The wall of the organ is distinctly thickened, puckered, inelastic and opaque. Mucosa is thrown up into coarse rugae. Japanese Macaque 6 (Macacus fuscatus). Chronic hypertrophic colitis. False diverticula of colon. The large intestine contains a moderate quantity of quite constipated feces. The serosa is smooth. The wall shows at several stretches enlargements of the normal saccu- lations, forming false diverticula. The wall of the gut in these herniae is thinner than in the surrounding parts; no ulcers exist; no local peritonitis is present. The mucosa everywhere is irregular in thickness, less translucent than normal and thrown into irregular rugaB; tenacious mucus covers it. No ulcers. TUMOES. Only one tumor was observed in the mammalian intes- tinal tract proper. Dasyure {Dasyuriis macidatus) Adenocarcinoma of the intestines. On postmortem there was a pale diffuse thickening of the coats of the small gut over a large area; numerous soft, light yellow, sharply circumscribed, elevated (like secondary tumors) nodules in the liver and spleen, and a pea-size whitish nodule around a bronchus in the right lung. Histological section of primary growth not made but a cross section of the intestine in the vicinity shows an adenomatous change with considerable increase in the connective tissue. The nodules in the liver, spleen and lung and the appearances of the abdominal IjTnph nodes, found microscopically, are precisely similar. They consist of irregnilarly arranged THE ALIMENTARY TRACT 221 epithelial nests and distorted acini, around which are sharply outlined spaces, filled with the remains of de- generated blood or a granular material. The metastases are always sharply outlined. Aves supply three papillomata which are interesting in that one occurred in the proventricle, and two grew in the duodenum in the vicinity of the upper biliary opening and presented within the lumen soft masses which, while not occluding the passage, offered some little obstruction as indicated by a slight distention above their location. In two, carefully studied, no indications of parasites or of cancer could be found. The birds concerned were an amazon, an owl, and a rhea. SECTION VII THE ALIMENTARY TRACT, PART 2. THE LIVER A consideration of the liver is anatomically and physiologically the next step in the discussion of diseases of the alimentary tract. While this organ may partici- pate in most of the pathological states of the tubal part of the system, it is comparatively seldom the primary seat of change and when damaged seems to be possessed of great accommodative and reconstructive power. This must be true, and fortunately so, since we ascribe to it the major detoxicating function of the body. Neverthe- less it is noteworthy that the largest solid organ of the animal body shows a relatively low percentage of changes threatening to life. In the sense of Pearl's method of statistics, it does not '' break down " easily. In pathology it is the custom to list with great care all the changes, gross and minute, in the liver, but with a few exceptions they are secondary or incidental. They do however reflect many things, especially referable to diet and to chronic infection from the intestinal drainage area. It is in these directions that the organ will be studied in the following pages. Anatomically the liver is situated in the right upper part of the abdomen subjacent to the diaphragm in both mammals and birds, being held in position by attachment to this transverse partition, by ligaments or folds of peritoneum, and by the other abdominal viscera. Its general relationships do not offer great variations since in all animals means are afforded for a dual blood supply and an outlet for the hepatic secretion, the bile, into the higher intestines. Naturally variations in the size of the lobes are observed and there has been considerable spec- ulation as to their independence and association. From the standpoint of comparative pathologj% little can be 222 THE LIVER 223 ascertained to assist in this matter unless the position of abscesses and hepatitis relative to cholecystitis have a bearing; some discussion of this will appear later. We have not observed any peculiar pathology of the lobes of Spigelius and Riedel. In so far as the size and arrange- ment of the organ is concerned a few general facts of significance may be mentioned. It was formerly thought that the liver varied inversely as the size of the animal but Magnan (1) and others have shown that the matter is not so simple. In the first place if there be an actual mathematical formula it is that the liver varies in size inversely as the surface area of the body, but this is not the whole story. It seems that the relation of size of the organ to its weight is not constant and that it is better to judge of the organic capacity by the latter. In herbivorous animals, both birds and mam- mals, the liver is lightest per kilo of body weight ; next in weight are in order, fisheaters, meateaters, insectivora, seedeaters, fruiteaters and omnivora. There is besides this a roughly inverse ratio between the size of the liver and the length of the intestine and in the class Aves inversely as the size of the lungs also. From the im- mediately foregoing statements it is apparent that a bewildering variation occurs and that only rough measurements of the relative volume of the liver are avail- able. An attempt was made in the Marsupialia, which present all the variations given, to discover if any peculiar pathology corresponded with the above groups ; as it was fruitless, no change from our zoological treatment will be made. Lobar arrangement varies from the relatively simple double avian type to the manifold lobulations of the seal or the marsupial but I can find no literature to indicate that lobes or lobulations have a direct effect upon func- tions. There must be a difference of blood supply for in (1) C. R. Soc. de Biologie, Paris, T. 73-526. Bull. Mus. Hist. Nat., Paris Ann., 1911, 492 et seq. 224 DISEASE IN WILD MAMMALS AND BIRDS certain infectious diseases like enterohepatitis and amoe- biasis, the cystic and extreme right lobes are more affected than the left parts of the organ. In the bird this is not so difficult to follow since the three divisions of the portal vein, while they combine at times in an ampulla within the hilum of the liver, seem directed to certain lobes, that from the left portal seeming to point toward the right side. The avian portal system differs from the mammalian in having a large branch from the renal area, the so-called renal-portal system, pass to the liver, and by having a free anastomosis between the portal area and the caudal vena cava whereby blood from the pelvic district may pass into the general circulation mthout going through the liver. There is no unanimity of opinion as to the func- tion or importance of this connection (2) and from the data collected here there is no pecuhar reno- hepatic pathology. The gall-bladder is not a constant organ in either mam- mals or birds and indeed it may be absent or present in very closely related species (Two-toed Sloth present, Three-toed Sloth absent). When present in mammals it is usually a dependent bag while in birds it commonly lies upon the cystohepatic duct, between the liver and the last curve of the duodenum, in some varieties filling from the bottom, the inlet being guarded by a valve. This cystic duct in nearly all birds, comes exclusively from the right lobe while the hepatic duct, with which the cystic has no connection, is formed by combination within the liver of radicles from both sides. It passes to the duo- denum well in advance of the cystic duct, in some birds, e.g., the Struthiones, very near the pylorus, that is on the descending limb of the duodenal loop. By this means obstruction to the biliary stream is rendered difficult. The common duct combines with one of the pancreatic outlets in most mammals but the abdominal salivary gland in (2) See Woodland, Proc. London Zool. 8oc., 1906, and MacLeod, Chemical and Physiological Medicine, Chicago, 1923. THE LIVER 225 lower animals has more often patent separate ducts or multiple ducts than it does in man. Birds have one to four pancreatic ducts separate from the biliary openings. The gall-bladder is missing in most varieties of the following groups: pigeons, parrots, wrens, ostriches, rheas, cuckoos, toucans among the birds ; most odd-toed ungTilates, hyraces, Indian elephants, all deer, peccaries, three-toed sloth, and many rodents. The varieties lack- ing this reservoir are herbivorous in the main, true carnivores seeming always to be possessed of such a structure. Among the important herbivorous ungulates, Bovidae, Tragulidas, Camelidse and Suidae have this bile reservoir almost without exception. Because of the interest now being shown in the pathology of the gall- bladder and its passages and of the pancreas, it was hoped that evidence of definite practical value for human pa- thology would be at hand in our study if we divided the animals into groups Avith and without a bile reservoir. The result is not unequivocal but worthy of note; it is discussed on pages 238 and 255. Microscopically the well known lobular arrangement of the liver is rather faithfully carried out among the mammals albeit the most systematic and complete archi- tecture is to be found in the pig while the marsupial seems the most disorderly, thus resembling the avian organ. In the latter class all the parts are indistinct, the cells having an unclear outline, the tubules being intricately wound and the interlobular connective tissue being scanty and not anastomosing in a definite framework. The intra- lobular reticulum is especially difficult to detect. Groups of cells are often found at portal spaces ; these are large and small mononuclears and granular cells, probably of the hematopoietic system. It is possible that blood formation is performed in the liver and spleen in some adult birds but such a function is denied for the mammal except under very unusual conditions of bone marrow atrophy. \ 226 DISEASE IN WILD MAMMALS AND BIRDS Glycogenic and fatty conservation is a function pos- sessed by both zoological classes as are the detoxicating and bile-producing powers. However it is highly prob- able that urea and creatin in metabolism is not cared for by the avian liver as it is by the maimnalian, judging by the researches of Paton and of Richet. Fat Deposits. The care of fat by the liver is very well shown by examining the incidence of fatty metamorphoses through the various orders. In the first place Mammalia show a slightly higher percentage of fatty change than do Aves and should show a greater difference were it not for the large number of cases in two orders of the latter. Among mammals, lemurs, rodents and marsupials store fat in the liver more than other orders but in, the second and third, it is chiefly the carnivorous varieties that have this property. Just why the slothful herbivorous lemurs should be first on the list is not evident especially since the grain-eating UngTilata are least apt to present fatty livers. With tliis exception, mammals with plentifully available hydrocarbons in their diet are most apt to show its deposit in the organ under discussion. Among the birds the gallinaceous varieties stand far ahead of aU others, the passerines following next. Galli show the con- dition in association with acute infections, chronic dis- eases and in health. Unless there be distinct reason for it at autopsy, it may almost always be said to be normal^ Passeres, especially the smaller forms, frequently come to autopsy with such excessively large livers, and indeed with a very large pad of abdominal fat, and nothing else, that one is compelled to look upon this over- burdened organ as incapacitated by the deposit. These two orders increase the percentage value for the birds. Striges, Anseres, and Accipitres also show a good number of cases but there is among the Aves no such clear rela- THE LIVER 227 tionsliip between food fat and fat infiltration as may be found in the Mammalia. Amyloid Deposits. Amyloid deposit is reported with reasonable frequency in domesticated animals, causing in them a fairly definite entity, being as usual related to the effects of long con- tinued or repeated infectious disease. Wild animals suf- fer from this condition but rarely and therefore to our few cases will be given a short discussion separately. An Indian Paradoxure {Paradoxurus niger) had patches of amyloid irregularly distributed through the organ. The animal had a carcinoma of the head of the pancreas, an obstructive biliary cirrhosis in a state of atrophy and a chronic nephritis with arteriosclerosis. There was nothing peculiar about the distribution of the deposit as there was in the next case, a Badger (Meles meles) where amyloid was found around the interlobular vessels and extending in the lobules along their canaliculi. This latter case seemed wdthout cause and we have considered it a primary amyloidosis, the spleen, heart muscle, kid- neys, intestines and other structures being affected. ( See Fig. 7.) A third mammalian case concerned a Dasyure (Dasyurus viverrinus) which showed distinct intralobu- lar collections. Its cause was a chronic suppurative process in the jaw bone. . Avian livers are somewhat more prone to show amy- loid deposits, eight cases being on record. Four occurred in the Passeres, one each in ColumbEe and Impennes and two in Anseres. Three were associated with chronic infectious disease and two with well established nematode parasitism. The remaining three, classed as primary, were not related to any other lesions, in two the amyloid liver being the only finding. The next abnormal deposition related to the physiology of the organ is blood pigmentation. Normally hemic pig- ment is dispensed with veiy rapidly but under unnatural 228 DISEASE IN WILD MAMMALS AND BIRDS conditions it accumulates. In only one order is there any- noteworthy percentage of hemosiderosis, the carnivores, the remainder showing a very trifling incidence. Degenerations. Going further into the physicochemical alterations of the liver brings us to consideration of those changes known as degenerations — parenchymatous, fatty, hy- dropic, hyaline, all of which we shall group under one heading. They occur in a great variety of conditions and do not appear to be specific, nor as the records are analyzed do they appear to occur preeminently in any one disease of the lower animals. The percentages are how- ever higher for orders and families whose diet contains relatively more protein, carnivores, the higher mar- supials, accipitrine, and wading birds. Acute Atrophy. , A very important degenerative disease of the liver is aciite yellow atrophy or, better expressed, acute degenera- tive atrophy for it is a total destruction of the w^hole or large parts of the parenchyma. It is apparently toxic in origin being related to the toxemias of pregnancy, to certain organic and inorganic soluble poisons ; some cases arise without discoverable cause. We have seen no cases in the mammal but two in birds. Both were females, one in active ovulation, while the other had no related pathology and the condition of the ovaries could not be deter- mined since they had been destroyed after death by rats. The macroscopic and minute anatomy offers nothing new. Jaundice was present but not intense. , Hepatitis. True inflammatory lesions are to be defined as some form of parenchymatous change to which are added con- gestion, infiltration of round or polynuclear cells, stagna- tion in the bile ducts or perhaps actual degeneration of THE LIVER 229 their lining cells. It seems necessary to stipulate these things because in the chronic forms, usually called cir- rhosis, it is necessary to have all of them, plus efforts at regeneration, in order to determine it as a chronic pro- gressive process. Acute hepatitis is a rare condition in mammals except when it is combined with septicemia or severe enteritis. In birds on the other hand the liver is, aside from the intestinal wall, perhaps the most frequent seat of pathology in the abdomen. This is because of its almost constant involvement in infective enteritis, and in such conditions as fowl cholera, fowl typhoid, coccidiosis and cecal amcebiasis, all of which we have sporadically. When one searches for special distribution among the orders, only one of them stands out as having a high per- centage, the Galli, an order which seems to have a very vulnerable liver. The macroscopic anatomy of hepatitis in birds is peculiar in showing a definite swelling with spots of gray or yellow color, sometimes coalescing to form irregailar areas. These are much more definite than in the mam- malian organ where swelling and hemorrhage are the commoner findings. These pale spots are of two origins. They may be focal necroses of the hepatic cells, with or without circumferential congestion or hemorrhage to make them stand out. In amoebic, coccidial and typhoid livers such is the type of change. In septicemia and cholera, the mottlings are made up of increased inter- stitial mononuclear areas, mth blood cells and shadow cells numerously present. I have seen what was in all probability a stage of repair after both these kinds of change. In the former, regeneration seemed to take place from adjoining liver cells, there being in the section no evidence of increased bile ducts to make new hepatic cells. It seemed also that phagocytes were derived from blood cells and not from Kupffer's cells. In the infiltrative lesion disappearance of the liver cells from the groups leaving compressed and deeply granular remnants was all 230 DISEASE IN WILD MAMMALS AND BIRDS that could be determined. Regeneration seemed to be progressing in the manner just outlined, i Neckoses. ,^ The degenerative and infiltrative areas of acute hepa- titis are simulated by focal necroses in livers not the seat of a general hepatitis from which they can be differen- tiated only by the microscope. These small areas of local tissue death are quite common in all pathological processes but are most common in the liver, possibly because of its exposure to toxins from the intestine. Their exact origin is not determined, various explanations being given. The somewhat distinct distribution in mam- mals versus that in birds may help in the final decision. In the former, focal necroses are more often encountered midway in the anatomic lobule and around the central vein whereas a perivascular location seems the usual position in the bird. Massive necroses of the liver may be of considerable importance in veterinary medicine. They take their origin in several different ways. The commonest in our records are those due to cecal coccidiosis and amoebiasis (quail disease and blackhead) while from the primary seat of these two infectious diseases, the cecum, may originate the virus of non-specific hepatic necroses. We have observed several birds, passerine, psittacine and galli- naceous, which at autopsy showed a distention of the cloaca, ceca, and lower small intestine with urates and slime but no mural inflammation and a large area of necrosis in the liver. This suggests perhaps a ** white diarrhoea " but it did not occur in epizootics and other morbid anatomy of this specific disease was absent. These frequent instances of association between the colonic area and the liver seem to suggest the transfer of necrotizing organisms, just as amcebae travel, and to indicate measures to clean out the tract when birds become '' plastered." Massive necroses also arise from mould disease, and from THE LIVER 231 infection with the necrosis bacillus, emanating from nearby infectious foci, or via the normal passageways from the intestine. Massive areas of degeneration may form by the coalescing of numerous foci, in any septi- cemic disease. Abscess. In man, amoebae, flukes, cestodes and biliary tract infec- tion are the commonest causes of purulent collections within the liver. In the lower mammals parasites play practically a solitary role at least as the major influence in localizing the collection, bacteria from the intestine doing the rest. We have one case of massive abscess in a porcupine suffering with septic pneumonia, the sup- puration in the liver being due to the colon bacillus, the general septicemia probably being from distemper. Monkeys have shown more abscesses than any other order, three being observed. One was due to infestation with trichocephalus which had apparently penetrated from the colonic wall into the liver through adhesions formed between these two structures. Another seems certainly amoebic but these protozoa could not be found, while the third followed an ulcerative enterocolitis of unknown cause. Two cats were seen with parasitic abscesses ; one harbored Distoma or ClonorcJiis sinensis, the other a nematode of ascaris type. The topographic distribution of these six hepatic abscesses was interesting. The position of the abscess is not mentioned in one case but of the remaining five three were entirely in the right lobe, one had the major lesion on the right side and smaller separate abscesses spread over the organ, and one with about equal distribution in all lobes. All three confined to the right side were solitary. Abscesses of considerable size are not met with in the bird as in the mammal perhaps because the former does not form real pus, necroses developing instead. 232 DISEASE IN WILD MAMMALS AND BIRDS Congestion of the liver is a matter of small importance from the standpoint of pathology unless it be of sufficient duration to cause cyanotic atrophy and induration. How- ever the facts that congestion of this organ occurs three times as often in the mammal as in the bird and that vascular cirrhosis has not been seen in the latter class, are interesting and noteworthy. [In addition ascites of hepatic origin has not been seen in the birds. The expla- nation for this lies in the rich anastomosis between the intestinal area and the caudal vena cava so that the blood does not have to pass through the liver to reach the heart. This arrangement would reduce the back pressure in passive congestion and relieve the liver in the congestion due to toxic or inflammatory distention of small vessels. \, ClERHOSIS. The chronic inflammations or so-called cirrhoses of the liver have been subjected to a great deal of study and many theories have been expounded as to their cause and classification. Here is not the place to discuss the aca- demic question of nomenclature but rather to adopt an acceptable worldng classification and to analyze our material thereon. A cirrhosis is a chronic inflammation of the liver indicated by increased connective tissue with evidences of degeneration and attempts at regeneration on the part of the hepatic cells. Certain cases of increased connective framework fail to show the last two features and, since they must be grouped near the cirrhoses because of the prominence of connective tissue, they are called fibroses, perilobular in type. Among the instances carry- ing out the full stipulations are livers with evidence of a perivascular fibrosis and obstruction, to which are added degeneration and regeneration of the lobular margins; such are portal, cierhoses in human medicine associated with passive congestion in the intestinal area, and ascites. In a second variety, fibrosis seems to succeed upon obstruction to the biliary lumina or upon peribiliary THE LIVER 233 inflammation, biliary cierhoses. The effects of this are to dam back bile with the production of varying degrees of jaundice and for the inflammation to spread into the lobules, thus distorting their internal architecture; this form is therefore unlike portal cirrhosis which alters the size and shape of lobules as a whole. Fatty change is very prominent in certain cases and it has been a custom, perhaps without warrant, to put such livers into a separate group. It may be that they represent a different chemical process. /When there exists for a long time a venous stasis in the liver, necrosis is apt to occur in the cells subjected to pressure and the absence of fresh blood. This gives rise to a '* nutmeg " liver upon which may succeed a definite perivenous fibrosis. This then is a working classification of the hepatic cirrhoses. Perhaps many slightly differing varieties might be constructed but this grouping will permit com- parison and contrast with human cases, and with instances in the various orders. Because of the relatively small total, thirty-two, it is perhaps unwise to attempt any con- clusions as to distribution but it is certainly noteworthy that twenty-six occurred in mammals. This means 1.6 per cent, in mammalian autopsies against .2 per cent, in avian. Among the former class the carnivores stand at the head of the list, followed in order by the marsupials, ungulates, primates, and rodents. Carnivora have sho^vn a few typical portal cirrhoses from a pathological standpoint but only one, in a badger {Taxidea taxus), was combined with the classical picture of intestinal hyperemia and ascites. Two of the cases were combined with chronic enteritis which may, of course, have been secondary but there was also a hyperplasia of the spleen which bespoke some grade of infection. None of the four showed involvement of the biliary tract. One animal, a skunk {Mephitis mesomelas), was jaundiced; it had anemia, nephritis and enlarged spleen but no intestinal inflammation; perhaps the associated anemia 16 234 DISEASE IN WILD MAMMALS AND BIRDS may have been responsible for the pigmentation. Biliary- cirrhosis occurred in two Carnivora, in both associated with enlarged spleen and nephritis. One showed jaundice and the other, with a huge liver from congestion and interstitial infiltration, had a small ascites. Fatty cir- rhosis was diagnosed in a raccoon but this is viewed with some reservation because this animal easily stores fat and in this case it may not have been a part of the process. In none of the foregoing cases did parasitism enter into the causation of the change and I shall always specify when such a factor was probable. The only vascular cirrhosis in our records occurred in a Gray Wolf incident to a longstanding myocarditis (Gray Wolf, Cams lupus mexicanus, Myocarditis, Adenomatoid goitre, Chronic gastroenteritis. Vascular cirrhosis of liver, Subacute dif- fuse nephritis. Edema of lungs, pericardium, and peri- toneum). Two examples of perilobular fibrosis appeared in this order, a raccoon {Procyon lotor) and a paradoxure {Trichosurus vulpecular vulpecular). The only note- worthy feature was, in the former, a very marked biliary stasis on the lobular margins and in the connective tissue ; this animal was not jaundiced. Ungulata are normally well supplied with definite interlobular strands which, in a few varieties, completely encircle the lobule but always show as clear fibrous septa going out from the portal areas. This richness of con- nective tissue renders more difficult a decision of increase so that unequivocal degenerations and regenerations with inflammatory changes have been demanded as cri- teria for cirrhosis. It has been recognized that cattle get a definite increase in their interstitial tissue without serious reaction in the parenchyma. With the knowledge of these facts in mind it has been possible to detect two distinct portal cirrhoses, two biliary cirrhoses and three perilobular fibroses. It is however evident by examining the rest of the autopsy notes that the chronic inflam- mations have had with one exception, little influence on THE LIVER 235 the animal's life and death and the associated pathology- is not instructive in etiology. One old deer with the definite portal type had ascites and intestinal hyperemia which hastened his end. The tjrpe of cirrhosis in the marsupial is progressively inflammatory and of the biliary variety. In two of the three cases there was active infection somewhere in the body, one a long continued streptothricosis, the other and more important a choledochitis with involvement of the pancreatic head. The third case showed a nephritis and a pericholedochitis and pericholecystitis. In all three there was definite evidence of biliary obstruction within the liver and in the occurrence of general jaundice. Monkeys have presented one portal, two biliary, and one perilobular cirrhoses. The London Garden reports a cirrhosis mth gall stones in a Chimpanzee. The case of the Barbary Ape is so good that it is quoted in brief. Barbary Ape 6 {Macacus innuus). Found dead. Never known to be sick. On exhibition nine years. Acute dilatation of stomach. Acute gastritis. Portal cirrhosis of liver. Acute parenchymatous nephritis. Chronic passive congestion of lungs. Chronic splenitis and perisplenitis. Ascites. Mild passive congestion of abdominal circulation. On open- ing the abdomen a dilated stomach occupies most of the anterior part, displacing the intestines downward and backward. The upper lobes of both lungs are uniformly deep red, soft, collapsed, subcrepitant. Sub- clavian vessels — veins distended with red clot, arteries with small amount of chicken fat clot. The heart is dilated on the right side, filled with currant jelly clot. The liver is small, surface hobnailed, edges rough, consistency tough, color brown. Section surface glistening, moist, granular and opaque, mottled by iiTegular brown areas separated by paler brown intercommunicating bands. Gall-bladder is small, contains viscid yellow bile and duct is patulous. Areolar tissue about the bile ducts is thick and opaque, the duct wall itself is thick and yellow. Gall- bladder tightly attached to capsule of liver. Spleen is slightly enlarged, soft and tough. Capsule is smooth, opaque and thickened on gastric surface. The trabeculee are prominent, pulp mottled gray-red, few recent hemorrhages. Capsule of the kidneys is smooth, strips easily leaving a smooth bro-\vn surface with dilated vessels. Organ is soft. Section surface is glistening, striae wide and indistinct, glomeruli faintly visible. Microscopic section of liver shows high grade of fibrosis almost entirely confined to portal areas with a marked increase in bile ducts although no place is found where these bile ducts are running 236 DISEASE IN WILD MAMMALS AND BIRDS into lobules suggesting attempt at regeneration. Liver cells show high- grade of fatty degeneration in some places, whole lobules being necrotic. There is no pigmentation and connective tissue is fairly rich in cells. Fibrosis quite well advanced. Cells about equally fibroblasts, round cells and polynuclears. Bile ducts very Avell preserved and cellular infiltrate rather less directly around them than at other parts of con- nective tissue. The section of kidney shows moderate congestion, granu- lar and vacuolar degeneration of epithelium generally distributed except in proximal tubules where there is swelling and desquamation. Detritus present in tubules and capsular spaces. Tufts swollen. The biliary forms of Primates were associated in one case with an undetermined parasite in the bile channels, in the other with tuberculosis and chronic enteritis. In all the cases the relative inconspicuousness of bile in ducts or in cells is worthy of mention. The perilobular fibrosis in a small cebus was trifling in extent but was associated with considerable round cell infiltration in isolated areas ; there was also nephritis, splenitis, and enteritis. The only representative of the rodents is a capybara {Hydrochoerus hydrochoRrus), their largest variety. This case was originally described as a typical Laennec or Pictou cirrhosis but I now class it as a portal form. The distinct insular arrangement of the lobules, the failure of involvement of the bile channels and the ascites are reasons for the present decision. The animal suffered also from tuberculosis (not in liver) and myocarditis. The Indian Elephant, ''Bolivar" {Elephas indicus), an old specimen, is the only member of his order to show cirrhosis. It may be considered as a senile process in part but the extreme distortion and compression of the lobules press the conclusion that it was a pro- gressive inflammation. Aves fail to show lesions which could be called portal cirrhosis, five of their six cases being biliary and one fatty with sigiis of continued infection. The macroscopic anatomy of the avian liver with chronic fibrosing hepatitis is fairly uniform and suggestive. In the first place it is grossly nodular, lumpy, not finely granular or ' ' hob- Fig. 18. — PORTAL CIRRHOSIS OF LIVER IN ATROPHIC STAGE APE (MACACUS INNUUSh THE DILATATION OF THE STOMACH AL: IN PHOTOGRAPH. THE LIVER 237 nailed." The sensation to the finger is resilient rather than tough. The color is variable but green and dull purple are the usual shades. On section no peculiarities present themselves unless it be that one can find pale spots on a dark background, which may correspond to the mam- malian connective tissue strands. Microscopically the increase of cellular groups at portal spaces and the exten- sive growth of connective tissue between the liver columns are the noteworthy features. There is nothing in mam- malian cirrhoses to compare with the intralobular growth of fibres in birds. There is of course no regularity so that the degree of replacement or necrosis of parenchyma is hard to estimate. Bile ducts do not proliferate but seem, once obstructed and surrounded, to succumb to the inflammation, The six cases in birds are : Psittaci, 3, Galli, Anseres, Struthiones each one. The cases in the last two orders were associated with parasites, to which bacteria or toxin may have been added. It is interesting to note that the two frankly progressive obstructive biliary cases in the parrots showed general jaundice. It was formerly customary in many quarters to speak of atrophic and hypertrophic cirrhosis. Now it is gen- erally thought that any form will be large or small as growth and regeneration on the one hand, or contraction, atrophy and degeneration on the other, may be predomi- nant at the time the organ is seen. It is perhaps mislead- ing to judge by our notes of what happens, but it is curious that in the thirty-two cases, the pathologist could state only in seventeen instances that the liver was larger or smaller than normal. This means therefore that the liver of cirrhosis need not deviate greatly from its customary size. Nine of the seventeen times the organ was considered smaller than normal, eight times it was greater. These variations did not strictly correspond to type, but the portal form, frequently called atrophic, was more often small than was the biliary form. 238 DISEASE IN WILD MAMMALS AND BIRDS Gastrointestinal disease accompanied cirrhosis in fourteen instances. Nephritis was present nineteen times. The spleen was enlarged six times, in all of which definite evidence of infection existed in the body. Choledochitis existed four times, twice with biliary cirrhosis, twice ^\dth perilobular fibrosis ; cholecystitis existed twice, once in a monkey, and once in a bird Avith parasites. Pancreatitis was seen in three biliary cirrhoses and once in a peri- lobular fibrosis. The relation of the existence of cirrhosis to the presence of a gall-bladder is interesting. Among the thirty-two animals twenty-one have gall-bladders, eleven have not. The exact number of animals in our whole list with and without this structure, unfortunately cannot be given with exactness. As nearly as I can figure it out, six- teen per cent, of our animal posts have been on varieties without a gall-bladder, eighty-four per cent, with it. This would make the absence of this reservoir a factor favoring the development of cirrhosis since one-third of the cir- rhoses are in groups devoid of this bag, yet these same groups supplied only one-sixth of the total postmortems. Gall Stones. Our experience wdth concrements in the biliary system is limited to six cases which can be detailed in brief. American Beaver 9 {Castor canadensis) showed a soft purplish liver with groups of tortuous yellow lines; these prove to be groups of hepaticola with fatty degeneration around them, but successful regeneration is going on ; bile ducts are not seriously involved over any great part of the organ ; the bladder is distended greatly with thin, yel- low-green fluid ; duct is not patulous ; common duct narrowed at middle and above this constriction lies a small concrement; bladder contains two large and several small pale yellow-green friable stones; mucosa injected and covered with mucopus; the pancreas is not affected. American Beaver 6 {Castor canadensis) shows a slight bile obstruc- tion and pigmentation through the liver but no pus or cirrhosis ; bladder is collapsed containing only a little limpid brown fluid ; wall is slightly roughened but not opaque ; there is a blue-black stone 1.5 x 1 cm. free in the cavity; duct patulous; pancreas and intestine not affected. Brant Goose 6 {Branta bernicla glaucogastra) liver shows slight fatty change; bladder much distended, contains twenty-six small, quite THE LIVER 239 hard, greenish stones; one is impacted in the cystic duct which is not patulous. Pigtailed Macaque 6 (Macacus nemestrinus) shows a normal liver; bladder contains a small black concrement, very hard, no cystitis. Polar Bear ? {Ursus maritimus) showed a chronic cholecystitis and cholangitis, the stone (?) in this case consisting of a solitary, black, friable mass, six mm. in diameter. Mongoose Lemur 6 (Lemitr mow^ro^) showed a normal liver; bladder of about normal size but the duct can be forced only by considerable pressure ; there is a small stone and a granule in the tortuous cystic duct ; no cholecystitis. The specimens that are preserved show these to be chiefly inspissated bile, those from the first beaver and the goose being the only ones to rise to the dignity of gall stones ; it would seem that there was plenty of opportunity for calculi to form in the bladder of this beaver. In no case is there a cholangitis or cirrhosis dependent upon cholelithiasis. While stones have been showTi as infrequent there is a condition of the bile which may be quite important. In Passeres, Accipitres, and Striges one frequently sees a very dense inspissation of the bile both in the cystic area and in the lesser independent bile duct. This need not be, indeed usually is not, associated with hepatitis or cholecystitis. There is no one thing more common than another in relation with it but the diagnoses most often made are enteritis, distention of the proventricle and gizzard, and constipation. Inflammation of the Biliaky System. >^ The biliary tract from its origin in fine intrahepatic radicles to the bladder and to the end of the common or intestinal ducts is the seat of many inflammations both acute and chronic, but since they are supposed to lead to damage to the liver and pancreas and to the production of gall stones, it is well to consider the system as a whole. As a matter of fact separate analyses of cholangitis, choledochitis and cholecystitis do not reveal different figures for each or for different orders. The vulnerability 240 DISEASE IN WILD MAMMALS AND BIRDS of this tract is found to be directly as the percentage of cirrhosis, to wit, the carnivores stand first, then the mar- supials, ungulates. Primates and rodents |among the birds the order is Accipitres, Anseres, Struthiones, Psittaci, and Galli.) It is difficult in most instances to evaluate the various possible etiological factors, but, due caution being exercised,;gastrointestinal inflammation could be held re- sponsible in seventeen of the total of fifty cases. In twelve of the seventeen this process was wholly or largely in the duodenum.) (The next factor was general infection, at the head of wliich pneumonia and ' ' distemper ' ' occupied about equal places. In marsupials, the streptococcal and streptothrical infections to which these animals are sus- ceptible, was the prime factor. This group almost always has definite signs of stasis both in the liver and, as indi- cated by jaundice, in the general tissues. Pancreatitis was present in seven of the fifty cases and in five of the seven, enteritis was also found. Common duct stones were not observed. I shall have sometMng to say about pericholangitis and pericystitis under the head of pancreatitis. TUMOKS. The liver presents a good share of the tumors appear- ing in solid viscera but, with the exception of a few points, they offer little of interest. In the first place three angio- mata have been seen and while they may not be tumors in the accepted sense of the word, may be considered briefly. A single cavernous angioma was seen in a goose. It occupied a large part of the right lobe but did not seem to affect mechanically the fmiction of the organ since con- ditions wholly foreign to the liver were the cause of death^ A leopard presented several small groups of telangiec- tatic angiomata lying mostly at portal spaces, a few also under the capsule. The liver of a thrush was likewise scatteringly beset with small angiomata. The original THE LIVER 241 notes and recent examination do not reveal parasites or perivascular sarcomatous change. : Simple adenomata were observed in a woodchuck {Arctomys monax). This diagnosis is made with the appreciation that nodular regeneration of the liver after damage and in cirrhosis sometimes suggests tumor, but with adenomata an increase of supporting framework may occur. The liver of this animal presented numerous .3 to 10. cm. irregularly spherical, encapsulated, firm or slightly resilient, bro^\^l masses which under the micro- scope consisted of large pale vacuolated cells in columns or strands not connected with bile ducts. The last feature speaks in favor of the diagnosis of adenoma. The damage to the organ was probably considerable and the portal circulation must have been impeded since passive con- gestion and ascites were present. Enteritis and nephritis seemed the causes of death. Adenomata or fibroadenomata of bile duct origin were seen in four animals, a Red Fox (Canis vulpes pemi- sylvanicus), a Gray Fox {Canis ciner eo-ar gent ens) , a Jaguar (Felis onca) and a Common Deer (Masama vir- giniana). The first two present similar pictures, pin- point to 8. mm., gray, well outlined areas some of which are clearly cystic, others opaque and more solid. In the first fox a larger mass was found near the hilum. Careful study and consultation has failed to discover parasites in these cases, although their presence was strongly sus- pected, so that we were forced to conclude, in view of the rather typical microscopic picture, that they are adeno- mata of bile duct origin. Their scattered distribution, but with a tendency to be more numerous beneath the capsule, corresponds with a human case just brought to my notice. The mass in the liver of the deer was single and resembled an infarct, with cysts exposed by cross section. This tumor was found on the diaphragmatic surface of the right lobe. 242 DISEASE IN WILD MAMMALS AND BIRDS Tumors of an atypical, therefore malignant, epithelial variety were found four times, in an Alpaca {Lama pacos) and three parrakeets ; these birds are very prone to have all kinds of tumors. The records of the first animal could not be as satisfactory as might be desired because of an advanced state of decomposition but there was a car- cinoma-like growth of the gall-bladder area and a large hard alveolated tumor occupying one-half of the liver. The colon had been involved by the former, with perfora- tion. Two of the parrakeets showed a simple carcinoma with well developed fibrous tissue bands running in all directions through the large mass. The whole growth was comparable to the usual picture of these massive tumors when they are primary in the liver. All these three cancers seem to take their origin in the liver cells but the third had such an interesting involvement of the connective tissue that its minute anatomy will be given; it was denominated adenocarcinoma sarcomatodes. Undulated Grass Parrakeet 6 {Melopsittacus unduJatus). Section of liver shows organic capsule normal. Nothing remains of the original structure by which it might be recognized, suggestion in places of granular cells resembling liver cells being only occasional occurrences and in small numbers. Where liver cells do occur they are highly granular in various degrees of atrophy and show various grades of nuclear retrogression. Greatest part of section consists of dense, white fibrous tissue in which lymphocytes are rather diffusely placed together with large numbers of epithelium-lined spaces. These spaces are often elongated after manner of imperfect ducts but are of irregular form, have single layer of low cuboidal epithelium and richly staining nuclei. Upon search certain acini are found to have especially hyperchromatic nuclei and penetration of basement membrane. In such localities col- lections of epithelial cells are to be seen in plug form in lymphatics and acini of imperfect development of lumen are found. In addition to these epithelial lesions connective tissue ones are seen, occurring generally in restricted localities. The interstitial framework is seen to consist of closely placed spindle cells, some of which are especially elongated after manner of imperfect ducts but are of irregular form, directed in a definite, purposeful manner, but interlace in the whoi'ling manner noted in fibromas. Nuclei are, however, entirely too chromatic for a connective tissue tumor. Whenever a vessel occurs in these re- gions its lining endothelium is always swollen and nuclei in its wall THE LIVER 243 will be proliferated and of embryonic type. This latter condition is apt to occur in patchy manner, part of wall appearing normal and other parts containing these peripherated elongated nuclei. Secondary tumors were observed in the liver seven times as follows: Red Kangaroo {Macropus rufus) from malignant papilloma of the stomach; Spotted tailed Dasyure {Dasyurus ynaculatus) from cancer in the small intestine; Dorcas Goat {Capra Jiircus) from sarcoma in lymph nodes in mediastinmn; Raccoon-like Dog {Cams procyonoides) from mixed tumor of thyroid; Undulated Grass Parrakeet {Melopsittacus undulatus) from a brain tumor probably glioma ; another of same species from a sarcoma of pectoral muscle ; European Robin (Erithacus ruheculus) adenoma of adrenal {hypernephroma) . . SECTION VII THE ALIMENTARY TRACT, PART 3. THE PANCREAS The pancreas, an organ functionating as a gland with an internal secretion and by pouring a digestive juice into the duodenum, remains a structure of constant anatomy throughout the zoological classes under discussion in that it is composed of compound racemose lobules whose outlets join to form large discharging ducts, and of interstitial bodies, the islands of Langerhans, without connection with the secreting acini but having some rela- tion with the blood and lymph vessels. The organ originates embryologically by sprouts from the side of the primitive gut just below the part destined to be stomach, and from an outbudding of the common biliary duct. These two sprouts or pouches combine to form one organ, but this does not necessarily effect a union between their lumina. In some birds and mammals (Accipitres and some Ungulata) the lobes of the pancreas remain distinct during life, and the discharging tubules seem to empty only their respective lobes. However, there is no uniformity in the matter, and indeed the anatomy of the ducts is subject to very great variation despite the rather similar beginnings of the organ. Those who are inter- ested in this point may consult Beddard,(l) Letulle and Nathan-Larrier,(2) and Opie (3) ; there will be given in the following pages the average findings of anatomy of the gland body and of its ducts. The region of the pancreas in lower animals, espe- cially those which travel constantly on four feet, is one of great activity, and the organs are more freely movable than in the human being. The only exception to the latter (1) Proc. Zool. 8oc. London, 1905. (2) Bull. Soc. Anat., 1898, 73, 491 (3) Amer. Med., 1903, 996 244 THE PANCREAS 245 part of this statement may possibly be found in the cats and dogs, in which there are firmer attachments of the duodenum and pancreas to the vertebral column and the liver; this is brought about by the short gastrohepatic omentum and the abrupt curvature of the duodenum toward the back, under the mesenteric stalk. In the Ungulata and Marsupialia and in some Rodentia, the pyloric, duodenal, and pancreatic attachments are rela- tively loose, and torsion of the pylorus seems to be allowed for, since in these animals great distention of the stomach is the rule. Among the Aves the anatomy is wholly different. The birds have no attachment of the duodenum and pancreas to the posterior abdominal wall, except indirectly through a narrow strip comparable to the gastrohepatic omentum, one division of which passes to the beginning of the duodenum, the other to its end, and by a thin tail of pancreas which goes toward the spleen. The bulk of the pancreas lies in the U made by the long free duodenal loop, the two organs being covered by the serous membranes forming the middle abdominal sac. It will be seen from the foregoing that the movability of the pancreas is considerable — a highly necessary provision, because the stomach and duodenum are also movable and subject to distention by food and alteration of position during flight. In the class Mammalia there are usually two ducts, one entering the duodenum in combination with the bile duct, the other variously above or below this common opening. As will be seen in Table 14, however, there are several exceptions to this statement, there being but one duct opening independently of the bile duct. The general anatomy is closely similar throughout this class, so I shall confine my notes to the exceptions from the general rule, especially where they seem to be of importance in the etiology of pancreatic lesions. In the class Aves the pancreas consists usually of two or three distinct lobes lying one in front and two behind 246 DISEASE IN WILD MAMMALS AND BIRDS the cleft between the limbs of the duodenal loop, and it discharges its secretion into the duodenum by two or three ducts separately, and almost invariably above the bile duct openings. One duct always opens near the top of the distal end of the duodenal loop, near the bile duct. In the gallinaceous birds that have a bile duct opening into the duodenum near the pylorus, there is usually a pancreatic duct opening there also. In some birds a third duct passes from the body of the pancreas to the duo- denum at different places along the loop. It does not seem probable that dislocation of the duodenal loop would seriously interfere with the passage of the pancreatic secretions, since the gland is so intimately related with the duodenal serosa, but obstruction to the biliary flow due to changes in position of the intestine is easier because the bile duct is separate and loose and arises from the end of the gall-bladder. The ducts of both these structures pass very obliquely through the duodenal wall a matter of importance, as will be seen when discussing the infiltrative forms of enteritis. The gall-bladder is not present in all birds, but this is probably of no importance, as the hepatic ducts are wide and run directly from the liver to the duodenum. The pancreatic ducts are short and are closely bound around by glandu- lar tissue up to a place quite close to their entrance into the intestine. The musculature of the gall-bladder and the ducts seems comparable in mammals and birds, and a con- strictor or sphincter usually called the muscle of Oddi, is present in all but pigeons (Oddi). There may be found also muscular fibres in the major ducts of the pancreas, but they are not so heavy nor distributed so definitely as similar tissue in the bile duct walls. The mucosa of the pancreatic duct is much more folded in birds than in mammals, seemingly, therefore, more adapted to obstruc- tion by swelling from any cause. THE PANCREAS 247 Passerine birds have two pancreatic ducts usually on the ascending loop of the duodenum, or there may be one ahead of the pyloric biliary duct. The picarian varieties possess three ducts as a rule, one near the beginning of the pylorus, one near its end and a third of inconstant location. Owls have a system like Passeres, but the rela- tion between the organ and the intestinal loop is looser and the ducts are wider. ColumbsB have two pancreatic ducts in the ascending limb of the duodenum. Gallinaceous varieties have a double biliopancreatic system, a duct of each kind entering the descending and the ascending duodenal reaches, with the biliary placed after the pan- creatic in each instance. Accipitres have always two and oftentimes three ducts as do Anseres, both orders frequently having the third duct opening at the bottom of the duodenal loop where stagnation can and does occur. FulicariaB have usually three ducts. The foregoing are the orders presenting pancreatitis and therefore those whose anatomy concerns this study directly. The irregularity in number and arrangement of ducts continues through all the avian orders which show a greater aberration from standards than do the mam- mals. Theoretically the birds should cast some light upon the unsettled question of the causes of pancreatitis, and as a matter of fact such a result seems to have been realized. In 1915 I published an article upon a study of this subject which indicated that acute inflammations of this organ may arise via the lumen of the duodenum and pancreatic ducts, while chronic processes were the result of periductal passage of pathogenic agents. Further study would seem to indicate that disease of the biliary tract is of importance in lesions of the pancreas since a decidedly large number of cases is found in mammals, where the relation of ducts is definitely more intimate than in birds. The work of Archibald, (4) Deaver and (4) Surg. Gyn. and Obst., 1919, 28, p. 529. 248 DISEASE IN WILD MAMMALS AND BIRDS Sweet, (5) and Judd (6) seem to agree with the findings upon our material. This need not be, however, in discord with the idea that acute inflammation is superficial in origin, chronic lesions deep or lymphogenic. The discus- sion will be resumed in a subsequent paragraph. The amount of pancreas to be found in birds is greater than that in manmials. According to our figures the organ represents ^/4ooth of the body weight in the former and Voooth in the latter. These figures are averages of a small number of instances and are not final. It is, however, obvious to casual daily observation that birds as a class have a large pancreas. The minute structure of the organ is governed by the same general rules throughout the two classes under con- sideration. Birds do not have as many interstitial islands as do mammals, but they are more compact and seem more definitely constructed of coiled tubules. In so far as the internal structure of the organ is concerned there has not developed in our study pathology peculiar to any animal. The importance of the ducts and position of the organ will be discussed later. Recognition of pancreatic disease during life is prac- tically impossible. In human medicine the signs and symptoms are vague and inconstant, (7) diagnosis often being a matter of exclusion. Veterinarians, except under the best hospital conditions make no attempt to diagnose pancreatic lesions but, since the improvement of surgical practice, at times operate upon cases of evident pain and distention which prove to be pancreatitis. These things were evident in a deer that I saw and that died on the following day from acute hemorrhagic pancreatitis; I made no attempt at this diagnosis, believing it to be acute tympanites. The feces were normal, according to the judgment of persons qualified to give an opinion. (5) Jour. A.M. A., 1921, 77, 194. ~ ~~ (6) Ibid., 197. (7) Garrod, Schorstein Led., 1920. THE PANCREAS 249 The condition of the pancreas at autopsy on animals not dying with lesions of this organ deserves some atten- tion since it may confuse the uninitiated. If the organ be seen in its normal resting stage shortly after death, it is not difficult to recognize the condition as normal for the species. Activity is indicated by a darker or redder color and an increase of consistency. In carnivorous or omnivorous animals and birds the pancreas in this state is a body with a distinct bulky character, whereas in strictly herbivorous varieties, especially ungulates, the structure is diffusely pink and doughy. This is impor- tant since the early stages of self-digestion and decompo- sition assume this same character in all varieties, while later stages present a deep red, swollen, wet organ. These appearances must be differentiated from acute hemor- rhages or inflammations, a distinction based upon actual local blood collections or extravasations and areas of degeneration in true disease. Oftentimes differentiation must be made under the microscope and in advanced decomposition, determination is impossible. When there is torsion of the stomach, notably in ungulates, the pan- creas is often found decidedly congested. This, it seems, is due to a twist of the duodenum and passive congestion of it and the pancreas — the only simple explanation despite the apparent provision for a high degree of mobility, as already explained. The organ is nearly always mildly congested in severe grades of acute duo-, denitis, although it need not be pathologically involved. It is, however, noteworthy that the pancreas is an organ with a low morbidity index, especially when one considers its proximity to a structure showing the highest disease index in the body, the intestine.} The succeeding para- graphs will reveal in comparison to other organs only a small number of cases of degeneration, inflammation and tumors. This has been ascribed to the freedom of blood supply and the power of tryptic digestion. 17 250 DISEASE IN WILD MAMMALS AND BIRDS An expression of this relative immunity to pathologic change is met in analyzing the data upon the simplest lesions, degenerations, to be expected in many states of disease. Only a small number of cases present them- selves, and they are under expected conditions, namely in association mth acute general infection, sometimes definitely septicemic in nature. About half of them were discovered microscopically, affecting the islands of Langerhans in vacuolization or granular disintegration. Focal necroses of the organ were met four times, three turkeys and a cockatoo. It is noteworthy that all these birds had some involvement of the liver, twice a complete acute hepatitis and twice a cholangitis. This is the more interesting since we shall learn that the liver is less often involved in avian than in mammalian pancreatitis. Hemorrhages occur occasion- ally in the pancreas in acute general infections and are seen in acute inflammations of the intestines ; the percent- age incidence with the latter is, however, very small. Pancreatic apoplexy proper has not occurred, for all the instances of large hemorrhage into the organ have been combined with changes forcing a classification of acute pancreatitis. Panckeatitis. Pancreatitis in the acute form is divided by many writers into exudative, hemorrhagic and necrotizing, while for the chronic variety an inter- and intra-acinus form has been described. It is questionable whether it is fair in acute cases to focus attention by special nomen- clature on different macroscopic pictures, unless it be for descriptive purposes solely, since there is nothing at hand to indicate that differing agents cause one kind every time. The physical findings seem to depend rather upon the speed of operation of the causation than upon its essence. Sudden obstruction of the pancreatic duct is believed to produce necrotizing processes to which hemor- THE PANCREAS 251 rhage may be added by digesting of blood vessels.^ Exudative cases seem due to extension of ulcerative inflammation, from a perforated gastric ulcer for example, to whicb digestive pancreatitis may be added. The interacinus chronic inflammations are usually con- sidered as due to obstruction or infection through the biliary or pancreatic ducts whereas vascular disease pro- duces intra-acinus connective tissue overgrowth. Analysis of the records of this laboratory would seem to indicate that necrotizing and hemorrhagic processes belong together, exudative in a class by themselves, and that chronic disease may be either interlobular or intra-acinar without regard to associated pathology. I have therefore studied our cases from this standpoint. Pancreatitis has occurred in thirty-eight mammals and birds among the 5365 autopsies, an incidence of 0.7 per cent. ; class incidence in mammals twenty-seven or 1.5 per cent.; birds eleven or .3 per cent. (Table 14.) Among the higher class all the important orders are rep- resented, but by no means in equal degree, whereas in the birds, less than half of the orders are listed, with the important Psittaci missing, despite a high death rate. It is perhaps well to be guarded in stating the relative vulnerability of the pancreas in various orders, but one cannot avoid the observation that Carnivora stand well in advance of the others (3. per cent, of autopsies), to be followed by Ungulata (1.9 per cent.) and Rodentia (1.7 per cent.). Nor can one fail to see that mammals have inflammations of this organ five times as often as do birds. Further analysis of the data leads into a consideration of the anatomy of the viscus in terms of the acceptable theories of the origin of the lesion. It is commonly believed that infection of the gland occurs by passage of organisms through the duct opening in the intestines, especially when there is swelling of the mucosa of both. For the human being the idea is current that infection or 252 DISEASE IN WILD MAMMALS AND BIRDS I- 'S'N ^1 1^ Bi^ua^ag oiuojqo Bijuajnji a^noy BBajonBj apis Bisoqjiio oi^udaH 8i:>iqoop8[oqQ JO sicfiaaBJoqo Bi^i^SjtoaioqQ 00+ oo+o+oo+o I OO OOO 00++0 o oo O + + O+O ++++0 o ++ rt^ oot-ajO-^oooooo+^o o o| I OOO ooot^oooo " o oo O +00 00++++00+ O +0 O e^-OO OOOc^-+OCOc^ O OO ana miM uadQ B:>onQ JO jaquinM Bpi^BaiouBj oinojqo siii^BaJouBj SaiziioJoaN ao oiaBqjjomajj a^jnoy as dsiss So 6 es <1<1 <6— FIBROMYOMA OF UTERUS. CORNUA AND TUBES. INDIAN ELEPHANT (ELEPHAS INDICUS)." ■ WHOLE MASS AS MOUNTED ON BOARD. IT MEASURED WHEN FRESH ABOUT SIX I-EET ACRObS_^^^^^ SECTION OF CORNU OF Fir.. 26 AT HIGHER POWER. A PIECE CUT OFF WHERE THE PALE AREA SHOWS ON THE RIGHT CORNU OF THE OTHER FKJURE. THE FEMALE REPRODUCTIVE ORGANS 309 spleen are negative. Kidney is large, greenish yellow, firm, smooth and glistening. The right uterine cornu is subinvoluted (the animal was delivered of a fetus before arrival at the Garden, which was sixteen days before death). Its walls are thick and distinctly congested, the congestion being of inflammatory type. Microscopic section of uterus shows a regular, not ulcerated serosa. Subjacent fibrous tissue is loose and contains a granular precipitate together with a few red blood cells. Muscular bundles under this are widely separated evidently partly by trauma, but certainly also by edematous interstitial tissue in which fibrillcB are widely separated and between which free red blood cells and plasma cells are seen. Capillaries ramifying through muscular bundles are greatly distended and congested. Numerous large arteries are present in addition. Some of these contain pink granular material within their walls together with diffuse collection of red blood cells. Fibroblasts extend into this necrotic mass from other sections of the walls. Lumen of such arteries is diminished and in places quite obliter- ated by recent organization tissue. At many places in muscularis are large cells of iiTegular rounded form and some cytoplasms incline to- ward the basic tint with one or several large hyperchromatie nuclei. They are especially likely to occur close to a capillary. There is an es- pecially large accumulation of these cells at that point of section farthest from fundus. Here these cells occur in chord-like masses which infiltrate the muscularis both internally and externally. This particular mass lies in the muscularis internal to great arteries and well removed from mucosa. In this mass are giant cells with multiple nuclei scattered through the whole cytoplasm together with smaller cells Avith exceptionally large hyperchromatie nuclei. Subepithelial tissue is especially congested and contains numerous fibroblasts together with a few well formed glandular acini. Lining epithelium is discon- tinous, of simple tall columnar type, in places becoming flattened or even lost. Lumen of organ is practically completely occupied by pus. The Indian Elephant " Empress " (Elephas indicus) showed calci- fied fibroids of the fimbriae; gross and microscopic notes and a photo- graph are given. The uterus is bicornate in type. In its body there are numerous fibrous nodules 1 to 4 cm. diameter. They can be traced from the cervix to the ends of both horns and tubes. At the end of each tube there is a gi*eat mass of calcified partly conglomerate tumors some of which are partly, others quite, pedunculated. One specimen measur- ing 2xlxli cm. has a peduncle 15 cm. long. The mass on the right side weighs 3,926 grams, that on the left side about the same. Section of tumor from uterine cornu shows the classical appearance of a leiomyoma with usual whorling and interlacing bundles of involuntary muscle fibres. Degenerative and vascular changes not seen nor is there any notable addition of fibrous tissue. This latter tissue is shown only in small amounts at one end of section. The preceding data record the discovery of three fibromata of the uterus and one of the fimbriae ; one fibro- 310 DISEASE IN WILD MAMMALS AND BIRDS adenomata of the cystic type, one malignant adenoma, one adenocarcinoma, and one chorion-epithelioma. They were found in Lemures 1, Carnivora 2, Ungulata 2, Pro- boscidea 1, Rodentia 1, and Edentata 1. Metastases occurred but once, to the lung. No secondary tumors were found in the uterus. While upon the subject of neoplasms of the female genitalia, it may be well to describe the three instances found in the avian organs. A very definite case of adeno- carcinoma occurred in the oviduct of an Undulated Grass Parrakeet (Melopsittacus undulatus), a variety of bird very susceptible to neoplasms. The notes in an abbre- viated form are given : Immediately under the ovary is an irregular mass measuring 2 cm. long, 1 cm. wide, 1 cm. deep. The lower part of the tumor thus comes to press against the cloaca. It is adherent anteriorly to the peritoneum. It apparently consists of two parts, an upper rounded larger, and a lower spherical smaller. Both parts are well encapsulated and separated from each other by a well defined constriction. The upper part has a pale opalescent appearance. It cut easily with moderate resistance. The lower portion externally has an egg-yellow color streaked with red. Upon section it has the same general appear- ance but contains in addition numerous small, irregular, yellow areas which mask the general opalescent appearance. The centre of this node contains an empty space (cyst) 1x2 mm. Microscopic section consists of an oval or elliptical mass showing over one convexity a depression simulating a constriction. A thin fibrous capsule extends over most of the section which is extra thick at the point of constriction. Constriction roughly divides the section into two. The upper portion consists of one or two coarse septa of fibrous tissue. From these central areas a delicate connective, tissue framework extends peripherally. In this framework are great numbers of irregular gland spaces. These gland spaces are so closely placed in most cases that room is afforded for but one nucleus of the bundle. The gland spaces vary in size, some large, some small, and show grotesque shapes. The larger gland spaces here contain granular debris and pyknotic nuclei. Compound granule cells suggesting colostrum corpuscles may be seen in this debris. The epi- thelium of the gland spaces consists of a single layer of columnar epithelium of low cuboidal type. In places it is heaped up so as to present several layers. In places too it is not applied in a regular manner to the basement membrane but breaks through and then the cells extend in most disorderly fashion into the lymphatics of the Fk;. 28. — PAPILLARY ADENOMA OF OVARY. WILD TIRKKY (MELLEAGRIS GALLOPAVO). THE FEMALE REPRODUCTIVE ORGANS 311 stroma. At these points the nuclei are hyperchromatie. The lower portion follows closely the description given above save that the glandu- lar spaces are much larger. They contain pink glandular material Avith admixture of compound granule cells. At the convexity of the tumor the acini are especially large. Here they contain a pink granular mate- rial which stains more intensely than the other granular contents and, too, inside of this intense pink material are sharply circumscribed areas of yet more intensely pink staining material. This latter sub- stance has a streaming appearance under the high power. This stream- ing appearance is due to elongated areas of less dense material which are placed with their long axes parallel. This lower portion shows, furthermore, even with the naked eye, two large cysts which are lined by epithelium and contain a very small number of compound granule cells. The capsule at the lower polq is worthy of note from the extreme dilatation of its capillaries. A papillary adenoma was found in a wild turkey (Fig. 26) and a mixed cell sarcoma in a King parrakeet {Apromictus cyanopygius). None of these tumors sent out metastases. Interest in the avian reproductive tract from a patho- logical standpoint centres around the tumors as already given, and abnormalities in egg-bearing. Among our specimens there have been many cases of soft shelled eggs apparently blocked in the oviduct, of ''egg-binding" and of the inspissated-egg-remains in the abdominal cavity. These conditions are well known to veterinarians and are explained on the basis of improper food, immaturity of the bearing fowl, injury and inflammations of the cloaca and oviduct. I made an attempt to associate these con- ditions with infectious disease incidence and with the normal egg size. The results are not harmonious. No relation existed between general or local infection and any of these conditions. Gallinaceous birds with their large eggs show the highest percentage (2.3 per cent.), but Anseres witli a somewhat larger relative egg size show 1.6 per cent. Passeres, with eggs of very variable size but relatively large pelves, have an incidence of .6 per cent. Struthiones' eggs are relatively small; their inci- dence is 1.5 per cent. \ 312 DISEASE IN WILD MAMMALS AND BIRDS Salpingitis occurred in seven cases, but there have been no evidences of an acute infectious disease such as is responsible for gleet. The Mammaby Gland. This structure has been the seat of three inflammatory conditions and four tumors. The former occurred twice in nursing animals, one of which seemed to be suffering with distemper, another from puerperal sepsis, while the third case was probably traumatic. The animals were in order, a raccoon, a coati and a skunk. Four cancers of the breast have been observed, all richly cellular or glandular in type; none of the scirrhous variety has been encountered. Two of them were ulcerating and one was about to become so. One gave extension to the axilla and lung, one had penetrated the abdominal wall and growm around the kidney ; the remainder had not yet extended. The animals were Black Bear {Ursus americanus) (thoracic mamma with extension), Common Opossum (Didelphys virginiana) (two sections of breast in pouch), two White-footed Mice {Peromyscus leucopus) (posterior abdominal right gland and whole side). These animals were all adult but not old. 29. — ADENOCARCINOMA OF MAMMARY GLAND (TWO SECTIONS WITHIN POUCH). COMMON OPOSSUM (DIDELPHYS VIRGINIANA). SECTION X THE MALE GENITALIA Affections of the penis are limited to phimosis and occasional inflammations on a traumatic basis. The lat- ter is best seen in carnivorous animals like raccoons and coatis, but has little pathological interest. Two cases of phimosis have been seen at the autopsy table and one in an animal (hyena) still living. The last was operated upon some years ago and has given no trouble since. The prepuce of this beast is seldom retracted, but no swelling or retention of urine has occurred. A Eed River Hog {Potamochoerus porcus) dying of a variety of lesions, was found to have contracted preputial opening, the edges of which were tight and adherent to the glans penis at various spots. The prepuce had been dilated with urine to a large size from which collection the fluid could be pressed dropwise only by considerable pressure. Opening the sac revealed forty to fifty gray white sand granules about the size of millet seeds. A gray fox {Canis cinereo argenteus) had a mild grade of phimosis; in this case probably traumatism had some etiological relation, for it is the one referred to before in which a terminal urethral stricture was followed by rupture of the bladder. The testes have been peculiarly free of disease, only a small number of lesions having been found. A raccoon had an acute inflammation, traumatic in origin, and a few passerine birds were noted as shomng involvement of this organ in the presence of some general infectious diseases. ( Two tumors were found in birds, none in mammals. The avian cases both occurred in Red-shouldered Parrakeets {PalcBornis eupatrius) and were round cell sarcomata, without metastases. Acute inflammations of the prostate and Co^vper's glands occurred only once as secondary to pelvic infec- 21 313 314 DISEASE IN WILD MAMMALS AND BIRDS tion. Chronic change was observed in these bodies on two occasions, and prostatic hyperplasia, commonly called hypertrophy, was seen thrice. The mammalian prostatic area is knowTi to become overfilled with secretion and to be affected by inflammation when it is not discharged. This occurred in a wood rat {Neotoma pennsylvanica) and a wild boar (Sus scrofa), the former having the condition so marked that prostate and seminal vesicles were swollen backward into the pelvis like a tumor. His- tologically one finds in these conditions an engorgement of the glandular and ductal spaces with a mucoid sub- stance and a mild round and plasma cell infiltrate in the connective tissue; there are no marked evidences of active inflammation. The testes were not abnormal in these cases. The three cases of ''hypertrophy" of the prostate are worthy of separate record. Indian Paradoxure {Paradoxurus niger) the prostate is greatly enlarged, of pale orange color, soft and does not exude pus on section or pressure. Common Opossum {Didelphys virginiana) The prostate is enlarged, soft, gray yellow. Urethra contains a little gray mucus but seems patulous. Seminal vesicles negative. Rhesus Macaque (Macacus rhesus). Glandular hyperplasia of prostate. The bladder is collapsed. The opening of the urethra is occupied by a firm, friable yellow-white cast of matter apparently in- spissated semen. Urethral mucous membrane nonnal. Prostate is 7 x 4.5 X 3 cm. firm, resilient, dull purple-gray, capsule negative. Section shows normal lobulations containing apparently normal secretion. Vasa negative. Testes negative. Microscopic section of prostate shows acini of various sizes lined by a single layer of tall vacuolated cells with nucleus at the bottom. Cells probably nowhere reduplicated. Acini nowhere grossly atypical but everywhere hyperplastic and dilated irregu- larly. Mass is adenomatous in general increase but no part is truly neoplastic. Interstitial tissue rather less than normal proportion- ately. No " amyloid " bodies. Still another case of accessory sex gland enlargement was found in a Ring-tailed Lemur {Lemur catta), this time, however, with more suggestion of a neoplastic change. The prostate of the lemur is normally large. IHK PROSTATE CJI THE MALE GENITALIA 315 pale gray-pink and rather firm. Histologically it is about equally glandular and fibrous. In this case the cellular activity is undoubted, and one must consider it adenom- atous. The cause of death was enteritis, being perhaps more serious in the presence of the urethral obstruction. The seminal vesicles are distended to 7 x 2 cm. with a thick boiled- starch-like material. Wall and mucosa are negative. Prostate large, tense and injected. Its cut surface is pale purple, homogeneous; character of fluid is normal although excessive. Urethra is occupied by a cast of rather tenacious starch-like matter which begins at neck of bladder and runs almost to meatus. Openings of excretory ducts are prominent. Mucosa and submucosa of urethra are deep purple and the former seems to be slightly opaque as if covered with desquamated epithelium. Testes and epididymes seem normal. Vasa deferentia are slightly distended with excess of normally turbid fluid. Microscopical section shows hyperplastic epithelial condition with accumulation of droplets of hyaline matter but there is no amyloid deposit. In places it is possible to see a hyperplastic and loosened epithelium with nuclei becoming vacuolated, and the whole being cast off. Less granular free globules suggest that this is the method of origin of the hyaline globules free in the acini. The picture is one of papillary adenoma. In some places there is surely reduplication of the lining cells. Connective tissue is deeply staining, compact and with adult nuclei. Growth is not very vascular. There are no corpora amylacea. A case of tuberculous prostatitis and seminal vesic- ulitis was seen in a Japanese Macaque {Macaciis fusca- tus). Judging b}^ the advanced stage of these lesions and their more recent character in other viscera, the disease was suspected as pelvic in origin, possibly due to infection by a thermometer. Whether or not such be the case cannot be established, but at all events, separate thermometers kept in carbolated vaseline were employed after this death. The females caged with this animal did not de- velop tuberculosis of the pelvic organs. SECTION XI THE DUCTLESS GLANDS The Thyroid Bodies. The ductless glands occupying the anterior cervical regions, knowoi as the thyroids and parathyroids are structures to be found in some form in all vertebrates, but increase in distinctness of outline and construction upward in the zoological scale. In the bird they present themselves as discrete rounded bodies lying well to the side of the midline resting usually upon the carotid artery or jugular vein or both. As one removes the skin reddish brown globular masses will be exposed to view, sometimes showing an irregular lobular outline, a variation due to separate but attached masses of parathyroid glandules ; the latter may be yellowish or even white. For the most part, however, distinct thyroid and parathyroid bodies are separated with difficulty, and one must discover the latter by microscopical section.) In the mammal these organs are by no means so readily found when removing the cervical integument, for they are usually buried beside the trachea and covered by sternohyoid and sternothyroid muscles. Their position, relative to the larynx and upper end of the sternum, varies considerably but this seems to have little importance in the enlarge- ments to which the gland is liable. It is, however, lower, that is more posterior, than in the human being, rarely rising as high as the lateral thyroid cartilages. The principal lobes are elliptical or roughly triangular masses with their long axis corresponding to that of the animal 's body and apposed mesially to the trachea. AVhen enlargement occurs it develops in the anterior or ventral direction, pushing through the cleft between the muscles and the trachea to present under the cervical skin. In 316 THE DUCTLESS GLANDS 317 quadrupeds it may become dependent and the swelling is nearer the sternum than is the case in man. The isthmus is a very variable structure, and its presence or absence cannot be said to be a constant character in any order, or indeed in any family. I have seen in old animals a fibrous band extending over the face of the trachea connecting the capsule of the lateral lobes, which might have been an isthmus at one time. From these few observations the idea of atrophic fibrosis occurred to me. Such may be the reason for the absence of this transverse link in some adult specimens. The amount of thyroid tissue possessed by an animal might be judged by measurement or weight. The former is misleading since the density might vary, as it certainly does in the two classes and between certain orders in mammals. Actual weights would afford little comparison, whereas the weight in terms of total body weight may supply a guide to the amount of gland normal to an animal. There are given in Table 15 the grams-per-kilo- gram-body- weights of the thyroid bodies (thyroid and parathyroids both sides combined) of twenty animals whose gland seemed entirely normal at autopsy. They are all adult specimens, free of cretinoid characters and of bone or heart diseases, conditions which might reflect abnormalities to these glands. The list is too small to warrant any conclusions, but in one respect confirms Murray's (1) observation and certain experimental work, notably of Vincent and Jolly (2) and Carlson, Rooks and McKie (3). The carnivores have more thyroid than ungulates (averages .55 gm. vs. .18 gm.), but the mar- supials on our list have nearly as high an average as the former, namely .44 gm. To these figures might be added others which I have worked out from the list given by Murray ; it is only possible to compute the gram-per- (1) Proc. L. Z. 8oc., 1919, p. 16. (2) Journ. Phys., Vol. 34, 295. (3) Am. Jour. Phys., Vol. 30, 129. 318 DISEASE IN WILD MAMMALS AND BIRDS Table 15 Showing Weights of Animals, of Their Thyroid Bodies and the Relation of These Weights Per Kilogram. Grams of Body Grams of Thyroid Grams of Thyroid per Kilo of Body Primates : tei Woolly Monkey Lagothrix lagotricha Carnivora: Wild Cat Felis ruffus Silver Fox Cams chama Kamchatkan Bear Ursus beringiana Lynx Felis canadensis Jaguar Felis onca Rodentia: Ground Squirrel Xerus capensis Ungulata: Zebra Equus burchelli Giraffe IKfeGiraffa camelopardalis Barasingha Deer Cervus duvanceli Kashmir Deer Cervus cashmirianus Urial Ovis vignei Marsupialia: Kangaroo Macropus robustus (?) Tasmanian Devil Sarcophilus ursinus Wombat Phascolomys mitchelli Wallaby Genus and species (?) Edentata: Anteater Myrmecophaga tetradactyla Accipitres : Wedgetailed Eagle Aquila audax Struthiones: Rhea Rhea americana Emu Dromseus novae-hollandise 2.370 8,180 2. 3,325 4. 31,800 22. 9,500 1 29,500 16 550 340,000 384,000 56,800 56,800 22,700 18,000 3,120 26,000 3,360 3,300 2,300 18,000 36,300 45. 8. 20. 2.5 10. 4. 7. 3. 2. 1. 5. 23. .84 .24 1.2 .63 .1 .54 3.6 .19 .12 .14 .35 .11 .55 .13 .27 .9 kilogram value for a few of his examples since the body weights are not given in all. Using 2.2 pounds as equal to THE DUCTLESS GLANDS 319 one kilogram, the seal of 432 kilos had .03 gm. of thyroid, lion had .18 gm., leopard had .46 gm., a serval .36 gm., a skunk 2.35 gm., (was this normal?) while a single herbivore with the weight given was a porcupine having .25 gm. per kilo. The average of Murray's carnivores is therefore .67 gm. per kilo of body weight, whereas our figure is .55 gm. That the incidence of thyroid abnor- malities stands in direct relation to carnivorous character has been recognized before, and is abundantly borne out by our statistics, as will appear at a later place. Anatomy. The minute anatomy of the normal thyroid is fully given in text-books, and is doubtless pictured in the minds of all but students of the subject as a fixed and definite affair. Such is not the case. There is certainly a variation in gross size under conditions of seasonal and sexual activity, and it would seem acceptably demonstrated that changes in diet, especially where meat is concerned, are associated with swellings or shrinkage of the glands. These gross changes must be due to alterations in his- tology. In youth also the bulk is larger, a condition due to cellular activity, while as middle age advances the thyroid becomes smaller and more balanced in its colloid and cellular proportions. I need not detail the ultimate constituents of the gland, but it is well to emphasize a few points which must be taken into consideration in micro- scopical diagnosis. In the first place, the lobules or acini are not all of the same size in normal organs. This is especially true in the normal adult gland but may be so in youth. The cells which form the inner lining of the acinus are low cuboidal in shape but the elements which lie under them are oval and do not change with hyperplasia of the former. The colloid which fills the glandular spaces is very susceptible to mechanical and chemical agencies. In material preserved in alcohol it may be shrunken away from the cells or heavily vacuolated, while in tissue 320 DISEASE IN WILD MAMMALS AND BIRDS preserved a long while in any liquid it may be found con- tracted or broken. Large vacuolated or vesicular cells are characteristic of the human parathyroid ; this type is by no means so prominent in the lower animals, and in the few examples studied closely the arrangement is less definitely glandular than in man. The avian thyroid is distinct from the mammalian in the delicacy of its fibrous framework and the flatter character of its epithelia. Acini are usually of more uniform size. Capsular vessels are prominent but internal vascularity is less in birds than in mammals and their goitres are not solid. Physiology. The physiological value of the thyroid-parathyroid complex has been the subject of extensive study and voluminous literature without exhaustion of the possi- bilities, but with the result that we are possessed of knowledge explaining certain phases of abnormality, even if the normal functions be not unexceptionally demon- strated. The accepted alterations of functions are hypothyroidism — inadequate physiology, and hyperthy- roidism — excessive activity. Absence or atrophy of the thyroid bodies is usual in hypothyroidism, w^hile enlarge- ments, collectively called goitre, commonly accompany excessive function. Exemplifying the former, cretinism is the result of failure of normal function and develop- ment during fetal life while myxedema is the expression of the disappearance of thyroid secretion after it has once been operative; the latter may occur in infancy after nursing has ceased, or at any time that the thyroid may atrophy, during some of the forms of goitre for example. Hyperthyroidism may express itself, with or without visible enlargement of the thyroid body, in nervousness, gastrointestinal disturbances, tachycardia, loss of weight and exophthalmos. The first group, which might be called athyroidism, is often associated with alterations in the THE DUCTLESS GLANDS 321 bony skeleton in the form of chondrodystrophy or of rickets, while some degree of osseous change is observed with myxedema. Goitre, be it simple or exophthalmic, may be foUow^ed by cardiac enlargement or by myocarditis. The character of the thyroid secretion is not kno^vn except that it is influenced by the availability of iodine in the diet, but there are some other as yet unexplained features. The normal thyroid fed to cretins or persons suffering with myxedema, has the power to improve the condition apparently by supplying iodine and the other essential elements. Iodine is an important constituent of the gland, being present in combination with protein.! Its quality varies indirectly with the amount of colloid and of hyperplasia (Marine). The administration of this element is beneficial in colloid goitre but is harmful in the toxic variety. Even though the administration of thyroid extract may relieve athyroidism, this procedure in normal animals fails to produce typical pictures of hyperthyroid- ism. Carnivora fed thyroid gland do not show toxic symptoms until excessive amounts are given, whereas herbivorous varieties are much more sensitive to this feeding. (4) Tachycardia, nervousness and exophthalmos are not produced by these experiments, an interesting observation since these signs are not recorded in wild animals, and only vaguely reported by veterinarians. Man is apparently very sensitive to thyroid dysfunction. On the other hand, meat-eating animals are more sensitive to excision of the thyroid body than are grain eaters. (5) Chemical studies have shown that the thyroid is concerned in basal metabolism since this is increased in hyperthyroidism and decreased in myxedema; nitro- gen output is much elevated. Thyroid physiology stands probably in some relation to the cardiac mechanism since in simple colloid or simple hyperplastic goitre if of long duration, cardiac enlarge- (4) Carlson, Rooks and McKie, Loe. cit. (5) Vincent and Jolly, Loc. cit. 322 DISEASE IN WILD MAMMALS AND BIRDS ment and disease may be greater than the excess work occasioned by the mere physical bulk of the enlarged gland in the neck would seem to warrant. A detoxicating function has been ascribed to the gland, but Murray dis- credits this on the ground that congestion occurs in infectious disease of warm blooded animals but not in snakes. He thinks the gland more likely stands in some connection with the thermoregulatory mechanism. It is evident from the foregoing that the thyroid is closely related to protein metabolism, and that this is in some way connected with the ability the body possesses to use iodine in the food if it can get it. Feeding of meat to fish was found by Marine to increase the size of the thyroid. With all this in mind it is not astonishing that two-thirds of our cases of thyroid abnormalities were found in the order Camivora. The anatomical changes of the human thyroid that precede or accompany the various clinical pictures cannot be said to be uniform to a degree that one can even approximately predict in every case what will be found at operation, at autopsy or by the microscope. Further- more, much discussion has existed upon the importance of the several changes, the association with clinical phe- nomena and the nomenclature. I shall not enter the academic discussion with our material because so much has depended, in human medicine, upon symptoms, signs and chemistry — data that we cannot adduce. Upon many occasions I have seen animals with very evident goitres, but have not been able to detect bulging of the eyes or especial nervousness. One striped hyena carried his mass for several years. It swelled up occasionally and seemed to cause dyspnoea. At one examination of the beast, to see if anything could be done for him, a large cyst broke under the examining hand, whereupon a deep inspiration was heard and reUef was apparent. Tliis handling was repeated twice, these times wdth the pur- pose of breaking cysts and when this w^as successful THE DUCTLESS GLANDS 323 disappearance of the dyspnoea was observed. However, a similar attempt upon another hyena and a wolf failed possibly because no large thin-walled cyst was present. These and the case of the lion cub (page 170) are the only instances in which the enlarged thyroid seemed to have given serious difficulty, and the symptoms were probably due to pressure. Many, indeed most, enlarged thyroids have been found at autopsy, when the Garden personnel was unaware of their existence. Interesting notes of familial cretinism will be found under the appropriate heading. Pathological Anatomy. Classification. And now to return to the question of morbid anatomy of the thyroid gland, I shall begin by outlining briefly the classification to be used in analyzing our cases, a system which combines those of many pathologists, yet which I believe contains the essentials of all. The changes in the gland being hyperplastic and recessive, at times to a stage of atrophy, no clear . cut definite line of demarcation separates all these pictures ; instead they must be thought of as merging into one another. When the thyroid enlarges more or less continuously with a maintenance of considerable colloid, the picture is that of colloid goitre. The gland is pale, gelatinous, tense but resilient and may show large cystic areas with fluid contents. Microscopically studied the acini are overfilled with col- loid yet the lining cells are retained but flattened. The cysts may show the ruptured septa of the original acini. Enlarged soft reddish glands are found at times to con- tain much colloid, nearly every acinus being distended with it, but in such organs the epithelia are high or even reduplicated; the amount of contents is the striking feature. These are termed hyperplasia with colloid. Hyperplasia may go on with the absorption of colloid, hyperplasia without colloid. The gland is then a darker body of more solid character, red, dull purple or uni- 324 DISEASE IN WILD MAMMALS AND BIRDS formly pale pink, somewhat dependent upon the blood content. By magnification one sees smaller acini wdth prominent high cuboidal or cylindrical epithelia and little or no colloid. The increase of cells seems to be due both to an increase in their size and number. As the pro- liferation increases the lining layer must be accommo- dated so that it bulges out into the lumen as a bud or papilla which, if it be extensive or universal in the thyroid, gives rise to the ADENOMATom goitre of papil- lomatous TYPE. This growth is accompanied by much congestion and small or large hemorrhages may occur, forming cysts containing a blood-stained fluid. Grossly such a gland has solid and cystic areas, is mottled red and gray or brown from old pigmentation and is usually of very irregular shape. These forms are more or less uniform and general, but in certain instances the hyper- plasia tends to remain in isolated areas or nodules, and in these develop solid masses of thyroid epithelium, some- times with a small lumen usually devoid of colloid, and a rather rich but loose fibrous supporting tissue, the whole picture resembling the microanatomy of the fetal gland ; to these the name fetal adenoma has been given, but since they are not fetal in origin and do not behave like tumors I have called them nodular adenomatoid hyperplasias. The next step in hyperplasia would remove it from benign to malignant in pathological character, and the term neoplastic hyperplasia is used ; this must of course be limited to the epithelial growths, since sarcomata, while they occur in the gland, come from other cells. The changes in atrophy consist in irregular distortion of the gland by fibrous tissue to which may be added large colloid or fluid cysts. There is no uniform finding in the thyroid for the diseases believed to be due to its atrophy, functionally at least, namely cretinism and myxedema. In the former there may be no thyroid, or it may rather closely resemble the normal organ while in the latter definite scarring and distortion is the rule. The gland THE DUCTLESS GLANDS 325 aciiii are compressed, the cells vacuolated or crushed out of existence or there may be colloid cysts. Inflammations occur as swellings of the interstitial tissues and of the acinus cells during many acute infec- tions. Repetitions of this may leave a definite increase of connective tissue with large cells in the acini, a lesion which many observers have looked upon as underlying certain goitres and myxedema. Hypeeplasias. The cause of progressive hyperplasias has been ascribed to infection, to chemical substances in water and food, endogenous toxins, heredity and many other fac- tors. While we can add nothing definite in this matter it is worthy of notice that all our animals are exposed to the same general climatic conditions, receive the same water, are fed from the same stocks and many varieties may be in charge of the same keeper. The influence of preexisting infections cannot of course be measured. Inbreeding or captive breeding seems to have a very definite effect upon thyroid insufficiency as is well known and so sharply emphasized by McCarrison in his reference to intermar- riage among certain Moslems ; I shall cite the history of a wolf bitch which gave birth to three cretin litters while apparently well but mated to a goitrous male. These facts concerning the etiology are given merely to emphasize the high degree of probability that the distribution of the lesions of the thyroid gland among our specimens indi- cates the susceptibility of the various orders. This per- haps needs no emphasis for the carnivores, but it does for the marsupials. The literature contains many references to goitre in domesticated ungulates ; this would give the impression that they are common among them, and so they may be, but this is not the case for wild ungulates. There being no doubt that the Carnivora have the highest incidence of thyroid enlargement, man being especially prone to it, and since goitre may be induced in fish by 326 DISEASE IN WILD MAMMALS AND BIRDS feeding meat, the inference is direct that high protein diet stands in some relation to this condition. Thirty- nine of our sixty thyroid lesions occurred in the order Carnivora; all the families of land varieties are repre- sented ; 8.1 per cent, of the specimens coming to autopsy showed definite thyroid alterations. However, marsu- pials have also a decided thyroid vulnerability as indicated by 4 per cent, of the specimens presenting abnormalities at death. Four of the seven cases were among the carnivorous opossums and ''devils," the remaining three being in herbivorous kangaroos. The influence of high protein diet is not evident in birds. A discussion of the essential pathology can be based upon the accompanying table. In making a diagnosis care was used to exclude mild swelling of the gland seen in acute infectious disease and under conditions of sexual activity. The gross diagnosis was checked by micro- scopical section, and all but a very few have been reexamined for the purpose of making the table. Table 16. Shoxving Distribution of Lesions in the Thyroid Body by Giving the Number of Cases Met in the Autopsies upon the Various Orders, According to the Classifi- cation Given in the Text. A ,1 R 2 X JS il E 2 V » » ^ia a 03 c« 2 Order Is •5 O T3 O ll 13 •5 II 13 ^1 il .1 a a catU 1 1 1 1 O o K K ^ '^ ^ s w < Carnivora 89 7 6 5 4 Q 1 1 1 7 Rodentia 1 1 Ungulata 2 1 1 Marsupialia . 7 1 1 1 1 s 2 2 1 2 1 1 1 Psittaci Gain 2 1 1 1 1 Alectorides Anseres 3 2 Total 60 12 11 7 6 12 2 1 2 8 Fig. 31.— hyperplasia with colloid. AMERICAN BADGER (TAXl- DEA TAXUS). MEASUREMENTS. 3 X 1.5 CM.; 3 X 1.5 X 1^ CM. J'% Fig. 32.— adenomatoid GOITRE. RACCOON-I.IKE DOG (CANIS PROCYONOII THE DUCTLESS GLANDS 327 This rather diversified group of pathological lesions would warrant one to expect a notable number of instances of disease observed during life, suggesting that the thyroid was at fault ; such, however, is not the case. In the first place, no case of exophthalmic goitre, as the symptom complex is known in man, has been observed, yet the anatomical alterations, hyperplasia without colloid, and with papillary or solid adenomatoid character, are abundantly represented. For the pathologist to accept a case as toxic goitre I would ask evidence of enlargement of the heart and perhaps in addition degeneration of the myocardium. The animal showing the closest resem- blance to the disease in man was a Raccoon-like Dog {Canis procyonoides) whose history and notes are given in brief. Raccoon-like dog {Canis procyonoides) 9. Acute hemorrhagic splenitis. Acute fermentative gastritis. Subacute catarrhal enteritis. Acute general infection. Hypertrophic cirrhosis of liver. Chronic interstitial nephritis. Hypertrophy of heart with acute myocarditis- infiltrative and parenchymatous. Chronic lymphadenitis with acute exacerbation. Epigastritic and gastric venous stasis. Nodular adenoma- tous goitre. The right thyroid is lower, both measure 4x3x2 cm., are soft, resilient with dense gray capsule. Section shows cysts filled with blood separated by pale septa of soft tissue of varying thickness. One similar mass under angle of jaw seemed like a lymph node but on sec- tion is like thyroid. Parathyroids not found. Pleurae negative. Lungs are gray, collapsed except lower half of lower lobes which are slightly emphysematous and edematous. No consolidations. Pericardium con- tains about 3 cc. clear, colorless fluid. Epicardium is glistening, trans- parent and pale. Heart muscle is soft flaccid, pale mottled gray-brown. All chambers are distended with mixed clot. The coronary muscles and columnas are mottled brown and gray. The tips at insertion of chordae are pale. Streaks of gray run through muscles. One area 1x2 cm. of softening found in middle of left auricular muscle. Aorta nega- tive. The liver is enlarged, surface rough and irregular, edges rounded and uneven, consistency firm and tough, color mottled deep red-brown. Section surface glistening, moist, granular, opaque. Lobular markings not lost but obscured. Connective tissue lines not clear but surely diffuse in lobules. Scars on surface leading to definite connective tissue strands about vessels and irregularly placed. Gall-duct patulous with limpid bile. Spleen is well forward in front and below stomach. It is much enlarged, soft, tough, has a smooth, tense capsule. Section 328 DISEASE IN WILD MAMMALS AND BIRDS surface shoAvs homogeneous purple pulp Avith faint, narrow but tough trabeculae. Follicles distinctly outlined, slightly large but merely of a slightly paler purple than pulp. On surface are many round 2-5 mm. sharply outlined gray thickenings of the capsule and immediately sub- jacent pulp. The right kidney's lower half has been replaced by a thin-walled clear cyst 3 x 2.5 x 2 em. The left kidney is small, capsule strips with difficulty tearing surface slightly. Surface and section are mottled pink and gray, glistening and opaque. It is firm, dense and tough, cortex narrow, medulla wide. Cortex has obscure, irregular mark- ings with few small cysts, striae and glomeruli, faintly visible, margin between layers iiTegular. Veins are distended over surfaces of stomach, under surface of diaphragm, in peritoneum over liver but not in ab- dominal Avail. Stomach contains sour gas and water. Mucosa especially near cardia is deep pui-ple. Rugae are large and pennanent but mucosa and submucosa are soft and on section congestion does not extend deep. The tips of the rugae near pylorus are infiltrated and the infiltration is surrounded by a zone of congestion. Mucosa seems about to slough but has not separated. At pylorus mucosa becomes deep broAvn-red, dense, swollen, opaque and coA-ered Avith a slimy, soft brown mucus. Folds are prominent but temporary. In jejunum and ileum mucosa is still swollen and opaque and rugae are still larger than normal and tem- porary Avith a dense sensation on compressing them. The color is not broAvn but deep pink and yellow Avith areas of submucous injection. Follicles not visible. Colon is negative except for slight thickening of mucosa unaccompanied by congestion or opacity. Lymph glands of small omentum are small, firm, yelloAV, homogeneous; those of the mes- entery are large, edematous, yellow brown and tense Avith lymph Avhieh escapes on section. LjTnph channels up to mesenteric stalk can be traced. Histological Notes. — LiA'er architecture much altered by passively dilated hepatic capillaries chiefly toward the centre of the distorted lobules. This distortion is in the form of irregular liver columns sepa- rated by irregular vessels and interlobular connective tissue. This latter is increased everywhere but is abnormal in distribution Avithin the lobules. The connective tissue at the portal spaces is not so much increased but it shoAvs most around arteries. Bile ducts seem not increased in numbers. Much bile pigment in large, coarse, dark brown masses chiefly settled Avithin portal spaces. The cells show slight fatty infiltration. No multinucleation. Organ is not seriously robbed of functionating tissue. Alterations are not equally distributed OA'er section. Spleen shoAvs enormous congestion with edema of the few chords and perifollicular tissue left unengorged Avith blood. Follicles are negative. No connective tissue increase. Blood destruction not noAv actiA'e but there are many hemosiderotic masses irregularly scattered. The subcapsular areas are loose edematous follicles. Heart muscle fibres have lost all transverse striations, some are hyaline Avhile others are fibrillar. The nuclei are decreased in number but there is no in- THE DUCTLESS GLANDS 329 crease of connective tissue nuclei. No pigmentation. There are several areas of round and polynuclear cell infiltration and one distinct abscess in section. The perivascular tissues are edematous. Muscle fibres are large and wide. Thyroid made up largely of slightly enlarged acini in most of which a slightly eosin-stained hyaline collection is found. There are a few cysts containing a thrombus and hemorrhage. There are no typical colloid cysts. Some scars from old hemorrhages may be seen. There is much free blood in and between acini. Blood pigment free and in granule cells is abundant. Some acinus cells show fat drop- lets. Lung shows old interstitial tissue increase especially about vessels and a few scars, some of Avhich are forming cartilage. These are deeply encapsulated. Mesenteric lymph nodes show trabecular thickening with active connective tissue formation which is also present about follicles and along edges of chords. Follicles lack germ centres, solidly lymphatic. About them and in and along chords are many tissue ceils some of Maximov type and a few eosinophiles. Many of these and endothelial cells are phagocytic of red blood cells. (Fig. 32.) Just at the time of completing this book another case strongly resembling exophthalmic goitre in man was encountered in a Gray wolf {Canis lupus). This animal had a history of enlarged neck and enlarging abdomen for about six months. His appetite and discharges remained about normal but weight was lost and activity reduced to a minimum. Attempts at removing the fluid believed to be in the peritoneum, by the use of diuretics, failing and the beast being in such poor shape, he was killed. An enormous adenomatoid goitre, concentric hypertrophy of the heart, passive dilatation of all cervical and thoracic veins, passive congestion of the liver and congestion of the portal area were autopsy diagnoses. It will be noted that no exophthalmos and nervousness were observed during life. Bone disease and atheroma are at times associated with thyroid insufficiency in man. The former, aside from osteogenesis imperfecta of cretinism, occurred only once in a carnivore and once in a marsupial. There is but one case of atheroma among the sixty cases of thy- roid disease. The reaction of the avian thyroid in its hyperplasias is somewhat different from that of the mammalian. The 22 330 DISEASE IN WILD MAMMALS AND BIRDS delicacy of the septa and the relative paucity of vessels is perhaps the reason that the gross and microscopic pic- tures differ from, those found in mammals. It should be emphasized, however, that while one can perceive a hyper- plasia of the gland of both classes when the testes or ovaries are active, there appears less participation of the thyroid in birds in infectious diseases than is the case for mammals. In simple functional hyperplasia the capsu- lar vessels are prominent, but the cross section need show no change. In the continued hyperplasias the organ remains more solid, being less apt to develop cysts ; large cysts are occasionally seen, however, and in one case the entire gland was composed of them. Microscopically the differences are largely of degree in that the process is less frank in development, but the essential changes of swollen epithelia and condensed colloid remain the same. Atrophies. The thyroid gland in its functional capacity, may be considered to undergo hyperplasia and then atrophy of the parenchyma cells. Normally this would leave the col- loid, the epithelia and the supporting tissue in proper balance, but in the presence of low grade inflammation or where an abnormally hyperplastic process retrogresses, the connective tissues may exceed their norm, the epithelia may be shed or remain high and the colloid be irregular in distribution. Such a state of atrophy may exist in fetal life, arise from unknown cause during a course of toxic goitre, or perhaps insidiously in chronic toxic conditions. When this occurs in fetal life cretinism or myxedema arises, when in later life, only the latter appears. Judged entirely by microscopic findings, eight instances of atrophic changes in the thyroid have been found. Three of these were in Carnivora and were secondary to definite goitres, but were not followed by myxedema ; one of these three was a cretin. A brother of this cretin but not himself a cretin, died at the age of Fig. 33. — COLLOID GOITRE WITH HEMORRHAGE FROM LEFT GLAND WHICH KILLED THE BIRD. BLACK AREA IS CLOTTED BLOOD. MUTE SWAN (CYGNUS OLOR6 ). Fig. 54. — AT THE LEFT, THE INSIDE OF THE CAI.VARILM SHOWING HEMORRHAGIC PACHYMENINGITIS. AT THE RIGHT IS A FEMUR. SHORT AND HEAVY BUT WITH THIN CORTEX; CONSTRUC- TION IS ORDERLY. CRETIN GRAY WOLF PUP (CANIS MEXICANUS). THE DUCTLESS GLANDS 331 five months from acute dilatation of the heart, and with a decidedly atrophic thyroid gland. A lion showed a dis- torted gland, the result of chronic inflammation, a condition also present in a leopard, in the latter possibly in association with a general infection of the heart, ves- sels and kidneys of long standing. A case in a bear can only be explained on the basis of chronic intestinal toxemia. A camel is the only other variety of animal to show this regressive change. The beast suffered with a marked anemia with marrow atrophy and hydatid disease ; calcification was found in the thyroid. As has been repeatedly stated, myxedema has not been seen, but hypothyroidism has expressed itself in these animals as cretinism. The most interesting pathological fact concerning the relation of the thyroid to this mal- development is that there is absolutely no uniform gross or microscopical change constantly present in the typical cases. This will become more evident as the following records of our cases are perused. In 1914 an apparently normal Gray Wolf bitch threw two normal young ones which died of lack of maternal care ; they were not posted. The father of this litter died shortly and was found to have a sarcomatous hyperplasia of the thyroid. A year after the first lot a second litter was born of an appar- ently perfect father which still lives. This animal was purchased in the same lot with the mother, and the two could be related. The first father was not related to the female. This litter consisted of seven, two dying almost at once and burned, the other five not being especially good specimens. They died at ages ranging from two to five months and were all cretins or cretinoid. Two showed hemorrhagic pachymeningitis, one external, one internal, and the usual bulky skeleton of cretinism (Fig. 34). The bones were constructed in a rather orderly and some- what graceful manner, the uncalcified epiphyses being only occasionally distorted. The same two animals, mated again in 1916, had as offspring seven pups. One 332 DISEASE IN WILD MAMMALS AND BIRDS evident cretin was killed while another runt was sacrificed and found to have fractures of both femora around Avhich no trace of callus was discoverable (Fig. 35). Two other cubs were apparently normal, while the remaining three did not develop and soon showed the cretin characters. When this group was about three months old they were fed chopped horse thyroid; one improved decidedly, one slightly, the third not at all, but it might have been too weak to get its share. These animals lived from eight months to three years ; the two good ones remain alive. In 1919 the mother was killed by her cubs, probably because she was weakened by long sickness. A papillary adenomatoid goitre, endocarditis, nephritis and chronic enteritis were found. The pathological changes in the thyroids of the cretins were as follows: In the second litter two cubs had hyperplasia with colloid, one had nodular adenomatoid change and the fourth showed dis- tinct atrophy secondary to colloid increase. In the third group two had distinct colloid changes, once pure and once as a secondary process with some evidence of atrophy to alter the fibrous tissue and shape of the acini. The remaining four seem to be all colloid in character, but I am not satisfied with the description or sections so that I shall not offer an unqualified diagnosis. The adrenals of these animals all showed some medullary congestion but no change in the chromaffin or lipoidal content. The deformative lesions of bones are frequently asso- ciated mth lesions in the pituitary body. Several of our thyroid cases have been studied for such changes without their discovery. Indeed no gross alterations have been noted among many hundreds of hypophyses seen in removing the brain nor in a small number studied histologically. Those examined under the microscope have seemed to correspond to the descriptions given by Stendell in Oppel's Handbook of Comparative Micro- scopic Anatomy. kk;. 55.— pathological kracturk of fk.vr'r. cretin gray wolf pup. THE DUCTLESS GLANDS 333 Tumors. True malignant hyperplasias of the thyroid epithelium in man are being more thoroughly studied in recent times so that similar lesions in the lower animals gain interest. The notes given below are of value as individual observa- tions only, but since three were in carnivores, another indication is at hand of the vulnerability of this order. One of the cases is admitted upon diagnosis alone, the slides and records having been lost, but since the deter- mination was made by Dr. C. Y. White, I am satisfied to accept it. The four of which notes are at hand are as follows : Raccoon-like Dog {Canis procyonoides) 6. Adenocarcinoma sarco- matodes. Metastases to liver and lungs. Fatty degeneration of liver and kidney. Acute diffuse splenitis. Submucous hemorrhages in stomach. At level of thyroid cartilage on each side and removed 1 cm. from same is a rounded encapsulated nodule measuring 2.5 x 2 x 1 cm. Portions are hard, others fluctuate suggesting cystic degeneration. Below these nodules are two bodies also bilateral, evidently lobes of thyroid, each measuring 5 x 2.5 x 2.5 cm. They are firm with some foci of cystic softening. From a ruptured cyst of the right lobe grumous, red, malodorous material exudes. Peripheries of such cyst show greenish discoloration. Bodies as a whole are greenish black in color. They are well encapsulated, do not meet in midline but are joined at lower pole by firm, apparently colloid, material. All through lung especially under pleura there are dark red, rounded, firm, well circumscribed foci measur- ing 2-8 mm. diameter. They project markedly on pleural surface. No capsule can be made out. Upon incising they have lighter red centres and deeper peripheries. They cut with resistance and have no inclination toward a wedge shape. Surface is for most part smooth except where tumors are present. Organ is soft and distinctly yellow. All portions of liver contain rounded and irregular nodules, some deep, others super- ficial. They vary in size from 2 mm. to 3.5 cm. diameter. No capsule can be made out, yet they are circumscribed. The central portions of larger nodules are dirty gray and friable. Peripheral parts dark red. Smaller lesions are solid red and of fleshy consistency. Spleen is deep dark red, homogeneous. Histological section of thyroid shows firm, old, dense capsule very irregular in thickness seemingly on account of the penetration of the enclosed tumor cells. Such infiltration gives the inner outline of capsule a very irregular, bizarre appearance, and at times thins the capsule until it is reduced to nil. In one place the tumor elements appear outside the capsule at a point where a large vessel is apposed to outside capsule. The appearances within this cap- 334 DISEASE IN WILD MAMMALS AND BIRDS sule vary; in places the picture is that of a carcinoma. Small, round, interspersed wdth larger irregularly shaped acini are seen lined by a single layer of low cuboidal epithelium. \ery frequently indeed the lining cells contain fine granules of golden brown pigment even where their lumina contain no blood. Some of the larger acini contain altered blood cells and a smooth, pink material, knife streaked and vacuolated peripherally. In some parts of section these acini are regular and well formed, in others they are very irregular and appear to be eroding the capsule. A second appearance concerns the connective tissue. Ap- pearing in almost any part of the section and bearing no regular rela- tion to the epithelial elements or the section in general are areas of closely packed large spindle cells with hyperchromatic nuclei. In another place such spindle cells are arranged purposefully to form irregular capillaries containing blood. A third appearance results from a combination of the first two. Here there are acini, between which run blood capillaries with remarkably rich and numerous embi'yonic lining cells. A section stained by Van Gieson stain proves that part of the pink intra-alveolar material is colloid. Every gradation can be made out in tint of this material from pink to salmon to orange. It is often veiy difficult or impossible to state whether a given blood- filled space is a blood vessel or an acinus with hemorrhage. In both structures the lining consists of flattened cells. In one there is the possibility of colloid, in the other of hematogeneous hyaline, both with peripheral vacuolization. Lung shows walls of alveoli thickened by young type of cells. Nuclei of cells lining bronchi are prominent, in good condition. Much coal pigment through whole section. Air sacs empty. There are several rounded nodes through section con- sisting of closely packed spindle and round cells. Blood is abundant in such nodules both in small lined spaces like capillaries and in larger necrotic foci where there is abundant blood pigment. In one place an irregrular, large acinus is seen containing a smooth pink material. A large part of the interstitial tissue is diffusely infiltrated by the large, round cells with hyperchromatic nuclei (Figs. 33 and 37). Prairie Wolf {Canis latrans) 6. Mixed tumor of thyroid. Metas- tases to lungs. The neck of the animal is enormously enlarged, the diameter exceeding that of the body. Thyi'oid is enormously enlarged to about the size of a child's head, rather firm before incision. When incised about 300 cc. blood stained fluid drained. It is rather soft and quite friable looking as if made up of fatty and hemorrhagic matter. The lung is of mottled deep red color with here and there on surface small hemispherical areas about the color of the surrounding tissue but of slightly increased resistance. They are raised above the surface and measure 2-7 mm. in diameter. Histological section of thyroid shows a mixed tumor. It is not possible to say that it is a pure thjToid gland tumor. It is largely sarcomatous, the round cell alveolar arrangement dominant at one place, at another the short spindle cell but not typical, so-called spindle celled type. There are THE DUCTLESS GLANDS 335 many areas of small and a few of large hemorrhage. Cartilaginous deposit is occurring at some places in the field showing the latter type of sarcoma. Section of lung contains a large sarcoma nodule. The cells consist of round cells without the large cartilage-like cells found in the original tumor mass. Coypu (Myocastor coypus) 6. Sarcoma of right thyroid. The thyroids occupy a position deep in the neck upon the anterior vertebral muscles, the left higher than right, being up to level of top of thyroid cartilage. Only a half-inch of lower pole of right remains and it is like the left which is soft, deep brown-red, delicately lobulated, closely bound to trachea but movable in fascia. It is 30 x 10 x 3 mm. The upper part of the right organ is occupied by, or at least within the same capsule as a 25 x 15 x 10 mm. encapsulated, pink mass with many small vessels on its exterior. It is soft and on section the surface is mushy, of gray-pink-yellow, and seems to have an exceedingly delicate trabecular network. Posterior and superior to this, lying near the salivary glands but back of them is a similar mass 15 x 12 x 5 mm. Still another lies anterior to what remains of the right thyroid and is about 8x5x4 mm. The adrenals are 30 x 13 x 8 mm. slightly hard and not unlike a long kidney in arrangement. The cortex is wide, regular, brown or tawny, the medulla rich in vessels and deep brown. His- tological section of thyroid is an almost completely cellular mass with here and there delicate and incomplete trabeculations. Small blood vessels are numerous and consist of a delicate line with a cell nucleus here and there, that is no true wall. It seems as if the blood channels jvere simply regular spaces through the cell mass. The cell type is mononuclear with definitely acidophilic " granuloid " somewhat vacuo- lated protoplasm. The nucleus is almost without exception eccentric, rather poor in chromatin but in places diffusely staining. Mostly, however, the nuclear skein is in spots or threads and fairly dense around margins, therefore not unlike a thyroid cell and a plasma cell. Here and there one finds compressed remains of thyroid acini. There is decided irregularity of size and shape in these cells. Its origin is not clear but this seems like a sarcoma of the thyroid. Undulated Grass Parrakeet {Melopsittacus undulatus) 6. Medul- lary carcinoma of thyroid. On opening the body a mass 10 x 6 x 4 mm. is found in the upper thoracic region on the right side. A similar mass measuring 5x3x2 mm. lies in similar position on the left side. They are identified as thyroid glands only from their position and from the numerous large vessels which radiate from them. An especially large vessel leads directly to the heart. Long axes extend anteroposteri- orly. The masses are of a firm gelatinous consistency, the color of carpen- ters glue in lower portions, shading off to a dirty canary yellow above. They have a translucent appearance in lower portions. The surface is fairly smooth, adherent latterly and posteriorly. They are well circumscribed. At one end of the histological section thyroid tissue 336 DISEASE IN WILD MAMMALS AND BIRDS is easily identified. It varies from normal in that its spaces are often very large, contain villus projections or may be completely filled by large compound granule cells with no colloid. Other acini are atypical and contain typical colloid. Continuous to such thyroid tissue is a very large, rounded tumor. It consists of round cells with round nuclei in which many mitotic figures may be seen. An arrangement into acini cannot be made out nor is colloid material abundant. In one or two places an irregular collection of such material may be seen with peripheral vacuolization but its confines are always indefinite. As far as section goes the mass is well encapsulated but the lymphatics are infiltrated by the tumor cells. The tumor, too, is sharply separated from the relatively normal thyroid. Irregularly scattered through section are remarkable cells with nuclei three or four times the size of other nuclei. They may be hyperchromatic or normally staining. (Figs. 38 and 39.) The THYMUS BODY is a structure encountered in our specimens with greater regularity than is the case in human autopsy experience. However, no great size of the gland is observed, and there is no record or recollection of anything which could resemble an enlargement suggest- ing status thymicolymphaticus nor has a tumor with this organ as its origin been observed. In one case only did the thymus present what was believed to be an unusual size. An adult Gray Lagothrix {Lagothrix lagotricha) died mth an acute intestinal infection. Its thymus was a large, soft, deep pink body lying in the anterior mediasti- num, running up to the clavicular joints and down along the sternum. The death had ample explanation without any state of this organ. The thymus body has not been found enlarged in association with thyroid disease. The SUPRARENAL, or ADRENAL BODY is an orgau of essen- tially the same general construction in the two classes here studied except that in birds the cortical portion may be imperfectly developed and in some of the lower groups is decidedly narrow. This outer zone may indeed be entirely missing since tissue comparable to it is dis- tributed elsewhere in the body, notably with ganglia along the vertebral column. The organ is infrequently the seat of alterations, detectable either grossly or microscopi- cally. Congestion and small hemorrhages are rather -MEDULLARY CARCINOMATOUS PORTION OF THYROID. UNDULATED GRASS PARRAKEET (MELOPSITTACUS UNDULATUS). THE DUCTLESS GLANDS 337 common in acute infectious disease especially when the respiratory system is involved, but these rarely destroy tissue or materially reduce the chromophilic cells. These circulatory disturbances have, however, been predomi- natingly among the mammals although birds have suf- fered with infections to a high percentage. The medulla is much more often the seat of congestion while, when hemorrhage has occurred, the cortex is apparently always involved. Lipoidal reduction has been seen in a few mammals. Primates and Carnivora, once to a state of complete exhaustion. More serious lesions have occurred eleven times, and since the cause and meaning of disease in this body are so vague it seems well to recite briefly each one. A Weeper Cebus {Cehus capucinus) suffered for several months with constantly but slowly increasing skeletal deformity of the osteomalacic variety. He died after moving him to a new cage, his end being hurried by a scalp wound. At autopsy the skeletal condition was determined to be of the above named kind. The organs were in good condition. The adrenals were decidedly enlarged for a monkey of this size, measuring 1.8 cm. in length. The medulla was a solid, brownish, homogeneous portion covered with a very narrow, barely discernible cortical zone. This was apparently due to a uniform hypertrophy of the cells of the medulla. The testes were slightly atrophied and fibrotic. A Black Spider Monkey {Ateles ater) had a history of stiffness of legs for six months. This was probably a sign of osteomalacia since at autopsy this condition was found together with a secondary anemia, chronic gastritis, acute enteritis and brown atrophy of heart. The adrenal was knob-shaped, the cortex was wide, brown, regular, the medulla small gray-purple. ''Histologically the capsule of the adrenal is thicker than is commonly seen in Primates and con- nective tissue bands between the units of the zona glomer- 338 DISEASE IN WILD MAMMALS AND BIRDS ulosa are somewhat stouter than common. The cells of this layer take the stain a little more deeply than usual, but are otherwise negative. The layer separating cortex and medulla is occupied by a band of well formed con- nective tissue which is not proceeding inward but outward and so encroaching upon the zona reticularis as to remove it completely in places, in others to make isolated islands of its cell groups. Fine lines of connective tissue are penetrating from this into the middle layer but not disturbing it as yet. The connective tissue septa pene- trating the medulla are somewhat wider than one would expect but show no activity in their growth. The medulla is somewhat broken up, vacuolated and the chromophilic cells are not especially prominent, indeed some of them seeming to have undergone necrosis." A puma {Felis concolor) died after a sickness of two weeks from acute gastroenteritis with its usual visceral associations, includ- ing acute nephritis, and with calcifications in the adrenals. These structures were quite firm and nodular, on section tough and resilient. ''The cortex is irregular, brown, with paler brown medulla. Areas of calcifications appear as small dots, as linear formations and in some places seemingly around blood vessels. Histological section shows marked vacuolization of cells, particularly of cor- tex. There is a diffuse overgrowth of connective tissue which has become hyaline. Here and there smaU cal- careous deposits may be seen but no massive areas as mentioned above. ' ' A Himalayan Thar {Hemitragus jemlaicus) came to his end, after a history of convulsions, from nephritis, which had resulted in general edema including the serous sacs, and an associated cardiac dilatation. His adrenal was egg-shape, of normal size, ^\Tth a wide, irregular dull brown cortex and a homogeneous opaque, darker brown medulla. ''Histologically the cellu- lar structure of the cortex is partly destroyed, partly THE DUCTLESS GLANDS 339 dropped out and partly disturbed by overgrowth of con- nective tissue. This connective tissue is quite prominent in the medulla where it is surely increased although it is made more prominent by absence of cells, some of which have been degenerated and some dislodged. ' ' A Japanese Macaque {Macacus fuscatus) after drooping three weeks presented at autopsy the following numerous lesions: anemia, chronic atrophic gastritis, atrophy of heart muscle with regeneration, hemosiderin pigmentation of liver, perilobular fibrosis of liver, chronic diffuse nephritis (subcapsular type), congestion of spleen, fibrillar fibrosis of spleen, hemosiderin pigmentation of spleen, local amyloid infiltration of spleen, calcareous infiltration in medulla of adrenal. Grossly the adrenal showed a thick, orange yellow cortex and small solid, brown medulla. ''Histologically the organ appears noiTual in all respects save for the presence of a few small irregular areas of calcification in the medulla. These occur apart from any recognizable necrotic or fibrous areas. In one place one appears to lie within the lumen of a blood vessel. No fibroses or special congestions found anywhere in the organ and cells show normal details and normal numbers of vacuoles. ' ' A California hair seal (Zalophus calif ornianus) which had been refusing food and having loose stools for about ten days presented after death the fol- lowing diagnosis: Hypernephroma of adrenal, chronic hypertrophic enteritis with acute exacerbation, hemor- rhagic splenitis, passive congestion of liver, congestion and edema of lungs with catarrhal pneumonia, acute fibrinous pericarditis, chronic lymphadenitis, chronic interstitial nephritis. His right adrenal seemed about normal, being 5x2x1 cm. with a narrow, dull yellow cortex and a large mottled gray brown medulla. The left one was 5x3x1 cm. The upper pole is swollen and contains in its centre a spherical tume- 340 DISEASE IN WILD MAMMALS AND BIRDS faction which is red, mottled, sharply outlined, with a suggestion of a capsule and slightly firmer than surround- ing organ. ' ' Histological section shows a capsule of very noticeable thickness but possibly not much in excess of normal. The cortex particularly in its deeper layers is much injected and in some places there has been hemor- rhagic diffusion. In many places in the zona fasciculata, more especially near the periphery, there is breaking up of the cell tubes with an infiltration of large round cells and some chromatophilic cells. In other places this seems to have gone on to fibrous tissue increase and necrosis of the cortical fibrous cell types. The coim.ective tissue layer below the cortex is wide and the spaces filled with blood. This connective tissue also surrounds islets of medullary cells which are not specially chromatophilic. The mass in the medulla is made up of varying sized alveoli sur- rounded by rather rare, highly vascularized connective tissue and enclosing islets of medullary cells. These alveoli may be subdivided by septa. Hemorrhage has occurred into many of them. The individual cell masses are made up of groups of rather large cells with illy defined margin, a granular, opaque but not vacuolated protoplasm. They have a bladder-like nucleus in which the centrosome is large and prominent. Definite mitotic figures could not be found, but mitosis is probably present. A few cells with double nuclei were seen and one mth four. In many of these large islands the centre has gone to pieces from hemorrhage or necrosis. Some of the vessels are thrombotic and one shows a ver^^ pro- nounced periarteritis. ' ' A brown cebus (Cebiis fatuellus) was killed because of a poor tuberculin test chart. His organs were negative except the right adrenal body which was 3 X 1.5 cm., or four times the size of its fellow. It was a tense body with a smooth, mottled, deep yellow surface. On section there were deep yellow islands separated by pale brownish septa; the structure did not THE DUCTLESS GLANDS 341 resemble adrenal. Histological section showed a hyper- nephroma of vacuolated cell type, roughly alveolar. Two other cases, w^hich because of their microanatomy are to be called hypernephroma can be added to those just cited. One occurred in an undulated grass parrakeet {Melopsit- taciis undiilatus) , the other in a black duck {Anas ohscura) . As illustrative of this tumor the former will be cited in brief. ' ' Hypernephroma of adrenal with hemor- rhage into body cavity. A tumor approximating in size the head of the host extends from the region of the internal genitalia and adrenals lying more on the right than on the left side extending fully to cloaca and shoving all abdominal viscera forward. It has a pedicle springing from between the two upper lobes of the kidney where adrenals and internal genitalia are not distinguishable. Tumor is coarsely lobulated, well encapsulated, nowhere adherent. It has a pale, dirty yellow color, richly marked by red lines of congested vessels. It is fairly soft, i.e., about consistency of normal liver. Upon incising, the cut surface bulges markedly, is a dirty gray-yellow, blotched with darker gray areas, shows no internal hemorrhages or markings of special import. No metastases noted to any other organ. Microscopic section shows a light cap- sule surrounding the tissue of the tumor. The latter has a very scanty fibrous reticular framework showing no orderly or purposeful arrangement. Upon and between the reticulum, irregular and for the most part elongated collections of cells are placed. At times these present an elongated fascicular form, but this is not by any means a prominent feature. The cells themselves are large, rounded or polygonal, have coarsely granular cytoplasm which only in rare cases contains vacuoles. Nuclei of these cells stain very poorly, but it can be made out that they are of large epithelial type and of vacuolar appearance. ' ' A somewhat unusual tumor was encountered in a Polar bear {Ursus maritimus), an adenocarcinoma of the 342 DISEASE IN WILD MAMMALS AND BIRDS adrenal, when judged purely by its histology but a second- ary tumor in the lung displayed the more familiar picture of large vacuolated cells as seen in hypernephroma. The diagnosis follows: *' Scirrhous adenocarcinoma of adrenals, secondary carcinoma of lymph glands, second- ary hypernephroma of lung, secondary carcinoma of diaphragm, acute mucopurulent bronchitis, acute catarrhal enteritis, chronic diffuse fibrous cholecystitis, choleli- thiasis, slight acute interstitial pancreatitis, follicular hyperplasia of spleen with fibrosis, hydrothorax, hydropericardium, chronic hypertrophic osteoperiostitis, encysted trichina in diaphragm, fatty infiltration of diaphragm, chronic diffuse nephritis, chronic productive lymphadenitis, pigmentation of lymph gland. Both adrenals are smaller than normal, of woodeny consist- ency, the pale cortex and medulla are poorly separated from each other. The cut surface shows gray white and tawny mottling and occasional calcareous points. Histo- logical section shows an extra capsule of fibrous tissue containing highly distended veins; the lining contains masses of tumor cells, many of which are necrotic. It is distinctly denser in type than normal and more abundant in places showing a proceeding fibrosis. Parenchyma shows but few irregular islands containing non-neoplastic cells, some of which are highly vacuolated, others are not. Interstitial tissues in peripheral parts are often grown together with the deeper portion where are intermixed tumor areas. The latter consist of small round acini of variable size lined by cells of active type. Nuclei are large and hyperchromatic, cytoplasm broad and disinte- grating. Parts show necrosis and hemorrhage. Upon search transitionals from non-neoplastic to neoplastic cells can be discovered in same fasciculus." SECTION XII THE SKELETON AND ITS JOINTS The bones with their articulations have been the subject of extensive study and research by zoologists in the direction of classification and evolution. Adaptation of the osseous construction to the needs of the animal is well appreciated biological knowledge. For example, the keel of the sternum in birds affords broad origins for the flying muscles, the pectorals, which also insert on the ala3 of this bone and on the clavicle, and in addition use these latter formations as fulcra. So too the extremities of quadrupeds are angular in their upper two segments for the purpose of supplying a direct action of the flexors employed in running and leaping. The thick masseter muscle of carnivores is accommodated in the deep zygo- matic fossa. Many other examples might be cited, but these serve to direct attention to the adaptation of function and construction. Pathological changes in our materials are however too few to permit con- clusions as to possible relation of zoological position and development except such as may refer to deformity incident to the degenerative processes — rickets, osteo- malacia and osteogenesis imperfecta, and in these conditions the alterations are merely passive accommoda- tions to weakened support in order to obtain comfort. To put the matter in other words, it would seem that, aside from the diseases just named, there is no outstanding change in the skeletal tissues peculiar to zoological orders that might indicate vulnerability of the system or the methods of response to injury or disease. Effects of Trauma. There must be considerable reserve or reconstructive power in the bones of animals since it is a common thing at autopsy to see unmistakable evidences of repair of 343 344 DISEASE IN WILD MAMMALS AND BIRDS fractures, dislocations and inflammations. Some illus- trations are introduced to exemplify this healing ability, one of which was found in an animal shot by a hunter, the other an incidental autopsy discovery. Even though there be no definite relationship between the zoological order and osseous disease, it is interesting to record a very simple observation. Animals with long extremities, especially when the bones are quite near the skin, have a rather high incidence of fractures and inflammations. Thus the ungulates have of all orders the highest per- centage of these traumatic and infective lesions ; herons and gallinaceous birds follow the ungulates. Marsupials, primates and carnivores, in this order, are susceptible to inflammations but not to fractures. Bones are often broken, among the Cervidse, Bovidse and Camelidae, when as they are chased by mates, they fall upon the slippery floor of the cages; or again the mounting of a small animal by a large buck may crush the former to the earth. Two cases of fractured pelvis have been seen in antelopes from a fall with extended hind legs. It would seem that repair is usually satisfactory if the animal have a quiet retreat where callus may form and union occur. A heron is known to have broken both bones of the leg ; at autopsy a very insignificant circumferential callus remained, the member being as straight and strong as normal. Figure 40 shows the femur of a deer shot by a hunter ; the shortening was considerable, but function was doubtless good because the hunter could perceive no limping as the animal ran. The most interesting fracture among our records was an intracapsular fracture of the hip in a Huanaco (Lama huanacos) shown in Figure 41. 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So far as the daily reports can be relied upon we have had no epidemic inflammations of the eye. The parrots and monkeys are the only animals that can be handled enough for treat- ment, and in them the applications have not seemed very efficacious. Two cases of conjunctivitis and iritis have had a tuberculous basis and two other specimens have had tuberculosis in the eye. Two parrots had, in asso- ciation with generalized tuberculosis, semisolid masses in the orbit which dislocated the bulb, thickened the lids and presented as yellowish granulating tumors under the conjunctiva; a Swainson's long-tailed jay {Calocitta for- mosa) had this lesion on both sides. The only case in a mammal concerned an Anubis baboon {Papio anubis) in which one eye had been enucleated, two weeks before death, for purulent ophthalmitis. The other eye became affected shortly after the extraction and the animal was killed; it was found to have caseous pneumonia. The tuberculous process had probably begun in the region of the optic nerve and involved the whole bulb. Secondary infection with pyogenic cocci had been superimposed upon the original process. Keratomalacia, encountered on a few occasions, will be discussed by Doctor White. Iridocyclitis was found in a white browed guan {Penelope superciliaris) the notes of which are condensed as follows : The internal organs show nothing of value pathologically. The right eye shows moderate conjunctivitis and a marked thickening of the cornea with complete opacity. The lens is destroyed and the 402 THE EYE 403 retina infiltrated by gelatinous material. Humors are watery, non-suppu- rative, but the fundal portion of the retina shows several poorly circum- scribed, yellowish white, gelatinous collections. Microscopical section of cornea shows replacement of normal bundles by wavy ones inter- mixed with small numbers of nuclei. These are never of inflammatory type, but always of connective tissue type. Conjuctival mucosa defec- tive in centre, puckered but shows no subjacent inflammatory features. Ciliary body richly infiltrated by Ijonphocytes and vessels distinctly congested. This condition extends over whole anterior surface of iris and for a short distance over posterior. Sclera shows much bone for- mation. Chronic interstitial keratitis, subacute interstitial iritis and cyclitis. Cataract. Opacities of the cornea are quite common among our specimens, most often due we believe, to local trauma; ungulates exhibit them more than other varieties. Cata- ractous opacities of the lens are frequently observed in senile animals but, while I have no figures for the state- ment, I believe they are not as common among our specimens as can be observed in domestic horses and dogs. No record has been made of streaky clouds or spots in the lens but only of complete opacities. There are three only, an aoudad {Ovis tragelaphus) a macaw {Ara macao) and a summer duck {Aix sponsa) the last of which alone is interesting. This bird, a fully developed adult female, was killed because of total blindness and found to have a low grade chronic pancreatitis and a bilateral Morgag- nian cataract, the lens capsule containing a thin cloudy fluid with the nucleus quite freely movable in it. Amblyopia. A very interesting case of amblyopia in a young monkey was studied and reported by Dr. H. M. Langdon and Doctor Cadwalader in the Journal of Comparative Pathology and Therapeutics, Vol. XXVIII, Part 4. Because of its unusual character and careful investiga- tion, the report is reproduced here : Pigtailed macaque {Macactis nemestrinus) was bom June 9, 1913, in the monkey house, a Avell developed baby. He thrived and was as 404 DISEASE IN WILD MAMMALS AND BIRDS good as any for his age. He was never known to have anything wrong with him until on the morning of June 3, 1914, when he was found on the bottom of the cage in the monkey house. He had clenched hands and feet, jaws tightly closed, lips drawn back, eyes staring and glassy, with convulsive shaking of the extremities. At intervals he would become limp, with fists still clenched, and Avith only occasional jerks in the extremities. This would last about a minute, and then convulsive movements would be resumed. The entire ''fit" lasted about ten minutes. He was immediately removed from the large cage in the monkej' house to a small one in the back room of the laboratory. When put in the small cage he staggered as if dazed, and groped about apparently blind. He never recovered his sight entirely, but at times seemed to see better than at others. He was not seen in a " fit " in the laboratory. On June 24th, a small piece of banana was offered by a person who stood directly in the sunlight. The monkey came to the front of the cage, reached out and grasped very firmly the thumb of the hand holding the banana but did not take the banana although he very plainly wanted it. The banana was thrown into the cage, hitting the monkey on the back. He turned very quickly, then smelled over the floor of the cage until he found the banana. On June 30th, he was examined by Doctor Langdon and the following condition was noted : " Pupils react to the light of the ophthalmoscope. Optic discs are normal. Arteries possibly a little small. No other fundus changes." On July 1, a cloudy day, he was laid facing a window. A coat sleeve was laid over his eyes for a minute and then quickly removed. His pupils were seen to react slowly but distinctly to the light. His gaze would not follow a finger moved in front of his eyes. "When put back in the cage he climbed up on the wire at the back and then tried to climb the plain sheet-iron side. He groped and felt for a support and then fell. This he did several times. About August 1, when the eyes Avere examined, there seemed to be more visual perception and very dis- tinctly prompter pupillary reflex, which condition remained about the same when examined October 1. He died October 10, 1914, of a com- pound fracture of the right femur inflicted by a monkey in the ad- joining cage. At autopsy the viscera appeared normal throughout. The animal was fairly well nourished. There was about 5 cc. clear, pale yelloAv fluid under the dura. It escaped upon removal of the brain. There were adhesions of the dura over the temporal lobe (inferior surface), poste- rior and external to the optic tract, so firm as to remove some periosteum and superficial bone. Rest of dura seemed normal. Examination of the brain. Sections were made from different parts of the cerebral cortex, all of which were more or less alike. There was SAvelling of the endothelium of the pial lymph spaces, with some separa- tion of the fibres of the pia itself which extended into the sulci. The perivascular lymph spaces of the larger arteries of the cortex Avere dilated, and the adjacent cerebral tissue was edematous. A well marked endothelial swelling and hyperplasia affected a number of the arteries THE EYE 405 and capillaries producing marked general or nodular thickening in some places. Accompanying these hyperplastic changes there was a marked calcification of some of the arteries. This was not confined to one tunic, but in some instances it extended almost completely through the vessel wall, and here and there the lumen of a vessel was nearly obliterated. The main features were endothelial hyperplasia, edema of the pia and of the subpial cortex with some calcification of the vessels. It was perhaps less well marked in the occipital lobes than in other parts. The optic nerve and other portions of the brain appeared to be normal. Moon Blindness. It seems also profitable to repeat here a report Dr. H. M. Langdon and I made in 1911 upon a horse with periodic ophthalmia or '^moon blindness," a widespread condi- tion and one upon which there is even to-day little known and much contradictory theorizing. It is worthy of record that Dr. J. H. W. Eyre of Guy's Hospital, had a case to study at the same time as ours. He did not find the protozoon-like body discussed below, but laid weight upon the isolation of St. aureus, an organism often men- tioned in the literature about this disease. I cite the whole report since our publication in the 1911 Report of this Garden seems not to have been quoted in any of the reference articles on ''Moon blindness." Those who are interested in the clinical and pathological sides of the question will find a good summary in Veroff. aus der- Jahres. Vet. Berichten der beamt. Tierdrzte Preussens, 1908, and the bacteriology of the equine eye by Karsten, Inaug. Disser. Giessen, 1909. ''During the latter part of 1909 and first part of 1910 we had a horse referred to us suffering with recurrent ophthalmia or moon blindness. This affection, suggested by its name, is supposed to have some relation to the lunar periods. Some points in our work showed that such may be the case. Attacks appear not infrequently at the time of the full moon, and in our only experimental infection twenty-eight days elapsed between inoculation and a general ocular inflammation. 406 DISEASE IN WILD MAMMALS AND BIRDS ''This affection manifests itself as a conjunctivitis early in the attack, but rapidly progresses to an iridocyclitis and lastly to a panophthalmitis. After each attack the ball is smaller until it is so shrunken as to be sightless from chronic thickening and opacities. The causation is not known. The disease behaves not unlike an infectious one, remaining in a stud for years at a time. Not every horse may be affected. It has been connected with damp- ness, bad fodder, overwork and the like. Again others have connected it with malaria or rheumatism. Potapenke, Vigezzi, Koch and others have found various microorganisms, no two of which seem to be the same. Even an animal organism like malaria has been described. (Whether or not malaria has anything to do w^th the disease, it must be said that our horse was favorably affected in regard to temperature as well as to the eye condition by repeated subcutaneous injections of Quinine Bisulphate, Grain xx daily.) The attacks last five to nine days. One or both eyes may be attacked and not uncom- monl)^ do they alternate. One eye may cease to have attacks while the other continues. The experiments here recorded were made with the idea of transmitting the disease to other horses. They were only partially suc- cessful. During eight months the affected animal referred to us had six attacks of ophthalmia. The attack was observed for study on the first occasion, but during the second his anterior chamber was entered by a needle attached to a syringe, the exudate aspirated and injected into the eye of a horse with apparently healthy eyes. The history of this second horse will be given later. The attacks of the first horse ranged from six to twelve days. Five of the six affected the left eye and one the right. In January, 1910, the left eye was used for further inocu- lation, and following this traumatism complete recovery never took place. The corneal scar left by the needle tract almost disappeared, but an inferior anterior synechia formed and was followed -by a spreading opacity of the THE EYE 407 cornea, much wrinkling of the iris and opacity of the depths. After the fourth attack in this eye it was com- pletely blind. Material was obtained from this eye during its last attack, but it was merely serous fluid containing a few blood cells and epithelium, but no bacteria. *'In transferring the affection from this animal, the conjuctival sac was washed with 1-5000 bichloride of mercury solution and well rinsed with salt solution. The anterior chamber was then entered with an aspirating needle and the exudate removed. This consisted of 0.4 cc. slightly turbid straw colored fluid containing a few shreds of lymph. Bacteriological cultures, moist and dry preparations were made from a part of this, while the remainder was introduced into the anterior chamber of the second horse. This animal's eye showed the effects of the traumatism for eight days, and then was normal save for a small opaque spot in the cornea left from needle puncture. After twenty-three days a small patch of lymph collected in the pupil. Tliis increased slowly accompanied by lacrymation until the twenty-seventh day, when a sudden and violent conjunctivitis arose. The lymph in the anterior chamber likewise suddenly increased and rapidly became pus, forming a hypopyon. The conjunctivitis became purulent. The violent stage lasted five days and slowly subsided, leaving an ectropion with a densely injected bulbar conjunctiva, almost com- plete corneal opacity and an irregular contraction of iris, apparently due to several small synechias. The depths could not be seen because of the corneal condition. This stage of affairs remained during the rest of the animal's life, two months. He was permitted to live to see if an exacerbation of this chronic process or involvement of the other eye would appear. Such not occurring in two months, he was killed and the eyes removed. Fluid removed from the left eye of the first horse when killed during the last attack was injected into the anterior chamber of a third horse. This animal's eye received 408 DISEASE IN WILD MAMMALS AND BIRDS the operation well and the trauma had entirely disap- peared when the animal died on the eighth day. ''Laboratory Examinations. — From fluid removed from horse eye (No. 1) anaerobic cultures made on milk and blood serum, blood agar, glycerine agar ; cultures were made directly from the fluid, while the coagula were dried upon slides and stained as follows: Loefifler's, Gram's, Giemsa. In all there are very few recognizable bodies. They are red blood cells, polynuclears and a very few small mononuclear cells. In regard to microorganisms three structures present themselves. A well staining Gram-positive, rounded end rod of fairly uniform size but tending to grow in pairs and stain rather irregularly with Loeffler and Giemsa. These forms are sometimes called ' dumbbell ' in that they are bipolar, or even seem to have a constriction in their centre. Another form is peculiar and cannot be said to be recognized as a bacterium. It is circular, of fairly regular size and contour and in many places looks like a very large coccus. In Loeffler 's stain it is colored deeply in the centre with a paler marginal zone and an unstained halo about it, which, however, is not like a capsule. In the Gram and Giemsa method it is deeply blue or purple with a refractile centre and very sharply outlined contour. These forms varied from 3 to 5 microns. The third form is a wavy delicate short mycelium-like thread. Smears from the cultures as made above showed chiefly a Gram-positive, rounded end rod but which did not grow on planting out. It grew on aerobic media, but was not found on anaerobic. The Gram-positive organism would not grow beyond the fourth generation. It was not identified with any known species by the characters manifested during the short time we were able to keep it alive but could be placed in the Hog Cholera group. The mycelium was found to be an aspergillus. In regard to the large coccus-like body, little can be added to the above description. Further examination did not reveal characters permitting us to THE EYE 409 place it among the protozoa. No evidences of division were seen. The body is quite uniform in appearance, varying only in size. Whatever this is it seems to be an organized body. ''Cultures from pus in the anterior chamber of the second horse showed the palely staining rod, an asper- gillus and Micrococcus aquatilis. The first was planted on horse serum bouillon, but did not grow after the first gen- eration. These cultures were made after death, but the cultures made during the acute attack direct from con- junctival sac contained such a host of organisms that no judgment could be formed of their relative importance. The polar staining rod was found in smears. No large coccus-like bodies were observed in the second horse. Fluid taken from the first horse 's eye at death was sterile. ''These observations are at variance with those of others but such results are not unique in this respect. It seems as if the polar staining rod deserves some consid- eration, and we expect to devote some attention to it if another horse suffering from recurrent ophthalmia come to our notice. The large coccus-like bodies are very inter- esting and may be protozoa. The finding of the amoeba in the cases of Potapenke, increases their importance. Before, during and after the fourth attack of the first horse twenty grains of quinine bisulphate were given hypodermically daily for twenty days. The attack was very mild. Before the drug was given his temperature had ranged from 99° to 101° F. Immediately after the first dose the temperature fell to below 99° F., and remained at a very regular level during the entire twenty days. No malarial organisms were found in the blood. ' ' The ear is without special interest except as a place of localization of sarcoptes, demodex and fly larvae. A few cases of acute catarrhal otitis media have been found in association with nasopharyngitis both of the non- specific variety and that which resembles distemper. One case which led to meningitis has been mentioned. 27 SECTION XIV CONSTITUTIONAL DISEASES Theee is a long list of diseases including among others such conditions as hyperthyroidism, osteodytrophies, diabetes and gout which are spoken of as constitutional but which in reality are usually dependent upon some lesion peculiar to a definite organ. Several have been discussed under systemic diseases so that there remain for consideration in this section only two, gout and diabetes. Constitutional diseases are recognized in mid animals either not at all or by some happy chance which permits of examination direct enough to elicit diagnostic criteria. Gout has been discovered for example in some parrots and herons because of their swollen feet and their move- ments. In veterinary practice fairly accurate diagnoses are possible but in wild collections they are nearly always hit or miss. Therapeutics naturally follow this rule. Gout. Gout in mammals has been observed in the London Zoological Garden but has not been encountered here or we have overlooked it. A\aan gout on the other hand in one of its forms comes to our attention not infrequently. ;'It occurs most often in parrots, gallinaceous and anserine birds and herons ; occasionally accipitrine birds will suffer with it, an observation more often recorded in European collections than with us. The figures show no pre- dominance of percentage for any order but the records indicate that the most beautiful examples of internal uratic deposits occur in the anserine birds and parrots, while the best specimen of general gout, including the joints, was found in a boat-billed heron {Cancroma cochlearia) quoted below. 1 410 CONSTITUTIONAL DISEASES 411 In so far as etiology of this disease is concerned in domestic stock, too rich food, especially in protein, and restriction of activity seem to be credited with the greatest influence. These factors, while doubtless of importance for birds as they are believed to be for man, do not seem to fill all the requirements since all our specimens are confined and, because of their lack of exer- cise, possibly receive too much food, /judging by our observations and by publications from other gardens, carnivorous birds are not conspicuous for the incidence of gout whereasj/grain- seed- and fish-eaters suffer more often. /This suggests that these varieties cannot dispose of dierary protein which might be excessive for their metabolism while in captivity, whereas carnivorous species have a digestive and chemical reserve to take care of excess protein. Some such accommodative power must exist in human beings since not every large meat-eater develops gout. Heredity, often blamed for the human disease cannot help us with these birds. Examination of the diet list at the Garden does not reveal a great per- centage of concentrated protein in the feed of the grain- and seed-eaters. The disease occurs too seldom to disturb the accepted dietary for its possible elimination. Studies now going on may indicate appropriate changes in the dietaries that might be responsible. Arthritic gout appears usually in the pedal joints but may be found in the wings. Irregular, sometimes very deforming swellings appear which must be tender judging by the quietness of the bird and by its behavior if the joints be touched, "i Most often the swelling seems greater upon the flexor (palmar) surfaces of the toes or in the end of the tarsal articulation. Aside from these few observations there is notliing peculiar about the at- tack or the specimen during its sickness. Chronicity seems to be the rule and little emaciation may be found. Appetite is normal or excessive, provided the food can be reached. 412 DISEASE IN WILD MAMMALS AND BIRDS Internal or serous membrane gout cannot be recog- nized during life so far as I know. The bird may seem in its usual condition of feather, activity, appetite and elimination, when suddenly it will fade in a day or so and die. At autopsy the serous surfaces of the heart and peritoneum will be white with uric acid crystals and the kidneys a pale yellow brown with, markings indicating that the pelves and tubules are choked with urates. The boat-billed heron (Cancroma cochlearia) had had bad feet for three months. The general condition is poor as to plumage and flesh. The tarsal and metatarsal joint areas of both legs are sur- rounded by firm tough swellings involving skin and periarticular tissue. That on left foot has ulcerated and bled. On section the swelling is found to consist of reddened fibrous tissue around tendons, the latter apparently running through smooth sheaths. At both ankles are urate deposits clearly seen in this inflammatory tissue but at the lower end of the tarsus there are no distinct deposits. The joint surfaces do not seem to be involved. Knee, hip, and wing joints seem uninvolved. Internally all surfaces are opaque by sprinkling of whitish or yellowish dots like urates ; this is especially marked over heart. Pleurae aside from urates are negative. Lungs very slightly uniformly congested through- out. Aorta and branches are stiff, intima smooth. The liver is soft, deep brown color, architecture seems normal. The kidney has a smooth capsule and a smooth pale yellow surface. Organ is firm. Section sur- face is glistening and opaque, every lobule clear, pelves filled with pale yellow material, cortical areas irregular. Alimentary tract nega- tive. Microscopical section of kidney shows general topography re- tained, vessels very much injected, some showing thrombosis. Cortex slightly irregular probably by swelling of medulla. Tubular epithelium swollen and granular or desquamating and degenerating. Glomeruli vary in size and shape, mostly fill out the capsule. Capillary cells show some vacuoles. Some urate collections in tubules; practically all pelvic tubules have some urates. Interstitial tissue not increased. Blood vessel walls somewhat loose. Endothelium prominent. No areas of degeneration seen. Diabetes. Diabetes is an infrequent but well recognized disease among domestic animals. Its detection depends on a rather vague chain of symptoms confirmed by the dis- covery of sugar in the urine. For the suspicion that a wild animal was suffering with diabetes one would have CONSTITUTIONAL DISEASES 413 to rely upon great thirst, loss of flesh, depression, exces- sive urination and possibly cataractous opacity of the eye. Such a chain of symptoms has not been detected. At every occasion at postmortem that the bladder is full of urine, a routine examination is made. In this way we detected one case which seems to have been diabetes, the diagnosis being based upon the glucosuria and the lipemia. For some unknown reason a section of the pan- creas was not made, a regrettable matter since a definite purulent gingivitis existed and may have lain at the basis of an infective pancreatitis, well known to be the cause of certain cases of diabetes. The case is recorded in full since it is unique, no other case in a wild animal being fully reported. The arctic fox [Canis lagopus) ate and appeared well the day before it was found dead. Diagnosis — Diabetes mellitus. The animal was in good condition. The left conjunctiva was reddened, congested, edematous, with slight mucopurulent discharge in canthus. Muscles have a cloudy appearance. Fat lacks rich yellow color. The general impression of anemia is present. Lungs and pleura are normal through- out. Heart muscle is pale, firm and tough. The tricuspid shows thickening of the edge of posterior leaflet, the mitral shows slight sclero- sis of edge of mesial leaflet. The auricles are distended with clot. Left ventricular wall is greatly thickened. Upon incising the heart the surface of blood shows fine fat globules. Peritoneum is normal. Liver is slightly increased, surface smooth, edges rounded, consistency soft, color brownish red with yellow mottlings which are without defi- nite boundaries ; the section surface is moist, granular and opaque. The bile is fluid, green-yellow and the duct is patulous. The spleen is slightly enlarged and soft. The kidney is slightly enlarged, capsule strips easily leaving a smooth, purplish red surface; section surface is glistening, moist and exudes blood; consistency is slightly softened; cortical striae very distinct. The bulging cut surface and poor demarka- tion of cortex and medulla characteristic of acute nephritis are present. The organ shows fat globules in the expressed blood. The adrenals are very small, firm, brown, bean-shaped bodies with a brownish medulla. The bladder is slightly distended with turbid urine. Urine shows dark granular casts, compound granule cells, spermatozoa and a positive Fehling's test. Prostate is large and firm and a turbid material exudes from external meatus. The mouth shows several decayed teeth. In the neighborhood of last molars on left side of upper jaw a bead of pus exudes; further pressure results in no greater flow. The stomach is 414 DISEASE IN WILD MAMMALS AND BIRDS distended with a great quantity of undigested food and gas ; no worms. Serosa and wall normal but anemic. Duodenum normal. Jejunum con- tains numerous worms about 1 to 1.5 cm. long ; it is distended with gas. The pancreas is large, soft, like fat, white ; it extends between the layers of mesentery along the course of the duodenum; at first the pancreas was mistaken for fat. Histological Notes. — Spleen shows a distinct overgrowth of trabec- ulaB. Beyond this there is nothing pathological. Liver shows distended portal venules in which there are chains of bacilli. There is no especial fibrous overgrowth of capsule of Glisson; capillaries are choked with shadow corpuscles and here too, long chains of bacilli may be seen ; parenchyma cells show postmortem change. Adrenal is the seat of postmortem degeneration, not congested, nor is there any evidence of bacterial invasion. The kidney shows no interstitial changes, in fact the section seems to be entirely normal save for moderate congestion. Vessels show no bacteria. SECTION XV THE RELATION OF DIET TO DISEASE BY Db. E. p. CoRSOISr-WHITE Food in the widest acceptation of the term, means every thing ingested that goes, directly or indirectly, to growth, repair of the body, or production of energy, all of which phenomena must continue when food is mthheld or supplied in insufficient quantities. Under the latter con- dition the processes go on at the expense of the body tissues as these are protected only when the diet is ade- quate in every way. A proper diet, therefore, must be one on which an animal will attain maximum development, maintain a normal weight curve, show a minimum sus- ceptibility to disease, live out a full term of life, breed normally, and rear healthy offspring, capable of normal independent life after they are weaned. It must fulfiU the caloric needs of the body, and in young animals it must also supply the growth impulse. In its physical prop- erties it must fit the morphological demands of each type of gastrointestinal tract. In its chemical content it must supply all the elements found in the body in usable form, and in amounts sufficient to cover the needs of the body for growth, repair and waste.' To evaluate fully the influence of food on the individual animal it is necessary to study its relation: (1) to the type of alimentary tract, (2) to the type of bacterial flora and their metabolic processes, (3) to the chemical needs of the body, (4) to the changes arising in the catabolism and anabolism of all types of food, (5) to exercise or its lack, keeping in mind always the constant interdependence of all factors. Our knowledge of nutrition has to a very large extent 415 416 DISEASE IN WILD MAMMALS AND BIRDS paralleled the advances in chemistry, especially the re- searches into the structural make-up of living cells, the intermediate stages in their upbuilding and degradation and the products resulting from their physiologi- cal activities. Incorrect feeding both qualitative and quantitative un- doubtedly plays an important role in producing disease. In the early works on nutrition, the proportion of fats, carbohydrates and proteins was regarded as the essential point of a normal diet. The researches on the composi- tion of foods marked the first real epoch in this .history and Fischer's (1) studies on the variation in the composi- tion of proteins from different sources first introduced the idea of quality. Later Mendel and Osborne investi- gated the biological values of purified proteins, while at the same time McCollum and others were studying the value of the groups of proteins occurring in a single natural food stuff, were calling attention to the so-called vitamines, and were emphasizing the need of balanced inorganic materials. These studies have practically revolutionized our knowledge, particularly of the effects of badly balanced foods. They have clearly demonstrated that dietary values can, in all probability, be discovered only by careful biological study of feeding experiments together with the finer analysis of the components of the diet, especially of the protein and fat radicles. At the same time a definite appreciation of the role of each ele- ment in metabolism must be kept in mind. These varied studies on nutrition have shown that the chemical requirements of a diet are in their ultimate analysis essentially the same for all species of the higher animals — that is all require approximately the same amount of protein, fat, carbohydrate, etc., per kilo of body weight, while the morphology of the tract decides the physical properties of the diet. (1) Chemistry of the Proteins, Mann. THE RELATION OF DIET TO DISEASE 417 Relation of Food to Alimentary Tract. Food derived from animal sources is high in protein, readily digested, and highly putrefactive. This type of diet is suited to an alimentary tract which permits rapid passage through its leng-th, and is fitted with sturdy walls. The gastric section is simple, the intestine short and narrow with ill-defined separation of its: parts into small gut, cecum and colon. This type is found in all land Carnivora. The fish-eating carnivores have a strong tubular stomach and an enormous length of intestine, but no cecum. The omnivores occupy a middle place. In them the alimentary tract consists of a simple stomach, a short wide intestinal tube, and a more complex, although still comparatively simple, cecum which is generally longer than that found in the carnivores. This tract is too small to manipulate the bulky vegetable masses neces- sary to pro\'ide their minimum protein requirement, and too long and complicated to dispose quickly of the putre- factive animal tissue. Among these animals colitis is common, due to the fact that the shape and position of this part of the tract favors stasis, or at least a sluggish movement of its contents at a point in the digestive scheme where the food residue is rich in protein by- products, ready for bacterial gro^vth. The herbivores mth food derived from plants which requires a long period of time for its digestion, have, on the other hand, voluminous stomachs, or large ceca or both; and very long small intestine. In this tract the concentrated food of the carnivores would provide an enormously excessive protein intake or if only the protein requirement is supplied would leave the tract so empty that it would be unable to functionate. All studies in comparative anatomy demonstrate the fact that while neither a complex stomach nor a large cecum is essential to the digestion of vegetable food, a capacious and complex alimentary canal, as a whole, bears a relation to vegetable diet, particularly in the mammals. 418 DISEASE IN WILD MAMMALS AND BIRDS Either a highly developed concentrated glandular appa- ratus is added to the stomach, as in the wombats, beavers and dormice, or the stomach is subdivided, sacculated, or otherwise amplified as in the ruminants and herbivorous marsupials. Sometimes both complexities are combined as in the case of the sloths. If the simple stomach is retained, it is supplemented by a large sacculated colon or cecum, as in the horse. In birds, the proventricle is larger in meat- and fish-eaters, while the gizzard is more muscu- lar in grain- and insect-feeders, and the intestines are longer in those devouring coarse green grass and leaves. The length of the ceca is related entirely to the diet, the long ones corresponding to the diet which needs pro- tracted periods of time to exhaust its nutriment, j The Bacterial Flora. ' The bacterial flora harbored in the intestinal tract is closely related to the type of food and to the character of the alimentary tract. Levin(2) found sterile intestinal tracts in white bears, seals, reindeer, eider ducks and penguins when in the Arctic regions; but these same animals when they are brought to a temperate climate rapidly acquire intestinal bacteria. The function of the normal inhabitants of the tract is, probably, to protect the body against invasions of obnoxious species. Herter found in man that a few species adapt themselves to the digestive tract and control the growth of new-comers capable of doing injury. These common varieties become a source of danger only when present in large numbers. Bacteria which produce decomposition of food in the digestive tract are of three types : (1) Pure putrefactive anaerobes, (2) organisms both fermentative and putre- factive, but tending generally to antagonize the putrefac- tive anaerobes, and (3) fermentative organisms. In the stomach, fermentation of carbohydrates vnth. the (2) Ann. Inst. Past., 1S99, XIII, 558, and Skandinavisches Arch. f. Physiol., 1904, XVI, 249. THE RELATION OF DIET TO DISEASE 419 production of organic acids is a frequent occurrence.!; Putrefactive types are very rare except with pyloric stenosis, a condition which favors excessive fermentation by diminishing the tone and motility of the stomach and the amount of hydrochloric acid. This condition is further increased by excessive carbohydrate food. In general the products of fermentation tend to restrict putrefaction, yet both may be operative. In the small intestines, bacteria are always present because of the pro- tein richness of secretions, the rapid digestion of food and the slight or ineffectual antiseptic properties of intestinal juice, bile and pancreatic secretions. The putrefactive bacteria rapidly increase and decompose any protein that is miabsorbed — a process most marked in the colon because its shape and position favor stasis or slow move- ment of its contents. In general the greater the amount of unabsorbed and digestible protein and the longer the material stays in the intestinal tract, the greater the putrefaction. The meat-eating animals develop Gram- negative bacilli, while the carbohydrate-eaters show a predominance of Gram-positive types. Ingested food never contains the enormous amount of bacteria found in the feces. The alunentary tract with its contents forms a most efficiently combined incubator and culture medium, in which bacterial growth exceeds that of any known location both in intensity and complexity. The range of reaction and composition of nutritive substances at different levels of the intestinal tract is such that a great variety of bacteria capable of growth at body tem- perature develop. The prominent types that appear in the flora of each order of mammals are fairly constant in their occurrence. They depend primarily on food ingested, and show well marked seasonal variations, dependent again on changes in food. Faulty feeding may itself give rise to a toxic condition of the gastrointestinal tube, and thus often prepares this soil for the develop- ment of organisms. . 420 DISEASE IN WILD MAMMALS AND BIRDS The intestinal flora also changes along rather definite lines as the diet of the host changes from the monotony of the infant to the variety of the adult. At birth the tract is sterile, but bacteria soon make their entry through the mouth in food and water. The majority of these organisms pass to the stomach where many are destroyed, but a number travel to the intestines where they may gain a foothold. There is always a mechanical transportation of intestinal bacteria from higher to lower levels. A con- tinued preponderance of protein in the diet of all animals leads to a partial or complete suppression of the Gram- positive acid-forming groups and an increase of the pro- teolytic Gram-negative types ; while on the other hand an excess of carbohydrate leads to diminution or suppression of proteolytic activity and an increase in the fermentative organisms. Therefore the most important normal factor in determining the intestinal flora in health is the chem- ical composition of the ingested foods. The nature of the dominant organisms which develop in diets rich in carbohydrates varies with the carbo- hydrate itself. In all ordinary diets there are (1) starches — forms not readily fermentable, and (2) sugars — which are largely absorbed from the higher levels of the small intestine, leaving residual starches and proteins in relatively great concentration in the lower levels. There- fore the obligate fermentative organisms are prominent only in the higher levels, the facultative appear in the intermediate places, and the obligate proteolytic organ- isms in the lower intestines. This accounts in a measure for the great increase of lower intestinal disturbances in omnivores. Complete proteins resist putrefaction, but the products of protein digestion and of the intestinal secretions constitute the main substrata for putrefactive bacteria. Animal protein develops more active proteo- lytic bacteria than vegetable protein, which accounts for the greater predominance of putrefactive infections in THE RELATION OF DIET TO DISEASE 421 There are two important factors to consider in dis- cussing the influence of diet on intestinal bacteria: (1) The substitution of types, which frequently follows a monotonous diet, and (2) the change in metabolism of existing types of bacteria when dietary conditions are such that the intestinal medium at one or another level fluctuates in its content of usable carbohydrate and other nutrient. The nature and extent of these modifications and their effects upon the host vary greatly, not only qualitatively but quantitatively. An invasion of the tract by exogenous bacteria, as the dysentery bacillus, cholera, typhoid, etc., in food or water may lead to a more or less pronounced replacement of some of the normal intestinal tjipes by these alien organisms, and to the production of disease. The importance of all the foregoing facts concerning the changes in the food, in the intestinal cultural substrata and in the advent of new kinds of organisms was emphati- cally demonstrated in the marked fall in gastrointestinal diseases in carnivores after proper screening of meats. The simple protection of the food given to these animals eliminated the air bacteria which, entering from dust and flies, alter the chemistry of the meat before consumption or change the flora of the intestine after consumption. Normal organisms, or types indistinguishable from them, may multiply, through unusual conditions, extend their normal habitat, and eventually lead to abnormal reactions detrimental to the host. These facts throw considerable light on the site and character of gastrointestinal lesions found in various orders, a subject to be discussed more fully later. There are many intestinal disturbances of unknowni causation, in some of which bacteria presumably play a secondary part. The primary disturbance is due to the products resulting from the action of bacteria upon food. (Many toxic bodies are produced either before or after ingestion by the bacterial decomposition of carbohydrate. 422 DISEASE IN WILD MAMMALS AND BIRDS fat or protein, independent of any actual infection. The s^Tuptoms arising from bacterial decomposition of foods depend largely on the organism concerned and vary from a mild intoxication to a severe toxemia.; Relation of Dietaey Groups to Autopsy Diagnoses. Analysis of the autopsies on file from sole point of view of dietary habits of the animals gives rather inter- esting groupings of disease states, which apparently and, in some cases definitely, emphasize the relationship between food, metabolism and disease. (Table 19.) From this table a few facts stand out prominently. It is definitely sho^vn that both birds and mammals on a diet of mixed animal and plant tissue show a low per- centage of disease in the gastrointestinal tube, liver, pancreas and kidney. The mammals on this diet give the highest figures for anemias and degenerative osseous conditions. Birds on this diet show very little osteo- malacia, but a fair amount of anemia. Possibly this may be accounted for by the fact that all of them pick gravel and may be able from this to supply some of the inor- ganic deficiency. Carnivorous birds and mammals, on the other hand, show an exceedingly large assortment of gastrointestinal disorders, diseases of the accessory glands of digestion, and of the kidneys. Disorders of the thyroid gland are almost entirelj^ confined to carnivorous mammals — 7.5 per cent., compared to 0.25 per cent, in all other orders. Gout, while common among birds, was not present in any mammalian autopsy, while arthritis in mammals reached its highest record among grass- and grain-eating herbivora. The percentage of rickets was highest in the young carnivores (2.6 carnivores as against .4 per cent, in all other mammals), and was very rare among all birds. The succulent vegetable diet was lowest in its relation to degenerative visceral disorders and highest in acute gastritis ; the latter fact was probably due to the fermen- THE RELATION OF DIET TO DISEASE 423 tation of soft moist food that requires rather a long time for its primary digestion. This type of food has also a high and easily available sugar content which makes it a Table 19. An Analysis of the Pathological Findings Described in the 5,365 Autopsies from the Point of View only of the Dietary Habits of the Animals. The Percentage Results Represent the Proportionate Number of Cases of Each Pathological Lesion Found in the Entire Group of Animals on Each Special Diet without Reference to Zoological Orders. Disease states Malnutrition Food Poisoning Acute Gastritis Acute Duodenitis . . . . Acute Enteritis Acute Gastroenteritis . Chronic Gastritis Chronic Enteritis . . . . Colitis Acute Pancreatitis.. . . Chronic Pancreatitis. . Acute Liver Disease. . . Chronic Liver Disease Acute Nephritis Chronic Nephritis. . . . Myocardial Degene- ration Arterial Disease Anemia pernicious.. . . Anemia secondary. . . . Thyroid Disease Adrenal Disease Diabetes Osteomalacia Osteitis deformans. . . . Arthritis Rickets Gout Sore Eyes MaUgnancy Tuberculosis Mammalia 1860 .1 .3 3.2 .5 2.5 26.3 1.1 2. 1.9 .1 .5 .8 3. 9.1 4.5 .1 .1 .3 4.2 1.6 5.2 .05 32.6 6.3 .3 3.4 53.2 6. 5.6 2.2 1.7 1.3 6.3 12.2 11.6 .34 3.1 .32 .32 7.5 1.3 .2 .4 .3 .9 3.5 9.3 3. 19.9 2. 3. 1. .3 3.3 12.7 6.7 2.2 2.5 3.1 .5 3.1 29.2 .8 2.2 3.1 1.4 6. 12.4 7.8 1.1 2.2 1.5 .7 1.5 .05 .05 .9 .1 7. 25.3 .2 1.1 .4 .2 4.2 1.1 5.1 2.9 .3 .3 1.1 .3 12. .4 .2 2. 1.4 1. 38.6 1.4 3.3 1.2 2.8 2.5 6.7 6.7 2. 3.1 2.5 .2 1.3 1.2 8. 35.6 .3 1.3 .5 2.5 1.6 4.1 2.1 2.8 17.2 13.5 5.4 5.4 64.8 5.4 13.5 2.7 13.5 8.1 13.5 8.1 1.8 5.7 very favorable medium for many of the fermentative types of bacteria. Most of the lesions in this group were around the pylorus and upper duodenum. 424 DISEASE IN WILD MAMMALS AND BIRDS Overeating is a factor that must be borne in mind when considering the hay- and grass-eating herbivora. Packing of the rumen is a not infrequent discover5\ This condi- tion is also found in certain seed-eating birds. As a supply of food is constantly at the disposal of these ani- mals and exercise is prevented by captivity, continuous eating becomes their principal diversion. In this group also food poisoning was highest, a condition which may be due to (1) spoiled food, (2) poisonous substances in the foods, (3) fermentation of grass foods (spoiled hay or musty fodder). Malnutrition also, is higher than with any other diet, due probably to the somewhat meagre nutri- tious value of the food. This group also shows a high per- centage of acute pancreatitis, degeneration of the liver, myocardimn and arteries. Arthritis was present in this group 2.2 per cent., against 0.2 per cent, in all other groups. A study of Table 19 demands a constant recollection of the morphology of the tract involved and its main points of vulnerability, the bacteria capable of living on the particular type of food or its constituents and the by-products produced during the digestion and absorp- tion of these foods. Not one of these factors can be ignored in evaluating the influence of diet, which to be correct must supply elements in proportions that are chemically available for body needs (for instance. Von Wendt(3) found that more iron was required if the diet was deficient in calcium). These proportions must be worked out by carefully combined chemical and biologi- cal experiments. Malnutrition. There was one omnivorous beast, a Hamadryas Baboon, which represented the only true case of starva- tion, probably induced by nostalgia, as it never ate after coming into the Garden. Thirty cases of partial starva- tion or malnutrition are listed in our records, the (3) Skandinavisches Arch, fiir Physiologie, Vol. 17, p. 211, 1905. THE RELATION OF DIET TO DISEASE 425 majority among the rarer specimens, ten carnivorous, seven herbivorous and one omnivorous mammals, ten carnivorous and two seed-eating birds, due possibly to inappropriate diet or to some unknown factor that ren- dered the diet inadequate. At the autopsy nothing was found to account for death except the draining of all storage supplies. Starvation". The reports of studies conducted during long labora- tory fasts have been among the most valuable records for the understanding of the chemical requirements of diet and of the close chemical interrelationship existing between the different food factors. In absolute starva- tion life is very short, primarily because water is necessary for respiration, for dissolving products of metabolism and for preventing changes in digestive intes- tinal secretions. The amount of water needed varies with different species of animals. If the water is supplied, the organism is enabled to maintain its energy for continued existence from the destruction of its own tissues. The length of life depends upon the amount of protein ingested before the fast commenced, and the amount of stored fat and glycogen, especially that stored in the liver./ The mechanism of the results is similar. The animal body uses first its available glucose, and when this is partially exhausted burns its stored fat and protein. The fat combustion is usually defective, ketone bodies appearing in the urine in large quantities. The change from fat to protein metabolism accounts for the pre- mortal rise in metabolism which occurs usually a few days before death. The chemical composition and corpuscular richness of the blood is tenaciously preserved; glucose and protein concentration are practically normal up to the day of death. There is at times a slight increase in globulins and always an increase in fat due to its trans- portation from storage depots. The cause of death is 28 426 DISEASE IN WILD MAMMALS AND BIRDS primarily due to loss of substance in organs necessary to life and to an acid intoxication. Wasting occurs first in stored substances, fat, glyco- gen, etc., then in the least used organs. The bones usually show some rarefication. The animal, as a rule, dies from acid intoxication before atrophy of the organs is marked.} In the wild, when animals are forced to seek their food with the expenditure of much energy and where feasts are often followed by fasts, this using up of storage supplies Table 20. Detailed Analysis of the Various Diets Used at the Philadelphia Garden on Basis of 100 Grams of Mixed Food. Omnivora Carnivora Herbivora Succu- lent Vegetables Herbivora Coarse Food S B 1 1 1 S a 0! 1 pq 1 >. K 1 1 Protein Fat. 14.3 9.5 26.7 .034 .058 .497 .103 .263 .117 .338 .0032 11.5 7.2 41.2 .068 .093 .713 .284 .484 .377 .486 .0063 15.6 18.8 .058 .118 1.694 .421 1.078 .378 1.146 .015 17.2 .3 .109 .133 1.671 .373 1.148 .528 1.119 .0055 6.1 2.6 18.5 .067 .164 .538 .08 .556 .038 .134 .0018 3.2 .5 25.7 .025 .119 .242 .291 .342 .044 .125 .0012 6.4 2.2 35.9 .071 .289 .644 .089 .692 .073 .217 .0022 7.1 1.3 Carbohydrate Calcium Magnesium. . Potassium . . . Sodium Phosphorus. . Chlorin Sulphur Iron 51.2 .044 .16 .324 .261 .458 .063 .163 .0012 cannot help being a factor in preserving the integrity of the storage and eliminative organs. In captivity this cannot occur. Food is supplied regularly, exercise is lacking, consequently overloading and disease of storage and eliminative organs is more or less constant — a situa- tion very marked in the Carnivora. A further study of Table 19 in the light of the finer analysis of the ingredients of the diets, shown in Table 20, explains, at least in part, the high percentage of certain types of disease in relation to particular diets. In the food of the first group, the omnivorous mam- mals, there is a moderately increased carbohydrate con- tent and an unevenly balanced inorganic content, the THE RELATION OF DIET TO DISEASE 427 last being the factor most at fault. The calcium and phosphorus are both so low that at the best the animal could only be in equilibrium, while any drain of the fixed bases would sooner or later have to be replenished from the calcium and phosphorus storage depots, the bones. Osteomalacia is most marked in the Cebidae, monkeys whose diet is even lower in these same elements : calcium .025, phosphorus .116, and iron .0008 per 100 grams of food. The inorangic composition of all animals is grossly similar ; the typical digestion developed from the habitual diet of the animal explains the more apparent changes and variations in their reactions to certain deprivations. Irkegularities of Inorganic Metabolism. (Twelve essential elements are present in the body, namely: carbon, nitrogen, hydrogen, oxygen, phos- phorus, calcium, sulphur, sodium, chlorine, potas- sium, iron, magnesium. Of these, five are furnished by the protein molecule and three of the five are duplicated in the fats and carbohydrates; the remain- ing seven must be present in the mineral ash. These elements functionate in three ways, (1) as constituents of bone, (2) as essential elements of organic com- pounds, (3) as soluble salts in body fluids. Chlorine, sodium, sulphur are supplied in sufficient quantity with most diets. In the case of chlorine, marked differences exist between the herbivores and carnivores. The meat- eating mammals easily acquire sufficient sodium chloride from the flesh and blood of their victims, while the herbivores on the other hand, find in their vegetable food large amounts of potassium and very little sodium or^ chlorine which must therefore be acquired separately.) Both omnivores and herbivores crave salt, probably because this large potassium content of vegetable food tends to increase the sodium elimination. A deprivation of salt always leads to a distaste for foods rich in po- tassium. So far as is known excessive sodium stimulates 428 DISEASE IN WILD MAMMALS AND BIRDS protein catabolisra, and through the overstimulation of the digestive tract, may interfere with the absorption of food. Sulphur is largely taken into the body in organic combination ^vith the protein, (a very little inorganic sulphur appears in the drinking water) therefore if the protein requirements are adequate the sulphur will usually be adequate. Magnesium is abundant in meat and most plant tissues ; so that except in diets of highly refined foods, it is more often excessive than deficient. The other elements, calcium, phosphorus and iron are frequently insufficient, especially for animals on omniv- orous diet (cf. Table 20). Phosphorus enters into every living cell, and in cases of starvation is excreted up to the last. It is involved in practically all the cell functions. In the body it is present (1) as an inorganic compound in the bone tissues and blood where it helps to maintain neutrality, (2) as phosphorus-containing pro- tein, phosphatids and phosphoric esters of a carbohydrate, all closely associated with the cell and its nucleus. In foods, phosphorus occurs in the same positions, that is, inorganically or combined with protein, fat or carbohy- drate. It is not entirely proved but is very probable that the phosphorus in organic combination has the greater metabolic value, inasmuch as there is greater storage of nitrogen and stimulation of tissue growth on foods con- taining phosphorized proteins, fats, etc. It has been shown, however, that the animal body can satisfactorily supply its phosphorus requirements by inorganic phos- phates. /The omnivorous diet, even the widely varied diet of man, is very often deficient in phosphorus, a fact which becomes very important when we consider that the omnivorous diet produces many acid residues wliich must be neutralized, and that phosphorus is largely responsible for the maintenance of tissue neutrality. Voit showed that the phosphates excreted during starvation were THE RELATION OF DIET TO DISEASE 429 withdraAvii from the bones ; and there is much proof that during the daily metabolism a certain slight movement of phosphorus takes place. The metabolized phosphorus is excreted by carnivores practically from the kidney alone ; by herbivores almost entirely through the intestinal wall, while in the omnivores it is excreted by kidney and intes- tinal tract. Whether these facts have any real influence on the phosphorus need of different types is not alto- gether determined. Calcium also enters into many of the essential func- tions of life, coagulation of the blood, contractility of the heart, etc. Omnivorous diet is usually deficient in this element, which is very irregularly distributed both in animal bodies and plants. Insufficient amounts lead to deprivation of body tissues and to the production of osteomalacia-like conditions. Voit produced marked thin- ning of the skull bones and sternum by a diet poor in calcium. Steenbok and his associates had the same results in cattle by feeding ' ' shorts ' ' a diet rich in mag- nesium. Etienne (4) showed that an excess of magnesium in an otherwise well balanced food caused a continual loss of calcium. Adults stand a deprivation of calcium much better than children or young animals. They often show no symptoms and retain a normal blood content as the losses from the blood and soft tissues are promptly replaced from the bones. Sooner or later all these animals show weakness and flexibility of the bones. Osteomalacia occurred in 5.2 per cent, of the animals on an omnivorous diet, that is this number showed gross evidence of absorption of bone salts. This condition occurring in man and the lower animals is a generalized softening of adult bones that were at one time normally calcified. Three clinical varieties are recognized in man : a mild form seen in pregnant, puerperal and lactating women, a senile form in which the lesions are usually limited to the peh^s, and a severe progressive form (4) Jour. Physiologic et Path., Vol. 14, 108, 1912. 430 DISEASE IN WILD MAMMALS AND BIRDS encountered in both sexes and at any age. This last form ends in marasmus. Its chemical characteristic is a loss of calcium and phosphorus with retention of sulphur and magnesium. The progressive type has occurred very frequently among the CebidaB whose diet on careful examination, showed a protein content low in quantity, poor in quality, and especially deficient in the phosphorus-containing pro- teins and total fat. The carbohydrate was high. The ash was small in amount and predominatingly acid. The daily ration often showed only an unweighable trace of calcium, phosphorus or iron. Sodium, potassium, sulphur and magnesium, on the contrary, were present in amounts sufficient for equilibrium or in excess. The Vitamines A.B.G. were present but were not always correctly pro- portioned. The fat soluble A was low and in some daily rations was entirely lacking. Diet has at various times been proposed as at least one factor in the production of this condition, a premise that has gained considerable weight through the increase in the number of cases, both in man and in the domesti- cated animals, reported from the war-famine district of Central Europe where the dietary was restricted and unbalanced. It has been shown that w^hen calcium is low in the diet, the amount excreted materially exceeds the intake. Benedict (5) has further shown that even during absolute fasts calcium is excreted. The requirements of this element for man have been fairly well worked out, but for animals we have no standards. Still it seems certain from the foregoing observations that storage supplies are called upon very early in cases of depriva- tion, while in pregnancy and lactation when the calcium requirements are greatly increased, a reason is found for a higher incidence of osteomalacia. Steenbok and Hart (6) have shown that the skeletons of cows and goats gave (.5) Carnegie Institute Publication, No. 203, p. 247, 1915. (6) Jour. Biol. Chem., Vol. 14, p. 59, 1913. THE RELATION OF DIET TO DISEASE 431 evidence of a drain of inorganic salts during the produc- tion of milk unless the calcium and phosphorus of the diet were liberally supplied. In osteomalacia it would seem that inefficient diet, if not the cause, was at least a very potent factor in pathogenesis. The disturbance of the calcium-phosphorus-metabolism may be due to the deprivation of the alkaline salts as in the famine osteomalacia, to a drain from the alkaline storage of the body associated with an inefficient diet as in the osteo- malacia of pregnancy and lactation or to the combined action of a diet faulty in more than its salt content, which by the production of acid in its oxidation and by favoring the development of acid-forming bacteria, causes a drain of the body alkali for neutralization of the acid ; or it may possibly be due to a combination of all these factors act- ing through their influence on the ductless glands. Paget 's disease or Osteitis Deformans is a chronic constitutional process which usually involves all the bones of the adult skeleton. DaCosta (7) believed it to be a disorder of bone metabolism probably dependent upon absence or perversion of some internal secretion. We have had the unique opportunity of observing three cases of this disease in Cebidse, the family of monkeys which has presented the highest incidence of osteomalacia. The experience is all the more interesting because of the typical picture presented by the specimens, and of the absence of references in the literature on the subject, to the occurrence of the malady in wild animals. The inter- esting point about these cases lies in the fact that the disease appeared in all three only after lime water was added to the diet to supply the deficiency of calcium. Search for literary record of the disease brought to light a case in a horse that Barthelemy (8) described, but this involved the epiphyses of the bone while osteitis (7) Publication of the Jefferson Medical College and Hospital, Vol. 6, p. 1, 1915. (8) These de Lyon, 1901. 432 DISEASE IN WILD MAMMALS AND BIRDS deformans is confined as a rule to the diaphyses. This case was probably more closely allied to osteitis fibrosa cystica. Goldman (9) described examples in fowls and Rossweg (10) refers to specimens in domestic goats and monkeys. Many of these cases first come under obser- vation through fractures, an accident common to osteo- malacia, but very rare in well developed osteitis deformans. The diet of our monkeys was exceeding low in those substances essential to bone development. Sherman (11) has shown that the calcium balance is reg- ulated to a certain extent by the calcium ingested, and that when the diet is poor in this element, the output materially exceeds the intake, a condition which is definitely changed when the animal is put on a diet liigh in calcium. So far as we could find there are no recorded studies of the mineral metabolism of beginning cases of Paget 's disease. It seems possible from the study of osteomalacia that the low mineral and otherwise faulty diet, added to the symptoms produced by that diet might so disturb the chemical equilibrium, directly through the neurotrophic mechanism or through the perversion of the ductless glands, that the mere addition of the lime water might entirely change the pathological picture. This is in accord with the histology where the initial lesion is resorption of bone followed by irregular proliferation. It is also in accord with the probable chemistry of cal- cification. These animals all showed a lowered carbon- dioxide-carrying-power of the blood, and therefore lowered ability to carry calcium in solution. It is possible that Paget 's disease is but a stage in a deficiency disease, a faulty reparative response through a disordered neuro- trophic mechanism, or through a perversion of the glands governing calcium metabolism. Such perversion could (9) Verein Freiburger Aerzte, May 30, 1902. (10) Vet. Med. Inaug. Diss. Giessen, 1913. (11) Chemistry of Food and Nutrition, Macmillan, 1918. THE RELATION OF DIET TO DISEASE 433 be caused by an improperly balanced diet, or by the addition of an excess of calcium to the diet of an animal whose body fluids were unable by reason of previous faulty diet or other disorder, to hold it in solution. In young animals the calcium demands are much higher than in adults, a need met in the high calcium content of breast milk, a content in excess of almost every other food, but apparently just sufficient to maintain calcium equilibrium. After it is weaned the young animal frequently shows disorders of its inorganic metabolism. Herter estimated that a child should store at least 0.1 gram of calcium daily and he described many cases of arrested bone develop- ment occurring during infancy and early childhood, because of an inefficient assimilation of calcium. One case, probably of this character, was found in a Hama- dryas Baboon (Papio hamadryas) a typical example of infantilism. The animal was an adult male about half the size of an adult female. His skin was fine and more delicate than normal, the bones were small and slender, contour of body was that of a young animal, genitalia were imperfectly developed, thyroid gland apparently normal, gastrointestinal tract atrophic, associated was a slight arthritis, portal cirrhosis of liver and diffuse nephritis. First among the results of inorganic insufficiency in youth stands Rickets. This disease occurs in children starting usually at about the sixth month and continuing with irregular remissions for several years. The bone changes, which are the most prominent, are always associated with more or less severe anemia, a general lowered resistance and flabby musculature. The excre- tion of calcium is very high in the feces and low in the urine. There is a frequent negative calcium balance dependent upon the great loss in the feces. Healing is preceded by a hyperretention of calicum and a relative increase in the urinary calcium. The excessive loss of calcium in the feces is not brought about through the 434 DISEASE IN WILD MAMMALS AND BIRDS agency of fats because fat could only remove calcium as insoluble soaps and these are not at all increased. This fact contradicts the idea of fat starvation as a cause of rickets. Rowland and Kramer found that the blood in active rickets bad a normal or slightly lowered calcium content, but a regularly reduced phosphorus content. The latter deficiency was extreme at times. They ascribe to this deficiency the failure of the bones to calcify. It can be readily understood that a decrease of phosphorus in the blood would render difficult the precipitation of calcium phosphate. Recently two series of studies, the first by Pappen- heimer, Zucher and McCann and the second by Shipley, McCollum, Park and Simonds have shown that rats fed on a diet low in calcium but with a sufficient amount of fat-soluble vitamine and phosphorus develop a bone con- dition with many fundamental resemblances to rickets. They were also able to produce the condition with an excess of calcium and deficiency of phosphorus. On the first diet, the condition differs from rickets in that the arrangement of the proliferating zone of cartilage cells is maintained and the evidence of bone resorption in the diaphyses is excessive. A diet deficient in both calcium and phosphorus leads to an atypical rickets. In the animals autopsied at this Garden rickets occurred very much more frequently in the flesh-eaters than in any of the other dietary groups. On closer analysis it was found that rickets in almost every case appeared in the carnivores which did not receive bones as a part of the food. Rickets occurred frequently in the omnivorous macaques which however did not show osteo- malacia, although they belong to the same dietary group as the Cebid^e. The reason they did not suffer the latter disease while adult but had rachitic young is probably due to the fact that this monkey group, which breeds best in our Garden, receives in addition to the diet given to CebidaB one raw egg. This increased the calcium content THE RELATION OF DIET TO DISEASE 435 of their food more nearly to the requirements of these mammals. These monkeys also have mouth sacs, which enable them to acquire more food per Mlo of body weight than the smaller Cebidae which are not so advantageously equipped. The food even in the amounts consumed by the macaques is low in calcium, phosphorus and iron. It is verj^ possible that there are enough of these ingredients present as a rule, to maintain the animal in organic equilibrium, during normal life, and possibly even enough to supply the needs of the embryo but not sufficient to maintain the young during the period of lactation. A few macaques dying during the delivery of young showed slight osteomalacic changes in the pelvis. This was notably present in one described in detail by E. A. Schumann. The calcium requirements of the female are always much increased during pregnancy and lactation due to the withdrawal from the mother to meet the needs of the embryo and nursling. Forbes and Beegle(12) found that lactating animals made heavy drains on their stor- age calcium even when the diet was liberal and the animal was storing nitrogen. Iron is the essential element of hemoglobin and chromatin — the body constituent most directly concerned with the process of oxidation, secretion, reproduction and development. The iron of the food is absorbed from the small intestines, enters the circulation through the l}Tnphatics, is deposited in the liver, spleen, and bone marrow and eliminated through the intestinal walls. There is very little iron reserve in the adult body; and as a result any failure of the intake to equal the output causes an immediate reduction of the hemoglobin. Voit found that the iron eliminated in the feces of starving dogs, or dogs on a diet low in iron comes from the body through the intestinal walls. Medicinal iron stimulates the production of hemoglobin and red blood cells but (12) Ohio Agricultural Experiment Station Bull., 295. 436 DISEASE IN WILD MAMMALS AND BIRDS whether it is directly employed in the production of hemo- globins has not been proved. Undoubtedly most of the extra iron given ^vith the food passes through the ali- mentaiy tract ^vithout being absorbed or metabolized. The greater the amount of iron in the food, the greater the influence of the inorganic iron. Anemia occurred in all the animals we examined at least four times as frequently in omnivorous as in all the other dietary groups, a fact probably explained by the low content of iron and calcium in this diet. Both Von Wendt(13) and Sherman (14) demonstrated that larger amounts of iron were required to maintain the iron equilibrium when the amount of cal- cium was low. Herter has sho^vn that many anemias are associated with intestinal putrefaction. The carnivores, however on a diet that putrefies very easily and on one in which the iron content is apparently of distinctly lower nutritive value than that of the iron found in milk, eggs and vege- tables, presented an anemic incidence of only 0.32 per cent. Tliis is probably due to the excellent hygienic care of the meat foods and to the morphology of the carniv- orous intestinal tract, which is short, straight and fashioned for quick elimination. The cases of anemia steadily increase among the animals as the conformation of the tract approaches the omnivorous type with the longer and wider hind-gut. Herbivora, obtaining their iron from vegetable sources, are much less liable to blood disorders. The iron needs of the female are greater than those of the male because of the drains of pregnancy and lactation. Young animals demand more iron than adults. All exclusively breast-feeding animals have a considerable storage of iron in the body at birth, while those that eat food immediately have no such supply. Bunge's(15) (13) Skandinavisches Archiv. f. Physiologie, Vol. 17, p. 211, 1905. (14) Bull, 185, Experiment Station, U. S. Dept. Agriculture, 1907. (15) Physiological and Pathological Chemistry, Blakiston, 1902. THE RELATION OF DIET TO DISEASE 437 experiments showed that breast-fed animals contained about six times as much iron as the milk that nourished them. The iron content of all these animals is lughest at birth, remains constant during the suckling period and then rapidly decreases to the adult standard. After this level is reached the iron metabolized must be supplied from the food if the hemoglobin is to be spared. The functions of all these inorganic substances are intimately interrelated and in places interchangeable. Calcium is capable of correcting disturbances of inor- ganic equilibrium in the animal body whatever the direc- tion of the deviation from the normal may be. These interrelationships are most involved in the maintenance of body neutrality. The normal processes of metabolism involve a continual production of carbonic, phosphoric and sulphuric acid which must be immediately disposed of if the neutrality of the body is to be permanent. The factors involved in this are carbonates, phos- phates, ammonia and proteins. Carbon dioxide is the chief excretory product but is at the same time a normal constituent of the blood and as such, is an important factor in this physicochemical reg-ulation. There is a tendency for the respiratory mechanism to hold its carbon dioxide tension nearly constant. Late investigations have sho^m that lowering of this tension is an early sign of beginning acidosis. When food such as protein, is taken in excess the strongly acid residues are neutralized by the sodium and potassium carbonates which are elimi- nated with a corresponding loss of sodium and potassium. The carbon dioxide tension diminishes, 37.2 per cent, on a high protein as against 43.3 per cent, on a vegetable diet. If this excess is long continued, the result may be, and often is, an increased elimination of the base-forming ele- ments which if not made good tends to diminish the body's reserve alkalinity. A diet with a preponderance of basic elements leads to an alkaline urine with an increased uric acid solvency and an increased carbon dioxide tension 438 DISEASE IN WILD MAMMALS AND BIRDS and reserve alkalinity. A diet with a preponderance in the acid-forming elements, on the contrary, leads to an increased urinary acidity and urinary ammonia, decreased ability to dissolve uric acid and lowered car- bon dioxide tension and alkaline reserve. Deficiencies of Vitamines, Recent investigations have shown that diets furnish- ing sufficient amounts of protein, fat, carbohydrate and inorganic salts may yet prove inadequate for growth or even for maintenance. Hopkins, (16) feeding rats on puri- fied food mixture was unable to obtain any growth until he added small quantities of milk or of the ether-soluble portion of milk but with this addition growth progressed in the normal manner, but it was out of all proportion to the energy or protein value of the addition. Five substances of this character, called by Funk (17) Vita- mines, have been described, two of wliich have definitely established a place as essential food factors. According to him, pellagra, rickets, scurvy and beriberi are the result of a lack of these unidentified but specific and indispensable food complexes. The first vitamine isolated was the fat soluble A, an adequate supply of which is necessary, not only because of its stimulating growth properties, but because its absence produces a serious condition of the eyes and, at times, marasmus leading to death. Xerophthalmia is a common condition in animals on experimental diets. The eyes are swollen, the cornea inflamed and often opaque while blindness and death invariably occur unless the die- tary error is corrected. McCollum(18) rescued animals almost at the point of death by butter or other fat rich in this vitamine. Opacities of the cornea are often seen in the animals in this and other gardens among ungulates — hay- (16) Joum. Physiol., 1912, XLIV, 425. (17) Die Vitamine und ihre Bedeutung fur die Physiologie und Pa- thologic mit besonderer Beriicksichtigung der Avitaminoses, Wiesbaden, 1914. (18) Newer Knowledge of Nutrition, Macmillan, 1919. THE RELATION OF DIET TO DISEASE 439 eating mammals ; four advanced cases were found, three in seed-eating birds and one in a fox on a diet made up solely of horse muscle. The quantity of vitamine A present in muscle, hay and seeds is very small. It is supplied in largest amounts in milk, eggs, glandular organs and leaves, substances which were very low or absent in the diet of all the affected animals. This xerophthalmia has been reported in man on several occa- sions, especially by Hrdlicka(19) in American Indians, by Mori (20) in 1400 Japanese during a period of food shortage (this epidemic was cured by the addition of chicken livers to the diet), by Bloch(21) in forty-seven children of Copenhagen fed on a fat free milk who were cured by the administration of cod liver oil. The disease is not however a fat starvation, as it is entirely unin- fluenced by vegetable fats which do not contain this Adtamine. Beriberi is an established deficiency disease, fre- quently seen among the poorer classes of the Orient whose diet is limited to polished rice and fish. It has appeared in Labrador coincident with the excessive use of bolted flour. A similar condition has been induced in pigs and cattle by a diet made up of an excess of cotton seed meal and tankage. Two forms of the disease are described: (1) acute or wet, characterized by marked edema, ascites, hydropericardium, hydrothorax, edema of the lungs, and a congestion of the spleen, liver, kidney, and heart muscle, (2) chronic or dry, characterized by polyneuritis. The disease was first produced experimen- tally in pigeons by Eijkman(22) in 1897 by means of a diet of polished rice. The paralysis appeared in 2-3 weeks after the diet was initiated. Eraser and Stanton (23) in 1907, found that it could be cured by an (19) B7ill., 34, Bureau of Amer. Ethnolo^. (20) Jahrbuch. Kinderheilk., 1904, LIX, 175. (21) Journ. Am. Med. Assoc, 1917, LXVIII, 1516. (22) Arch. Path. Anat., 1897, CXLVIII, 523. (23) Lancet, London, March 12, 1910, 733. 440 DISEASE IN WILD MAMMALS AND BIRDS alcoholic extract of rice polishings. Funk (24) later determined the vitamine character of this extract. In pigeons and fowls experimental feeding usually results in the chronic or polyneuritic form, expressed by a typical degenerative inflammatory condition of the peripheral nerves. In pigs, on the contrary, Rommel and Vedder(25) produced both types, though the acute or wet beriberi appeared more frequently. In rats the same deficiency causes multiple hemorrhages in the cerebellum and midbrain followed by a degeneration of the associ- ated nervous structures. It is possible that the pathol- ogy following a lack of the vitamine B or in fact any of the vitamines will vary with the different species or with varying demands of different individuals. This antineuritic vitamine affects more than the nervous system, and it is possible that all vitamines may have mder effects than are at present described. Scurvy was the first condition to call attention to diet as a cause of disease. It occurs in man when deprived of fresh vegetables. That faulty diet was in some way the cause of scurvy has been known for many years, but only since 1905 has there been any systematic attempt to determine the peculiar value of the curative foods. At this time Theobald Smith (26) called attention to a disease suggestive of scurvy^ which developed in guinea- pigs fed on a diet of oatmeal. This observation was con- firmed by Hoist and Frolich(27) who stated that the disease could be prevented by the addition of fresh milk or cabbage, because in these foods there was present an antiscorbutic or C vitamine. This unidentified substance was easily destroyed or diminished by heat or an alkaline medium. It was found in rather large amounts in succulent vegetables and fruits. McCollum(28) and his (24) Lancet, London, 1911, II, 1266. (25) Bull, Dept. of Agriculture, Dec. 13, 1915. (26) Bureau of Animal Industry, 1895-96, 172. (27) Z. Eyg. u. Infektionskrankh., 1913, LXXV, 334. (28) Jour. Biol. Chem., 1917, XXXI, 229. THE RELATION OF DIET TO DISEASE 441 coworkers showed that the oat kernel was low in inor- ganic salts and vitamine A and poor in the quality of its protein; but with these faults corrected it proved to be a complete food for rats. McCollum also found that scurvy developed more readily in animals if the physical properties of the diet favored constipation. He was able to delay the onset of the disease in guinea-pigs for a considerable period by the addition of mineral oil which has no food value, or phenolphthalein, a cathartic. At the same time, Jackson and Moore, (29) found the cecum of all guinea-pigs dying of scurvy, packed with putrefying feces. They were able to produce a mild type of the dis- ease by the injection of the diplococci isolated from the swollen joints. From these observations it seems safe to conclude that scurvy may not be purely a deficiency disease, or even a simple dietary one, although the presence of a vitamine influence is not excluded ; but it is probably the result of a bacterial invasion of tissues debilitated by a faulty diet and by the toxins produced by the putrefactive bacteria developing in a diet unsuited to the anatomical demands of the alimentary tract. This theory receives support from the fact that pasteurization destroys all aciduric bacteria, allowing only the spore-forming putrefactors to develop ; and from the fact that scurvy develops more frequently in children on stale pasteurized than on stale raw or boiled milk. In this Garden no suggestion of scurvy has been noted. Pellagra is very definitely a disease of poverty en- demic for years among the poor, especially in the moun- tains of Northern Italy, It has been under observation in the United States since 1907. So far as is known no cases have been observed among animals. Opinions dif- fer as to the role of diet in the etiolog}^ but the results of recent studies seem to show that uncomplicated cases of average severity clear up entirely on a diet rich in " (29) Jour. Infect. Dis., 1916, XIX, 478. 29 442 DISEASE IN WILD MAMMALS AND BIRDS animal protein. No vitamine deficiency has so far been determined. Wilson's careful studies of the diets known to have produced the condition show that the etiological factor lies in a deficiency of the protein molecule. The results of Goldberger(30) corroborate this fact, and he concludes from his latest studies that '' the dominating role of diet in the prevention and causation of pellagra is referable primarily to the character of the protein sup- ply or to the specific quality of the aminoacid makeup of the protein supply. ' ' Just what aminoacid or combina- tion of aminoacids it is, has not been determined, nor has the possibility of a vitamine alone or in combination Avith the aminoacid factor been absolutely excluded. The principal influence of the omnivorous diet is toward those degenerations arising primarily from im- balances in the inorganic makeup, or to insufficiencies of certain necessary factors. The vitamine deficiencies are markedly less prevalent in animals than in man whose food is less often consumed in its natural state. It is now known that much of the injury and loss of nutritive value in foods is produced by the processes involved in preparation, preservation, refinement and storage. Whenever the choice of food is not restricted, vitamine deficiencies do not occur. The vitamine requirements probably differ in different species and in individuals from the same species according to their environmental and individual variations. It is very possible that if the diet is low in vitamine content there may arise conditions of relative deficiencies ; and McCarrison has sho^vn that a vitamine deficiency associated with a high fat or car- bohydrate content may disturb the balance of the endo- crine glands. It is however to the inorganic content of the omnivorous food that most of the disturbances pecul- iar to this diet are to be assigned. With the flesh eating animals and birds the records present a very different picture. Disorders of the diges- (30) Jour. A.M.A., 1922, 79, 2132. " THE RELATION OF DIET TO DISEASE 443 tive tube, of the storage organs, of the organs of elimina- tion and of the endocrine glands predominate. Their diet is low in carbohydrates and, at times, in fats and very- high in protein. Bone supplies the inorganic salts, which in this Garden is fed only to the larger mammals. The carnivorous birds get their inorganic supply from mice which are eaten entire. The carnivores are as a rule large and are given to active fighting or to long flights. In the mid, very probably there are long periods between feasts, while in captivity the food is always plentiful and regu- larly supplied. This regularity added to the lack of exer- cise, particularly among the larger animals, must lead to excessive demands upon the storage and eliminating organs. Storage is always promoted by rest and liberal diet, and cleared away by exercise and starvation. The life of these birds and mammals, moreover favors inac- tivity of the bowels, which, together with the highly putre- factive diet adds another serious factor to a problem which in gardens is almost insurmountable. Irregularities of Carbohydrate Metabolism. The carbohydrates are derived from the glucose and glycogen of the meat and from the protein molecule. They are absolutely less than in the diet of herbivores but be- come a factor in the disorders of this group because of the lack of exercise and the regularity of feeding. In digestion the carbohydrate becomes available for absorp- tion and bacterial growth in the upper small intestine and appears on the other side of the intestinal wall as blood glucose in which form it is burned for energy or stored as glycogen for the future maintenance of the blood glucose. The blood of different animals has a glucose concentra- tion between 0.05 -0.1 which for each species is quite con- stant, as it is regulated by the coadaptation of four factors: combustion, fermentation of glycogen, forma- tion of fat, and elimination from the kidney. In excessive 444 DISEASE IN WILD MAMMALS AND BIRDS feeding the amount needed for energy is burned, the remainder is stored in the liver up to its capacity, then in the muscles and other cells, after which fat is formed and all further excess is eliminated by the kidney. Over- feeding causes an immediate overloading of the oxidative mechanism mth symptoms of gastric disorder, achylia, and at times acid fermentation with irritation of the stomach walls and the development of bacteria in the organ. This is frequently followed by glycosuria, several types of which are described: (1) associated \rith an increased concentration of glucose following excessive ingestion exceeding the normal glycogenic function of the liver, a form common among the Herbivora, (2) that due to a reduction of the glycogenic function of the liver, (3) that associated with disease of the ductless glands in which the resulting glycosuria probably depends upon the influence of these glands upon the pancreas, (4) that dependent upon the defect of glycolysis or to an over- stocked liver seen in gout, obesity or hypertrophic cirrho- sis, and (5) renal glycosuria due to a lowering of the renal threshold and usually associated with gout, arterio- sclerosis or chronic nephritis ; this last is best explained on the ground of increased renal permeability. Normally when the blood sugar concentration rises above a certain level the elimination via the kidney begins and continues until the blood has again reached its normal concentration. The relation of the kidney to glucose concentration is not constant and variation is always toward the side of lesser elimination while the kidneys become accustomed to the higher level. Diabetes, a disease of the islands of Langerhans in the pancreas, is essentially a disturbance of sugar metab- olism always associated with an exaggerated and defec- tive fat and protein combustion. It is not only that the diabetic has lost the faculty of combustion but these abnormalities all establish states of intoxication to which the diabetic must sooner or later succumb. Among lower THE RELATION OF DIET TO DISEASE 445 animals the disease is rare. Dogs are most frequently affected (about 1 in 12,000 deaths). It has also been described in horses, cattle and monkeys. In our records there was one case an Artie fox {Canis lagopus) present- ing a typical picture. Degeneration of the islands of Langerhans was seen in three other animals, but there was no other evidence of diabetes. This disease is not due to diet but to the absence of a normal ferment (pan- creaticozymo-excitor) for one particular type of food. iRREGULAErriES OF FaT MeT ABOLISH. Disorders of fat metabolism are very rare among lower animals notwithstanding the fact that fat even in the carnivorous diet, represents about 13 per cent, of the whole intake. It plays two important roles in the body, storage for energy reserve, and as a most essential structure in cellular protoplasm, in which position it joins with protein in complex combinations of still unknown composition which present to a striking degree the phenomenon of absorption. Very marked biological differences exist in the value of fats from different sources, due to the presence or absence of vitamines. The body fat is derived from the fat of the diet or is synthesized from glucose. The former is specific to the fat consumed while the latter is specific to the animal. In omnivores the type depends upon the varying extent to which animal fats enter the diet, in carnivores it depends almost entirely on the fat intake, while in the herbivores practically all the fat is synthesized from the carbo- hydrate. On digestion, fat splits, yielding a glycerol and fatty acid which are collected in the lymph spaces of the intestinal mucosa, there changing to some complex com- bination which is not only soluble but diffusible. Fatty infiltration and fatty degeneration are condi- tions of much pathological interest and of great frequency in captive animals. The researches of Mansfield (31) have (31) Pfluger's Arch.,\^m (129), 63. 446 DISEASE IN WILD MAMMALS AND BIRDS throAvn considerable light upon these conditions. He found that the total fat content in cases of most marked degeneration was normal or reduced. The proportion of fat free from protein was increased and the firmly bound fat decreased. This increase is due to neutral fat brought from without the organ by the blood when for any cause the oxidative powers are decreased, and setting free of the previously invisible intercellular fat and lipoids, which are normally present in the cells, by autolytic or physicochemical changes. This condition is pretty evenly distributed among the dietary groups, the liver being most commonly involved. The hepatic cells are easily degenerated by the toxins or other harmful substances passing through the organ and become passive and unable to throw off or to utilize the deposited fat. In all prob- ability the same general situation occurs in the atheromatous changes in arteriosclerosis which on this diet shows a high incidence. The causative agent is prob- ably some poisonous substance, possibly a protein degradation product, indol, pressor substance, acting on the intima over long periods, or at irregular but often repeated periods causing first destruction then fat accu- mulation. It is also possible that it may be caused by repeated absorption of some sensitizing protein. Arteriosclerosis in these animals is often closely asso- ciated with nephritis. Obesity may result from excessive ingestion of food in individuals whose habits are sedentary and whose digestions are active or it may come from an inherent abnormality of metabolism dependent upon ductless gland disease. It is very common in castrated animals. The obesity of overeating is always of milder type than that associated with endocrine disturbance. So far as is known there are two main disorders of fat metabolism — the failure of the diabetic to form fat from glucose, and acidosis, the inability of the organism to burn fat beyond betaoxybutyric acid, acetoacetic acid, or ace- THE RELATION OF DIET TO DISEASE 447 tone. The symptoms are unsteadiness of gait, stupor, coma, air hunger, in all of which the essential features are due to the impoverishment of the body in available bases. In infants this frequently follows an excessive fat diet. It is also common in starvation due to the deprivation of sugar. It is associated with phosphorus poisoning, nar- cosis, carcinoma, liver disease, inanition, etc. It has been produced experimentally by the administration of acids or by foods deprived of their bases. The excess of acid in the body whether produced in the body or introduced from without must be neutralized in part by the ammonia manufactured in the ultimate metabolic transformation of the protein and by the alkaline salts of the blood and tissues. When alkali is reduced the carbon dioxide accu- mulates in the tissues, blocking oxidation. The urine immediately shows an increase of ammonium salts, a decrease of the urea and an increase in the output of sodium, potassium, calcium and magnesium, which last two are dra^vn from the bones. Symptoms do not arise until the fixed alkalies are exhausted; and they are immediately relieved by the administration of alkalies, except in those cases of starva- tion where the administration of sugar and the subsequent sparing of the fats relieves the situation. In herbivores, acidosis does not follow starvation, but, on the other hand, it is markedly easier to excite it in herbivores than in carnivores whose heavy protein diet produces more ammonia, which better enables the animals to protect their fixed alkalies. The acid intoxication of infections arises from different causes and is dependent on the intensity of the type of infection; but ultimately it also depends upon the depletion of the fixed alkalies. Ireegulaeities of Protein Metabolism. Fat and carbohydrate disturbances are not infrequent in carnivores, but it is with the protein fraction of the diet that most of the trouble is connected. Natural foods 448 DISEASE IN WILD MAMMALS AND BIRDS contain several proteins or groups of proteins, whose biological adequacy depends upon their yield of amino- acid. Experiment has shown that many proteins are entirely lacking in one or more of these essential radicles ; and no food can be adequate unless it contains at least all the aminoacids that the individual animal is unable to manufacture for itself. So far as is known, no animal can produce in itself either lysin or tiyptophane. Gliadin, the principal protein of wheat and lacking in lysin, is unable to support gro^\i:h even when given in amounts sufficient to insure the storage of nitrogen, and is associated with a diet adequate in all other factors. Absence of tryptophane prevents not only growth but maintenance. Any of the aminoacids, whose radicles are contained in tissue pro- teins, may contribute to the maintenance of adult equilibrium ; but no growth occurs unless all the necessary groups are present. Except in laboratories, diets are never made up of isolated proteins, but they are often composed of proteins derived from one plant and are often deficient. McCollum and his associates in their studies showed that while there were pronounced differ- ences in the composition of many foods used by men and animals not only in their protein content but in water, fats, carbohydrates, etc., yet in the combinations found even in rather restricted diets, the errors, as a rule, cor- rected each other. During digestion the protein molecule is broken down into the component aminoacids which are absorbed and synthesized in the intestinal walls, and appear on the other side as the specific blood protein, which serves as the substrate for the anabolism of all the special tissue proteins. Excessive protein is stored to a slight extent as aminoacid for the future maintenance of the blood pro- tein, the integrity of which is tenaciously protected during hibernations, sexual migrations, and even during starva- tion. The animal body tends to adjust its nitrogen THE RELATION OF DIET TO DISEASE 449 metabolism to its nitrogen supply; the adjustment requires an appreciable amount of time. A diet changed to a lower nitrogen level results in a continued loss of nitrogen, increased combustion of fats and carbohydrates. The animal makes no apparent effort to reestablish equilibrium, and sooner or later digestive disturbances and loss of strength occur. If, on the contrary, the protein is steadily increased after an animal has established equilibrium, the nitrogen metabolism increases and the level of nitrogen equilibrium rises to higher and higher levels. There is, at the same time, a lowering of the fat combustion, an increase in the respir- atory quotient and in the heat production. The excess protein must be split, deaminated, burned and eliminated. Fifty-five per cent, of the intake is converted into glucose which is burned and the excess stored as glycogen. The sulphuric acid formed during the protein cleavage is neutralized by the body alkalies. In these cases the liver is often congested and enlarged. The urine shows excess of urea and ammonia. At times the excess, being so great that it cannot be absorbed, undergoes chemical and bac- terial decomposition which causes digestive disturbances, torpor and constipation. The organisms associated with protein food are usually the putrefactive types which break the protein molecule into the aromatic bodies, phenols, indolacetic acid, indolpropionic acid, skatol, etc. These bodies on absorption are believed to give rise to hypertrophy of the adrenal, interstitial changes in the kidney, and arterio- sclerosis. Another group of substances, pressor bases and amines, are manufactured by certain anaerobes acting on proteins. These, when fed by mouth, are detoxi- cated by the liver cells, but when formed below the portal circulation, give rise to anaphylactic phenomena — urticaria, etc. Certain other organisms give rise to soluble toxins as in botulism and thyrotoxicon poisoning. 450 DISEASE IN WILD MAMMALS AND BIRDS All these types of toxins will destroy if they act acutely in sufficient concentration; or as is more common, if they act persistently over long periods, or at oft recurring inter- vals they will cause serious injury to the tissues coming in contact with them, and have a part in the production of cirrhosis of the Hver, chronic nephritis, myocarditis, arteriosclerosis, etc. All foods have a limit beyond which they are excreted untouched or imperfectly oxidized. Many of these partial oxidation products of protein are in themselves toxic and may also be a source of these degenerative organ condi- tions. The pathological material studied by us showed a marked decrease in gastrointestinal diseases in close asso- ciation with the more hygienic care of the meat foods. Always associated with the protein foods are the nucleoprotein complexes, which are split by both bacteria and digestive juices into globulins and nucleic acid, and then through the agency of a special enzyme, into purin bases and uric acid, in which forms they are excreted in the urine and feces. The oxidation of purins is never complete. Gout, representing the pathology of purin metabolism, is a paroxysmal inflammatory disturbance, due to the deposition of sodium urates in the joints or in the internal organs, usually accompanied by a fibrosis especially in the liver, kidney, arteries, etc. The disease occurs almost exclusively in birds. Isolated cases have been described in dogs, horses and hogs, but among lower animals it is undoubtedly very rare. In birds it is most frequent in the carnivores — 4 per cent., as against 0.02 per cent, in all other groups. It is higher in fish-eating birds than among the flesh-eaters. The avian gout is usually of the visceral type and was most marked in its distribution over the organs in the Anseres and Psittaci, birds whose diet apparently is not unduly heavy in nucleoproteins, but whose tract approaches the carnivorous type. The only THE RELATION OF DIET TO DISEASE 451 arthritic cases occurred in Boatbilled herons {Cancroma cochlearia), fish-eaters. Our records show examples in Accipitres, Galli and Columbae, although the number of cases in the last order were few and slight in extent. Tliis disease stands in close relation to diet, as it develops on generous protein food, high in nucleoprotein or hypo- xanthin, especially if this be associated with restricted activity. The carnivorous mammals lead in the disease of the thyroid glands. Thyroid disease occurs among the birds, but is equally distributed among the dietary groups. Thyroid activity has a marked influence on metabolism probably through the influence of the iodine-containing protein of its secretion. There are some experimental evidences in favor of a detoxicating function of the thy- roid, of which the following are quoted : (1) The effects of thyroidectomy are most marked in the carnivores; Herbivora are often capable of several years of life with- out thyroid tissue; (2) administration of meat to thyroidectomized omnivores or herbivores caused a marked increase in all symptoms. The importance of the relation of the meat diet, detoxication and thyroid disease receives considerable confirmation from the fact that among the 1,860 mammalian postmortems thyroid disease occurred in 2.6 per cent, of all mammals, 94.9 per cent, of which were found in flesh-eating varieties.] Wells (32) suggested that possibly this could be inter- preted as an indication that toxic materials found in the meat in the intestinal tract were, under normal conditions, detoxicated by the thyroid. Against a local neutraliza- tion, however, is the improvement following the administration of dried thyroid substance. The function is either neutralization of toxic substances or the stimu- lating action on intracellular metabolism, both of which might be called into play by an excessive protein diet. (32) Chemical Pathology, Philadelphia,, 1918. 452 DISEASE IN WILD MAMMALS AND BIRDS The Carnivorous Diet. The pathology of the more prominent diseases developed in carnivores points at least to diet as a pre- disposing or determining factor. This diet is very high in a distinctly putrefactive protein and yields products, chemical and bacterial, which are toxic and which give rise to acute or more often chronic diseases of the ali- mentary tract and its adnexa. By reason of the amount ingested, excessive because of lack of exercise, there is a severe tax on the storage organs and on the detoxica- ting glands, as the liver and thyroid. The constant absorption of these toxic substances gives rise to chronic degenerative or fibrotic changes in the organs through which they pass : liver, kidneys, arteries, heart. In birds the degenerative diseases are even more marked than in mammals on the same diet. The ultimate fault of this diet, especially for mammals and birds with restricted activity, lies in the production of toxic bodies, produced either in the incomplete degradation or oxidation of the protein molecule or as the result of bacterial action on the protein molecule, a poisonous quality which is probably enhanced by the chemical changes occurring while the digested protein is passing through the intestinal mucosa. Garden conditions are such that these factors are almost un surmountable unless the substitution of vegetable pro- tein could be accomplished. Failure is often caused by limited feeding to carnivores of muscle and bones, whereas they should be supplied with glandular organs and blood. The Herbivorous Diet. Herbivorous diet must be divided into two groups, (1) that composed of succulent vegetables, and (2) of grasses, grains and seeds. In the first group there is an apparent variation in the results found in mammals and birds. In both there is a marked decrease in the chronic degenerative pathology. In both, acute gastritis is more THE RELATION OF DIET TO DISEASE 453 prominent, far outstripping the incidence of this condi- tion in other classes. This diet yields a large and quickly available amount of carbohydrate which in conjunction with the moisture, heat and bacteria which are unavoidably associated mth raw vegetables, makes an ideal situation for infection. These foods carry many saprophytic bacteria, moulds, etc. In birds the conditions are aggravated by the injuries that may occur from the sharp objects picked up with the gravel. The incidence of acute infection is higher among birds than among mammals of this group, and often there is involvement of the whole tract. The explanation of the other pathological findings occurring among birds must be found in the frequently repeated low grade infections which result finally in the production of chronic lesions in the digestive tract, liver, pancreas and kidney. Toxins as an etiological factor cannot be altogether excluded, but as a rule they are not important because the by-products of vegetables are distinctly less toxic than those derived from animal sources. Arteriosclerosis is much less fre- quent and less intensive in herbivorous birds than among the caiTiivorous, probably because of differences in the concentration and character of toxins in the two groups. Soft Herbivorous Diet. The diet of succulent vegetables is composed of tubers, edible roots and leaves. The tubers and edible roots are high in water and carbohydrate and poor in the amount and quality of the protein, most of which is not even a true protein but a mixture of aminoacids. The leaves, on the contrary, are rich in organic ash, especially calcium, sodium, chlorine, and fat soluble A vitamine, and as a rule contain a good quality of protein. They often, however, contain injurious substances. This diet, wliile measurably less nutritious than that of the carnivores, can satisfac- torily nourish many animals with an extensive intestinal tract during growth and even throughout their entire life. 454 DISEASE IN WILD MAMMALS AND BIRDS but proves entirely inadequate when fed to an omniv- orous tract. Seed Diet. Closely allied in general character to the diet of suc- culent vegetables are the seed diets, eaten only by birds and having no parallel among mammalian foods. All seeds, in contradistinction to tubers, contain true proteins which, however, are of poor quality because of the defi- ciencies in the aminoacid content. They are as a rule low in the fat vitamines and in the amount of calcium, sodium and chlorine carried. In three pathological conditions only do these birds show any oversusceptibility : (1) Sore eyes, (2) acute enteritis, (3) osteomalacia. Sore eyes were frequently noted in this group. The lesions were very like those described in animals deprived of the fat vitamine, which was present in this food in very small amounts or entirely absent, thus giving a very plausible explanation of this condition, especially as in some of the cases no other cause could be found. Gastric disease of any type is rare in this group because the food at the gastric stage is highly resistant to bacterial action. In the duodenum, however, the conditions are early changed because the bacteria carried with the food through the stomach become active in the presence of available carbo- hydrate and protein decomposition products. Osteomalacia is confined almost as exclusively to the seed-eating birds as it was to the omnivorous mammals, and it is also associated with the same deficiencies, cal- cium and phosphorus (cf. Tables 19 and 20). It is also interesting to note that these two diets, the omnivorous and seeds, yield the greatest number of cases of tubercu- losis. Mammals showed 32.6 per cent., as against 5.8 per cent, in all the other dietary groups, an observation which becomes more striking when man is added to the omnivo- rous group. Seed-eating birds showed 17.2 per cent., as against 6.4 per cent, in other groups. In both diets the THE RELATION OF DIET TO DISEASE 455 fat, fat-vitamine and inorganic salts, especially the calcium, are deficient in amount. In the wild, birds vary their diet of seeds with insects, worms, soft fruits and the tender shoots of plants, and at the same time they increase their inorganic intake by the minerals picked up with the gravel and from the water which has penetrated the soil. ) Geain and Grass Diet. The hay-eating animals constitute a large and well studied group — indluding practically all the domestic varieties. Table 19 shows that these animals yield the greatest number of cases of malnutrition, food poisoning, acute pancreatitis, acute degenerative conditions of liver and myocardium. Recent literature describes many cases of osteo- malacia, especially among horses and cows, in the famine districts of Europe. In our collection of 1,860 post- mortems only one case was found, that of an Isabelline gazelle {Gazella isahella) , a hay-eating animal, and in this case it was secondary to infection. Arthritis, occurring in 3.4 per cent, of all the autop- sies, was almost entirely confined to the hay-eating animals. The literature describes many cases of arthri- tis almost entirely confined to ungulates, of which many were associated with calving and subsequent infec- tion. Bacteriological researches have found it most often associated Avith streptococci, staphylococci, or Bact. per- fringens, organisms that require a certain amount of carbohydrate for their proper development. The relation of diet to this condition probably lies only in the fact that it provides an excessive carbohydrate substrate suitable for the optimum development of these organisms. Folin and Bergland, noting glycoresis in Herbivora, thought that it represented the absorption and excretion of unusable carbohydrate, present in grains, vegetables, fruits, etc., and that it was sharply separated from the main carbo- hydrate metabolism. These products were absorbed from 456 DISEASE IN WILD MAMMALS AND BIRDS the blood exactly as they were ingested like lactose, dextrose, etc., are absorbed, but do not enter into the economy although they might cause disorders, especially forms of arthritis. The grain foods are composed largely of carbo- hydrates (principally in the form of cellulose and starch) small amounts of protein and little or no fat. They have a very low nutritive index so that large amounts must be consumed to supply adequate calories. This food is constantly present, and during the enforced idleness of capti\4ty is almost continuouly eaten. Despite these facts, however, malnutrition is present in 2.2 per cent, of the animals on this food. Associated with the plentiful food and lack of exercise are overeating and pica. Overfilled stomachs occurred thirty-four times. They were limited to these mammals and to the seed-eating birds whose environmental conditions are practically the same. Pica or excessive appetite for abnormal food, is also more fre- quent in these groups, but is usually associated with badly balanced diets, and thus represents an effort on the part of the animal to supply its own deficiency. It is present in osteomalacic monkeys and has been reported in cattle from regions where osteomalacia is common and following crop failures where the rations are restricted. In cattle it very often accompanies food poisoning, espe- cially that produced by ingestion of peat hay. Disturbance of the alimentary tract and its adnexa occurs in two forms: (1) Infection which is quite com- mon and involves the duodenum, pancreas and liver, and (2) toxic. Compared with other diets alimentary dis- orders are not frequent among grain feeders, despite the ease with which grass foods ferment and the great variety of organisms found in them such as moulds (aspergillus), Bact. coli, paratyphosus, enteritidis, suipestifer, oidium lactis, etc. Few bacteria can attack whole protein, cellulose or starch, and the decompo- sition products, peptone, glucose, etc., are not available THE RELATION OF DIET TO DISEASE 457 in any quantity until the lower stomacli and duodenum are reached. The inflammation of the alimentary tracts of these animals is confined to the fourth stomach and duodenum, with, in many cases, extension to liver and pancreas. Acute and chronic degenerative changes occur very frequently, and as a rule are the result primarily of absorbed toxins. After ingestion of new hay this often appears. The toxic substance probably is a terpinol ester, cumarin, which is produced by an enzyme in the cut grass. The result is a gastroenteritis with jaundice, thirst and marked flatulence. It is very probable that many of the gastrointesinal and degenerative lesions are the result of the combined action of toxin and bacteria. Food Poisoning. ;^ Food poisoning occurs in all diets, but especially among the grass-eating mammals. To-day under the general heading of food poisoning are included those cases due to (1) some injurious substance inherent in the food itself, true food poisoning, (2) those due to toxic substances liberated or produced in food contaminated by parasites or bacteria, (3) those due to bacteria that are carried by food and develop into true infection after ingestion. Most of the cases of meat poisoning described in literature undoubtedly belong to this third class, i.e., flesh is infected during the life of the animal or during its preparation for food and the virus develops in the host after ingestion. A fourth and more rare class of food poisoning is due to the condition of the individual consuming the food — ^protein sensitization. Injurious constituents of normal flesh foods are very micommon. There are a few poisonous fish, notably the balloon, puffer, and Fuga fish of Japan, which when eaten give rise to cholera-like conditions ending in con- vulsions and paralysis. A marked intoxication has been described in dogs which have fed upon the Greenland ■' 30 458 DISEASE IN WILD MAMMALS AND BIRDS shark. Some fish are poisonous at certain periods as spawaiing season, the poison then being confined to the roe. Still others are harmless miless rendered toxic by some injurious food. This poisoning of muscle meats is seen in quail and partridges fed on mountain laurel, in some fish after consuming certain marine plants, and in cattle poisoned by amanita. The most common sources of poisoning are spoiled meat and flesh of diseased animals, both of which are serious factors in the production of the gastrointestinal disorders of omnivores and carnivores. Practically all the reports of meat poisoning from the literature have been traced to the use of raw or insuflficiently cooked flesh, and have yielded on bacteriologic examination Bad. paratypliosus, Bad. enteritidis, Bad. suipestifer, Bad. coll, or Bad. proteus. The bacteria may produce toxin in the food previous to ingestion causing in the host only a severe intoxication. This is the situation developed after eating sturgeon infected with Bad. piscidus agilis, an organism which manufactures a highly poisonous alkaloid. A similar intoxication follows the ingestion of potatoes infected with Bad. proteus or containing the poisonous alkaloid, solanin, which is produced in diseased and sprouting potatoes. Other examples of this are (1) ergotism — due to an infection of rye and wild grasses with Claviceps purpurea which produces three poisonous bodies, ergo- tinic acid, which is not poisonous when taken into the stomach, sphacetinic acid and cornutin which act on the nervous system, brain, cord, vagus and vasomotor centre giving rise to toxic polyneuritis, and (2) favus, an acute febrile anemia with jaundice and hemoglobinuria prob- ably due to a bacterial infection or fungus growth of the bean. Infected food may also produce soluble heat- resisting toxins that produce immediate symptoms and increase the animal's susceptibility to infection. This is the more common finding in cases of poisoning with milk THE RELATION OF DIET TO DISEASE 459 and milk products. Non-pathogenic saprophytes carried in milk produce (1) a poison closely allied to tyrotoxicon, (2) a toxalbumin which in itself causes serious disturb- ances. Botulism, also probably of this group, is a disease initiated by a toxin elaborated by Bad. hotulinus acting on a protein. There is, however, some evidence that Bad. hotulinus can also establish a real infection. The toxemias from food infected with bacteria may not occur until the food is ingested or the bacteria implanted. This result occurs in infections with Bad. bovis morhif- icans, Gartner's bacillus, etc., or after the feeding of meat from animals infected with Bad. paratyphosus and enteritidis. The plant poisons are more frequently due to inherent injurious substances, although even among them, bac- terial and fungus diseases play an important role. Among the 16,673 plants indigenous to North America, almost 500 are more or less poisonous and about 30 are of great economic importance. The toxic factor may be confined to the leaf, seed or root, but more often it is associated with all parts of the plant. Through the efforts of the Department of Agriculture a more or less complete list of the plants implicated in the poisoning of stock has been compiled. This list includes the following : Amanita muscaria; A. phalloides; Veratrum viride; Phytolacca decandra; Agrostemma githago; Delphinium, 25 varie- ties; Astragalus mollissimus; Aragallus lamhertii; Crotalaria sagittalis; Euphorbia lathyris; E. marginata; Rhus radicans; R. diver siloba; R. vernix; Aesculus pavia; A. hippocastanum; A. glabra; A. Calif ornica; Cicuta maculata; C. vagans; Conium maculatum ; Kalmia lati- folia; K. augustifolia; Leucothoe catesbaei; Rhododen- dron maximum; Pieris mariana; Datura stramonium; Solanum nigrum; S. dulcamara; Helenium autumnale; Asclepias pumila; A. verticullata; A. galoides; A. mexi- cana; A. eriocarpa; A. speciosa; A. fremonti; Eupa- torium agertoides; E. urticar folium; Isocoma wrightii; 460 DISEASE IN WILD MAMMALS AND BIRDS Dauhentonia longifolia; Senecio jacohia hurchelli latifo- Zw5.(33) Some of these as the Amanita are only occasional sources of disaster, but as they frequently involve man they are important. The Amanita muscaria symptoms appear very soon after eating the fungus and consist of a deepening stupor. A. phalloides, on the contrary, starts with severe abdominal pain, cramps, discharges of blood and mucus and later convulsions. The meat of animals dying from fungus poisoning is distinctly poisonous. This transfer of poison to the muscles of the animal par- taking of these plants occurs also in poisoning with Kalmia. The other plants of this list are closely associated with the grass foods and are consumed usually when the food on a range is scarce. Some groups as the Asclepias contain a distinct neurotoxin and give rise to a condition known as trembles or staggers. It affects mostly cows and sheep, causing staggering, trembling gait, bloating and salivation and death with convulsions. There is marked congestion of alimentary tract, liver and kidney. In the cerebrospinal axis there are marked changes in the nerve cells of the medulla and spinal cord. The Purkinje cells show the effect of extreme fatigue. Other plants causing stiffness or weakness of the extremities, show on microscopic examination no definite lesions in the cerebrospinal axis. Loco weed — Astragalus mollissimus and Aragallus lamhertii — causes maniacal disturbances but no gross lesions. This weed in Colorado costs the state enormous amounts of money yearly. Helenium poisons domestic animals by means of a toxic glucoside, dugaldin, which produces stiffness, sali- vation and nausea mth mild depression ('' spemng sickness ") . The alimentary tract shows severe inflamma- tion of the rumen and reticulum which may at times be hemorrhagic. The Hver usually presents an interstitial (33) These botanical names are taken from Chestnut's Poisonous Plants of America. THE RELATION OF DIET TO DISEASE 461 hepatitis. This toxin is decidedly hemolytic. The effects of this plant are always permanent, total recovery being very rare. The larkspur (25 different varieties), on the contrary, shows prompt recovery after treatment, but no establish- ment of toleration. These plants give rise to nausea, vomiting and great agitation and destroy many animals yearly. The poisons are included in four alkaloids, all spinal cord depressants resembling aconite in general character. These poisonous plants all produce more or less gas- trointestinal inflammation and practically all are destructive in their action on the liver, pancreas and kidney. It is impossible to form even approximate esti- mates of the damage done by them because of the general ignorance of the subject. The Division of Botany has been collecting for the past few years specific information concerning these plants, but the individual plants are not equally poisonous, and all animals do not show the same susceptibility to the poison. Veratrum viride, for instance, is eaten with relish by sheep and elk and is decidedly toxic for the horse. In many the toxic factor has not been isolated. Some, as Euphorbia, are poisonous only when fed in honey derived from its flowers. The influence of diet on the general health of animals is very far reaching and very inclusive. Metabolic dis- turbances are undoubtedly at times the result of unbalance — deficiencies on the one side, excesses on the other, at times are probably much more the results of bacterial invasions aided and abetted by the food admin- istered, at still other times are poisonous either in their own content or from the degradation products resulting from digestion or bacterial decomposition. SECTION XVI NEOPLASMS The occurrence of true neoplasms in domesticated animals has always been well known and thoroughly studied while for beasts in the wild the data has been fragmentary. That tumors exist in natural environment has been accepted upon the testimony of hunters but there is an impression, and nothing more, of their extreme scarcity probably because only younger vig- orous animals come to the attention of the sportsman or collector. This matter will of course not be settled until some natural historian with a knowledge of pathology, makes a survey of a large number of speci- mens taken during a collecting expedition. Observations in menageries are valuable to the extent that they show what tumors may occur, the orders most commonly affected and the incidence under captive conditions. It is unfortunate that too seldom do we know the history of our specimens in regard to the age, manner of capture or breeding, data which if at our disposal would permit of a very fair idea of the probable incidence in the wild and of the effect of captivity. Some observations in this direction are however possible by using the figures of known captivity and breeding. The facts gleaned from a study of neoplasms under captive conditions may be of interest to the experimental pathologist, especially when considering the relation of the origin from the embryological layers. I have tabu- lated this with great care, using Jordan's (1) table for the source of the various tissues, and further have studied the destination of metastatic emboli in terms of the blastoderm. The following observations are based entirely upon our own data for while it might be valuable to include the cases in the literature their descriptions are often so (1) Textbook of Histology, 1920. 462 NEOPLASMS 463 meagre that they would not combine readily with our records. Plimmer, Seligmann, and Murray have pub- lished in the Proceedings of the London Zoological Society since 1903, their annual report of the pathological service in which they have recorded very many interest- ing tumors. So too from time to time Harlow Brooks and W. R. Blair in the Annual Report of the New York Zoological Park, have presented cases occurring in their ser\dce. Joest (2) discusses tumors in the lower animals in a broad way and analyzes their incidence and characters. C. Y. White and I (3) have already published articles on this subject. Numerous single references may be found in the Jahresher. der Veterindr-Medicin. In so far as the incidence of tumors in wild animals is concerned this literature can scarcely give an adequate measurement but it would seem that they are less than in domestic varieties. Exact figures for the occurrence of tumors in the latter seem not available in the literature, but one can find that in the Prussian army horses about one hundred are obser\^ed each year. In our 5,365 speci- mens collected during nineteen years, 94 tumors in 92 animals have been found, 1.7 per cent, or about one in every sixty specimens, not at all a low figure. If one were to include all fibromata of the feet and the blood collec- tions to which the name angioma might be applied, this incidence would be greater; they are excluded because few in number and vag-ue in history; only one true angioma was seen. The gross and microscopical appearances of tumors in the lower animals are essentially the same as one encounters in human beings or at least it is possible to apply the pathological nomenclature used in human medicine to all the neoplasms we have discovered. There is appended a list of all the animals and their tumors, a table of zoological orders, tumors and organs (Table 21) and an analytical table of the histological data. (Table 22) (2) Zeitch. fiir Krebsforsch, Vol. 15, p. 1. (3) Proceed. Phila. Path. Soc, 1910, and Journal of Pathology and Bacteriology, Vol. XVII, 1912. 464 DISEASE IN WILD MAMMALS AND BIRDS c a a2 nMonjpifj 1 IBuajpv 1 ^ -H ^ sa;sax | IRS 1 SBajouBj 1 ^^r-, XlBAQ 1 BOIUIBIM 1 ^ (N ansBix qdmXi | r-> mnano^uaj | •oy 'jaAt^ 1 ^ C^ - ■joBJx icnnsa^nioJc^SBo | ^ aacn^iBO puB anog | -H -^ AaupiH 1 ^ enjajxi | C^ P!o-i^qx apgnj^ I aun^ ejouinx paxip^ BUiOjqclanjad;{jj I BoioiiaqiidanouoqQ I ^«0 I CO.-H (N Buioaapv '"' (M COr-^ Bcaoiiaq^ida BcaoiiidBj BUIOOJBg I Binoiiaq^opua ■Binoianv I HO I BcaoXuiojqij puB buioXjv | Biuodiq I Biuoa^eo I Buioaqii[ NEOPLASMS 465 mI w ?s w r? S3 3 •s I i o rr Oi wir-t (N »0-H 1 2 -^ -H 1 ^ (M 1 '^ -H 1 ^ 1 cc -* --H 1 ^ -^ 1 '* r-> 1- ^ 1 "^ ,-<^ CO 1 "^ ^ ^ ^ ^ -^ ^ ^ rHrH •« ^ ^^ C^J^ »c ^ ^ 1 "* ^ -H ^ ^-H ^ « -< '-< .c (N -H CO ^^ ■<*< -H did ■■^ altiP«l|la, . -? '^ o ^ -g^ =3. =3 ■3^ li-i T3 C of CO r- 466 DISEASE IN WILD MAMMALS AND BIRDS - 1| 03 3[39nj^ anittsa^n; saponqdm/tq XaupiH ^°g ^ ^ t>. CO -^ oot^ '-^ ^ c^ Q Q £ 00 >> r. E° >.t; CO oc^ ^22 00 § S2 »3 IN CD -1 •^ 1 (N -H |co r^ . (N «0 ^ Tt< 05 lapjo lad -»a30 jaj ■^coco 05 ®} 8[BaitnB iB^ox iN^r>|C^oo rHOt-^t-O 00 -o^o a 6 c« « S3 . ■■■^a ee3'6fe:^T)c3c3.2g O M)3 u fl „ ^ ... „ > 3 £ >> c-c ^ o I^.Ip^ NEOPLASMS 467 I- ^ 1^ (N I IN |co CO I t^ ■^1 ^22 M CO '-' t^ (N^^ -^ -^ •<* s (N ^^ CO 2 r- (M ^ -^ t^ g ^s?§ 00 (N eo ^ t-- (N-H^C '^ OOOOOOOO^ -° ^ g S 9J • s a -O 03 ._ >i^ . „ tp QJ O g< : 468 DISEASE IN WILD MAMMALS AND BIRDS Incidence of Tumors. Examination of the table, (21) from the standpoint of differential percentage reveals that mammals have 48 tumors giving an incidence of 2.58 per cent, whereas birds have 44 newgrowths equivalent to 1.23 per cent. Were it not for the high figures for one single variety of bird (Undulated Grass Parrakeet) this value for Aves would be still lower. At all events our figures would indicate that the mammal is at least tmce as productive of neo- plasms as is the bird. In our material the latter class has had a better chance than Mammalia to show its susceptibility since there have been nearly t-\vice as many autopsies. Within the classes the comparative figures have less value because of the smaller and varying numbers. Such high percentages as are shown by the elephants and armadillos cannot be taken as indicators for their orders since too few specimens were examined. Judging by orders with more than one hundred autopsies the rodents stand at the head of the list followed by the marsupials and carnivores. It is interesting that the animal nearest to man, the monkey, and with greatest number of autop- sies in its zoological class has the lowest tumor incidence. Psittaci lead the avian orders, followed by the Fulicariae, but as there are but thirty-five autopsies upon these, the second place rightly belongs to the Anseres. All the principal orders are represented but the only one of importance is the leader. The Psittaci are very prone to have tumors in the renal area, sometimes of the kidney, at others of the adrenal and occasionally of the sex glands. Some remarks have already been given to this matter in the sections devoted to the Iddney and genitalia but it mil be discussed again under tumor morphology. Among these ninety-two animals, one bore multiple tumors, a Jaguar {Felis onca) with adenomata of the liver and uterus and angiomata of the mesentery. Care- NEOPLASMS 469 ful study failed to reveal any parasitism as the cause of the growths and since the first two were of slightly vary- ing structure it is not believed that one is a metastasis from the other. The sex incidence stands in direct relation to the pro- portion of total males and females posted or in other words it is the same for the two. The figures might be somewhat affected were the gender of all the parrakeets available but the tumors growing in the upper renal area frequently destroy the sex gland. Definite statements concerning the importance of breeding in the causation of neoplasms cannot be made since we cannot quote figures for the percentages of wild- and captive-born of our entire autopsy list. The data are confused by scanty information concerning the twenty-six parrots, the history of which is vague and I am perhaps too severe in accrediting the birth of sixteen of them to capti\T.ty. Tliis was done because of a lack of exact information concerning these specimens and, be- cause their variety is lmo\\Ti to breed when captive by the residents of their habitat(4), the distribution into wild- and captive-born is based upon what information we have. If the order Psittaci be subtracted entirely, it leaves a total of 62 tumor-bearing animals of kno^vn breeding, 49 of which were born in the wild, thirteen in captivity, a fact which strengthens the thought that unnatural breed- ing increases the chance of neoplasms. The kno^vn length of capti^ity has also a direct bearing on this point. The figures' given in the columns " known captivity " and ^' average for tumor bearers " were compared with figures obtained by averaging the lives of fifty others (when possible) of the same order or of at least three times as many as bore tumors. Animals dying from injury were excluded. With one exception the average for '^ tumor bearers " exceeded that for "non-tumor bearers"; the exception, the Ungulata, had (4) See Gould's Birds, Vol. II, p. 83. ((' 470 DISEASE IN WILD MAMMALS AND BIRDS the same average for both groups. It seems then that tumors occur in animals in captivity longer than the average for their order, or in specimens that have the power to live under confined conditions until neoplasms develop. In this respect I recall the state- ments made by Harlow Brooks (5) that tumors will prob- ably be found more commonly in animals when they live in a manner comparable to that of urban man and that racial degeneracy will favor their development. There is adduced here perhaps the first definite evidence that long captivity allows tumor tendency to express itself but it does not prove that confinement increases tumors. Nor does the expectation of life, average or potential, stand in any direct and definite relationship to the fre- quency of neoplasms. The only clear case of long life and high tumor incidence is to be found in Parrots ; we feel however that some miknown factor increases tumors of the renal-adrenal region in these birds and that unquali- fied statements about age and tumor growth are not per- missible. Since tumors grow in many wild-born specimens, a high percentage of which become known in the first few years of captivity, is it not highly probable that tumors are reasonably common in the wild and that we do not observe enough purely natural specimens to assume an immunity on the part of free li\dng beasts. One of the undesirable features of captive breeding is consanguinity of parents and there is good reason to believe that tumor susceptibility can be bred into or out of a line of animals by mating tumor bearers and non- I tumor bearers, the tendency following the rules of Mendelian inheritance (Slye). Is there any proof that inbreeding does not occur in the wild and if it do, it is perfectly possible that tumor tendency may be trans- mitted as a dominant character ; the effect of artificial or intentional inbreeding in captivity would only offer an opportunity for a summation of these influences. (5) Am. Jour. Med. Soc, 1907, 133-769. NEOPLASMS 471 If injury and animal parasitism have any importance in neoplasmata then this opportunity certainly occurs under natural conditions, Fibiger observed gastric tumors in rats arising under the influence of nematodes while Slye and Wells report facial neoplasms in mice apparently arising at points of old injuries. It seems to me that we have no right to assume an immunity of wild animals, in their native environment, to tumors; the incidence is another matter but it may be considerable. It was thought possible that there might be some light shed upon the matter by an analysis of our sarcomatous and epitheliomatous tumors in wild- and captive-bom animals. In our second paper (6) upon this subject I ventured the statement that sarcomatous growths occurred more frequently in captive-born, epithelioma- tous in wild-born specimens. Greater data have not borne out this conclusion and information was sought as to the embryonal derivation of tumor-bearing tissue. Analyz- ing the cases in which all the factors could be obtained, it seems that among seven tumors of captive-bred animals, five came from the entoderm, two from the mesoderm, whereas in wild-bred animals, of the fifty-seven tumors, five came from the ectoderm, thirty-two from the mesoderm and fourteen from the entoderm. These figures do not include the parrots. The sex values have no significance. It is interesting and noteworthy, that, as in the human being, the majority of the tumors came from tissues aris- ing in the mesoderm and that the entodermic derivatives received the largest number of metastases ; no ectodermic tissues were sites of secondary tumors. The visceral seats of metastases are probably of little value for com- parison in so small a number ; the lung and liver however occupy the prominent places. Interesting as the foregoing facts may be, they do not shed light upon the question of breeding and degeneracy (6) Jour. Path, and Bact., Vol. XVII, 1912. 472 DISEASE IN WILD MAMMALS AND BIRDS in the causation of neoplasms. Attention is arrested how- ever by the paucity of tumors in derivatives of the ecto- derm since in man new growths are common in the breast, at the rectal and labial mucocutaneous jmictions and on the skin. The immunity of the ectodermic tissues to secondary growths is very distinct ; this holds true in man. Special Tumoes. The diagnosis of fibroma offers the same difficulty in the zoological material as it does in man and even more care must be exercised for solid tumors in certain locali- ties. The bird often presents hard nodular masses on the palmar and lateral aspects of the feet, sometimes sur- mounted by callosities, to which the term fibroma or fibro- matous corns might be applied. Section of some of these will reveal areas of granulation tissue about points of inflammation so that we have considered them as infec- tious or the result of incorrect perches and excluded them from the tumors. True fibromata have been encountered thrice but in combination with muscle tissue as a fibro- myoma thrice in addition. The '' fibroids " seen in the elephants and armadillo have already been described. The nodular growth sometimes accompanying degen- erative disease of the osseous system followed by attempts at repair as discussed under osteitis deformans, leontiasis ossium and actinomycosis, are often produc- tive of masses to wliich it is easy to apply the term osteoma. If one demand that an osteoma shall be a dis- tinct neoplastic, localized bony growth of unnatural or greatly exaggerated structure, then the tumor is quite rare. We have seen one growing from the vertebrae and clavicle of a gerbille and a fibro-osteoma on one jaw of an Isabelline gazelle. The chondromata have been limited to one case, a unilateral mass growing from the nasal cartilage of a caracal. ( Lipomata are localized collections of fat consisting of cells with greater fat capacity than normally, sometimes NEOPLASMS 473 surrounded by an indefinite capsule. Judging by the A observations of Joest and Johne they are reasonably \ common in horses and cows. We have not seen a single ' case in mammals but eight cases appeared in the birds. These were with one exception disposed under the skin mostly over the abdomen and chest and once under the scalp. In a hawk the tumor grew as a pehdc mass sur- rounding the cloaca and apparently caused decided obstruction to the lumen. The lipomata of the Psittaci usually grow as pendulous masses on the abdominal wall covered by thin, featherless, delicate skin, often show- ing dilated veins. Upon section they are rather rich in blood supply, " angiolipoma," but fail to show any angiomatous or solid cellular areas under the micro- scope. The frequency of the growths in one variety (Roseate cockatoo — Cacatua roseicapilla) led to an attempt to transplant the tumor. The plant seemed to thrive in the recipient for a while but soon disappeared, j Breeding experiments on the tumor-bearers are now under way. Angiomata of lymph channels were observed in the omentum and mesentery of a jaguar (Felis onca) ; this is the animal with three apparently separate and dis- tinct tumors. " The omentum is normally fatty and slightly congested. In its meshes are myriads of tiny cysts containing gray fluid. The main peritoneal area is negative but in the pelvic region on anterior rectal wall, in the superior edge of the broad ligament and in Douglas' pouch, are cysts from a few millimetres to several centimetres, with clear contents. The microscopic section of omentum shows the multiple cysts as cavities of varying size, from that of an arteriole to the diameter of a two-third lens field. They are lined with flat, closely placed pavement cells with well stained but vesicular nucleus. The septa are adult connective tissue. No con- tents or gTanular eosin-staining material. No swollen cells like in adenomata. No parasites seen." 31 474 DISEASE IN WILD MAMMALS AND BIRDS Two endotheliomata have been found, one of the flat variety with warty excrescences common on serous sur- faces, located in the pleura of a leopard {Felis nebulosa), and one of the nodular variety, growing from the clavicle of a Moorhen {Gallinula chloropiis). The sarcomata present their usual morphology grossly and minutely and with the exception of the cases arising from the pectoral muscle and from the genital area offer little of interest. Two instances in the former location, observed in parrakeets, presented several puz- zling features. The component cells were spindle in shape, similar to a muscle cell but were fitted with the round or elliptical nuclei of embryonal cells. In a few places they were exceedingly large and had shadowy outlines like a syncytium or they would be so arranged as to suggest a glandular structure. The dominant type of cell was, how- ever, everywhere the spindle as it is seen in sarcoma. The sarcomata when they occur in the genital area usually assume the alveolar arrangement and are of the round or mixed cell variety. Only three of the sixteen sarcomata gave metastases. Papillomata of minor character appear occasionally on the skin of animals as warts, but only one instance of any greater importance has been found. The duodenal mucosa of an owl {Bubo virginianus) presented a soft growth which partly obstructed the intestinal lumen. Papillary adenomata, on the other hand, have been observed several times, but since they have more importance as irregular hyperplasias of glandular origin have been included in the next group. An interesting case was seen in a baboon {Papio hamadryas) in which a large part of the gastric wall was the seat of adenomata, presenting in addition several distinct papillary out- growths. ^ A similar picture was found in the duodenum of the rhea {Rhea am eric ana). ) The greatest interest in the adenomata centres around these growths in the renal area in parrakeets, and as they NEOPLASMS 475 have much in common with all the glandular tumors of this region, a general discussion of this subject may be introduced here. We have observed seven tumors con- structed on a glandular basis of renal or adrenal character. Grossly these tumors develop as irregular masses usually of distinct brown color, constructed on a lobular plan, delicate barely visible septa dividing the growth. They seem devoid of large vessels, a gross observation confirmed microscopically. There is no cri- terion to the naked eye, which will distinguish the variety of epithehal hyperplasia or permit separation of these neoplasms from some sarcomata; the latter are usually gray but need not be so. Minutely studied, three of these tumors proved to be adenomata, all papillary, one cystic as well. Three had to be denominated carcinoma because of their distinct separate crowded nests and incomplete acini. The cells comprising these growths are comparable to the lining elements of the collecting tubules of the renal lobule in that they have relatively large nuclei and a tendency to basic staining protoplasm. The adenomatous picture is, however, more comparable to the cortex than to the medulla. The remaining tumor was a hypernephroma of the usual large cell, acinus-forming type and seemed to originate in the adrenal. None of these tumors in the parrakeets sent out metastases. Other hj^Dernephromata have been diagnosed, to the number of six. Upon review of their descriptions and sections, the determinations are to be confirmed. However, it must be recorded here that none of the three in mammals gave metastases, while two of the three in birds did so. They are all of the usual type with large vacuolated cells in glandular groups or strands. Three rather interesting examples of epithelioma have been observed. The first and most important was a baso- cellular growth of the tongue in a black bear (Ursus americanus). The local damage — ulceration and infiltra- tion — and swelling sufficient to interfere with deglutition, were quite considerable. The basal cell nests had pene- 476 DISEASE IN WILD MAMMALS AND BIRDS trated deeply into the muscle, but extension had taken place only to a single submaxillary gland. A squamous epithelioma was found on the skin of the thigh of a Tas- manian devil {Sarcophilus wsiniis). The construction was somewhat unusual in that it was cystic but lined with squamous and keratinized plates. It could not be decided that it originated from glands like a trichoepithelioma; it was not like a basal cell cancer. No metastases had occurred. The third case was that of a tumor mthin the abdomen of an Amazon {Chrysotis leucocephala). It con- sisted of an illy defined basement membrane upon which were irregular stratified squamous epithelial cells. Upon the surface were wavy bands of horny material, very much like dried and cast-off epithelial scales, except more compact and extensive. These latter seemed to form the bulk of the mass. Beneath the membrane a few irregular accumulations of cells bearing a similarity to those on the surface could be found, but they were prob- ably large plasma cells. *The epithelial layer dipped down like in epithelioma. No pearls or separate nests were found. Wliile this mass was not localized, it was doubt- less an epithelioma, and should be included in this series. Its possible origin in the small intestine has been considered. The question of the occurrence of tumors in ^\dld animals seems fairly well settled when twenty-five examples of malignant epithelial neoplasms can be dis- covered in fifty-three hundred autopsies. It is interesting to note the incidence of these tumors in wild- and park- bred animals. Exclusive of the parrakeets there are twenty-one cancers, seventeen in known wild-bred, two in known park-bred specimens, and two with breeding un- certain. The average known duration of captivity of the wild-bred animals is about four years, while the two park- bred animals lived eight and eighteen years. Thirteen of the twenty-one cases were males, eight females. Adeno- carcinoma was discovered twelve times, simplex nine Fir,. 49. — BASAL-CELL CARCINOMA OF TONGUE. BLACK BEAR (URSUS AMERICANUS). NOTE ULCERATION WHERE PIECE HAS BEEN EXCISED. AND ALSO NODULAR THICKENING OF WHOLE BASE OF TONGUE. Fig. 50. — MICROSCOPICAL APPKARANCE OF TIMOR IN Fin. 49. NEOPLASMS 477 times, medullary and squamous each twice. Three tumors of the pancreas and mammary gland were seen in which fibrotic or scirrhus areas were found, but in no case was there detected that hard cicatrizing cancer so conomonly found in the human breast. All the interesting cases of carcinoma have been recorded in the discussion of organs from which they took origin. The only case of chorion- epithelioma has been reported in detail on page 308. The two cases of mixed tumors are as follows : Mixed tumor of the thyroid and adenocarcinoma sarcomatodes in the liver; they have been discussed in detail on pages 334 and 242 respectively. Analysis of the incidence of tumors according to organs is disturbed by the large number of cases in Psittaci. Including this order the first place is taken by the kidney, followed by the liver, uterus, muscle, gastro- intestinal tract, bone and cartilage, thyroid, adrenal and lung in this order. Curiously enough, if these birds be subtracted the degree of organ susceptibility to new growths is not greatly altered. The lead is still held by the kidney, the uterus occupying the second place and then in sequence the liver, gastrointestinal tract, muscle, thyroid and adrenal. Examination of the figures for mammals shows the uterus to lead in numbers, followed by the liver, thyroid, and mammary gland. For the birds the kidney takes the undisputed head of the column with a total of twelve tumors (27 per cent, of all avian tumors) ; the next figures are shown by the liver, gastrointestinal tract and muscle. ZOOLOGICAL AND PATHOLOGICAL LIST OF TUMORS MAMMALIA Primates (2) Cercopithecidse — Hamadryas Baboon {Papio hamadryas) Papillary adenoma of gastric mucosa Cebidae — Brown Cebus {Cebus fatuellus) Hypernephroma of right adrenal 478 DISEASE IN WILD MAMMALS AND BIRDS Lemures (1) Lemuridae — Ring-tailed Lemur (Lemur catta) Papillary adenoma of prostate Carnivora (17) FelidaB — Clouded Leopard {Felis nebulosa) Endothelioma of pleura Caracal [Felts caracal) Osteochondroma of nose Lion (Felis leo) Malignant adenoma of cervix uteri Metastases to lung Jaguar (Felis onca) Fibroadenoma of uterus Fibroadenoma of bile ducts Lymphangioma of mesentery ViverridsB — Indian Paradoxure (Paradoxiirus niger) Adenocarcinoma of pancreas Malayan Civet (Viverra tangalunga) Carcinoma of lung Canidae — Corsae Fox (Canis corsac) Adenoma of pancreatic ducts Red Fox (Canis vulpes pennsylvanicus) Cystic adenoma of bile ducts Raccoon like Dog (Canis procyonoides) Adenocarcinoma sarcomatodes of thyroid Gray Fox (Canis cinereo-argenteus) Papillary cyst adenoma of bile ducts Prairie Wolf (Canis latrans) Sarcoma of thyroid region Metastases to lungs Prairie Wolf (Canis latrans) Sarcoma of thyroid region Procyonidae — Common Raccoon (Procyon lotor) Adenoma of pancreas UrsidaD — Polar Bear (Ursus maritimus) Adenocarcinoma of adrenals Metastases to lungs, lymph nodes, diaphragm Black Bear (Ursus americanus) Medullary carcinoma of breast Metastases to lungs Black Bear (Ursus americanus) Epithelioma of tongue Phocidae — California Hair Seal (Zalophus calif or nianus) Hypernephroma of adrenal NEOPLASMS 479 RODENTIA (12) Sciuridae — Beeehy's Spermophile {Citellus grammurus beecheyi) Osteoma of sternum Gray Squirrel {Sciurus carolinensis pennsylvanicus) Hypernephroma of kidney Woodchuck {Arctomys monax) Adenoma simplex of liver Muridae — Waltzing Mouse [Mus wagneri rotans) Adenocarcinoma of thigh muscles White footed Mouse {Peromyscus leucopus) Carcinoma simplex of mammary gland White footed Mouse {Peromyscus leucopus) Spindle celled sarcoma of leg White footed Mouse (Peromyscus leucopus) Carcinoma of mammary gland Larger Egyptian Gerbille (Gerbillus pyramidum) Fibrosarcoma of shoulder region Heteromyidae — Kangaroo Rat {Perodipus richardsoni) Sarcoma of urinary bladder Octodontidae — Coypu Rat {Myocastor coy pus) Sarcoma of thyroid Hystricidje — ^Canada Porcupine {Erethizon dorsatus) Chorionepithelioma uteri DasyproctidaB — Azara's Agouti {Dasyprocta azara) Squamous carcinoma of larynx Proboscidea (1) Indian Elephant (Elephas indicus) Leiomyoma, uterine cornua and fimbria Ungulata (7) Equidas — Chapman's Zebra (Equus burchelli chapmani) Fibroma peritonei with sarcomatous and osseous change and metastases to lung Bovidae — Isabelline Gazelle {Gazella Isabella) Osteofibroma of jaw with mucoid degeneration Nylghaie {Boselaphus tragocamelus) Fibroma uteri Dorcas Goat {Capra hircus) Lymphosarcoma of mediastinum with metastases to liver, kidney and lymph nodes CervidaB — Common Deer {Manama virginiana) Fibroadenoma of bile ducts Camelidae — Alpaca (Lama pacos) Carcinoma of liver or bile ducts with extension to intestine SuidaB — Wild Boar (Sus scrofa) Carcinoma uteri 480 DISEASE IN WILD MAMMALS AND BIRDS Edentata (1) Dasypodidae — Nine-banded Armadillo (Tatu novemcinctus) Fibroma uteri Marsupialia (7) Didelphyidae — Common Opossum {Didelphys virginiana) Adenoma of kidney Common Opossum [Didelphys virginiana) Adenocarcinoma of mammary gland Dasyuridae — Spotted tailed Dasyure (Dasyiiriis macidatus) Adenocarcinoma of intestines with metastases to lymphatics, liver, spleen, lungs Tasmanian Devil {Sarcophilus ursinus) Cystic epithelioma of skin of thigh Peramelidae — Rabbit eared Bandicoot {Thylacomys lagotis) Carcinoma of lung Macropodidae — Red Kangaroo (Macropus rufus) Malignant papilloma of stomach Metastases to liver, spleen, kidney Red Kangaroo {Macropus rufus) Carcinoma of lung Mestastases to spleen and gastric wall AVES Passeres (7) Turdidae — American Robin {Planesticus migratorius) Hypernephroma of kidney, metastases to intestine Crateropodidse — Jungle Babbler (Crateropus canorus) Adenoma of kidney Tanagridae — ^Palm Tanager (Tanagra palmarum) Lipoma of abdominal wall Fringillidae — Saffron Finch {Sycalis flaveola) Adenocarcinoma of kidney Chestnut eared Finch (Amadina castanotis) Adenocarcinoma of kidney with metastases to lung Chestnut headed Bunting (Emberiza luteola) Lipoma of scalp leteridae — European Blackbird {Meriila merula) Hypernephroma of kidney region with metastases to liver Striges (1) Bubonidae — Great Horned Owl (Bubo virginianus) Papilloma of duodenum Psittaci (26) Loriidae — Musky Lorrikeet (Glossopsittacus concinnus) Carcinoma of lung NEOPLASMS 481 CacatuidaB — Roseate Cockatoo {Cacatua roseicapilla) Lipoma of abdominal wall Roseate Cockatoo {Cacatus roseicapilla) Multiple lipomata of abdominal wall Psittacidae — Undulated Grass Parrakett {Melopsittacus undulatus) Glioma of brain with metastases to liver Undulated Grass Parrakeet (Melopsittacus undulatus) Hypernephroma of adrenal Undulated Grass Pan-akeet {Melopsittacus undulatus) Papillary adenoma of kidney Undulated Grass Pai-rakeet {Melopsittacus undulatus) Cystic papillary adenocarcinoma of kidney Undulated Grass Parrakeet (Melopsittacus undulatus) Adenocarcinoma sarcomatodes of liver Undulated Grass Parrakeet (Melopsittacus undulatus) Papillai-y cyst adenoma of kidney Undulated Grass Parrakeet (Melopsittacus undulatus) Adenoma of kidney Undulated Grass Parrakeet (Melopsittacus undulatus) Adenoma of kidney Undulated Grass Parrakeet (Melopsittacus undulatus) Adenocarcinoma of oviduct Undulated Grass Parrakeet (Melopsittacus undulatus) Carcinoma simplex of liver with metastases to liver, spleen Undulated Grass Parrakeet (Melopsittacus undulatus) Carcinoma simplex of liver Undulated Grass Parrakeet (Melopsittacus undulatus) Multiple lipomata Undulated Grass Parrakeet (Melopsittacus undulatus) Multiple lipomata Undulated Grass Parrakeet (Melopsittacus undulatus) Sarcoma of pectoral muscle with metastases to liver Undulated Grass Parrakeet (Melopsittacus undulatus) Round cell sarcoma in region of liver, spleen, kidney Undulated Grass Parrakeet (Melopsittacus undulatus) Carcinoma simi)lex of thyroid Blue fronted Amazon (Chrysotis cestiva) Adenocarcinoma (?) of pro ventricle White fronted Amazon (Chrysotis leucocephala) Epithelioma in peritoneum ( ?) All Green Parrakeet (Brotogerys tirica) Sarcoma of pectoral muscle Red shouldered Parrakeet (Palceornis eupatrius) Sarcoma of testes Red shouldered Parrakeet (Palceornis eupatrius) Sarcoma of testes 482 DISEASE IN WILD MAMMALS AND BIRDS King Parrakeet {Apromictus cyanopygius) Sarcoma of ovary Crested Ground Parrakeet {Calopsitta novcehollandi-ii KMCvKM-^r^—ci-HrtC^ -^ I 05'-HeO'-H-«iCiCfO'-i^OO(M^iOOO ec . ■* »c t^ 00 ic o co^ ^5: — ^CCirJc^OO-^TjJlOciM^OOTjJdN^df^Cr^lOoici ^ —I CO— I r^cOOilO^Oi) c^ ■3 K III c 8 o M<00 -d CI 0H()\VS THE BOVINE PEARL DISE.iVSE ON THE COSTAL PLEURA, AND SERVES AS WELL TO ILLlS'IR-VIE THE NODULAR TUBERCLES COMMONLY FOUND IN THE SPLEEN OF ALL PRIMATE TUBERCULOSIS. THE COMMUNICABLE DISEASES 497 probably primary as it is not known that the retropharyngeal glands drain to or from the larynx. About the lesions on the laryngeal mucosa there is an area of congestion probably due to the tuberculin injection. Local lymphatic tuberculosis of comparative interest was encountered a few times. Three cases of cervical adenitis, large enough to be visible, were seen, of which one broke down about two weeks before death, and dis- charged. The others did not ulcerate through the skin but, contrary to the usual rule for the human being, remained as isolated glands only lightly adherent to one another where they lay adjacent. There was also seen an ulcerating tuberculous lymph node in the groin of one monkey, the animal ha\dng rather pronounced abdominal and pelvic tuberculosis. Two instances of tonsillar tuber- culosis are recorded, in both of which the lesion was of some duration and associated with caseation in the lymph node lying immediately behind and below it. A Guinea Baboon {Papio sphinx) had as an unusual part of his gen- eral tuberculosis, an active caseopurulent collection in the antrum of Highmore, which attacked the upper maxilla and immediately adjacent muscle ; tubercle bacilli could be demonstrated. One of the cases of nodular or massive peritoneal tuberculosis is quite like the tumor-forming variety of human adolescents ; it is as follows : Reddish Macaque (Macacus rufescens). Caseous tuberculosis of mesentery and spleen; miliary tuberculosis of lungs, pleura and liver; chronic myocarditis. The lymphatic glands of the posterior and supe- rior mediastinum and bronchi are slightly enlarged, soft and anthracotic but do not show any tuberculous change. Both lungs are riddled with small, firm, gray miliary tubercles, some surrounded by a clear mantle of connective tissue. The intervening lung is practically normal. Pleura over base of right lung on both surfaces shows small, pale miliary tubercles. The liver contains various sized miliary tubercles. There is a large caseous mass in posterior end of spleen with adhesions to kidney, stomach and colon. Retroperitoneal glands are much enlarged, firm, homogeneous — probably tuberculosis of a different type. Lesser omentum contains one caseous gland. Few caseous glands in great omentum. In the right iliac region there is a large mass involving many coils of intestine. It is found to arise probably from the ileocecal 498 DISEASE IN WILD MAMMALS AND BIRDS glands and can be traced along the mesentery to the central lymphatic stalk. The mass involved the tissues of the mesentery and surrounds many coils of intestine. Cecum and first part of colon can be traced over its right side. Rectum is free except on right side where it is lightly attached to the mass. Epicardium is grray and irregularly thickened and the muscle just beneath serous membrane is pale and streaked with red lines. There have been three cases of tuberculosis of the internal male genital area, one of which was suspected of having been the primary seat of the disease ; it was de- scribed on page 315. The other two could have been sec- ondary since other points of morbid change were as old or older. One of these cases formed a tumor as large as a goose egg at the vesical neck, obstructing the flow of urine and blocking up the seminal vesical, in consequence of which paralytic distention occurred in the bladder while the vesicles were tightly filled with inspissated semen. A case of Fallopian salpingitis has also been cited. Two instances of cerebral tuberculomata have already been described. Camivora. This order has the reputation of being quite resistant to the tubercle bacillus, based upon the relative infrequency among cats and dogs in contrast to cows and swine. Some veterinary statistics cite the incidence up to 5 per cent., and occasional references may be found to tuberculosis in circus lions and tigers. Our records would suggest that in gardens the wild varieties of this order have about as much of the infection as the domestic carnivores, 3.5 per cent. The group is made of six FelidaB, one Viverridae, three Canidae, six Procyonidae, and one Ursidae. The first family includes a lion, tiger, a jaguar and three smaller cats. The Canidae are all small foxes. The Procyonidas are all coatis. The features of this order are the occurrence of the fibro- ulcerative variety with cavitation in the Felidae and the caseous nodular abdominal and glandular disease in the coatis. All these animals, even those of the last named variety and pathological type, tend to show some tissue THE COMMUNICABLE DISEASES 499 resistance to the tuberculous disease. Connective tissue activity is characteristic of the process, considerable dis- tortion being produced by the fibrosis. This feature is borne out where the tissues are studied microscopically. Definite milia are sometimes found, but they consist of epithelioid and round cells with imperfect caseation, giant cells being often missing. About the miliary tubercles a diffuse and not essentially specific tuberculous granulation tissue is found, mixed with which is much connective tissue growth. The fibrotic adhesion-form- ing serous membrane tuberculosis of carnivores seems worthy of emphasis by the citing of a case in point. White-nosed coati {Nasua narica), was received in poor condition and died in a few days. Upon dissection a slightly turbid yellowish fluid was found to occupy what remained of the peritoneal cavity which was reduced in size by dense adhesions of the intestines into an inflam- matory mass. The omentum was a diffuse thickened apron, also beset with fine tubercles, lying over the mass. Fine young tubercles could also be found upon the intestines and liver while the mesenteric lymph nodes were early in caseation ; thoracic organs not infected. One of the most interesting cases concerned hyper- trophic osteitis in a chronically tuberculous lion ; the feet are discussed on page 346. This process was described by Marie for human beings many years ago, and was reported in dogs by Cadiot(l) in 1912. This beast was one of five large cats which have died from tuberculosis in its chronic ulcerative form. Three of the cats, one fox and the bear showed definite cavitations of a ragged locu- lated form. The cavities were usually of the multiple variety and were found in the posterior, that is lower lobe. Rodentia. The paucity of cases in this order permits little information to be drawn from the form of tubercu- losis. The total seems to have been swollen by a group of three beavers, all of which came in one shipment. The (1) Rev. de Med. Vet. T. 89, p. 221. 500 DISEASE IN WILD MAMMALS AND BIRDS remainder were a Capybara and an Agouti. The general type is that of much caseation with little or no sur- rounding fibrosis. One illustrative case is cited : American Beaver {Castor canadensis). General tuberculosis. The animal presents generalized tuberculosis. The regional lymph nodes show caseous nodules. The right hip joint shows caseous material about the acetabulum with necrosis and pathological fractures in the OS innominatum immediately above the acetabulum and including its cavity. Lungs show almost no normal respiratory tissue, the proc- ess being a diffuse precaseous, partly gelatinous pneumonic phthisis. The superior and posterior mediastina show caseous glands. There is miliary tuberculosis of the liver. Nodular caseous tuberculosis of the spleen with small tubercles and some cirrhosis of the intervening tissue. There are caseous nodules in all perirenal glands and in the kidney cortices. The psoas muscle glands are densely caseous. The pelvic organs except about the right acetabulum escape involvement. Adre- nals not opened but probably not involved. Ungulata. This order shows the most definite figures among those for the order of mammals. Nearly one- tenth of the whole number of specimens have had some form of tuberculosis and of a very definite character. It is well at first to mention, however, that only four of thirteen families are represented, from which four came 328 of the total 365 autopsies. The remaining thirty- seven were such animals as tapirs, giraffes, swine, and peccaries, in all of which tuberculosis has been reported from elsewhere. Pulmonary disease with less prominent lesions in other organs, especially the intestines and their related glands, speaks in favor of the aerogenic route being the common one. This of course has been a bone of conten- tion among veterinarians, and I do not presume to settle the matter with these figures. This order resists tuberculosis to a certain degree as attested by the fibroulcerative character of the majority of the cases. Two instances, one in a buffalo and one in a deer, showed very highly fibrotic puhnonary lesions with a partial attempt to surround and wall off numerous areas of caseation. So too in this order there is a greater THE COMMUNICABLE DISEASES 501 tendency to calcification, both in the intra- and extra- pulmonary nodes. It is to be emphasized that in our material the thoracic lymph nodes are affected more than the abdominal and regional as 3 to 1. The apparent immunity of the spleen of this order is well illustrated. The paucity of serous surface involvements in the wild BovidaB and their prominence in the Cervidae cannot be ignored in the figures, but it seems misleading since pleural growths and adhesions are quite common in the domestic Bovidae. The case in the Equidae was that of a Zebra with a large tuberculous abscess in the retro- peritoneal glands forming a tumor in the left renal region. It was quite well surrounded by fibrosis, and the infection had not extended ; it seemed quite recent. Analysis of the figures for the remaining three families of ungulates offers little for contrast and much for comparison; it is the usual picture as seen in the domestic cow. Some special cases are worthy of review. An interesting specimen of softened glands chiefly on one side of the neck was found in a Fallow deer {Cervus dama). It resembled the juvenile human cases that require surgical attention. Although palpable lymph nodes can be found in practically all cases of generalized tuberculosis in the Ungulata, this is the only case in our records in which they have presented a large tumefaction and broken down. Pulmonary cavitation is recorded but thrice, one for each of the last three families. Fibro- caseous tuberculosis of the testes was discovered in a Nylghaie {Boselaphus tragocamelus), but there is no knowledge of mating or offspring. An ischiorectal abscess was found in an American Bison {Bison bison) showing nodular precaseous tubercles of the lung. The former was the cause of death. No tubercle bacilli could be found in the abscess contents, so that the tuberculous basis is inferred, not proven. Tuberculous salpingitis in a Nylghaie was discussed on page 306. 502 DISEASE IN WILD MAMMALS AND BIRDS Proboscidea. Eber mentions in the article already referred to that there are three reports in the literature of tuberculosis in elephants. When looking for an explana- tion of tuberculosis in this animal it must be remembered that it is one of the most attractive objects in a zoological garden and receives perhaps more attention, including feeding, from visitors than any other specimen. The beast while possessing some tissue resistance to tuberculosis, is by no means immune thereto, as has been thought by some persons on account of its reputed longevity, and therefore he is to be protected from infection just as much as other animals. It would appear that he may present caseous pneumonia or nodular caseous dissemi- nated lesions. Our two cases, in animals at the Garden twenty and thirty-eight years respectively, were both of the fibrocaseous variety; the lesion was confined to the lungs. A brief description of their lesions is as follows : Indian Elephant {Elephas indicus) 6 . Chronic polyarthritis. Chronic myocarditis. Chronic hepatitis (cirrhosis). Parenchymatous nephritis. Chronic tuberculosis of the lungs, partly encapsulated. Pig- mentation of the spleen. The pleurae are very fat but the surfaces are smooth and devoid of adhesions. The lymph nodes of the mediastinum are about 10 x 20 cm. for the largest while the smaller ones vary around 2x4 cm. They are firm, deep red-brown Avithout clear divisions into medulla and follicular cortex. There are several large, firm, pale rather cheesy follicles in all the large ones and a few of the small. These do not appear like tuberculosis. The lungs are flaccid and soft; gray and red mottled. The bronchi are firm and stand open. Around one in the upper lobe of the right lung, there is a large area of cheesy degeneration around which a zone of connective tissue has formed. This extends about the bronchus about halfway in a sheath-like man- ner. There is also a separate nodule the size of a cherry with a cheesy centre. The trachea appears normal. Tubercle bacilli could be demon- strated in the cheesy material. Microscopic section of lung around the cheesy area shows a low-grade chronic granulation tissue in some places enclosing cheesy masses with giant cells on the margin. The neighbor- ing septa are slightly thickened and in some places broken, fonning emphysematous cavities. Some of these cavities are edematous. Indian Elephant {Elephas indicus) 9. Miliary and conglomerate caseous tuberculosis of lung. Edema of lungs. Endarteritis deformans of lung. Cloudy swelling of liver. Chronic passive congestion of liver. THE COMMUNICABLE DISEASES 503 Hemosiderin pigmentation of liver. Acute parenchymatous nephritis. Chronic passive congestion of spleen. Chronic hyaline perisplenitis. Multiple calcified fibroid tumors of uterus. Leiomyoma of uterine cornu. Senile atrophy of ovaries. Acute catarrhal enteritis. There are some adhesions of the upper lobes of the lung to the ribs. The lungs are large, increased in weight, color pink and mottled red, air content diminished. There are several masses of tubercles, each as large as a cocoanut, in both lobes. In one such the tubercles are yellow and caseous ; some are fibroid but none are liquefied or calcified. The fibrous tissue of the lung parenchyma here is much overgrown. In one instance the terminus of a bronchus is solidly plugged by caseous material. Mucosa of bronchi is reddened, markedly ulcerated, ulcers overlaid by mucopus. Individual Features of Avian Tuberculosis. The avian form of tuberculosis is somewhat peculiar iu its physical appearance as well as in its distribution.) The isolated nodular type is far and away more common by more than 100 per cent, than all the other types com- bined. These nodules are usually well circumscribed, and to the naked eye suggest that they have a restraining fibroid wall. This is, however, not the case, the impres- sion being due to the dense but actively growing fibro- cellular cortical zone of the tubercle. The centre of the nodule, instead of having the soft character like Camem- bert cheese, resembles the firm but brittle American dairy cheese. Upon opening such an area the central necrotic mass may split away from its cortex and even shell out, lea\ang a cavity lined by a graj^-yellow membrane. These characters are best displayed in nodules of moderate size, the small ones being like the yellow mammalian analogue, the large being like indefinite cheesy masses. In the sur- rounding tissue evidences of inflammatory processes seem decidedly greater in our material than I am accus- tomed to see in human and veterinary pathology. This, it seems, should be emphasized since secondary infection with pus cocci and other pathogenic germs appears less often in birds than in mammals. The difference speaks, therefore for a difference either in the tubercle bacillus of birds or the avian physi- 504 DISEASE IN WILD MAMMALS AND BIRDS ology. Judging by the limited morbid processes produced by injecting avian bacilli into rabbits and guinea-pigs the reaction of the bird itself would not seem wholly responsible for the difference. The local tissue reaction in all avian lesions is mononuclear and fibrous, softening and pus being rare. It would seem from this and similar operations that the bird expresses its resistance to the bacteria by a fibrocellular reaction which goes on to fibrosis without softening; perhaps this means also that their polynuclears are not sufficiently active, but the pathogenic power of the bacillus itself doubtless is individualistic. The character of the cheesy degeneration is likemse different from the mammalian. It seems like an abrupt hyaline necrosis of a large central mass and not the slower cell death seen in the other types of tubercle. At times the degenerated area, instead of having the yellowish color of caseation, will present what we have designated ''gelatinous tuberculosis," the whole infiltrated area resembling boiled sago or tapioca. This seems to be a complete homogeneous coagulation or hyaline necrosis of the whole mass out to the delicate fibrous mantle sup- plied by the tissue in which the tubercle lies. The organic distribution of tuberculous lesions has already received some attention and is to be discussed with the orders. There are, however, some localities affected conspicuously in the bird. The skin lesions often attract attention during life. They occur around the eye, at wing joints, on the cresta sterni and on the legs. Parrots and jays have shown nodular or diffuse growths around the eye, originating both in the lids and orbit, which on section have proved to be tuberculous. These seldom ulcerate, but those upon the skin of the breast and wings tend to have superficial erosions or deep ulcers. The latter lesions are more common upon pigeons but have been seen in Psittaci and Galli. Toucans and pigeons when -MASSIVE TUBERCULOSIS IN LIVER AND SEVERAL MURAL TUBERCLES OF INTESTINE COMMON PEA FOWL (PAVO CRISTATA). Fig. 53.— nodular HEPATIC LESIONS IN THE LIVER OF A DOVK. THE COMMUNICABLE DISEASES 505 pinioned, have on three occasions shown a tuberculous mass on the stump. Two parrots with hyperkeratosis of the beak and of the skin of the feet, have also had tuberculosis. These have been mentioned in literature as of tuberculous origin. One case well studied failed to show tubercle bacilli in the corns. In the absence of tubercle bacilli, one is inclined to think that this might be explained on the basis of a circulating toxin such as is assumed to be responsible for hypertrophic periosteitis. This latter condition has not been seen in birds. Still another type of occasional occurrence deserves mention. While most of the lesions in birds correspond to the description given in the preceding pages, some lesions fail to degenerate in the centre, retaining instead a solid homogeneous fleshy character of dull gray- yellow color. Upon section these have been found whoUy cellular in construction. To distinguish them from the ordinary nodules they have been designated tuberculo- mata. Lesions of this kind may occur along the lymphatic paths, indeed seem more common in the lateral cervical and thoracic chain, and upon bones and nerves. When they are numerous the nodular caseous type is incon- spicuous. , They suggest the bovine infection (Pearl dis- ease), but one attempt to prove this failed. We are of the opinion that this is the avian lymphatic form, as our examples correspond to the literary descriptions of cervical tuberculous lymphatics in birds. No especial variety of bird is more often affected by this process. Intestinal tuberculosis among the Aves may be said to assume three forms. The best known, indeed the form usually spoken of as representing the common picture, is that which produces varying sized nodules upon the serosa, sometimes associated with adhesions to neigh- boring intestines. Just how this type develops is not known. In some quarters it is believed to originate by the penetration of the tuberculous granulation tissue 33 506 DISEASE IN WILD MAMMALS AND BIRDS from the mucosa through the intestinal wall by following lymphatic channels and that irregular contractions of the musculature squeeze the exudate outward under the serosa. Other observers think that the bacilli are car- ried; -1;^ the lymphatics to the superitoneal tissue, there starting the tubercle. The truth of the matter will probably be that both methods are operative although we have seen more cases suggestive of the second than of the first explanation. When these peritoneal nodules are numerous and prominent, mucosal ulcers are uncommon and vice versa. The second form is the ulcerative, flat ragged or crateriform defect situated in a diffusely thickened wall. This was well illustrated in cases of pulmonary infection in doves and guans, suggesting reinfection of the gut tract from swallowed tubercle bacilli or a backward development of the disease after the lungs were nearly solid. The third form of tuberculous enteritis is quite inter- esting and striking. It is best seen in the duodenal loop but may occur anywhere. Diffuse thickening of the enteric wall is noted, and when palpation is practiced a resilient but leathery sensation is obtained. Careful inspection reveals the mucous surface to be velvety, a con- dition due to a swelling, that is widening, of the villi which retain their erect position and, when washed in flomng water, will be seen to move like a field of grain in a breeze. The serosa may be, usually is, negative. Studied microscopicaUy the peculiarity of this form is in the development of tubercles and diffuse cellular exudation in the villus stalk, sometimes extending into the submucosa also. Round cell masses like lymph follicles are some- times prominent. This form is not associated with any peculiar organic distribution so far as my studies go. Passeres. The peculiarity of this order seems to be in the predominance of the pulmonary route as origin of tuberculosis. Perhaps in no other order has there been Fig. 54. — TUBERCULOUS MASSES OF INTESTINAL WALL SHOWING OUTWARD GROWTH. SOMETIMES THESE MASSES OBSTRUCT THE LUMEN. THE COMMUNICABLE DISEASES 507 sucli extensive and advanced lesions as in these little birds. Sometimes one whole lung will be solid while its fellow will be half occupied by caseous material. The doves alone seem to approximate the Passeres in ability to live with so much tuberculous exudate. Picarise. Specimens from this order illustrate well the intestinal origin and distribution of tuberculosis. There was, among these birds, one case sho\ving tuberculomata which was, because of its gross anatomy, listed as the pearl type. Its description is as follows : Lesson's Motmot (Momotus lessoni). The region above and behind the right clavicle in front of the brachial plexus on the internal sur- face of the thorax, exterior to the first and second ribs, and on the internal surface of the ribs at the junctions of ribs with the alae of the sternum, there are many small, irregular, smooth, firm, yellowish Avhite nodules varj-ing in shape from spherical to sweet potato and in size from 3x3 mm. to 3 x 7 mm. These are found quite homogeneous on cross section. They do not resemble tubercle or mould infection but make one think of neuromata. There are also a few present in the left lateral air sacs, close to but not joining the intestine. The lungs are apparently normal. Histological section of the masses de- scribed as distributed along the nei-ves consist of sharply outlined but not well encapsulated masses made up of irregularly disposed bunches of large cells with vesicular nuclei in a stroma of loose connective tis- sue very inconspicuous in amount. There is also quite a number of small round cells and a few leucocytes. The large cells first described have the nucleus eccentric for the most part. Many of them have two nuclei and a few three and occasionally a giant cell is observed. Blood vessels have a very delicate wall and are frequently encountered in the centre of these masses. Atypical mitoses can be found. Here and there a seal ring placement of the nucleus can be found. A few eosinophiles are present not definitely placed. Necroses, with large quantities of nuclear fragments, are scattered irregularly through the mass. The diagnosis rests between an infectious granuloma, false neuroma and sarcoma. Tubercle bacilli were found by stain in great numbers both within and without the cells. Psittaci. Tuberculosis occurs in this order somewhat more frequently in the varieties whose habitat is the Eastern world, although South American birds also suffer from it in the characteristic manner. There seems to be no difference in the pathology of these two groups. Par- rots present very beautifully the separate solid or semi- 508 DISEASE IN WILD MAMMALS AND BIRDS solid nodules of avian tuberculosis, whether they be in the lungs, liver, or spleen. "VYlien the lung becomes rid- dled with masses, coalescence occurs and the whole mass turns into a cast of the hemithorax. Lesions in the liver are mostly isolated, but the spleen often appears like one large pink tuberculous nodule. The liver occupies as usual the first place in organic incidence. Striges. Owls (and Struthiones — see below) present the interesting exception to the rule of intestinal origin of tuberculosis in birds. Perhaps the platting is incorrect but the birds in the order under discussion had older and much more advanced lesions in the lungs and thoracic serosa than they did in the abdominal organs. That this was true in all three examples is in itself noteworthy. Perhaps they possess less pulmonary and more intestinal resistance. One of these birds showed a small recent cavitation in the posteroinferior angle of one lung. Accipitres. With one exception the cases of this order occurred among the Falconidse, that is in hawks, buzzards, and eagles. Their lesions are usually gen- eralized as indicated by the figures for visceral distribu- tion, but that half the number should have the oldest, most prominent lesions in the lungs is curious. Their intestinal tuberculosis seems mostly of the diffuse infil- trative type.^ Columbae. These birds are obviously the most sus- ceptible of all the varieties of which there are sufficient autopsies to make a comparison. Generalized nodular lesions emanating from the intestinal tract comprise their usual form, while most of the hepatic lesions are small miliary and nodular; occasionally one sees caseous masses destroying large sections of the organ. Their intestinal lesions may assume any of the three forms described. Galli. This is an order of something over the average percentage incidence for the birds but containing families that seem very susceptible to tuberculosis.; The small THE COMMUNICABLE DISEASES 509 immber of Brush Turkeys (Catheturus lathami) had 60 per cent, of the disease, while South American Cracidae had 44 per cent. These two groups raise the incidence for the order. Galli as a group have generalized nodular tuberculosis originating by the intestinal route. This is especially seen in the Phasianidae, while the very sus- ceptible Cracidae have much more prominent lesions in the lungs, often of a massive caseous type. It is really astonishing at times how much of the pulmonary tissue is occupied by infiltrate before death has supervened. » Fuhcarias are represented by a special contingent of rails and gallinules. Avian characters are well illus- trated in the order. So too the succeeding order, Alector- ides, another variety of shore birds, run true to the avian form. It is interesting to note that in the two cases from each of these orders tuberculosis and aspergillosis have been combined. The former has assumed the firm nodular type, while the mycosis has been of the air sac variety. The following case is worth citing as possibly illustrating infection per cloacam. There is, however, no trace of this bird having been with a male with the disease. Demoiselle Crane {Anthropoides virgo) $. General tuberculosis including the oviduct. All organs are thickly beset by caseous tuber- culous nodules except the lungs which have only a few scattered ones. The oviduct is, for its lower two-thirds, much enlarged, firm, tough, pale yellow, thickly beset with caseous nodules; upper parts uninvolved. The kidneys are definitely enlarged, in-egular, almost mulberry-like, brownish yellow, firm and tough. On section the lobules are irregular, connective tissue increased, urates in pelves. Tubercles in intestines seem to be wholly peritoneal. This seems like a tuberculosis of genital origin judging from condensation of tubercles in the lower abdomen (mass around cloaca). The ovaries are not involved. Lungs and tho- racic air sacs relatively free. The pericardium shows a whitish thicken- ing of both layers due to the presence of whitish granules like urates. Histological section of kidney shows the capsule not greatly altered. Glomeruli largely negative but a few show hyaline capsular thickening of vacuoles in tufts or fibrosis in tufts or obliteration of whole structure. Tubules largely degenerated, distended or distorted. Interstitial tissue between the tubules definitely but irregularly increased. No real attempt 510 DISEASE IN WILD MAMMALS AND BIRDS at regeneration. Few vessels show perivascular fibrous change. One tubercle seen. Anseres. These birds present no especial features so far as percentage or organic incidence are concerned. The individuals are mostly geese and swans, ducks being somewhat more often affected by mycosis than by tubercu- losis. However, both these diseases tend to assume the nodular type in Anseres so that the diagnosis should be supported by bacteriological discovery of the respec- tive organisms. Struthiones. The marked feature of this order is the prominence of the isolated and confluent nodules in the lungs, of apparently greater age, certainly of great size, than similar lesions in the abdominal viscera. Case- ation of the avian variety is well illustrated in these birds. The thyroid body was involved in two of the three cases, the ovary in one. The representatives of the Crypturi, two tinamous, came at the same time and lived only a few months. Miliary tuberculosis of the small pre- caseous variety was the form exhibited by both specimens.*) Histology of the Tuberculous Lesions. The initial and characteristic unit of tuberculosis, the miliary tubercle, seems to be constnicted upon the same general principles in all cases of the disease and in all members of the zoological groups in our study and in a manner entirely comparable to that well known for man and for the domestic animals. There are, however, cer- tain minute differences which are interesting and may at some time become important. It is customary to speak of the bovine tubercle and of the human variety, but there are also slight variations of the microanatomy of each of these, while one may find on occasion a tubercle of the human type in a cow and vice versa. Not all the domestic animals show the bovine form, although in sheep and swine it is approximated very closely. In the horse there is much greater tendency to a central softening and Fig. 55. DIFFERENT VARIETIES OF THE MILIARY TUBERCLE. M :j^^' «• ^* '^'^ ^•-«. •'« *>« A 'i * * t %r'.^ 'O: ■i'J] •f*: ^%«i: V/fef/t -.i-*- '*?'.:•**?'. .•• •^ Kr*f, 'f i" * » ** J* ?* ■- - i-:o ,^^ fi'/» /.A: :<4 /:• ?^a: "' to*?:.*^ ^^-'^^tV :'/o»f-r • * .^^ H.— THK HUMAN TUBERCLE WITH CENTRAL COMPLETE NECROSLS, TYPICAL CJIANT CELLS. ABUN- UANT EPIIHELIOIl) CELLS AND THE RELATIVELY NARROW SMALL ROUND CELL MANTLE. Fig. 55. DIFFERENT VARIETIES OF THE MILIARY TUBERCLE. 'V.*,#*v ;^//f< •.:%:• \^y;''- ">•'-. ^.: '^ •:-. :^c'v- .•• C. — THE TUBERCLE FREQUENTLY FOUND IN MONKEY TUBERCULOSIS. WITH RAPIDLY ADVANCING CENTRAL NECROSIS ENCLOSING MUCH CHROMATIN DEBRIS, THE ABSENCE OF LANGHANS' GIANT CELLS. THE PRESENCE OF LARGE. PALELY STAINING EPITHELIOID CELLS OF LANGHANS' TYPE AND THE VERY SLIGHT CIRCUMFERENTIAL REACTION. Fig. 55. DIFFKRF.NT VARIKTIES OF THK .MILIARY TUBERCLE. D. — AN AVIAN TUBERCLE WITH CENTRAL SHARPLY MARGINATED NECROSIS CONTAINING MUCH CHROMATIN DEBRIS, THE IRREGULARLY ARRANGED POLYNUCLEAR CELLS TYPICAL OF AVIAN TUBERCLES. THE SMALL NUMBER OF REGULARLY ARRANGED EPITHELIOID CELLS. THE PAUCITY OF SMALL ROUND CELLS AND THE PRONOUNCED CONNECTIVE TISSUE MANTLE. THE COMMUNICABLE DISEASES 511 fibrosis is not so common as in the bovine tubercle. I have attempted to study the histological anatomy of each of the zoological orders, but it has not resulted in any profitable discovery. It is, however, possible to contrast the type commonly found in monkeys with that character- istic for man and the ungulates and also to emphasize the construction of the avian tubercle that it may be dis- tinguished from mammalian tuberculosis and from avian mycosis. The tubercle of the Primates is a loosely constructed affair lacking the fibrous mixture of the bovine and the close cellular packing of the human form. Studied from the periphery to the centre, there will be found very little fibrocellular reaction in the immediately surrounding organ, while the mantle of round cells, rather prominent in the human tubercle, is often quite inconspicuous. The principal cellular component of the miliary granuloma is the large pale endo- or epithelioid cell, which is abundant, loosely arranged and without apparent purpose. In the centre is an irregular necrosis usually retaining some chromatic matter, probably the remains of recently destroyed nuclei, but this caseous midpoint does not assume the dense acid staining common for many milia. Giant cells of the Langhan's or foreign body type are often entirely missing, and when present are scanty. There may be large cells, resembling the aforementioned epithelioid cells, with two or even three large palely stain- ing nuclei, but these latter are arranged irregularly and not like the spokes of a wheel near the cell wall. This picture suggests a rapidly growing inflammatory mass and indeed this is the type that tuberculosis follows in monkeys. In a few cases gross evidence of fibrosis in the serous surfaces and in the lungs has been observed, but they are too rare to permit one to think that con- nective tissue activity is an important part of the reaction of this beast to the disease. 512 DISEASE IN WILD MAMMALS AND BIRDS The avian tubercle as it rests in the tissue seems like a sharply outlined almost encapsulated body. This is in part due to the homogeneity of its structure and in part to the fibrocellular condensation around the caseous part of the growths. Examined from without inward, there is a round cell mantle, between the elements of which course fine but easily perceptible fibrils ; elastic tissue has been seen among them. The small cells continue more deeply than the fibres, to be succeeded in prominence by epithe- lioid cells of rather dense character, the nuclei especially seeming quite rich in chromatin and round. The rotundity of the nuclei remains not only in the single separate cells forming the middle zone of the cellular cortex, but can be found in the nuclei of the compound or giant cells which comprise the internal layer lying upon the necrotic centre. These giant cells are characteristic for the avian tubercle in assuming a form like syncytia with nuclei arranged in irregular radiating columns. This internal large cell area may completely surround the central necrosis or it may be interrupted by the large single cells. Tubercle bacilli are more common in and between single cells than multinuclear ones. Within the cellular zone lies the necrotic centre, often, indeed usually, full of chromatic debris. This centre is commonly quite amorphous but occasionally one will see what is probably the remains of a coarse coagulum. Between the necrosis and the cells one usually finds a split, a sort of separation of the gangrenous from the living part. Old tubercles with denser fibrous capsule retain this microanatomy in part, but the cellular zone gradually becomes thinner and thinner until all that remains is a narrow cortex of round cells and imperfectly retained multinuclear cells.) Tuberculous granulation tissue without definite milia consists entirely of the round cells with small vacuoles and a fine but definite fibrosis. Tuberculomata consist of cells of varying sizes with small round nuclei. Interstitial fibrosis is delicate and barely visible unless especially W^\;i^l\ K-? 55 —PHOTOGRAPH OF YOUNGEST AVIAN TUBERCLE AFTER COMPLETE FORMATION. Fig. 57. — TUBKRCULOMA. .\ SOLID IL'MOR-I.IKK MASS, CONSISTING OK CLOSELY PACKKI) LARf;E CELLS KITTED WITH RKLAIIVELV SMALL ROUND LOOSE NUCLEL THESE CELLS ARE CROWDED WITH BACILLL THE COMMUNICABLE DISEASES 513 sought. Giant cells may be encountered but are not so large as in milia. Necrosis occurs but not in an orderly manner in relation to cells as in an isolated tubercle. Tubercle bacilli are very numerous. Types of Bacillary Infection. According to experimental and statistical research, all the tubercle bacilli of the higher vertebrate classes can be infective for any member of these classes. Thus, for example, human bacilli have been found in many orders of mammalia and in birds. The bovine form has been found in swine. The lesson from this is that while the special predilection of a variety of the tubercle bacillus may be for one kind of animal, it is potentially a vinis for other kinds. Hygienic principles have therefore been laid down at the Garden which aim at the protection of all specimens from every variety of tubercle bacillus. For this reason and because the laboratory has not attempted extensive research on bacteriology, few type determina- tions have been made and those at hand oifer nothing new or unusual; they are noted here as a matter of record. Bovine bacilli have been judged by their slow growth and inf ectivity for rabbits, human bacilli by the reverse of these characters. Avian tubercle bacilli can be culti- vated with reasonable ease directly from lesions not bearing a mixed bacterial flora, and grow in a yellow, moist, even, spreading colonization. In our two attempts at infection of guinea-pigs, no success was had, although Rabinowitsch and others had no difficulty in so doing ; this strain may vary in virulence as do other tubercle bacilU. No avian culture was obtained from a mammal, but a bovine was found in a parrot and a human in a duck. Bovine bacilli were isolated once from a monkey (see page 496) and in another case of lymphatic type, bacilli of the short heavy blunt shape, supposed to be character- istic of this variety of the germ, could be stained. Monkey tuberculosis in our experience is usually due to the human 514 DISEASE IN WILD MAMMALS AND BIRDS tubercle bacillus, judging by the staining characters and two successful cultures. Discovery of Tuberculosis During Life. Fully developed chronic tuberculosis may be recog- nized with reasonable ease in the human being and some domestic animals. The diagnosis rests largely upon the history and symptoms and partly upon the appearance of the individual and upon signs elicited by physical exam- ination. There is good reason to believe that these latter methods are entirely applicable to certain wild animals, notably those that can be caught and held quiet, but because of their naturally great reserve many specimens offer little reason for suspicion as to their tuberculous condition until near death. Certain ungulates with chronic pulmonary disease get thin and weak but remain on their feet with good appetite and satisfactory dis- charges for many months. Primates, Camivora, Roden- tia and Aves not uncommonly come to autopsy with very good coats and without great emaciation and yet are heavily infected. It can be stated with fair positiveness that no chain of historical data or gross observations are certainly known to us as indicative of tuberculosis in the wild beast. Coughing is not necessarily characteristic of chronic pulmonary infection, although when continuous it rouses considerable suspicion, especially in the Ungulata. It is to be interpreted with care in all animals that have loose bedding as bits of straw or seeds get into the throat causing irritation; the dust of hay may cause coughing in horses. However much chronic or fatal tuberculosis may be interesting from the standpoint of pathology or of zoological or visceral incidence, the most important fac- tors in our knowledge of the disease are its early recogni- tion and treatment, either for curative or hygienic purposes. Since we have learned that advanced lesions may exist in an animal without materially affecting its THE COMMUNICABLE DISEASES 515 external appearance and behavior, it naturally follows that early cases, possibly of an ''open" or infectious character are still less likely to give evidence of their existence. This is well recognized by veterinarians as being true of cattle, but is perhaps less well known, or possibly admitted, by those who handle the very sus- ceptible monkey. Upon a visit to a foreign garden I was told that experience alone is suflBcient to enable an observer to detect tuberculosis, and that the disturbance entailed in physical examination and tuberculin tests is preju- dicial to the well being of all varieties, but especially the delicate ones. I learned later that they had the disease in their exhibition cages all the time but decided to put their method to the test. Shortly after my return from abroad a splendid specimen of Grivet Monkey {Cercopithecus sah(Bus) was condemned by the tuberculin test. He was well studied by the superintendent and two very experi- enced keepers, all of whom pronounced him one of the finest specimens they had ever seen, and stated tliat he was behaving quite normally. Despite their protests he was sacrificed, tuberculosis with early cavitation being found in the upper lobe of the left lung. Incidentally vague physical signs were found by auscultation, but as the monkey was unruly and had long pectoral hairs little weight was placed on the observation. However, it is frequently possible to make very thorough physical examination of the lungs of the more tractable specimens, diagnoses of pneumonia and bronchitis being frequently made in this and other parks, so that treatment may be instituted. Importance of Transmission and Known Susceptibility. Some light upon possible reasons for the poor condi- tion of an individual animal is of course shed by a knowledge of the disease to which that particular variety is most susceptible, to which may be added the data 516 DISEASE IN WILD MAMMALS AND BIRDS obtained from previous deaths in the same group or enclosure. Thus, for example, a sickly monkey would be suspected of having tuberculosis or early osteomalacia, whereas no suspicion of these diseases would fall upon the marsupials. The same position would be assumed if a dove and a heron were out of condition. In so far as enclosures are concerned, the matter is somewhat different. "WTienever a case of tuberculosis occurs in a cage, the remaining specimens if any are removed and the place cleaned by soap and water and disinfectant and paint. The naked flame from a blast lamp is used when possible. Out-of-doors enclosures are vacated, spread with lime and allowed to lie fallow for as long a time as practicable. These methods have been in the main successful in clearing a cage of the disease, and all our experience demonstrates the effect of cage hygiene and the selection of non-infective replacements. A few places such as those occupied by doves and guans have not been freed of infection, if one judge by its appearance when new specimens are placed in them, but they may of course be due to the infection from elsewhere. The his- tory of seven years in the new bird house where the hygienic conditions are excellent, seems to indicate that a cage thoroughly cleaned is no longer a source of danger, and that a repetition of tuberculosis in such an enclosure is due to its importation with new exhibits. The spread of the disease to nearby cages seems to depend upon two factors. If the number of cases has been large and the infection virulent, immediately adjoin- ing cages are involved, but the tendency to spread is directly proportional to the proximity of orders or families that have a high susceptibility for tuberculosis. This second factor seems to be the more important and is illustrated by our experience in one corner of the new bird house. In this area are exhibited certain doves and pheasants, among which are many cases, while the pas- serine varieties nearby are little affected. So too in the THE COMMUNICABLE DISEASES 517 flying cage the disease has occurred in varieties with high general susceptibility. There are at the present writing ten orders on exhibition in this large enclosure and there have been more. An occasional case of the disease occurs, but only in the orders which show it elsewhere. The Herodiones, of which we have had nearly one hundred autopsies and many now are on exhibition, are always well represented in this cage and yet show no tubercu- losis. In the ten orders mentioned above three show no cases of the disease. These observations illustrate the spread of tuberculosis, especially to the most susceptible varieties, and how non-susceptibles under good hygienic conditions fail to become infected even when infected animals are near them. The freedom of activity in the large enclosure is doubtless an important factor. The history of the past three years with regard to the control of tuberculosis in the small cages shows that twenty-nine were infected, but by the measures employed nineteen have remained free of the disease for one year ; three of the remaining ten are known to have received newly arrived and possibly infected specimens. The accredited method of transmission in birds, the swallowing of material soiled with the feces richly laden with germs, is the principal reason why infected en- closures and their immediate environment are the prin- cipal breeding places for tuberculosis. To be sure air currents may blow the virus around, allowing it to light upon food in other cages but this cannot be a great menace if for no other reason than that we have had no epizootic outbreak of the disease, when there were groups of deaths in doves and guans. Evidences with which to trace transmissions are much clearer in the birds than in the mammals with the excep- tion of monkeys and some ungulates. Of course cases are perhaps too few in the carnivores and rodents to permit correct deductions but it is very rare that more 518 DISEASE IN WILD MAMMALS AND BIRDS than one case occurs in the same enclosure containing groups of these varieties. Nor do animals in adjoining cages seem to '' catch " the infection. This observ^ation does not suggest that any relaxation of hygiene need be allowed but probably it implies that not many bacilli are excreted by these animals; they cough very rarely. Groups of ungulates (bison and deer) are often known to be infected but just how it has arisen is seldom clear. Transmission from monkey to monlvey has been observed so frequently that it cannot be doubted, nor mil anyone wonder at it if reflection is given to the close personal contact of these animals during their natural behavior. They huddle, pluck lice from one another, take food from the mouth of another, bite and perform many other actions greatly facilitating the transfer of any virus. Bacilli may also be disseminated by cougliing, drooling and with the fecal discharges, for which latter there seems ample opportunity since a notable percentage of cases have intestinal lesions.^ Monkeys do not seem to raise sputum and expectorate it but they do eject saliva from their lips. Contraction of the disease from infected cages is be- lieved to have occurred at least once in our experience but the lesson of complete sanitary cleaning of the enclosure learned from that happening, seems to have enabled us to forestall its repetition. Tjhe Tuberculin Test. Tuberculosis presents the greatest single problem among the specific infectious diseases which the director of a menagerie must attempt to solve. Even though one may possess a knowledge of its zoological distribution, clinical characters and pathological effects, these are in- sufficient criteria for its detection at a stage when the animal might be saved by treatment or, what is most important, removed from its companions that they might be protected. To this end there remains but a single THE COMMUNICABLE DISEASES 519 procedure for the discovery of the existence of tubercu- losis — the use of tuberculin in one of its forms by one of its methods of application. The use of this test in veterinary medicine needs no commentary, having made its place in clinical and hygienic practice for a quarter century or more. Armed with the knowledge of the satisfactory use of the toxins of tubercle bacillus in cows, Dr. Penrose, Dr. C. Y. White, Dr. A. E. Brown and Dr. Leonard Pearson began in 1901 a series of experi- ments with old tuberculin of Koch which have led to the development of a technique for its use in the detection of infected monkeys. These interesting and instructive animals, being known as highly susceptible since most of the collection died of the disease in those days, and being handled with reasonable ease by experienced men, were investigated as the most important specimens upon which to perfect the method. Other varieties have been studied since and I shall refer to them individually. The greatest amount of work and the most conspicuous suc- cess attended the observations upon monkeys and the results of this study are now in daily use in this Garden. The work, conclusions and results, originated by Doctor Penrose, Doctor White, and Doctor Brown can be described as one of the most completely satis- factory series of observations in scientific medicine. Applying the principle that a tuberculous animal reacts to the injection of tuberculin by a temperature rise, the normal temperature curve of the monkey was studied, that of the tuberculous monkey determined by killing many specimens. This enabled them to state which animal was infected, which was not and to place on exhibi- tion only healthy specimens. Added to tliis, strict hygi- enic principles in the housing and handling of the animals have resulted in the elimination of the disease from our exhibition house. Occasionally a case may develop, per- haps from feeding by visitors, but the matter is no longer a problem. I know of no more complete and satisfactory 520 DISEASE IN WILD MAMMALS AND BIRDS experiment and its practical application than this work, which is condensed in the succeeding paragraphs. (2) The Temperature of Monkeys. The success of the tuberculin test in the lower animals as in man depends chiefly upon the alterations in tempera- ture following the injection of the toxin. It is generally admitted to-day that a healthy animal's temperature will not be affected by the introduction of this material. There are in addition changes in the pulse and respiration rate and in the physical signs but these are detected with difficulty and are much less definite than thermometric records. The first essential was therefore a thorough familiarity with the normal temperature of the monkey, a requirement which met with considerable difficulty from the beginning since the earliest observations revealed puz- zling irregularities. This necessitated the establishment of certain regulations of teclinique which, after the pre- liminary tests, have been found satisfactory enough to continue until the present day. All monkeys are received in the quarantine rooms of the laboratory where they are observed by the officials of the Garden and of the labora- tory and there they remain in separate cages until passed, as free from tuberculosis, to the exhibition house. The handling of monkeys for the purpose of taking temperatures is a matter of no small importance since excitement will quite definitely increase the registration. We have been fortunate enough to have in charge of this work since its inception the same man, Keeper McCrosson, who is thoroughly experienced in the care of these beasts and who can catch and hold them with a minimum of dis- turbance. To him and to the interested laboratory helpers much credit is due. Small specimens like capucins and spider monkeys are caught with the gloved hand or with the protection of a piece of heavy cloth. Larger specimens may be caught in a net while strong monkeys are fitted (2) White and Fox, Archives of Internal Medicine, 1909, Vol. IV, p. 517. THE COMMUNICABLE DISEASES 521 with a collar and chain by which they are pulled into the corner of the cage and held, while the door is opened to permit a helper to catch the feet and arms. Two experi- enced men can take the temperature of any monkey that can be handled at all safely. The knowledge of how to do such work reduces the excitement of the animal and renders more accurate the observation of its tempera- ture. During the period of temperature-taldng food is given in small quantities and only after the record is made. Temperatures are taken in all animals by rectum, (3) the thermometer, a. separate instrument but always the same for each animal, well greased with plain vaseline, being passed along the anterior rectal wall and allowed to register for twice its indicated speed. During the pre- liminary work, special instruments of officially standard- ized accuracy were obtained by Doctor Brown but once the normals were obtained, ordinary good thermometers registering from 94° F. to 108° F. have been employed. If the record vary very much from the expected, such as the figures obtained at the same time on the preceding day, or if the rectum be crowded with feces, the instru- ment is shaken down and reintroduced. In order to facilitate timing of exposures we use sand glasses of three minute run. After some experimentation by taking records at vari- ous times of day it was found that monkeys as a group do not have a uniform temperature during twenty-four hours but register a higher figure during daylight than during darkness. This is probably due, as I shall dis- cuss, to the period of activity, not to the time of day. Figures obtained at various hours indicated that the highest and lowest temperature would be obtained if records were made at four-hour intervals at three, seven and eleven o 'clock AM. and PM. To give the normal tem- (3) The temperature in the axilla is often .5° F. hi^rher than by rectum, but the difficulties of the axillary method render it impracticable. 34 522 DISEASE IN WILD MAMMALS AND BIRDS perature of a moiikey, the kind and the time of day are necessary adjuncts. Reference to forty-eight hour charts which are used for the illustration of normal records, and for contrast with tuberculin reactions later, wdll convey to the reader a better idea of the normal daily rhythm M E M E M r M I M 1 ^ E M E M E M E M E Ta\ E M E M E M E M E M E M E M E M E M E M E 'J~^. z^^z r* F 107" 106° gios- |l04° K)a° 102° I0>° 100° 90° 1 °°° PulM. Sop- Dau. •4l» '3fl - L - M* •37« ■3«» •o ,ri K- n (N ^ -o <\ <; 'O fs ^ ^^ ■:j -^ .•-■ ..■■' y ,-••■ y y .---■■ ..--■ y' .-■-■ ,-•-■ ,-••■ ^--' ..-■■ .■■' iM., »m-l C- , 107" |- 1- 103° 102° 101° 100° 99° = 08° 1 -^ ^ 97° 1 - Duu. -4l» -38 ■38» -37« l-38« -^ '0 fs ^ n C- ^ N-j rs :^ r- ::; (r, ./ ^■■' ..-•^ ,-• ,'•• ..■■' ,..' X' ..•■ •■ i Chart C. Composite chart of twenty-two non-tuberculous Lemures. in these illustrative curves but these latter offer a guide as to what is to be expected of the different varieties. They show unequivocally the V-shaped curve of tke temperature of the monkey during twenty-four hours. The CallitrichidaB or Hapalidae have failed to show tuberculosis in our Garden and little has been done upon them. As a matter of record there is reproduced the only satisfactory chart at hand (K) taken very early in THE COMMUNICABLE DISEASES 525 the researches. It shows a similarity to those of the higher monkeys ; because of its very high afternoon record the animal was killed ; no tuberculosis was found. The Lemures, being close to the Primates zoologically and presenting a high incidence of tuberculosis, were M E M E M E M E (1 Jememememe- /.EM- : M|E M E M E ^EMEMEMEME / E M E s: ^^•,. 5 y " ~^ \ --^ :-::---V : j- — s — l-^. ^-- -Tij 'oo° 5;--, \^_ --. -1- ^ — xi- -1^ 89°^ _L-- 1 ~* =^-^-^-5- -^-r t_^^_- q -IK-JH-jK-^- -f«=F ^H^4^ — ± z-±. DauQfDi, ^ Cn -ij /rj t^ ^ 'rj N ^ t^ ( \ •s' 'O FuUe. ,.■■' ,.-■■ ...-' ,X y y y y -■■■' y- ,.- y y y ,,.-■ .-••■ y- y n,v>. y' y y ,,••■' y ,y y y y ,--' .-■ .--'' ---'' .--'■ ..■■ y "^s* Z)««. Chart D. Composite chart of eighteen non-tuberculous Cercopithecus. included in this study. Observations upon their normal temperature were hampered more than upon that of monkeys and even to-day we cannot feel the same confi- dence in the records. Irregularity is most marked and they seem easily disturbed by handling. Chart C shows a composite temperature for forty-eight hours of twenty- two proven non-tuberculous Lemures. The tendency for the '' night drop " is certainly existent but with much less definiteness than in the Primates. 526 DISEASE IN WILD MAMMALS AND BIRDS Conditions Which Modify the Temperature. Observations by A. E. Brown (4) and by Simpson and Galbraith (5) would seem to indicate that the diurnal vari- ation in monkeys is due to periodicity of activity. Doctor Brouni found that the temperature of a night monkey is t; ■e M E M E M E M £ M E M E M E M E M E M E M E M E M E M E M E M E M E M E M E M E S?^^ „,5;rr.., ^■T F. 107° 2 roa" .0.= 101° .00° 09° 1 ®^° ^ 97° 1 -^ Pulu. Dau. •4I« - _ — -37» -3e* V ^ ■s^ ^<^ fs ^ 'r> N ^ ^ N t> PitUe. Dau. Cn f^ ^ ./ y' y' .y y ,--• y .-•-■ .■■■ y' ,.-•' .'-- ,..-' y y y y' -•-• ..-' ,.'■ ,-'' .,-• Chart J. Composite chart of twenty non-tuberculous Cebus. be valuable ; such animals however rarely survive the dis- turbance incident to the test as they are usually suffering with some serious disease. Illustrative charts of several positive reactions serve to elucidate their character better than description. If comparison and contrast of the nor- mal and post-injectional temperature be made, the con- clusions are definite. It cannot be said that any type of reaction indicates a particular form of disease although the last type, the 534 DISEASE IN WILD MAMMALS AND BIRDS falling of the temperature beyond the thermometric regis- tration point, usually means advanced lesions especially of the caseous pneumonic form. A very small lesion may give a definite reaction as in Cebus (V). The examples given are those of a definite character but there are many charts that vary from the normal M E M TTm E M E M ^ M E M E M E M E M E 7^ - M E M E M E If, E M E M E M E M E li E M : '"r"^ ^•v 107° 1106" 1,^ 103° 102° 101° IOO° o©» 1 98° f.; PuU. Scsp. Dale. -40* - -;:; — — — — - — — fT y- — - - — ^ — — -C; — — — — — — - - - ■^ ^ i^ ^ — ^ ^ J -^ — N — l^ - fl — ^ — ,t; — t.i — ^ L J^ fl$ — k ^ g'-a«» r 1^ ^ fO T T r<^ ^_ '0 •n 0«. ^ -^ (^ ^ nS U '^ \^ U 0| ^ y' ./' y ,-•' y. ..--' y--' .-'' /' X IfO f>-'" ,.•' •' .■■■' y ,.'- .'-' ,/' y-- y^ 'A y. '/t n 5/ 'AW- % 'Ju '^/Li. — iii Chart K. GEOFFROY'S MARMOSET (Leontocebus geoffroyi). Non-tuberculous. upon which a decision is extremely difficult to make. Such animals are held in quarantine to be retested after the lapse of three months. Early in the work a suspected specimen was reinjected after two weeks, failed to give a reaction but died in about two months of tuberculosis. The nullification of the test by previous injections of tubercu- lin is well kno^vn. Three months ' interval permits a dis- THE COMMUNICABLE DISEASES 535 appearance of the non-sensitivity and allows any latent tuberculosis, possibly stimulated by the toxin, to develop. Eepetition upon the same monkey has occurred as high as ten times without apparent harm. There are sometimes in human beings local reactions at the point of injections. These have been entirely lack- M 1 M E M E M E M E M E M|E M E M E M E M E M E M E M E U E M E M E M E M E M E M \z ;;h^ Iji^d, 107° ^106° |l04° 103° 102° toi° 100° ee° 1 98° 1 ~* PuUc B»p. Dull. r" -4f ■38» -37* / - L- _ ^ _ -< _ _ _ _ _ _ _ _ _ _ _ _ _ 1 , _ _ _ _ _ _ _ _ -jf- — — M — — — — — — — — — - — — — — — -A — — — — — - -ae» -0 ^ ^ Tl ^ N -n ,,■ ,' y y .,--- y ,,--• .-••• .-•• /-•• m 7-'' .-^ ./ ,/ y y y' y .-'• 'lu "Jk _J Chart L. POTTO (Perodicticus potto). Healthy. ing from our monkey specimens. Nor have we ever seen secondary tuberculous lesions appear at the point of the needle-stick. Aseptic syringes and generally cleanly tech- nique have also protected against local abscesses. When an animal is injected he may scratch or pick at the spot for a minute or two but thereafter seems to ignore it entirely. 536 DISEASE IN WILD MAMMALS AND BIRDS Results. The value of the test can best be estimated by a recital of the mortality of monkeys, from tuberculosis, since its inception. Before the test was started practically every monkey in the collection for sufficient length of time to U E M E M E M E mT? M \ "m T VI E M T M E M E M - M E XT E M E 77 E m1 E ^ E M E M E V E Oct;*;™-! ^•. 107° ji: J 104° 103° 102° 101° 100° 99° 1 1 88° 4»» ■-JO* -3«« -a?' _ , _ _ _ 1 _ —J 1 1 _ ^ J _ _ _ 1 - _ 1 1 _ _ _ _ _ j_ 1. -, FuUe. Sup- Butt. "1 /? - 7^ — ^ ^ r^ - s t. --A te ^ ^ ^ ^ ■_ ^ '- ■A A E ^- ^ j- '- ^ 1, It^ - |- - L \\ r. ^1 f^ ^ CJI N~) ^ ;s N\ N •^ r JS ^ T^ fs N, .-'■ y' .---■ ..■■' ,.-•-' .y .--' ^-•- ..- ,--' y y y' m. r.-'' y .--' ./ y ,.--' y y ,.-•• -■' ,--'' .-" y .--' -A % % '2^ "/r. 'n ..••• .-''' ,-'' y .■■ y' /' 0^ .--' y' / y y' y .--' ^2 '¥/ 'if. % % % !?k % Chart N. BLACK-HANDED SPIDER MONKEY (Ateles geoffroyi). Tuberculous. quarantine, they do not properly belong to the exhibition collection. Up to 1906 when the test technique was perfec- ted nearly all deaths were due to tuberculosis, the figure for 1906 (including experimental animals) being 78 per cent. However from February 1906 to October 1907 and from then until May 1910 no case of tuberculosis occurred in the exhibition cages and both deaths at these given times seem like infection from ^dsitors. During the next three years thirteen monkeys died of the disease in the ex- 35 538 DISEASE IN WILD MAMMALS AND BIRDS hibition and many more in quarantine. By 1913 the out- break was stamped out. Its explanation is not so very far to seek. In the fall of 1910 we obtained some suspected monkeys which were kept in one of the quarantine rooms. After repeated testing two were passed. From them five U E M T m" T 17 - M T M E TT E M ~ M - M E M F. ■^ E M E m" E M E M E M E M E M E M E M E :^" ::^;r"^' rSa- F. 107° llOB" * _^ 1- 103° 102° .01° 100° 69° Pubc Dalt. -4I» ■38* -87» «y rr K- N- fD N ;s <^ N, -r. ^N N^ ^ (S ij frx ,,-•- ,,•' ,/ ,-•• .-^ y .--- .-•-■ y y /' l?f3 --' ^' .--' y ,.--' y' .,---' y y ^,-' .-■' f/y u Vm 'h ^M ^.. ^ 'A Chart O. RHESUS MACAQUE (Macacus rhesus). Tuberculous. cases are known to have originated and it was not until in 1912 when the whole exhibition house was cleaned of speci- mens, thoroughly disinfected and fumigated and until every specimen was retested, that the infection passed. In 1914 no cases occurred, while in 1915 a case either slipped through undetected or was a visitor infection; 1916 two cases, 1917 one case, 1918 one case (see orang- utan charts), 1919 and 1920 none and 1921 one case, 1922 no cases. Since 1912 the whole monkey collection has been THE COMMUNICABLE DISEASES 539 tested every two years, a method which enabled us to catch a small group in 1916 and has protected the collection since then. Three of the six monkeys specified above were never placed free in the general cages of the exhibition house, they being segregated in smaller cages. One, the M E MEMEMEMEMEMEMEV EMEMEMEMEMEME^ E V E M E M E N (1 E M E t^^^, ^..,-- .__L _. "S r' V. -- 107° -: : -- '-\\ : zb : _ -40" 103° z: 102° — 101° z 100° ^ 99° Z =5?E=5E-EEEE=-= % \-^---.'--- -380 -ae* -37" 1 °^° - |-° = 1 -* = ^ ,^ ^ ,^ ,r rr ^ ^ ^5 ;y- ^ ^ r^ Cn <■ 1 zr -3e« ■ .-•' •••' X y ,•' y .-•'■ ,-'■ .''■ •-'' ,--'"' ..-'' .-'■ y'' • ••' ,--' .--' •'' -^ ■ Atfl P.cip. / ' iffi y ■■■' .-'' ■■■" -^ y' ,.■' y- ,,-■' y y _.•-' ..-•• y ,'-'■ -•-'' ,.. ,,- 1-^8 tt^^'A^h::/^^ /"■ A Chart P. ORANG UTAN (Simia satyrus). Tuberculous. orang, was Avith its mate in an isolated cage. The other two were in larger cages and their history suggests visi- tor infections. We have never underestimated the possibility that an occasional very early case might evade detection by this test but we believe the history just outlined warrants us in depending upon it for the protection of the exhibi- tion. By the tuberculin test we have detected the existence of the disease in twenty-three per cent, of specimens. 540 DISEASE IN WILD MAMMALS AND BIRDS Every condemned specimen, forty-one, showing tuber- culosis, gave a positive test. Fifteen monkeys condemned on their temperature charts failed to show the disease. Eight per cent, of the tests resulted in suspicious charts, and the animals finally died of the disease in quarantine. M E M E M E M E M ■i M £ M E M E M E M E M E M E M FeI M -E E M E M E M E M T M EM- : ^iSL oSi^a'mi 0. 107° 2 1 .03» 102 » .0.^ lOO" 00 » i 1 - ! 07 = 1 - Dai/q/Dii PulHt. Dale. _l _ _, _ _ _ _ -i h" ^ ^ -S i_ _ _j _ _ _ _ _ _ _ _ _ _j _ _ M _^ _ _ — — '-— — ^ H - - — — — — — — — — ■^ ■<■ " - — ' -( — — - _ _ ^ ^ _ _ _ _ _j _ _ _ _ _ _J _ _ _ _ _ !- _ _ _ -< — -^ ^ — — f— — V — — — -< — 1 — ' — V — — — - '^ Tl -^ -^ f^ '-, 'O N C; ^0 «\ c: f^ CN ::: '^ •f\ ^ ro ..-•■ ^.■■■ ..-•■ ,..'- y' y .-•' ,.-'' X /' "rfi ^--' .^' / y' ,.--' ,' ' .--'■ ,---■ y' y ..'' % i y, 'A '^s- i/^ y? ± >/V Chart Q. VARIEGATED CEBUS (Cebus variegatus). Tuberculous. Fifty-six tuberculous monkeys died on exhibition, of which thirty-one were original there and twenty-five their contacts. Twelve of the thirty-one were in the early stages of the work, thirteen due to our misadventure of 1910 and the remainder, six, scattered over nine years. Another interesting experience concerns the exliibi- tion of a group of Ehesus Macaques in an open '' band stand " cage. The idea arose in an attempt to find a separate exhibition space for some good specimens that THE COMMUNICABLE DISEASES 541 gave unsatisfactory charts, with the purpose of applying at the same time the *' open air " treatment if tubercu- losis existed. The experiment has been entirely success- ful since in the eleven years during which this enclosure has been used there has been but a single case of tubercu- M E M E M E M E M E M E M E M r M £ M ? M E M E M E m' E M E M E M E m1 E M E M E M E s ^■. 107° 106° 1 104° ,03° 102° 101° .00° 99° 1 ®°° DayofDi, PuUt. Eup. Datt. -*f - - = ~ - /- = :: ^ - z = = = :^ = = r = r < ^ < >■ Z = Z, Z z Z z — j P s = b - 1^ F z y r r r z z =: - = r ^ Z ~ = - z ZJ = z 38» — - - _ _ 1 — _ _ _j _ J _ _ _ _ _ _ _l _ , _ -< — — — — —\ — — — -^ - — — 1 - — — - - — - - — -1 - - - - — - - \- - ^ - L - f- L - - - ^ -1 - - k- J -ST - - — — — — - —J - — 1— - - — - - - r - - — - ^ H - ~4 — - - ^ - ^ >^ ^ ^ u k ^ r =$ — 4 ^ -^ ^ J ITJ — ^ H ^ - -^ r< ^ IS ■< A E M E w "z-~." I'kL \ r' y. 107° I06° §105' 103° 102° 101° IOO° ■4I» 'o' '39* 1 98° 1 — 1 67° 1 -* DauofDU PuUt. DaU. _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _. _ L _ _ ^— \ — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — — ._ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 1 _ _ _ _ _ _ _ _ _ — _ — _ — — — — — — — — — — — — — — — — — — — — — — — — — — — — — \ — ^ ' _~ ^ _ _ ^ _ f _ _ _ _ _ _ ^ _ _ _ ■r~ _ -f" ^ -r _ -r T- ^ _ — - -oe« —f-^ — — — — '-' — — — 5 ^T-< — ^ - — — - — z ~ - t- -\- =P- — ^ — r=~ — — K — "*: — ^ -r- -r — -^ i^ — \— f* — r~ — - + ^ ^ re tN ^ n N -^/". ^ Bff. /' iftf '.■'" .-•■'' .■^' ■■■' -^ y' --'' .--' -'' ^^'^ ii'hWK%.iiA'jii m 1 Chart T. GRIVET MONKEY (Cereopithecus sabaeua). Tuberculoue. the hidden or unrecognized case is the menace. There is little or no problem when an unequivocally good or bad temperature record is obtained ; it is when there are slight variations from the standard for the group that decision as to the disposition of the specimen must be made. Nearly always such specimens are retested until the records are definite. If they be constantly irregular the animal is either sacrificed or exhibited in a separate cage far from other monkeys. It is by the sacrificing of 544 DISEASE IN WILD MAMMALS AND BIRDS infected specimens or the segregation of suspected ones that our collection is kept clear of disease. Hygiene of a general character must be maintained also. Our quarantine rooms are disinfected by f ormalde- hj^de and mechanical cleansing after every case detected M E M E E M E M T m1 E iJ E M E M E M E M E M E M E M T ^ E M T 'm E 7J T M E M fE M -E i^-lr-,", :!?;•: r* 107° 106° MOS" J 104° 103° 102° 101" 100° 99° |07° DaU. -4I» ^ ^ Ls«» . ::j C> V ^s ^ -^ ,• ^--' ..■■' .,.-'■ ■-'' .■■' y y ^'' ir/.i ..^ X' ,,-' ./ ,--'' ,.'-• X'' y' y ,-' .--' %.. 72^ 'Jk. 'Jj. "k. 7^. _ Chart W. CHACMA BABOON (Papio porcarius). Tuberculous. hawks and spits in public places or at home, unaware of his malign power! The Skin and Eye Tests with Tubeeculin. The first of these can be dismissed briefly, for in a few cases it was absolutely of no value. A kno^ai tuberculous monkey was injected into the skin of the chest mth 0.5 mg. of old tuberculin. The small bleb disappeared in a few hours and was followed by no reaction whatsoever. Other THE COMMUNICABLE DISEASES 547 attempts likewise failed, some of them I believe due to the technical difficulty of injecting into the skin. This tissue is very thin, delicate and loose at the less hairy places where a reaction might be read — arm, chest, abdomen. The hairless parts of the rump might be used, but are so M E M i M E M E- M \ M E ra E M E M E Fm E M E M E M E M E M E fw m M E M E M E w, m M E 'rX^m, :>Xr^-, ^U. 1070 |l04° 103° JOZ" 10) » 100° 99° |9S° 1 -- 1 97° 1 -* DaiiQ/DU Fulu. Sap. J> — — ) - - — — — — ■30' 3B» -87» «~ '^ re '<■ «-. ": ^ ^ ^ ^ ^ ^ 'O JN ^ ,-•■' y y" ,.--' X y y' /' /--■ / 1.^2 ,/ .-■' y ..-'' / y-- ,-•-' ,---■ /' y .-'■ y .--' y' ^. ^6. Vi % 'A 'A. i/7., '/f Vf 'h Chart X. BLACK APE (Cynopithecua niger). Tuberculous. often scratched and soiled with dirt that readings might be misleading. The Von Pirquet test was done on the first mentioned specimen and was likewise negative. His tuberculin test was afterward positive. The ophthalmic reaction is highly spoken of in the New York Zoological Park and has been used elsewhere. It was tried by me at the time Doctor Blair first discussed it, but with variable results. One set of two monkeys was treated with Calmette's purified tuberculin into the 548 DISEASE IN WILD MAMMALS AND BIRDS conjunctival sac and given a subcutaneous dose of old tuberculin. Another set received 1 per cent, old tuberculin into the conjunctivae and the usual subcutaneous dose. Although all these monkeys gave a temperature reaction only one gave a conjunctival reaction. Fearing that the two tests simultaneously might be an unfair trial, another poor specimen was given an eye test which resulted negatively; a later subcutaneous test and autopsy revealed the disease. Because of these experiences and the fear that any reacting conjunctivae might become sec- ondarily infected from the uncertain personal hygiene of the beast, we decided to omit this method and rely upon the temperature test. Pathological Effects of the Injection of Tuberculin'. It is generally believed that tuberculin injected into tuberculous animals, in doses large enough to produce a marked reaction at the site of disease, may stimulate the process to growth and spread and that certain parenchym- atous organs in such bodies undergo degenerative changes. We can give little information concerning the first point because known infected animals have not been sacrificed during the test and we are not informed of the degree of morbid lesion in those dying, since we had no previous knowledge of its existence. An interesting and practically useful observation has, however, been made upon the kidneys of several monkeys dying shortly after tuberculin injection. It consists in a marked cloudy swelling of the renal epithelium and a con- gestion or even thrombosis of the glomerular capillaries, accompanied sometimes by increase of nuclei in the tuft and by amorphous material in the space of Bo^\^nan. Grossly such kidneys are but little changed, albeit the cortical zone may be dull and opaque and swell out slightly on section; very occasionally bloody streaks may separate the cortical and medullary striae. In a few kid- Fl(,. 58.— EKFKCT OF TUBERCULIN ON THE KIDNEY. AN UNUSUALLY SEVERE REACTION IN THE RENAL TUBULES AND INTERSTITIAL TISSUE FOLLOWING A TEST DOSE OF TUBERCULIN. SUCH A MARKED CHANCE SUGGESTS THAT IN THIS CASE PRE-EXISTING RENAL DAMAGE WAS AGGRAVATED. THE ANIMAL WAS TUBERCULOUS BUT HAD NO LESIONS IN THE KIDNEY. THE COMMUNICABLE DISEASES 549 neys there have been suggestions of preexisting nephritis but usually the findings are confined to those given above. At all events true glomerulonephritis is not often found. Monkeys which have this condition may or may not exhibit a behavior suggesting its existence. Sometimes it will be noted that the animal is dull and eats little, at other times the keeper mil report that the cage is seldom wetted and we know of cases in which only an ounce or two of urine has been passed in a day. Two monkeys were distinctly ataxic and incoordinate and one of these had a convulsion. From one a specimen of urine showed albumen but no casts. These signs of renal affection are not always alone nor are the kidneys necessarily the only part diseased since postmortem records show a variety of accompanying lesions, bronchitis and enteritis, for example. There are, however, several cases dying in a few days after tuber- culin injection, both with and without tuberculosis, in which the renal changes were quite prominent; two examples, without tuberculosis, exhibited the damage to the kidneys very well and with no other evident visceral pathology. The relation of cause and effect may not be unequivocal, but these findings suggest that the condition of the kidneys deserves attention when tuberculin is to be injected. My associate, Dr. Corson-White, is firmly convinced that the substance whips up a preexistent parenchjTuatous disease and wants to see a urinalysis from every monkey that is in any way abnormal. The Tuberculin- Test in Other Animals. Two cases of tuberculosis occurred in White-nosed Coatis {Nasua narica) so that it was decided to test their neighbors in the next cage. There is reproduced a com- posite (Y) of the temperature record of three of these animals after receiving 2 mg. of tuberculin under the skin. No tuberculosis was found in them at death, all dying 550 DISEASE IN WILD MAMMALS AND BIRDS mthin two years. The similarity to the primate type of temperature curve is striking. Chart Z shows the course of temperature before and after tuberculin injection in a Bactrian Camel (Camelus hactrianus) 2 which lived for some months and showed no infection at autopsy. Charts AA and BB show the temperature ranges of respectively a healthy and a tuberculous Bison [Bison bison.) Chart CC is that of a Malayan Sambur Deer (Cervus equinus) which died a week after injection, showing fibro- caseous tuberculosis. The failure to make an initial rise is noteworthy, but the fall in temperature may be explained by the severity of the lesions and the approaching death. Chart DD represents daily and post-injection records of a Virginia Deer (Cervus virginianus) which at death was found free of the disease. Cliart EE is that of a healthy American Elk {Cervus canadensis) . THE COMMUNICABLE DISEASES 551 M E M ■e M E M E M E E M E M E M E M E M E E M E M E M E M E M E M E M E M E M E ^srjr,. 0. r. I07 1 i- >os° 102° 101° 100° 00° 1 98° |e7° PuUi. Date ■4I« ■SB* -a?* -^ tv. ^ •^ <\ <; -n H :^ T ^ ^ X '■■■ ,-••■ _.y y' y- /-' ^' /• y y y ,.--■ y .X' ..' ,.-•• '■■^ y' _ Chart Y. Composite chart of three non-tuberculous coatis. 552 DISEASE IN WILD MAMMALS AND BIRDS d? ? s ? s s ? ' ) - :::i:::::it:::::::::::::::it:::::::::::i-:::::-:t:: \ = t X i i--^ ±..t---- i^J ■\, \^ 2 _ . -|-.H>---4--^ : "' ""'li 7 ■'■. ^, :::::!^::::: ::::::: :: :;: i; 4 w..._ ^[ T ""n^ 5 1. _...-., t '■., ^?7 s T ^^"~ + ;<>■ f ":"" ---T-- .-- ^ ■\, :.:::::::::::::: ::::::::i:::::: :: i _ \. w^ \, 5 . '*-•. ]n ..:■:;: w^ \ s P rn^ ■^ ,2 'mM'^ '\, \=fl 5 ^' it X^'T \ \ :.:::::::: ^:::::::: i . ^:,, ""-^c / 5 ^^; T — ; fZT '\ "^ 5 ::: :: :::::::::::::;:^: :::: : T .. ,,^'' WT \_ = ^'-^ f"" ..::::!:i;;::::::::::'"ii:7 \ '\, £ ^^[:>,... i j^ "^■■j::^ I ,,;..:::;;:::::^::i::4 ^ i - ^v ---^ "'-^ :::::::"": ^s; ---x-vi \^ s - ^'==:: - ::::::::::::::::::::' ■ = ;;-- ---«;. j \^ ^ '^•-, 5 ^!:, j 5 :) ±.. ::'" : wi'j '~\ '"\ N^ 2_ ::;.:::::::::::;::::::::: ::z :::;::; r . ill y \ s _ :::::::::::::;::::::::: :^:::::::::::::r:" t 5. ^^ T \, \ = _. T ^^;; y- " t=;; n'''J '\ \^ 2 _ :.::::::::::::::::::::::::::-' i = :::::::::::::::";"""" 'K X"^ "^ :i. fm. / '\ 2 " "T ----1^ -*^ . _.-.:.::::::::::': ''=:: ~"kyj - :.::.:::::::::::::;:^:: :::: I :"■ ,,-'' *< 7 "\. ■\, 2 _ 'I I ""n \, s 1 'v f^' '■. f5^ s ,-- ' T ...I :::::::::::;;== ""^^:t \ \.^ 5 t -"^^ T'"" :::::::::::::""" ■ n- ~'m^~^ \ \. _:::::::::::::::::::::::] t , W-V J \ \i^ 2 :<"' : ...:.::::::::::::: :i'' "^n:T '•■. '1 2 '5 ::::::::::::: te;;: ^h^t Ci 5 ...I .1 1 THE COMMUNICABLE DISEASES •? '^ 8 ? S ? ^ 553 ~ \ \. u S UJ s tu t UI Ji 2 1? "•■^ UJ *■ \ Ui '.:'.'.:'.: \i\: i \ \ i^ s "" "^ 'Zol S \ \ S ui )r S , u ^i ' 5 '\ ^ S UJ ^ ~\, UJ 'Z ' : , S' UI 'i 1 2 a \ \. r^ s u il < s 5 u t '■\ \ UI if \ :^ S UJ ^l":' ^ \ 2 UJ ^ 1 s z ' ■. i' i ■\, \ ^ UJ \ ■^.^ UI UJ ^■, UJ ^;:5 i \ s^ S UJ ' kv ^ S UJ ^ ■^ ' t . r UI i' ' i n^ UJ ■J" ' 3 ■■., \ UJ ^z:c: ): \ S ■' ' UJ i \, \ ^ S UI ., '.'::::.k\h ^ ' UJ V\l c \, u ^ , r\-' i \ ^ UJ 3 ■■■., UI i\\\ T \ \ UJ ...,{. __ >f S< UJ ^ ^ •., \ s - nv 2- \ UJ " s \'V ^ \ ^ """ " ;^ " UJ / \^ UJ 7 r \ ;i^ 36 554 DISEASE IN WILD MAMMALS AND BIRDS M E M E M E M E M E M E MJE M E M E MfE M E M E M E M E M E M n M T M fE M E M E M I '^!r:-", eJir-' F. 107° 106° glOS" |.04° 103° 102° 101° 100 ■> 00° 1«. 1 - \z Dagn/ZH, i>uZ.«. JJ«p. _ _ _l 1^ J -i- _ _ _ _ _ _ _ _ _ _j _ _^ _ _ — — - — — ^ -^ — — — — - — — — — - — - — — - — — -^ — — — — - y^ ^i tx (x "^ Cn. cx "> u bo ^ (>v u uj 0^ "i u Uj ,--■ ,-•-• ..•• X y- y ,-•'■ ..--• ..-•' y- .,-• y- y /' V// .'^ y- ,/ x;' y y .-•'' ,.'■ y .--' ,.■■' y % J^.. %i 1^^ ^h %i y.7 Chart BB. AMERICAN BISON (Bison bison). Tuberculous. THE COMMUNICABLE DISEASES 555 m" iQ 00 001 id B □ B □ Q □E id B □ BE SB WL ica B □ B □ B □ B □ B D B E3I id! inrnii^H 0- = : EE = i: - E - = = z: !2 E - E = : E E E E E = 1 e! e: - <, 39' - I Z C3 : - I 1 »* JX 3 II So j \ i 1 3 c \ ^ E E c E i g: E r z ;5 E 5 i En ^_ E^ .30. -e7» -3S* -• y ..-^ y- ,--• y' .--■ ,-•' , • ~- y- ,-•' .,• .--' _^-' / w'o .■ - .■■■ / ./ ,,.-- .-•• ,/ .-■ ,,.-• .--' .--• --' .-•' '-A \ JL i^ S^. .i^ ^ i4 ^ Chart CC. SAMBUR DEER (Cervus equinus). Tubercuk 556 DISEASE IN WILD MAMMALS AND BIRDS 6? 5 ? ? S S 1 ^ ^ ::::::::::: i i- \ "■ = _ :::.: :::::::::::::::::: :r "^ _ _. .V ^2 / •-., , . " ^Vn'-T ^\, 5 ., ::::::;:;;: %^ .. z^:'T \ \^ ^ 5 T 7^'^ ,,: :..:::::": ^'^s: ? 2 * •. "" ::::::::::::::::::::::::::::::::•:::::::::::::::: a: T 2 ^\ F " ■;: -- rcTj \, f 2 '^ : ^ ::::::::::::::::::::::::::::::;"::::::::::::: ^^ft '\_ \ s : L """^ZFT '\ 2 '" ""F :■"■ F: " ''^"' K \, \, ^ 5 i\'.v.v::v. F ^ ""^ i ? = : t..--^v.v^v:.vw I '^ _ - _ I <- i^ / \, ::"::::::" ff ' '"^Ti JV> s .. J.Z i" ■^ ^' ^F' 9 \ \. 2 "F ' ^ "::::::::: f:" -"tiui ■--,, ''\ 2 ^^ .:::::::''::: '^: " ir^^j \_ 2 _ ,..^:\: ::::::: 1 ,,,'-' '^^iy \, \ 2 ^K ;--li '^ F. ^ll'T ~-.. 5 _ ..: f:ffff ff: f F, iz:'j ■■■ 1 H 2 ;-'' ----^ ^^ ~ ;;:= ^F'T '•■, 5 :..: f: : fffff :ff ff f .:;,..""""" .:^'± / '\ \ 2 ^'^s, ; :"::::::::::::::::" '::::::::\'.::::\]]:::\::::'::''''^:r J ~\ '-\ N^ ^ .' J 4^ _ :.:::.^,.i: .:::": ^iej \ \, = ,.. I^ .. _ ^. s?iT 2 H -- - F ■ : si , \. >^ ^ . ^^ "/ ^i'J \, = FF F FFFFFFFFFF FFFl : F FFF FFFF: ' V 5^?T ''\ '\, 5 F F_ \ ^'WJ \ ^^ ^ ^. 'F^ "FIT ■\_ \, ^ ^.. ... i 'Fc^ / ^. ^? 5 F F F FF FFFFFFFFFFFFFFFFF.F; ^ "^ ^^^k- <.rT \ ^ 2 I THE COMMUNICABLE DISEASES 557 ;; T M - M '^ M ? M r M r J? E M r 17 E M ^ M £ M E M E M E 17 E M E M E M E M E 17 - M :m E I" r? Im fE J [e 2-. E I t E j s E E E E I E E E E ^ I = E 1 § ^ 7 \ - ^ 1 E ^ E E z E E - E :E 1 i E E 41* 38* 3 S E \ 5 i 1 E E 2 ft A i = ^ i 4 ft I i ft; 1 g > \ E E ft \ I I - = P c ^ -< E = 1^ ^ ^ E \ E^ j E i s E •3«' ^0- ». -- ^ ^ Sj tx ^^ V* >>5 VI l> "^ >3 >« •s< t>« ^ ,'•■ ..-•' y .^ ^- ■ ..-'■ X ^--' .-••■ ,.-' y ,.--• ^-■' y' Ytf-i ,,-' ,.--' _,•'' ,.--' ./ ,.-- .-•' y ,-' ..•-' ,--" .--' y- .--' ,.' ,.--■ 'A i^ 'A ^ — iz. i^ 'A ^^ Chart EE. AMERICAN ELK (Cervus canadensis). Non-tuberculous. SECTION XVII— PART 2 MYCOSIS This is a general term applied to the infections with Hyphomycetes, but in the zoological material discussed here it refers chiefly to the growth of aspergillus in the air sacs and viscera of birds. ; A few cases of cutaneous mould growth have been seen in mammals but are of trifling importance ; brief mention will be made of them on a later page. Under the names of Aspergillosis and brooder pneu- monia, the infection with Aspergillus fumigatus, A. glau- cus and others is well known to breeders of chickens, ducks and ostriches. Literary references to its occurrence in zoological collections are numerous, but there seems to be no record indicative of its frequency in the various orders nor discussion of the pathological types best exhibited by different birds. There has indeed been some discussion of the actual entity, mould disease, the picture found at autopsy being referred to secondary con- tamination with fungi in the presence of bacterial infec- tion. Experiments by DeLong and others have been indifferently successful in the production of the disease by iiilialation of mould spores. The appearance of groups of cases in breeding places, apparently all exhibiting the same organism, in the absence of other varieties of avian epizootics seems to warrant the conclusion that the hyphomycetes can at least be associated with a fatal morbid lesion of quite uniform character whether or not they be the original invaders. Judging by our experience it would seem probable that the aspergillus can, under conditions not fully explained, cause inflammation of the avian air sac and tubercles in viscera, in the absence of other evident causes of illness and death. Over three-fourths of our cases have no other 558 MYCOSIS 559 diagnosis than ''mould disease"; this may be in part due to overlooking other things, but to a much greater extent to the very extensive mould growth which obscures all other changes. That unexplained preparatory condi- tions may exist is indicated by general observation and some experimental work. Whereas in breeding estab- lishments this disease occurs in epizootics, or in groups of cases, with us it is enzootic, constantly present, never, however, bursting forth in virulent form with high mortahty. Nor is the condition highly contagious. In an attempt to explain its source I examined over forty varieties of feed and found therein several strains of aspergillus and of mucor ; the latter occurs occasionally in the avian air sac. The infective material is therefore constantly present, and it would seem that if it were capable of initiating a fatal disease many more cases should come to our attention. Perhaps these moulds do gain access to the avian air sac and are killed off, or only assume a pathogenic role when they are in large num- bers or a preexisting disease assists them. Since our records do not support the idea that a preceding condi- tion must exist for a growth of mould to be successful, and yet pathological and experimental observations sug- gest that something helps its colonization in the air sac, what are such conditions! Moulds grow on feed and lit- ter in which birds pick ; from this it is quite possible for a piece of grain or even inorganic matter laden with spores to be inhaled and lodged in a secondary alveolus near the air sac whence extension into the air spaces could occur. In addition I think it quite conceivable that a whole colony of mould might be inspired with the same result, the mechanical obstruction being sufficient physical damage to incite inflammation. The continuous moist surfaces of the bronchial passages and air sacs afford conditions favorable to the growth of mould and as inflammation is not vigorous, little resistance is presented to its spread. 560 DISEASE IN WILD MAMMALS AND BIRDS The mode of operation of these hyphomycetes has usually been assumed to be a mechanical one, local colonization replacing healthy tissue or spreading along surfaces so that function is physically impossible. A support of this idea is to be found in the fact that inflam- mation, as produced by schizomyces, is trifling or absent; the necrosis that occurs is due to choking off of tissue by the intricately tangled masses of mycelia and blocking off of air or blood supply. The existence of an infiltrating and necrotizing form in some parrots and gallinaceous birds, suggested to me that a toxin might be responsible for some part of mould action. Proof for this speculation was sought by injecting into the pectoral muscles of pigeons an emulsion of a dead mould and a filtered broth culture. Necroses occurred but only to an extent which I interpreted as due to the physical destruction of muscle by the injected material ; they were larger with the dead mould than with broth filtrate. I concluded there- fore that aspergillus perhaps has no toxin as usually described for bacteria. Types of Mycosis. Avian mycosis occurs in three different forms, two of which are probably of similar nature and two are fre- quently combined. (The first variety, most often seen in gallinaceous and anserine birds, consists of thickening and opacity of the air sac walls, upon the surface of which either a curd-like pseudocoagulum or a velvety or fluffy mould growth appears. This variety usually begins about the anteroinferior pulmonary stoma on the right side extending thence to the related sac, upward toward the wing and downward to the abdominal spaces. Occasionally the middle thoracocervical space is involved, probably via the opening in the syrinx. Extension takes place by the way of normal passages, but when the growth is dense it also seems to occur by continuity through tis- sue. This variety may or may not be associated with the Fig. 59. — NODULAR OR TUBERCULAR MYCOSIS IX THE LUNGS OF A DUCK. ONE-HALF OF THE LUNG IS CUT AWAY AND LAID ON THE INTESTINES TO CONTRAST THE PLEURAL AND SECTION SURFACES. MYCOSIS 561 second, an infiltrative type of lesion best seen in the lung. Under what seems to be a true picture of mycotic pneu- monia, dirty gray consolidated areas will be found around the bronchial space, infiltrating in all directions and with- out definite boundaries. A similar lesion has been seen also in the liver on rare occasions, but the lung is its usual seat. Judging from microscopic appearances this is a process complicated by the addition of bacteria. l^ The third variety is nodular or tubercular mycosis, a process of probably more chronic nature since around the isolated lesions connective tissue is perceptible, it being absent or inconspicuous in the other types. The formation of gray or yellow-white nodules from a few millimetres to a centimetre in cross section, is the charac- teristic production in this variety. Lungs, liver, spleen, intestines and air sacs are involved in about this order. Attempts at explanation of this peculiarity of growth were made in the direction of identification of the species of mould, kind of bird and probably degree of resistance. No conclusions could be drawn since the same variety of mould was found in this as in other types ; no bird showed a special susceptibility or resistance to it. It is quite diflBcult to obtain a culture from nodular mycosis, it being necessary to crush or grind the solid masses before making cultural implants. This variety should always be differentiated from tuberculosis by staining for the organisms, j Histologically studied these three types are not as easily separated as the gross appearances would warrant one to expect. The original mould nodule begins in essentially the same manner in all, a small colonization of mycelia and spores which grow centrifugally, but ever becoming more intricately wound in their first location. As the tissue is invaded, total necrosis takes place, no recognizable cellular architecture being left. When fully formed the mycotic tubercle consists from the centre outward of a necrotic mass, in which spores and mycelia 562 DISEASE IN WILD MAMMALS AND BIRDS stain indifferently well, surrounded by a dense zone of fully formed mould beyond which mononuclear cells and a few loose fibrils may be found. The circumferential tissue of the viscus supplies a mild congestion and per- haps a moderate connective tissue capsule. Giant cells, as known for tuberculosis, do not appear. Where the process involves loose tissue like the lung, especially when growing rapidly, the sporulating heads of the mycelia stretch out in advance of the main mass and resemble rays. So too in a rapidly gromng nodule radiating mycelia are sometimes seen but never with the regularity of arrangement typical for actinomycosis. Diffuse and irregular mycelial spread is characteristic of the second or infiltrative type of lesion and between the mould stalks one may discover well-preserved tissue cells and at times bacterial forms like cocci or bacilli. This picture, suggesting as it does bacterial admixture and more active inflammation, leads one to the conclusion that the morbid process in which it is found does not represent mould disease per se, but a mixed infection. Whether or not the bacterial disease exists first and paves the way for the mould I am unprepared to say; I am inclined to the view that mycosis can start by itself. iNcmENCE IN Avian Orders. Mycosis may be said to occur in all birds although the appended list fails to show cases in a few of the orders included in this study; the sum total of autopsies from missing orders is only 45, so that they may be ignored. Picarise. . . Psittaci . . Striges. . . Accipitres. Columbae. Galli 3.7 1.2 3.9 6.7 5.1 .6 2.7 Fulicarise . . Alectorides. Gaviae Impennes. . Herodiones . Anseres. . . . Struthiones . 8.6 2.7 10. 40. 2. 6. 9.4 For meaning of italics see foot note Table i. 8y8SBMHpC^:. •' t^i^ ^^^^^^^^^^1 f %.^^ ^^^VH^kL^^^'^ "^^^^1 ^^^^^^ ^^^^m^i IG. 60. — MICROSCOPIC APPEARANCE OF ONE OF THE NODULES. NOTE THE CENTRAL NECROSIS AND THE CLOSE FIBROCELLULAR PACKING IN THE RESTRAINING CAPSULE. MYCOSIS 563 These figures hardly permit conclusions as to rela- tive vulnerability unless the large percentage of cases for the small number of Impennes, Gavise and Struthiones be permitted to stand. Judging from orders upon which there are at least one hundred autopsies, owls, ducks and eagles are most likely to suffer with mycosis. The per- centages are, however, not very convincing, and it would seem better with these data to conclude for the present that any variety of bird is susceptible to mould. Ostriches have long had the reputation of succumbing to this affec- tion, so that their outstanding position in the list is more easily credited. It was to be hoped that the various susceptibilities would assist in an explanation of the genesis of mould disease, but the result of the analysis is suggestive only in one direction which can be stated quite briefly. Ostriches, owls, shore and swimming birds have large stomata between the lungs and the lateral air sacs and show a high mould disease incidence. The natural thought is that access of mycelia and spores to the air sacs is facilitated. More comparative data is being assembled upon this point. Water birds are more susceptible to mould than land birds ; the percentages based upon cases and totals for orders is 5.4 to 3.7. The hygiene of mycosis is that of scrupulous cleanli- ness. Being hampered by incomplete knowledge as to its genesis one can only apply common sense measures. The germs having been found upon all the vegetable feeds, it naturally follows that they cannot be eradicated, but their colonization in large numbers can be prevented by repeated cleansing or sterilization of bins and pans so that no mouldy or musty material is given to the animals. At times of serious outbreaks sterilization by burning all old feed, starting fresh with good material in bins painted or saturated with disinfectant and then deodorized, is the only salvation. Where the blast lamp can be applied, it is the safest procedure. Autoclave 564 DISEASE IN WILD MAMMALS AND BIRDS sterilization should be thorough if practiced. B}' con- stant vigilance we believe that our large exhibition house is protected, but the sanitation of the pond, where many ducks have the disease, can never be so satisfactory. Mycosis among mammals as an organic pathological entity is certainly a rare condition, indeed almost always to be considered an accidental or secondary one. In so far as human pulmonary disease is concerned it is among the pathological curiosities deserving of individual report. Moulds of several varieties have been found in intestinal ulcers, in cranial sinuses and in the ear, while generalized mycosis from thrush and favus are reported (cases of thrush in kites with fatal outcome has been described on page 168). Cutaneous infestation with mould is quite another matter for ring worms and similar conditions are now believed to be due solely to the pene- tration by spores and mycelia into the superficial dermis and into hair follicles. Numerous small lesions have been detected on dogs and cats that coiTespond to the ring worms described for them ; to these we have devoted little study, because when discovered the animal is removed, treated or killed to protect others. Doctor Weidman has discovered several varieties of hyphomycetes which he will report upon at a later time. No case of generalized mould disease has been found in mammals, but Doctor Weidman has discovered a hitherto undescribed mould in ulcers and cutaneous abscesses of seals; no ex- tension beyond subcutaneous areolar tissue occurred in these cases. Botryomycosis, while not strictly belonging to the foregoing group, may be mentioned here because of its nodular tumor-forming superficial growths. The case to be cited certainly belongs to this illy defined group of diseases even if the organism was not isolated. It cannot be accepted as wholly demonstrated that the disease described under this name is always the same or that it has a single cause. MYCOSIS 565 California Hair Seal {Zalophus calif ornianus) 9 . Had sore spots on side for several months. Ate well up to three days before death but ate something to time of death. Diagnosis. — Chronic enteritis Avith acute hemorrhagic exacerbation. Low grade chronic diffuse nephritis. Botryomycosis. General condi- tion poor, subcutaneous fat practically absent. On the left side of the thorax there is a warty and nodular thickening of the skin overlying diffuse and flat thickening of subcutaneous tissues. There are warty, pustular, fistulous communications in four places between surface and deep mass. They are covered over with light crust. On dissection mass is found to be in subcutaneous tissue well outlined and encapsulated and consisting on section of dense, white, firm trabecul83 forming a mesh around yellow, soft areas which can be squeezed out. An adjoin- ing lymph node is much enlarged, dense, tough, resilient, on section showing great connective tissue increase and solid brown medulla. The thyroids are solid, brown, 3. x 2. x .6 cm. and 3.7 x l.S x .6 cm. Trachea and bronchi contain pink froth but mucosa is negative. Anterior edges of lungs are distinctly emphysematous. Remainder of lung is uni- formly congested and lobules are quite prominent. No consolidations. Bronchial lymph nodes are slightly large, anthracotic and wet. The heart is dilated, filled with mixed clot, muscle firm and deep brown color. The liver surface is smooth, edges sharp, size normal, color dull brown mth greenish cast, consistency firm and tough. Section surface is glistening, smooth, moist, lobules indistinct but probably normal. The gall-bladder is full of fluid yellow bile. The common duct is patulous. The spleen is of normal size and shape, capsule opaque gray. The trabeculae are prominent, the pulp stippled rusty brown. The kidney is of normal size, capsule is smooth, strips easily leaving a smooth purple surface. The organ is firm. The lobules and lobular markings are distinct. The adrenal has a narrow regular brown cortex and gray homogeneous medulla. The stomach contains whole fish. The mucosa shows digestion, congestion and mucus formation. Begin- ning at the pyloinis and extending to the colon the mucosa is swollen and edematous, yellow brown. In the lower part it is quite firm and opaque. In the upper part it is more translucent except where there are difl'use hemorrhagic mottlings of the submucosa. Here and there are shallow erosions but no ulcers. The lymphatics of tlie mesentery are definitely enlarged, white, pale and very firm. Smear from the surface of the growth in side fails to show any definite yeasts or moulds by Loeffler's or Gram's stains. It is largely made up of polynuelear cells mth many large mononuclears, many of which are pha^oeyting polynuclears and nondescript bodies. Cultures failed to grow. Micro- scopic section of liver shows marked congestion with slight hydropic degeneration of the epithelium. The kidney capsule is not thickened. Interstitial tissue not grossly exaggerated but connective tissue nuclei fairly numerous. Some tufts have decidedly more enlongated nuclei 566 DISEASE IN WILD MAMMALS AND BIRDS than others and connective tissue around the stalk vessels seems hyaline. Capsule for mcjst part not thickened but space contains cells and detritus in many instances. Epithelium of the tubules for the most part swollen, loosened and without nuclei. Some imperfectly formed casts. The intestine shows distinct congestion of the whole mucosa with here and there definite small hemorrhages well out in the villi. Slight round cell increase but no definite fibrosis. In submucosa arounK I loN OF V'\^', ,f I',",\" "^.V, ,\ nVv\ « v'^rissflF^ NECROSKS. ONE WITHIN A BRONCHIA, (.\1 IN SKMICON^OLIDA 1 I I > IMI.M..\AR^ I ISSUE. iRDIAI, STRANDS COUI.D BE FOUND IN BOTH AREAS. THE STREPTOTHRICOSES 583 opaque, architecture probably normal. The gall-bladder is distended with viscid green bile; the common duct is patulous. The spleen has a rough, thin capsule, consistency tough and resilient, the section sur- face is mottled red with purple points ; on section tAvo small, pale objects seem to be squeezed out. The kidney capsule is smooth, strips easily leaving a smooth, deep red surface, the consistency is soft, the cortex is deep red, then a purple line between it and the red medulla, striae invisible. The stomach contains frothy grayish mush. The mucosa is finely mammillated, deep pink until the last third when it becomes deep red, deeply injected and somewhat thickened. The pylorus is closed. Externally the gut is congested, in places translu- cent but for most part seems thickened by reddish swelling of both external layers and mucosa. The mucosa is granular or pebbly with here and there a small bloody suffusion. The histological section of lung shows alveoli open, septa relatively thin but somewhat congested, bronchi mostly open and connective tissue not increased. Some few bronchi, especially the larger, show a slight catarrhal bronchitis but mostly an infiltrative peribronchitis. The nearby veins and arteries show the most striking change, there being in nearly all of them a distinct thrombosis without circumferential pneumonia. In one place a distinct peribronchial pneumonitis was found. The kidney shows very marked congestion of all parts, causing compression, cloudiness and granularity of the epithelium. Glomeruli and connective tissue about normal. The intestinal serosa is negative save for congestion. Sub- mucosa is densely infiltrated with mononuclears, some in definite groups. Section does not show areas mentioned in notes but these could be accu- mulations of cells Avith congestion. No streptothrix forms. Section from the infectious focus of face consists of active granulation tissue, densely injected and filled with mononuclears of two types, one the lymphoid cell, the other of the young connective tissue type. Areas of grouping like abscesses are seen and some necroses. Streptothrix in small numbers in the cellular collections. Nail tailed Wallaby {Macropus unguifer). Kangaroo disease of nasal region. Necrotizing bronchopneumonia (Aspergillus fumigatus and Micrococcus albus). Acute diffuse splenitis. Congestion of liver and kidney. The general condition of coat and nutrition is good. The face is wide just below the eyes. About the " hare lip " and the nose the soft tissues are soft, gray, necrotic. All the internal nasal tissues seem swollen, gray-red. There is subcutaneous edema, bloody in places, around the right face, eye and jugular angle. Tissues of nasopharynx swollen, deeply injected and covered by a thick mucus. Pharyngeal and buccal cavities negative. Tonsillar areas pink and flat. Larynx and trachea slightly swollen but pale on mucosa. Salivary glands and cervi- cal glands normal in size and pale pink. Pleurae pale and empty. Lungs swollen out uniformly, quite cottony except at lower right base where there is a nodule about 3x5 cm. firm and doughy. On section it is found to be a peribronchial consolidation of pale reddish gray color 584 DISEASE IN WILD MAMMALS AND BIRDS and indefinite outline. The bronchus itself is deeply congested and eon- tains a grumous mass. The peribronchial lymph nodes are small, soft, pink, homogeneous. The heart is negative. The liver is large, surface smooth, edges sharp, color deep purple, consistency soft. Section surface is glistening, smooth, moist, very dark purple with obscure markings. The gall-bladder contains fluid brown bile; common duct is patu- lous. The spleen is soft, tough, capsule pebbly, section surface is mottled, light and deep pink, follicles and trabeculae not distinguished. The kidney capsule is smooth, strips easily leaving a smooth purple surface, section surface is glistening, deeply congested, striae obscure but seem normal, glomenali not visible, organ is soft. The gums and teeth are not involved in the mycosis. The stomach contains mushy digesting food. The mucosa is mottled pink, soft, digesting, at lower half submucosa is deep pink, a few small ecchymoses. From pylorus to ileum, serosa is deeply injected, edematous, mucosa swollen and edema- tous, deep pink, loosened in places, but translucent. Below this the mucous membrane becomes smooth, flat, pink-yellow. Lower ileum and colon contain rather firm fecal balls. Follicles nowhere prominent. The pancreas is small, soft, yellow-pink. The follicles of the mesentery are small, pink-gray and homogeneous. Smears from the broncho- pneumonia show a thread-like Gram-positive form and a few Gram- negative rods. Cultures from lung show Aspergillus fumigatus and Micrococcus alhns. Nose too foul for culture. Histological section of lung shows the alveoli mostly open but the septa widened by congestion. Blood vessels are open and contain recent clots; one vessel near lesion below is thrombotic. The two large bronchi in section show catarrhal bronchitis and infiltrative peribronchitis of which the latter is more severe and advanced. Beside the larger is a necrotizing pneumonitis from which nearly all the architecture has disappeared. The exudate is chiefly mononuclear around the edges; centre no cellular identity. Another mononuclear process not connected with bronchus in section is found with an early necrosis. Streptothrix strands may be found in the bronchial exudate and near the margin of the necrotic patch. They do not grow in colonies however. The spleen shows general congestion without pigmentation. Follicles large, solidly lymphoid. Connective tissue about normal. The kidneys show marked congestion everywhere. Capsule and intrarenal fibrous tissues about normal. Very severe con- gestion which seems to have caused compression and granularity of the epithelium. The last case, judging by stained smears, is one of pure nasosinusitis from streptococci and streptothrices. Cultures were not tried because of the enormous bac- terial flora. Robust Kangaroo {Maeropus robustus) . Acute purulent ethmoiditis. General acute purulent anterior cranial sinusitis. Acute necrotizing ^^T Jx',^ 67— KANGAROO STREPTOTHRICOSIS. LOW POWER PHOTOMICROGRAPH OFANOCARDIA COLONY WITH NECROSIS WITHIN AND AROUND IT. THIS WAS FOUND IN A SECTION FROM THE LIP OF THE SPECIMEN SHOWN IN FiG. 62. THE BLACK BORDER CONSISTS OF PARALLEL THREADS SO CLOSELY PLACED THAT THEIR SEPARATION UNDER THE CAMERA IS PRACTICALLY IMPOSSIBLE. THIS TYPE OF COLONY RESEMBLES THE "RAY" COLONY OF ACTINOMYCES THE STREPTOTHRICOSES 585 glossitis and pharyngitis. Cloudy swelling of kidney. The face seems a little full and the subcutaneous tissues slightly edematous. The naso- pharynx contains a thick tenacious mucopus. Ethmoid and frontal sinuses and turbinate spaces contain a thick purulent matter, the mucosa being densely injected, swollen and velvety. Pharyngeal wall and right half of posterior half of tongue are involved in a dull brown and necro- tizing process, quite sharply outlined by zone of congestion. This process is comparable to the necrotizing gingivitis seen in front of jaw in kangaroos. Larynx, trachea and lungs seem uninvolved save for slight generalized congestion. Cervical lymph nodes especially those about the larynx are definitely enlarged, soft, moist and brown. Medias- tinal nodes slightly enlarged, soft and pink. The heart is negative. Liver normal. Spleen is soft, homogeneous dull red. The capsule of the kidney is smooth, strips easily leaving a purple surface. The glistening section surface swells slightly, vasa recta are congested, striae wide and pale, glomeruli not visible ; consistency is resilient. The mouth and teeth are not involved in the process mentioned above. There is a small quantity of properly digesting food in the stomach. Stomach and intestines negative. Brain not involved. No extension from anterior cranial sinusitis. Smears from the mucopus confirm the gross appear- ance and contain short chains of streptococci and large diplococci. Smear from cut surface of tongue shows innumerable small bacUli and diplococci but especially mycelia with rather heavy clubbed ends but without true branching. One group was found arranged like ray fun- gus. It is noteworthy that there is no aspiration pneumonia and very slight evidences of septicemia. Biology of N. macropodidarum. The original discovery of the streptothrical forms was made in stained smears from necrotizing lesions. They were considered as secondary invaders until repeated observations of a similar character aroused the suspicion that they stood in some important relationship to the lesion. Early attempts at their cultivation were made under anaerobic precautions, a method now known to be almost certainly doomed to failure because a strain long under cultivation requires two to three weeks to make an appreciable growth in the absence of air. Finally in 1911 a successful cultivation occurred by searing the surface of an unopened mass in a freshly dead animal and planting bits of the interior upon aerobic blood serum plates. Colonies grew after three or four days and from them the first strain was started. It grew for several generations, 38 586 DISEASE IN WILD MAMMALS AND BIRDS long enough for the preparation of a vaccine, which will be described later, when by mischance it was lost. In 1920 another successful cultivation occurred, this time by incising a mass in the soft sublingual tissue and plating in the same manner; upon this culture the biology is described. Smear preparations offer no more than has already been mentioned. Colonies develop upon blood serum plates as opaque, pale yellow, circular, discrete masses with a slightly depressed uneven centre, but without umbilication. They remain smooth and slightly glistening for several days, then become slightly wrinkled and twisted with a more definitely raised edge and a tendency to an uneven sinking in of the centre. Transfers to agar slants show wrinkled continous opaque, dull yellow, sharply outlined growths which soon wrinkle, fold, and twist like certain tubercle bacillus cultures. Spreading occurs, but is slow after forty-eight hours. As medium becomes drier it is possible to see a thin, colorless, wrinkled film stretching out from the main growth. If the medium be dry or old or if only a small portion of seed material be used and this scattered over the surface of the slant, discrete colonies arise. These are circular, seldom exceeding 3 mm., dirty yellow-white, distinctly umbilicated and without clear film of spreading around them. In nearly all quite old cultures, a white chalky efiSorescence appears over the surface. The morphology of the young agar culture is chiefly mycelial or filamentous, whereas from a culture on dried media and those show- ing efflorescence, the organisms are short, heavy, deeply granular and of the mycobacterial type. Glycerine agar. — Corresponds to agar. Blood agar. — Similar to agar but much less luxuriant. Blood serum. — Limited dirty yellow, raised, dull, wrinkled and granular, tightly adherent to the medium. Potato. — Spreading, dirty yellow, much wrinkled, friable, tightly adherent. Gelatine. — Limited growth as a wrinkled, tough scum only on surface. Fig. 68.— kangaroo STREPTOTHRICOSIS. HIGHER MAGNIFICATION OF EDGE OF STREPTO- THRIX COLONY, FiG. 67. IT SHOWS THE DEEPLY STAINING MYCELIA SEPARATING MUSCLE FIBRES WHICH ARE DEGENERATING. THE STREPTOTHRICOSES 587 Litmus milk. — No change for six days, then beginning slight alka- linity which increases very little, shows digestion of the caseinogen, slight, thin filmy growth on surface. On media such as litmus lactose agar and old Endo it grows slowly on surface and assumes the color of the medium. Broth. — Only surface growth appearing during early generations, after 3-6 days as a wrinkled, pale yellow scum very much like the tubercle bacillus growth; later generations grow perceptibly in one to three days. Medium perfectly clear. If a large mass be seeded into neutral broth there is a perceptible increase in the growth after ten days. The medium thereafter tends to a faint turbidity. Titration of broth growth after twelve days shows alkalinity requiring 0.3 cc. decinormal acid, while the control tube incubated same length of time showed an acidity requiring 0.57 cc. of decinormal NaOH. On the following sugars there is a slight surface growth without change in the color, Andrade indicator — dextrose, lactose, saccharose, maltose, mannite, dextrin, galactose, salicin. Cultures observed on two per cent, neutral agar. A. — Stained by Loeffler's stain. Twenty-four hours. — Shows threads growing out from a central amorphous mass, but the whole does not retain the regularity or parallelism of actinomyces. Threads are poorly stained and rather disconnected but not jointed. Small number of metachromatic bodies apparently in older individuals, certainly in the better formed ones. No intercalary spores, unless the metachromatic bodies be so con- sidered. Individual threads measure from one-third to one micron in width. Metachromatic bodies measure on the average one micron. The threads in the forty-eight hour preparation seem distinctly wider, up to one micron and possibly become heavier toward the end, but do not have a distinct bulbous extremity. In three days the threads are much longer, show distinct branch- ing and a tendency to transverse segmentation. More than one meta- chromatic body may be present in one segment. Four days. — Still coarser, short segments have appeared separately. Metachromatic body is coarser and blacker; some of the masses have gone to pieces and show only a diffusely staining smudge of metachro- matic bodies. The short segments show a tendency to grow out into threads. Fifth day. — Condition is much the same plus many young, delicate, poorly staining threads. Sixth day.^ — The same but all seem to be somewhat wider and diffusely staining. Seventh day. — More diffuse staining and decidedly fewer meta- chromatic bodies. A. — Stained by Gram's stain. Twenty-four hours. — All forms are light purplish. The threads stain much more clearly than by Loefifler's and show distinct transverse 588 DISEASE IN WILD MAMMALS AND BIRDS segmentation of rather uniformly long bacilliform shape. Metachro- matic bodies not so distinct but seem larger Avhere found. Coarser threads have swellings iji some of the areas which are not segmented and this type seems to have more branching and metachromatic bodies; in other words it would seem that this is a form that reproduces by budding or intercalary spore formation. Forty-eight hours. — Much the same, more long threads with trans- verse division, somewhat more delicate, generally fewer coarse threads with swellings and spores. Still pale purple and not distinctly Gram- positive. Three days. — ^Condition much the same. Four daj-s. — Two forms present — definitely Gram-negative delicate slender threads, nearly Gram-positive, and heavier, curved and twisted long bacillary forms, some streptococcoid threads and a few bulbous short threads. Very few metachromatic bodies. Five days. — Condition much the same except that the delicate threads are inconspicuous and the darker purple bacilli have increased. Metachromatic bodies increased as have swellings in coarser threads. Six days. — Much the same but for the appearance of young, deli- cate definitely Gram-negative threads. There are fewer metachromatic bodies and internal spores. Seven days. — The same. B. — Grown on Loeffler's blood serum. — Loeffler's stain. Twenty-four hours. — Delicate, poorly stained short threads, few tiny metachromatic bodies. Two days. — Not well stained, relatively short threads show numer- ous metachromatic bodies varying from exceedingly tiny dots to coarse granules wider than the thread. These may be numerous in the same segment and form a row from six to ten. Many short bacillary forms. Three days. — Poorly stained, metachromatic bodies apparently more numerous but much smaller. Four days. — Almost entirely short, heavy bacillary forms, some of which are very like diphtheria bacillus in the irregularity of width; many metachromatic bodies, distinct branching, some of the small heavy ones have fusiform swellings ; practically no long, heavy threads. Five days. — Essentially the same, individual elements slightly larger, fewer but coarser metachromatic bodies, more numerous round forms suggesting large pale cocci. Six days. — Much the same but elements shorter, smaller and some more segmented. Seven days. — ^More long forms of uniform staining but still a majority of coccoid or short bacillary forms with irregular staining and metachromatic bodies; no long threads. B. — Gram's stain. Twenty-four hours. — Pale purple, almost Gram-negative, long, slen- der but well outlined threads, a few coccoid forms, practically no granules. -KANGAROO STREPTOTHRICOSIS. PHOTOMICROGRAPH SHOWING THE SEPARATE THREADS OF NOCARDIA IN A SOFT NECROTIC LESION. THE STREPTOTHRICOSES 589 Two days. — Very pale, almost Gram-negative threads, very many eoccoid forms and short rods, considerable segmentation of the longer threads. Three days. — Increase in short, heavy bacillaiy forms with bulbous ends, deeply stained ones and the granules being lightly Gram-positive ; long, slender threads are disappearing. Four days. — Almost exclusively short, heavy forms with bulbous ends with eoccoid forms, heavier forms almost definitely Gram-positive, granules Gram-positive. Five days. — Much the same but more segmentation in the bacillary forms, eoccoid forms become more numerous. Six days. — Individuals are somewhat longer but there are many rods with fusiform swellings containing granules; eoccoid forms pres- ent in chains sometimes. Seven days. — More long rods or short threads, pure eoccoid and bacillary forms. The morphology upon bouillon depends somewhat on age and upon the location. Upon the surface the long branching mycelial type appears early and persists until the whole surface is covered whereupon the segments divide into eoccoid elements with metachromatic bodies. If heaping-up develop the coarse grains on the mass consist of granular or eoccoid rods. When growing in the depth the eoccoid form is the predominant one, only a few delicate mycelia, usually Gram-negative, being found. The Gram character of the organism should be emphasized. The young, delicate mycelia are negative or take a very feeble blue stain. The hea\^ bacillary forms are Gram-positive. Like the ray fungus the heavy ends are sharply Gram-positive, but unlike it, there has never been seen a Gram-negative bulbous capsule around this end. The determination of this organism was undertaken from the classifications of Petruschky (Kolle-Wasser- mann), of Castellani in Castellani and Chalmers' Tropical Medicine, and of the Society of American Bac- teriologists. In the first classification it corresponds in some ways with Streptothrix Jwminis, and in some ways with Streptothrix caprce. As for the second authority it falls into the Nocardiacese, section parasitica, subsec- tion I, in that a distinct earthy odor is absent and that there is no liquefaction of coagulated protein. It resembles several of the species given in this subsection, but does not correspond exactly with any of them. Con- 590 DISEASE IN WILD MAMMALS AND BIRDS sultation of the classification of the American Bac- teriologists would place it among Mycobacteriaceae. The facts that it is strongly aerobic, produces whitish efflo- rescence which may possibly be aerial hyphae and breaks up into short segments, place it in the genus Nocardia. It seems, however, to belong to a division of Nocardia which is close to the Mycobacterium since the short ele- ments are swollen, cuneate and usually heavy, wliich is imusual in the more typical Nocardia. It is not, however, acid fast and therefore cannot be classified among the Mycobacteria. This culture seems to be a variety not heretofore described, and since its association with the disease is so definite, whether or not it be the cause, the name Nocardia MACROPODroAEUM is proposed, because the kangaroos belong to the order Marsupialia, family Macropodidae. The discovery of these organisms within tissues is by no means easy even though the larger colonies may be located by staining. If Loeffler's method be used the central mass stains quite diffusely and the spreading mycelia around the edge stain faintly. For study pur- poses this stain is preferable to Gram-Weigert, since despite the positivity of the cultures, the blue dye can be removed very easily from sections and only with great care will enough remain to permit tracing of the separate threads ; with Gram stain no detail can be made out in the centre of the colony, it being a diffuse blue. Careful search near the edge of these necroses will usually succeed in the discovery of a few mycelia stretching in between the mono- and polynuclears of the low grade inflammation. This is best seen in the margin of gastric ulcers, but may also be found in the cervical masses. When searching in the pulmonary tissues the organisms are to be found in the bronchial exudate or at the edge of pneumonias. In one nasal mucosa the mycelia were dispersed, not gromng in colonies as in localized inflammations. THE STREPTOTHRICOSES 591 Experiments at the Repeoduction of the Disease. When the first culture was isolated it was injected into gTiinea-pigs ; its loss stopped further work because it could not be regained from the animals. The present culture had been injected into g-uinea-pigs, rabbits, opos- sums — all with negative results; such an experience is not unknown for actinomyces. Intraperitoneal, intra- venous methods having failed, inoculation was made into the gums of rabbits and of opossums with no result, even after traumatizing the mucous membrane. The injection of about 5. mg. of a twenty-four-hour agar culture was made directly into the masseter muscle of an opossum without producing even a lump at the site. Atomizing a culture into the nose and throat of an opossum seemed also without effect. Injection of cultures into the nose, gums and labial tissues of a wallaby have been negative ; nor has any perceptible effect followed the atomizing of a heavy nocardial suspension in broth into the trachea of this animal. The results of these experiments are in accord with those of many similar attempts to reproduce actinomy- cosis. Perhaps in Kangaroo disease the small Gram- negative bacillus is a necessary factor. Specific Prevention and Treatment. Encouragement that we were upon the right track was, however, found in another direction. Improvement in human and bovine actinomycosis having followed the use of vaccines, it occurred to me to try this method as treat- ment and prophylaxis. The first culture to be isolated was just at hand, so that it could be used at once. Five injections were given under the skin of the thigh to a recently developed case of the ulcerative gingival variety, a noticeable improvement occurring almost at once, and at death there was an apparent cure of the local lesion. However, the accompanying protocol made at the time 592 DISEASE IN WILD MAMMALS AND BIRDS tells the whole story, no adequate explanation being at hand. Red Kangaroo [Macropus rufus). Disease of the mouth first noticed March 31, 1912, died September 13, 1912. Necrotizing osteitis, arthritis and periarthritis of left ankle, subacute fibrinous right pleuri- tis, hemorrhagic bronchitis with atelectasis in right middle lobe, abscess of right middle lobe; passive congestion of lungs, liver, kidney, chronic splenitis, chronic general lymphadenitis. The animal is in general good condition except for a fusiform swelling about the left heel with evidence of fracture. The necrotic process in the hare lip, nose and palate has entirely disappeared. One front incisor has gone and the other is loose. There is a scar on the under part of the soft palate in a small healed channel between palate and floor of nose. There is no evidence of pyorrhoea. Cervical and axillary lymph nodes are much enlarged, pale yellow, firm and of the appearance like early stages of Hodgkin's disease. Faseias of cavities congested. The lungs are mottled purple, air content decreased, section surface purple, exuding frothy blood. The whole right lung is covered with a thick fibrinous exudate, most intense over middle lobe at site of atelectasis. There are light scattered adhesions. The anterior margin of the lung is adherent to the pericardium which is covered in the front by exudate. Upper and lower lobes show hypostatic congestion. Middle lobe has separate bronchus filled Avith necrotizing blood clot extending into a smaller bronchus with complete occlusion. The alveoli supplied by the last show atelectasis like hemorrhagic infarct. There is a small sub- pleural abscess near the margin of this atelectatic area. The bronchial lymph nodes are slightly enlarged, mottled yellow and pink, firm with large, diffuse follicles. The pericardium contains 2-3 cc. clear fluid. The heart muscle is pale, purple and soft. All the vessels are full of currant jelly clot. On the posterior surface of the aorta internally about an inch above the valves there is a patch of roughening with a sugges- tion of thickening and opacity. It is comparable to the early stages of syphilitic aortitis. The liver is noiTual in size, surface smooth, edges sharp, consistency firm and friable, color purple. The section surface is glistening, smooth, moist, and shows passive congestion. The gall- bladder contains fluid brown bile and the common duct is patulous. The spleen is slightly enlarged, firm and tough, capsule Avrinkled. Sec- tion surface is mottled red and purple with irregular gray trabeculoe and faint scattered follicles Avith diffuse margins. The kidney capsule is smooth, strips easily leaving a smooth brown surface. Organ is firm. The section surface is glistening with a line of passive congestion with distended vessels between the cortex and medulla which are of noi-mal -n-idths. Intestines seem normal throughout. The pancreas is firm, pale pink, slightly edematous. The mesenteric lymph glands are moderately enlarged, yellow, firm, homogeneous Avith congested centres. About the left ankle joint there is a necrotizing infection Avhieh has THE STREPTOTHRICOSES 593 involved the bone causing a pathological fracture of the lower end of the tibia. Smears from the periarthritis, pleuritis and blood clot in the bronchus show streptothrix, a short colon-like rod and a coccus in fours — a picture precisely like that obtained from the jaw bone cases. In addition to the above there is a very distinct encapsulated pneurao- coecus form in smears from the blood clot in the bronchus. This is the animal which was vaccinated with a culture made from the depths of a necrotic mass, upon which treatment she rapidly improved and as seen from the above notes recovered from the palate condition. Why she should have a second infection apparently with the same organism is difficult to determine. Possibly the second batch of vaccine was not sterile, it not having been controlled because the fii'st batch of vaccine was sterile after one hour at 60° C, Possibly the animal was sensitized and a few bacteria settled in the leg. It was along this leg that the inoculations were made. We permit ourselves the facetious observation that that vaccine prevented the labial and cervical variety for five years, because during that period it stood in the ice- box, and there was no case of that particular form to which to give it, although a few of the nasal and gastric varieties occurred. It was recontrolled and did not show living organisms. That it should have cured the disease in the jaw and apparently later permitted a lighting up of a septicemic and pulmonary form with necroses in the leg is difficult to explain. Just recently we have used a vaccine from the current culture upon another case beginning in the gums and jaw bones. This case was detected early and was treated with ascending doses beginning at 0.3 mg. and running up to 10. mg. At first there was some improvement, but the animal finally died from pulmonary complications. The course of the disease, however, instead of being three weeks, as is the customary duration, lasted two months, an extension of the course which has made us hopeful. These two experiments, indefinite though they be, have offered encouragement and seem to supply a little addi- tional support to the idea that the organisms stand in etiological relationship to the disease. The employment of the vaccine has been extended to its use as a prophylactic in animals exposed to the disease 594 DISEASE IN WILD MAMMALS AND BIRDS or specimens that have slight reddenings or erosions on the buccal mucosae suggesting possible early stages of streptothricosis. Five animals have now had a course of vaccine injections, ranging in number from 5 to 10 and in quantity from 0.3 to 2.4 mg. over a period of a month. Fourteen months have elapsed at the time of writing and only one case has developed, but this of course cannot settle the efficacy of the method ; perhaps it would be safer to demand that no case should ever appear in a treated animal, while the disease did appear in the untreated. The preparation of the vaccine is by no means a simple matter, since the surface growth upon solid media is so tenacious. Methods such as are employed for the tubercle bacillus have to be used. The first two vaccines were made by scraping off surface colonies from agar and grinding with glass balls. One successful batch was made recently by simply triturating the colony directly on the agar slant, but the latest method seems to offer the simplest and most generally satisfactory way. Neutral broth is placed in flasks containing glass beads and sterilized in the incubator. This is seeded mth the Nocardia, incubated at 37° G. until the surface is covered, heated to 60° C. in a steam sterilizer and tested for sterility. If growth occur it is reheated until dead, whereupon the broth is syphoned off, the growth emulsi- fied by whirling the flask, thus grinding the bacterial mass by the glass beads. Sufficient saline is added to make a workable emulsion, and the fluid then poured into bottles. Control by reculturing is again done, and if the fluid be found sterile, 0.5 per cent, trikresol is added to keep it so. These organisms cannot be counted accurately because of the variation in length, their budding and coccoid forms. Standardization is done by weight. A definite equal quantity of the suspension and of the saline used to make it are evaporated to dryness in weighed vessels and the whole then weighed. The difference is the weight of the organisms suspended in the saline. Such a fluid can THE STREPTOTHRICOSES 595 be diluted so that a given bulk will contain a convenient weight of germs. The one now in use contains 8. mg. per cubic centimetre. Dosage as indicated above usually begins at 0.5 mg., a quantity which does not produce any local inflammation at the site of injection. It is perhaps well to adopt a quantity of 0.1 mg. per kilo as the initial quantity. The Garden has encountered no case of the remaining important chronic infections, glanders, lymphangitis, and infectious abortion. SECTION XVII— PART 4 ACUTE DISEASES RESEMBLING THE SPECIFIC INFECTIONS OF DOMESTIC ANIMALS Specific communicable diseases are sometimes divided into those most often encountered as ''herd diseases" and those which appear as single cases or in small groups. This would seem to imply that the first behave as easily disseminated epizootics, their \drus passing from animal to animal simply by proximity or by casual contact where- as the transfer of infective material is less readily ac- complished by the second group, often demanding special assistance. Foot-and-mouth disease, pleural pneumonia, cattle plague, and influenza illustrate the epizootics while tetanus, rabies, quarter-ill, malignant edema, and infec- tious vaginitis are examples of less easily transferred processes. It is not intended that these remarks shall cover all possible means of transmission but instead they are in- tended to focus attention upon the sources of viruses whereby animals become infected. An original case must always be present in order for spread to occur. Where animals are being added to a herd a new comer may be diseased or the carrier of a virus; when animals are transported for sale or other reason, infection may be met in a new stall, conveyance or pasture ; contaminated food may be offered. In menageries, with specimens, single or in small groups, and arrivals always quarantined before other animals are exposed, acute specific infections seldom appear. It is also improbable that a wild animal, infected at its source or in some dealer's place, would survive the journey and arrive in an infective condition. Consultation with the reports of other gardens fails to discover records of any serious outbreaks of epizootic disease except for fowl cholera and distemper, examples 596 MISCELLANEOUS INFECTIONS 597 of infection with the bipolar organisms of the Pasteurella group, believed responsible for the hemorrhagic septice- mias ; instances of the occurrence of the group specified secondly — anthrax and the like — are also reported. This represents fairly well our own experience. The bacteria variously named Bac. avisepticus, ovisep- ticus, hovisepticus, canisepticus, etc., grouped by Ligniere under the name Pasteurella, are doubtless of considerable importance and are probably quite widespread in natural surroundings. The viruses of the epizootic conditions like cattle plague and influenza are apparently more definitely parasitic, requiring for their persistence ever renewed transfer from host to host. The former infec- tions we have met in repeated single isolated cases and in small groups, whereas no cases of the specific epizootics have been diagiiosed. Hemorrhagic septicemia, a denomination very descrip- tive of its pathological picture, has been encountered in many varieties, carnivores, ungulates, primates, rodents, and birds. The diagnosis depends upon the presence of hemorrhages with edema, degenerations of the parenchy- matous organs, more or less respiratory catarrh to which may be added relatively mild gastrointestinal inflamma- tion; the bacteria are found in the circulating blood and in exudates. A description of these organisms is not profitable, they being well known in veterinary pathology. "What is more important, significant and supportive of the opinion expressed above concerning the mdespread distribution of the virus, is the incidence of the infection. Exclusive of the condition known as fowl cholera, it has appeared among mammals and birds as single cases with one exception — that of two Barbary apes which had been in separate cages side by side. The total of cases with determined bacteriology is eleven, with undecided bacte- riology but suggestive pathology nine additional. No per- tinent history in common can be found in the records of the determined cases, except perhaps that they were 598 DISEASE IN WILD MAMMALS AND BIRDS all animals which had been in the collection at least three months, a period which would seem to exclude the proba- bility of an imported infection. Because of the isolated character of the cases and impossibility of making a clini- cal diagnosis, no attempt at specific nomenclature as used in veterinary medicine has been made, hemorrhagic sep- ticemia seeming to cover its identity and nature. The disease knowm as fowl cholera is practically al- ways associated ^dth the bacteriological discovery of a member of the hemorrhagic septicemia group while its pathology corresponds with that of mammalian infection with these germs. Enteritis is a prominent feature. This disease has appeared thrice among our parrots carrying off from six to ten birds before hygienic measures became effective. ) In all three our cultures showed the bipolar organisms. Besides these specific outbreaks numerous isolated cases of acute general infection have occurred among small passerine and picarian birds which could not be determined as hemorrhagic septicemia by bacte- riological methods although superficially resembling it in gross pathology ; they yielded to the same hygienic meas- ures. Perhaps we were dealing with fowl plague, a disease believed to be due to a filterable virus. That this is the case is strongly suggested by an outbreak of fowl typhoid in the parrots, from some fatal cases of which we were able to isolate B. sanguinarium, and by a group of deaths in small parrots from which no specific organism could be recovered. The identification of these supposedly specific diseases — plague, typhoid, septicemia, leucemia — by pathological criteria is by no means simple even if we have at hand the complete description of Moore, of Hutyra and Marek, of EUermann and of Ward and Gallagher, Bacteriology must decide and cultures should be made upon bodies recently dead. In addition to the above infections we have had two small outbreaks of psittacosis in parrots from which it was possible to isolate the specific organism. On MISCELLANEOUS INFECTIONS 599 both occasions there was more than one death before the specific nature of the disease was identified yet, note- worthily, no spread to the other birds in the same exhibi- tion house occurred. Distemper, a disease variously held as due to cocci, to influenza-like organisms and to a filterable virus, may ap- pear in sporadic or epizootic form. The diagnosis during life is not so easy unless all the cardinal features are present, while after death the same thing holds good. I am inclined to think that from the standpoint of diagnos- tic accuracy, the term is used much too loosely, a ready excuse for such laxity however being that it stimulates to greater care in hygiene. Whether or not B. bronchi- or canisepticus be the cause of the disease, organisms cor- responding to it can be found in stained smears from nearly every case in which the respiratory, cutaneous, nervous and internal signs suggest the disease. To make a diagnosis of distemper it is my practice to require at least three of the cardinal clinicopathological features, whereupon, if the bacterial findings be as described, the denomination is permitted. This was dictated because during the period, now happily well in the past, when the cats and dogs suffered frequently with enteritis, naso- pharyngeal signs occasionally presented themselves or spasms were reported, but no skin eruptions appeared, yet seldom were all of these signs combined nor could we find the bipolar organisms. I note that in 1915 Doctor Blair of New York observed a toxic enteritis resembling but not identical with distemper. As with our cases he failed to find that the condition was communicable. We ascribed our cases to spoiled food — fowl heads or dirty horse meat (see page 179). Our acceptable examples of distemper number three, two ferrets and a lynx, but very suggestive cases were found in foxes, wolves and raccoons. Since writing the above notes, sixteen wolves, foxes and wild dogs died in an outbreak of distemper imported by a newly arrived specimen admitted to the 600 DISEASE IN WILD MAMMALS AND BIRDS colony by mistake. When we were aware that the disease had appeared antiserum was administered therapeutically to all that were sick and prophylactically to all the rest — large doses, 25-35 cc, were given for treatment, smaller quantities, 10-20 cc, being used as a preventive. Seven sick animals recovered and no animal (8) given serum prophylactically became sick. This experience en- courages us to think that with antiserum and rigidly enforced quarantine rules, distemper will not be a serious matter to handle. The hygiene of the foregoing conditions is of a general character — removal of the specimens when kno^\^l to be sick, thorough cleansing of the cages, segregation of mates or of neighbors when this is practicable, burning of refuse, liming of the gromid and such other measures as the local conditions may indicate. Diphtheria. Although no cases of mammalian diphtheria have been observed, three and possibly four birds have suffered with tliis disease. The three acceptable cases were in cassowaries {Casuarius occipitalis) occupying adjoining cages and sickening within a few weeks of one another. Just how the infection was brought to them must remain a mystery since no additions had been made to the group for some time previously. All three birds were observed during life, and from the first case the Bac. diphtheria avium was isolated ; in smears from the other two simi- lar bacteria were seen but isolation was unsuccessful. The two acutely fatal cases showed large pseudomembran- ous collections on the nasopharyngeal mucosa and beneath the tongue while the nares were occluded by the same material. Plaques of membrane were also found on the surface of the esophagus and proventricle. The exudate ran out of the mouth and formed dried crusts upon the cervical skin. Pseudomembranes of a continuous charac- ter were lacking in the third bird, their place being taken MISCELLANEOUS INFECTIONS 601 by small yellow or yellow-piuk nodular elevations, appar- ently just beneath the surface, here and there upon the reddened, slimy buccal, lingual and pharyngeal mucosae. Crusts upon the skin of the neck also formed in this case. ) These cases are of interest not only because of their appearance without satisfactory explanation but because one improved very much after injections of human diph- theria antitoxin, this remedy being used because we were then unaware of the existence of an avian diphtheria anti- toxin. No claim can be made post hoc ergo propter hoc that the human antitoxin helped the attack — it may have been mild — but th^ experience is worth recording. Dosage was as follows: '.December 3, 3,000 units; December 8, 1,500 units ; December 21, 5,000 units ; December 27, 5,000 units. Shortly after the inception of the treatment the bird was noticed to eat better and to be more lively ; this was followed by a reduction in the mucous strings in the mouth and the crusts upon the skin. This improvement con- tinued and the bird seemed well in about two months but, after the lapse of three months more, a mucous naso- pharyngitis was again observed. Despite two injections of 5,000 units human diphtheria antitoxin the bird suc- cumbed five days after the beginning of this attack. Autopsy revealed much the same condition as was found in the first birds and from the larynx the Bac. avium was isolated. Another case suggestive of diphtheria was seen in a hornbill but antemortem observation being imprac- ticable and postmortem decomposition being advanced when autopsy was performed, the diagTiosis could not be confirmed. ;■ (An unusually well developed case of molluscum con- tagiosum was seen in the Wild Turkey (Meleagris gallo- pavo) recorded here by photograph and in the form of notes upon the histology made by Doctor Weidman. The bird's head was affected universally from beak to ears by horny nodules up to the size of a pea. They were so large and numerous around the eyes as to com- 39 602 DISEASE IN WILD MAMMALS AND BIRDS pletely close them. There were no lesions elsewhere on the body, none of the other turkeys were similarly affected and though watched, none have since developed a similar condition. Histological examination shows a keratosis, many of the cells shomng characteristic *' molluscum bodies " which appear the same and behave the same tinctorially as the human examples. Tliis turkey case differs from the human, however, in that there are none of the pocket-like epithelial extensions deep down into the corium and this turkey case may be very useful in the further study wliich is contemplated to show that such things as molluscum bodies are not sufficient of them- selves to stamp a dermatosis as a pathological entity, but that they are general pathological processes wliich may occur in a number of different diseases. The disease has been reported in sparrows, pigeons, but never so far as I can find, in turkeys. ) A few isolated cases of infectious disease are included here as a matter of record although they may not be espe- cially significant or important. Rabies was found in a pair of deer which had been bitten by a stray dog. The period of excitement was relatively long, while the para- lytic stage was only a few hours. Negri bodies were found. Tetanus killed a Persian Wild Ass {Equus onager) the infection wound seeming to be a bruised and abraded area on the rump. From the contused muscle tetanus bacilli were isolated. A gas-bacillus infection, emanating from the vagina which was protuberant and lacerated because of injury by mates, was seen in a preg- nant llama (Llama lama.) On two occasions nodular masses have been found under the skin of seals, not unlike the one studied by Doctor Wiedman and thought by him to be due to moulds. These two have, however, failed to show mj^celia or yeast-like bodies, and one thinks only of placing them in the group of botryomycosis. I have never seen a case of this disease, so that I am forced to rely upon literature, a method that inspires no especial confidence MISCELLANEOUS INFECTIONS 603 in the diagnosis. The bacteria usually held responsible for botryomycosis could not be isolated. Just what can be done for the condition is difficult to state, since seals are scarcely tractable animals. The following case has some features like paralytic hemoglobinuric fever and is reported as a matter of record. The long standing gastroenteritis may have been the basis for the intoxication which led to the paralysis and muscular degeneration. This laboratory has now under way studies upon the laming of ungulates, accom- panied by weakness of the hind-quarters, but no con- clusions have been reached. It is interesting to note that Hutyra and Marek quote Johne as having seen a case of hemoglobinuric paralysis in a zebra in 1879. Burchell's Zebra {Equus burchelli burchelli). The only symptom observed in this animal was gradually increasing lassitude which was first noticed about three months ago; toward the end he habitually stood with tucked tail and nose to the ground as if asleep. He ate well and digestion appeared good, but he became very weak as shown by his inability to rise when he got down on the third and second day before he died, although on both occasions he was able to stand when lifted. Injury, hemorrhage in thigh muscles, chronic gastritis, sciatic neuritis. CEstrus larva in stomach, ascaris in intestine. Both lungs are widely distended and the caudal half of both is the seat of passive congestion. Upper lobes are slightly edematous. No consolidations. Heart normal. Abdomen contains about two quarts of clear straw colored fluid. Liver is of normal size, smooth surface, sharp edges, firm, friable. On section it is very bloody, veins distended, some with clot. Architecture normal. Spleen is of normal size, soft, tough, capsule rough. Section surface is homogeneous, pulp purple, trabeculae normal, follicles not visible. The kidney capsule is smooth, strips easily leaving a smooth brown surface, firm. StriiE nonnal, rather wide, glomeruli not visible. Stomach is (filled but not distended -with partly digested straw. Mucosa of cardia dry, roughly irregular, some irregular mammillations. Two flat papil- lary growths. (Estrus larva attached to a smaller elevation. The mucous membrane of the fundus is soft, moist, irregular, in some places, translucent, in others opaque ; near pylorus mucous membrane is swollen edematous, pink, slightly eroded at pyloric valve. Small intestine has smooth, flat, pale yellow translucent mucosa. Lumen filled with muco- purulent matter like mixed egg. Ileum slightly congested but mucosa firm and translucent. Pancreas is soft, slightly uniformly congested. All mesenteric lymph glands are slightly enlarged and edematous but with normal architecture. In the posterior thigh muscles beside the 604 DISEASE IN WILD MAMMALS AND BIRDS sciatic nerve, most marked on the right side, is a large hemorrhagic infiltration. There is edema of muscles and intermuscular septa all about this area extending upward as Avell as to pelvis and psoas muscle. This latter within the abdomen shows slight blood stained edema. No other muscle shows this hemorrhage. Microscopic section of liver and kidney are negative aside from congestion. The stomach shows very irregular epithelial covering, in some places wholly desquamated. Where this is most marked there is a dense round cell infiltration in the villi with some increase in the connective tissue cells. This chronic inflamma- tory reaction is present in all fields, most marked, of course, in upper layers of mucosa. Glands are distorted and upper epithelium of them is polychromatophilic. The intestine shows similar changes in less in- tense manner. Waterfowl Epizootic. There is reproduced here an account of an unexplained epizootic among ducks and geese from the Annual Report of the Zoological Society for 1916. Nothing additional has been learned and no repetition has occurred since the drainage and cleaning of the lake. There began on August 27 a series of deaths among the waterfowl and in one month there were lost forty-one specimens including both ducks and geese. Four additional cases were scattered through the next four months, the last case dying January 11, 1916. All of these came from the lake, none being from the adjacent stream for rare waterfowl or from the more distant stream into Avhich the lake drains. The sj^mp- toras were most marked and striking. In the early stages the wings drooped, then the legs became weak followed by inability to raise the head. In the latest cases the voice (ducks) lost its normal character and became hissing. The mind appeared clear for the eyes were bright, feathers unruffied and the bird attempted to escape when approached. Diarrhoea was present, dejecta thin, watery white, no admixture of mucus. Autopsy findings were not frank. At most some swelling of the spleen and a little pale thickening of the intestinal wall constituted the picture. Smears from intestine and nasal mucosa showed no protozoa. The blood taken from the living sick ducks showed no parasites or anemic changes in either raw or variously stained preparations. From the spinal cords of three ducks a 50 per cent, glycerine emulsion Avas pre- pared and Avas injected into the cerebral substance and abdomen of domestic ducks with negative results. A variety of different bacterial cultures was obtained from the liver, spleen, blood and congested nasal mucosa of several birds dead with the disease and injected into domestic ducks with negative results. Histological sections were cut from the important organs of thirteen birds. The kidneys, lungs and pancreas shoAved no abnormalities. The heart muscle in some cases and also MISCELLANEOUS INFECTIONS 605 some of the skeletal muscles showed Zenker's hyaline degeneration together with minor hemorrhages and edema. Several of the proven- trieles showed low grade inflammatory signs toward the gizzard. The intestines regularly showed lymphatic infiltrations of the villi most marked toward the tips but without congestion. The lumen showed no parasites, bacteria or protozoa. Liver showed in almost every case pigmentation by hemosiderin at times as heavy as that seen in per- nicious anemia. The finer bile ducts here showed peripheral round cell infiltrate, which was not continued into the major ducts as determined by serial sections. Parenchymal cells were cloudy and swollen. Spleen showed in early cases polymorphonuclear infiltrate of the follicles, in later cases atrophy of follicular splenoeytes and more or less pigment occun-ed in both stages. The spinal cord and various peripheral nerves showed no inflammation or degeneration as determined by the appro- priate special nerve stains. The above clinical, histological, protozoologi- cal, and bacteriological examinations having failed to detect the cause and the epizootic now being over, its nature becomes a matter of deduction. The only constant features of any importance were the paralysis, the intestinal round cell infiltrate and thickening, the pigmen- tation of the liver and degeneration of skeletal muscles. Of the various possibilities, beriberi was early considered. This is not possible because the food of the birds was a varied one and furthermore none of the neiwe degenerations of beriberi were noted. Second, acute bacterial or protozoal infections are unlikely because no constant primary lesions were discovered at autopsy, the numerous cultures failed to produce the disease and other birds living on the stream draining the lake were not similarly affected. Third, a food poisoning. This is possible first because paralytic symptoms were present such as are seen in vetch and mussel- poisoning and secondly because the epizootic ceased when the birds were taken from the lake and placed upon the grass. If this be the case the toxic material produced the paralysis by direct action upon the muscle fibres just as that of typhoid fever does and must have caused hemolysis as shown by the hepatic pigmentation. The source of this food poison- ing is conjectural. Perhaps a dead fish decomposed in the water or there were some algae with poisonous properties present. The outbi-eak has a resemblance, but only a superficial one, to infection with one of the group of botulism bacilli. The cause of the trouble must be con- sidered as undetermined. Enterohepatic Disease. Since the normal drainage from the intestinal tract passes so largely through the liver, there is little to wonder at in morbid lesions of the latter organ consequent upon disease in the former. Not only does this succeed upon bacterial infection of the digestive tube but also upon infestation with animal parasites, under the latter condition forming changes of 606 DISEASE IN WILD MAMMALS AND BIRDS much more considerable extent, at least in gross bulk, than in the former. Changes in the liver secondary to enteric disease from bacterial infection take the form of cholangitis, thrombosis, degenerations and probably cirrhosis while abscesses and necroses succeed upon pro- tozoal or metazoal parasitic involvement. JThe latter is exemplified by amebic abscess in man and other mam- mals and by ''blackhead" and ''quail disease" in birds; it is to the latter conditions that attention is now directed. The chapter upon the cause of these diseases has yet to be completed, although many reams have been written about it, while the transmission is fairly well understood and the pathology well described. My purpose here is to discuss our experience with the two above mentioned diseases which, while far from conclusive, may assist somewhat in explaining their etiology. There is also reproduced our original report upon quail disease from the Society's Report of 1915, giving data and figures. ( Blacldiead has been found in five wild turkeys. An unusual case in a Berwick's Swan is recorded since it bears a striking resemblance to the disease. The points at issue in the determination of the etiology of blackhead are the importance of Heterakis papillosa in the ceca and the frequency and activity of ameba or histomonas. ' In three of the five cases of the disease in turkeys the nematode was found macroscopically in the ceca, in two it was not; in one its absence was confirmed microscopically. In two of the turkey cases, forms corresponding to the ameba or histomonas were discovered while the descriptions of the he- patic lesions in two birds use the term coccidia which, from a revision of the slides, is probably incorrect although some of the parasites seem to be possessed of a doubly contoured refractile margin. The larger, more diffuse and ameba-like forms in the intestinal wall sug- gest that the hepatic inclusions belong to the same group. In only one case was exhaustive search made for coccidia, MISCELLANEOUS INFECTIONS 607 and without success ; the material was not preserved. In two turkeys entirely free of lesions distinctive of black- head, cecal nematodes (one heterakis, one unknown) are recorded, and in the intestinal wall of another, also free from the disease, forms indistinguishable from ameba could be discovered. (1) The protocol of the Berwick's Swan is interesting because the full fledged disease is not known in this bird. While this case is not by any means typical, the chronic cecitis and ameba-bearing necroses in the liver stamp it as of a kind with the true infection of turkeys. Perhaps the resistance offered by the swan effected a modification of the disease, preventing the usual necrotizing enteritis and turning it into a chronic interstitial variety. Berwick's Swan {Cygnus herwicki). About a month before death passed several large clots of blood. Acute catarrhal enteritis, mural endocarditis, chronic colitis, chronic nephritis, passive congestion and necroses in liver, acute follicular splenitis, edema of lungs, chronic peri- carditis, chronic salpingitis, hydrothorax, hydropericardium, hydro- peritoneum. Tissues generally are slightly yellow. In serous cavities of thorax is about three ounces of clear fluid. Lungs are distended, subcrepitant, pale red and gray, highly edematous. The pericardium contains about one-half ounce of clear watery fluid. Epicardium is glistening, congested, irregularly thickened especially near the blood vessels. The heart is contracted, slightly large, pale brown-red muscle. On the posterior surface of the right ventricle extending from the auric- ular opening to the pulmonaiy valve is an irregularly curved line of grouped, recent red vegetative granulations. Valves negative, they and chambers competent. Aorta negative except heavily blood stained. Liver is slightly large. What of the liver remains undamaged is homo- geneous deep purple. Major portion of right lobe badly contused ; this seems to have been partly antemortem because there is blood staining and mottling under capsule. In view of colon finding and history of possible injury it is probably the result of degenerations in the liver plus slight trauma. There are several small, pale gray, well outlined, homo- geneous areas probably necroses in the liver. The spleen is slightly large, soft, egg-shape, capsule smooth. Section surface shows bright red homogeneous pulp with clearly cut, large follicles. The kidney capsule is smooth, surface smooth brown, consistency firm and tough. The section surface gives a dull gray-brown appearance, seemingly from ( 1 ) Those interested in the investigation of the cause of Blackhead are referred to the recent literature by Tyzzer and by Smith, in the Jour, of Exp. Med. and Jour, of Med. Research, 1918-1922. 608 DISEASE IN WILD MAMMALS AND BIRDS connective tissue. Markings indistinct. Oviduct is negative except over a distance of an inch near the cloacal opening. Here there is a com- pound curve with constriction to almost obliteration of lumen. This does not seem to be connected with the colonic trouble. The stomach is negative containing only a few small pebbles. Beginning at the pylorus and extending through the whole of the small gut is a recent, moderately severe catarrhal enteritis with so much exudate as to form almost a cast of the tube. Colon and cloaca show an infiltration of submucosa with areas of hemorrhage. Mucosa swollen as if by edema, glistening and covered by bloody mucus. Ceca negative except that they seem to have been closed as their contents are scanty and firm. Histological section of cloaca shows it to be the seat of a chronic inflammation which has constricted and distorted the tubules into simple masses of nuclei. Marked polynuclear and round cell infiltration of mucosa and submucosa. This is apparently due to ameba-like bodies — a large vacuole with a delicate limiting membrane and a piece of diffuse chromatin in the centre — a few of which may be found deep in the mucosa. Liver shows marked passive congestion, here and there areas of necrosis with some fatty infiltration. Small groups of ameba-like bodies can be found apparently lying in sinusoids of liver and in neighborhood of necroses. Quail disease, since the careful work of Morse in 1907, has been thought by most observers to be due to an organ- ism of the colon group, but I am informed recently by the Pennsylvania State Board of Animal Industry that coccidia have been found often enough in the droppings and in the morbid lesions to warrant a suspicion of their etiological importance. Although they were not espe- cially sought in the work about to be reported, their pres- ence probably would not have escaped detection during that investigation. I have recently had occasion to examine three birds with lesions identical with those accepted as characteristic of quail disease, one of which was subjected to the proverbial ''fine tooth comb" methods ; no coccidia were found in the liver or intesti- nal lesions. The idea that quail disease, with its ulcerative typh- litis and necrotizing hepatitis, is identical with blackhead or at least that if the latter be due to protozoa, the former is also, requires no special stretch of imagination to one famiUar with the morbid lesions. A decision is the more difficult because of one's inability to reproduce quail MISCELLANEOUS INFECTIONS 609 disease at will and the none too great certainty of the intentional production of blackhead. At all events the transmission is potentially the same, gromid or food soiled with droppings, indicating that hygienic measures should take the form of segregation and disinfection. Here follows the report of our original observation : '^ An epizootic disease has decimated three newly im- ported lots of quail, Scaled quail (Callipepla squamata), GambePs quail {Lophortyx gamheli) and Texas bobwhite {Colinus texasus virgimanus). On January 5, 1915, the first lot of twenty-four quail arrived from northern New Mexico via Kansas City; on Januaryllth a second lot of twelve bobwhite arrived from Brownsville, Texas, via Kansas City ; the first of this lot died the day after arrival with lesions of this infection. From this lot of birds the first lot was probably infected, the first death occurring on January 20th, no other deaths having occurred in the first lot since arrival. On January 21st the third lot of twelve quail arrived direct from Mexico. The first of this lot died of the disease on January 24th. Some birds were also sent at the time of the arrival of the third consign- ment, to Doctor Kalbfus of the State Game Commission. It is to be emphasized that to date no cases of infectious enteritis have occurred in the lot sent to Doctor Kalbfus. The first case appeared at this Garden on January 12th, more than a week before the third lot arrived. It would seem that the disease was brought to the Garden by the second lot of birds, and that they picked it up on the way from Texas to Kansas City to Philadelphia. The birds made a stop at Kansas City. The birds died at long intervals for the first two weeks, but late in January and early in February several died each day. The last death with characteristic lesions occurred February 11th. After the epidemic reached its height it subsided very quickly. ''During the illness the birds exhibited very few symp- toms, indeed some of them were not known to be sick. A few sat huddled in a corner mth ruffled feathers and 610 DISEASE IN WILD MAMMALS AND BIRDS drooping head; the stools were little if any altered as far as could be determined among so many in the enclosure. At death the birds were in good condition, feathers fairly smooth, skin clear, body plump and fat in good amount — not abundant, nor were the animals emaciated. The principal lesions were enteritis, degenerative necroses and abscesses in the liver, congestion of all the viscera and plastic peritonitis in a few. A small number showed congestion of the Imigs and two had patches of pneu- monia. Many but not all of the birds had Heterakis in the ceca. The process seemed to start as a focal necrotizing lesion in the mucosa or submucosa of the ileum just above the ceca and colon ; many had lesions in the ceca and as far down in the colon as the cloacal dilatation. Among the animals dying late in the epidemic several showed lesions involving the whole small intestine, a few indeed with greater involvement of the duodenum than of the lower parts. (** Judging from the gross and microscopical appear- ances it seems that the virus causes at first a cellular infil- trate in the mucosa or submucosa upon which necrosis shortly supervenes. The overlying mucosa soon degen- erates, and the surface is covered with an indefinite slough. In other cases, especially early in the epidemic, the process extended outward and appeared as muscular or subperitoneal necrotic areas before the mucosa was much involved. At all events necrosis was an early change in every case. The blood vessels were usually thrombotic. In the cases that spread toward the peri- toneum a plastic peritonitis of varying severity was present. The focal liver lesions were not present in every case. They took the form of focal necroses or abscesses. Some fatty or parenchymatous degeneration w^as always present. The liver lesions probably started as inflamma- tions of the veins from which necrotizing or infiltrative lesions spread. The splenic lesions were those of lymphoid hyperplasia, only distinctive in the enormous MISCELLANEOUS INFECTIONS 611 number of large lymph cells. Typical microscopical changes are as follows, quoted from one of the autopsy protocols : The lung showed moderate congestion with here and there a little epithelial swelling and a mild bron- chitis and peribronchitis. The type of bronchitis is infiltrative rather than catarrhal. The heart muscle showed granular degeneration of the fibres with breaking up or irregularity of the striae. Some increase in inter- fibrillar nuclei and especially those of the capillaries. There is moderate congestion. Epi- and endocardia are slightly raised as if by edema. Here and there slight fragmentation of fibres. The liver cells are granular and some show fat droplets. There is moderate conges- tion and more than the normal number of round nuclei between the columns. Here and there are focal necroses of varying sizes without circumferential reaction. Here and there are also some small collections of round cells near to which the liver nuclei are large and show attempts at regeneration. In these collections but not in the necroses, bacillary forms may be found. There is no reaction on the part of the bile ducts. The larger vessels are thrombotic, and in one section a thromboangiitis was found. One stretch of early plastic perihepatitis was found. The kidney showed slight granularity with slight cloudy swelling of the epithelium. The nuclei of the glomeruli are prominent. There is moderate congestion. The spleen showed distinct large lymph cell hyperplasia with relative inconspicuousness of small round cells. The follicles are very diffuse, their centres filled with large lymph cells. The cords are hyperplastic and the sinuses compressed. Moderate congestion; no unusual blood destruction; one area of hyaline necroses found. The proventricle and gizzard are negative with the prob- able exception of active desquamation on the surface of the former. The outer coats of the duodenum are nega- tive except for slight richness in nuclei. The deep mucosa is very rich in nuclei and red blood cells. The outer parts 612 DISEASE IN WILD MAMMALS AND BIRDS of the villi are either swollen with a cellular infiltrate or by an area of granular necrosis, or have disappeared. It would seem that the surface of the mucosa rapidly degen- erates and desquamates. Bacteria are very numerous. The adjacent pancreas is negative. The ileum showed round cell infiltration of the deep mucosa, swelling of the villi and a desquamation of the surface. One ulcer was found having its base on the swollen muscularis and being covered with necrotic slough. Adjacent peritoneum is slightly infiltrated, but chiefly congested and edema- tous. This ileum lesion seems to be the characteristic one of the disease. Bacteriological observations were made upon cultures obtained from the intestinal mural lesions, the peritoneal exudate, the liver necroses, and the heart's blood in eleven cases. In seven cases I was able to isolate a motile rod like the B. coli communis and in four cases a non-motile rod of the Bad. aero genes type. The former is quite similar to the B. scoticus (Migula) reported in Grouse disease. *'We obtained from Doctor Kalbfus of the Pennsyl- vania State Game Commission, four perfectly healthy birds for experimentation. A culture of the isolated germ was injected into two of them and mixed with the food of the remaining two. It does not seem profitable to cite the details of the work as the results were entirely negative, no lesions resulting that bore the slightest resemblance to the spontaneous disease. . The birds either lived indefi- nitely or succumbed to wholly foreign conditions. This negative experiment is of course no proof that the organ- ism is not the cause of quail disease, for the methods employed might not be the correct ones to propagate the virus or the germ may have lost its virulence during the laboratory culture work. However, as some observers have not reported this bacillus in the disease this germ loses something in importance by the negative inocula- tion experiment. MISCELLANEOUS INFECTIONS 613 "Judging from reports and based upon the observa- tions of Morse upon Grouse disease it would seem that the incubation period of the disease is about eight to ten days. However, one of the tliird lot of our birds died within three days of its arrival at this Garden, and there- fore within three days of its exposure to the second arrivals ; if it be. correct that this second lot brought the disease and the third lot did not have it, it would seem that the incubation period can be as short as three days ; how long it may be is only suggested by the fact that some of the third lot did not die for three weeks after arrival and exposure. All the Gambel's and scaled quail suc- cumbed to the disease, but two of the twelve bobwhite survived. It would seem that although these last birds probably introduced the disease, they still possessed more resistance than the others, for the second death among them occurred seventeen days after the first death./; The epidemic as we have seen it here seems to be the same as Grouse disease of Scotland and as the Grouse disease in this country as reported by Morse (Bureau of Animal Industry Report 109, May 18, 1907). "The means of transmission of the disease is not exactly known, but is in all probability by a pollution of the food, the water supply or the ground. Since the lesions are so marked in the lower ileum, cecum and colon, a possible transmission by cohabitation must not be entirely overlooked. There does not seem to be any means of limiting the epidemic in a flock by segregation or sacrifice of the infected birds, because symptoms are few and do not appear until shortly before death.^ Each bird would have to be put into a separate cage until proved infected. Scrupulous cleansing of the enclosure is desirable, but its efficiency is difficult to estimate." SECTION XVIIl THE ANIMAL PARASITES, THEIR INCIDENCE AND SIGNIFICANCE Feed D. Weidman, M. D. It is quite to be expected that animal parasites would be found in the animals of zoological gardens, garnered as these beasts are from all parts of the world, tropical and otherwise. It inevitably follows that many of the forms should be strange and new, enticing one to the fascinating determination of their identity, life history and hygienic importance ; and, developing from all this, one can easily imagine how limitless the opportunities are for scientific work in parasitology in a laboratory like ours. As in other biological fields, the taxonomic range of parasites here is vvide. It extends fr'om the lowly proto- zoa to the insecta, and, dropping to the smaller subdivi- sions, includes not only most of the genera familiar to human parasitology but many knov\Ti only among the lower animals. From the standpoint of the host, the bio- logic state of parasitism extends from the lowest protozoa to homo. The above vnll suffice to indicate the wide range of parasitism in animals, but the extent of work actually done thus far in wild animal material is a different story. Collated, consistent studies, so far as I am aware, have been undertaken only at the London Garden, here at Philadelphia, and at Wasliington, D. C, by Dr. Charles W. Stiles and Albert Hassal. The data collected by the last mentioned workers are incidental to the Index Cata- logue of Veterinary and Medical Zoology, and embrace only the (index) phase indicated by the title, but it is so valuable, and vvithal so altruistic, that it must be credited. 614 THE ANIMAL PARASITES 615 What other work there is is scattered where-not in lit- erature — general biological, medical and veterinary. That at London has been conspicuous through the observations of Plimmer and of Beddard on filariae and cestodes respectively, while the w^ork of Nicoll must not fail of mention. That the reader may the better appraise the sections of our own work which are to follow I wish at once to indicate their material basis. Ordinarily only the larger parasites are looked for at the autopsy table and there must be special indications to demand search for the finer ones. Those of microscopic size, or so minute as to be overlooked in the guise of seeds, vegetable fibres, etc., have not, both here and elsewhere, been routinely studied as have macroscopic ones.(l) From our autopsies there have accumulated records of nearly 900 parasites — some determined generi- cally, others but as to order. The parasites have in greatest part been preserved and are available for fur- ther study ; in the past, special groups have been culled out from time to time and examined. Where conditions have been pressing, as in certain epizootics, investigations have amounted to more than observations and descrip- tions, and received detailed laboratory examinations with more or less animal experimentation as the occasion demanded. The foregoing may suffice to apprise the reader that the subject of wild-animal parasites has been but broached so that data are especially incomplete on life histories — a phase most important in relation to hygiene ; but in spite of this and although the statistics are only approxi- mate, as is the case in most parasitological work, these data have attained to sufficient proportions to justify at least a beginning in the matter of collating and general- ' There are certain exceptions to this, as with Nicoll's (Proc. Zool. «Sfoc. London, 1912, p. 858) careful search for trematodes with sieves, but this means a separate research, and is incompatible with the all-round, general policies of present routine laboratory organization. 616 DISEASE IN WILD MAMMALS AND BIRDS ization. At any rate the time has arrived to establish at least a nucleus for the accretion of data, which can be later subjected to conjSrmation or correction. We draw just a grain of comfort from the knowledge that the more fully worked field of human parasitology is also -vulner- able to criticism of very much the same order. The Value of Parasitological Studies in Zoological Gardens. The foregoing chapters have made clear two fields of practical usefulness of any study in such gardens. These — hygiene in relation to the animals and comparison in relation to human beings — need therefore only to be mentioned at present since it is ob\T,ous that both benefit by our parasitological work. But there is yet a third — a scientific phase of parasitology which may be considered purely academic. It consists in morphological and other studies necessary for the identification of the parasite, the determination of its life liistory, etc. These last studies may still in a restricted sense include a modicum of the practical in so far as they have a bearing on the disease with which they are associated. But on the whole they are a source of danger for us since such things as studies on the finer structures of worms, taxonomic arrangements, descriptions of new species of commensals, etc., being alluring, are likely to lead one so far afield that eventually an attitude of stubborn resistance will have to be assumed in order to conserve that precious, volatile laboratory asset — time — for the more crying, practical problems ever reaching out to us. However, in parasitological investigations as in other scientific work, immediate abstract information may at some time prove to be of greatest practical value. Thus for example if we can discover the exact facts concerning one phase of the life history of a certain parasite, it may be possible by hygienic measures, to break the cycle of development of the parasite at one point thereby prevent- THE ANIMAL PARASITES 617 ing its completion. This information is perhaps obtained most readily in experimentation upon the role of lower animal forms in the pathogenesis of disease but where reliable evidence is lacking, help may be had by compari- son with others in the same taxonomic group. Undoubt- edly systematic classification will go far to help solve many of these riddles. Pathogenicity of Animal Paeasites in General. The first question which arises in this connection con- cerns the actual ability of animal parasites to produce disease in mid animals. At once it will be seen that this must be a relative matter, for no one on one side would contend that every symbiont in an animal is harmful — parasites sensu stricto — nor on the other that none could possibly be, i.e., that all are always commensals. It is evident that the issue boils down to questions as to the extent to which they are harmful. Before attempting the answer let us consider the means by which the para- sites may conceivably produce disease. Modes of Disease Production (Pathogenesis). The medical reader is familiar enough with the patho- genic powers of some animal parasites, but may be suffi- ciently interested to glance over specific wild animal instances illustrating them while they are being listed for those less familiar with this subject. 1. Mechanical Obstruction. ; I refer here particularly to simple blockage of normal body passages as the result of bulk or mass. This occurs more commonly in the intestines than elsewhere on ac- count of the greater frequency, greater numbers and larger size, in general, of parasites inhabiting this tract. Thus, we have recorded a liothrix {Liothrix luteus){2) where the combination of a small host and consequently (2) Phila. Zool. 8oc. Rep., 1920, p. 28. 40 618 DISEASE IN WILD MAMMALS AND BIRDS narrow gut and comparatively large parasite induced obstruction. Plimmer(3) records microfilaria clogging the brain capillaries. Shipley (4) mentions two specimens of Ascaris lumbricoides obstructing the nares of a chim- panzee {Pan niger). Blockage may also be produced sec- ondarily to the presence of the parasite, even in the absence of notable numbers of them, and quite apart from the element of verminous bulk. This occurs through in- flammatory swellings which the worms excite. We saw many serious grades of this in our spiroptera epizootic, the lumen of the proventricle being narrowed by swelling of the mucosa and more or less occluded by exudate and necrotic mucous membrane. Yet another direction wherein a mechanical rationale pure and simple obtains is by the production of divertic- ula. Worms encysted in the gut wall may, by weight alone or by excitation of peristalsis, cause the wall to bulge outwards (or inwards even) like a pocket. Such a diverticulum has been noted in the gut of a Pale Cebus {Cehus ftavescens) (5) parasitized by acanthocephalus, but in this case there were adhesions to the nearby stomach, and it is possible that in this individual case the diverticulum was a traction one, i.e., pulled out by the anchorage of adhesions externally. 2. Mechanical, Irritation. — In those instances where inflammation is the manifestation which reflects the simple mechanical effects of parasites it will be difficult indeed to prove, in the present state of our knowledge, that it is not rather the effect of associated toxic sub- stances or excreta elaborated by the parasite. But instances of a purely mechanical irritation there must be, although one can scarcely put the finger upon them and say that this or that individual inflamed mucosa did not become so from a toxic cause. Omitting these then, (3) Proc. Zool. 8oc. London, 1910, p. 134. (4) Proc. Zool. Soc. London, 1905, p. 252. (5) Phila. Zool. Soc. Rep., 1920, p. 29. Fig. 71.— ACANTHOCEPHALUS (THRKK SPECIMENS) PROJECTING FROM THE IN- CISED INTESTINES OF A PIGMY MARMOSET. COMPARE THE SIZE OF THE PARASITES. WHICH MAY BE DISTINGUISHED BY THEIR ANNULATIONS. WITH THAT OF THE INTESTINES. Fig. 72. — BLOOD-RED NEMATODES PROTRUDING FROM FRONTAL SINUSES OF COMMON OPOSSUM (DIDELPHY8 VIRGINI ANA). IHE SKULL- CAP HAS BEEN LIFTED OFF AND THE POSTERIOR WALLS OF THE SINUSES BROKEN. THE ANIMAL PARASITES 619 the more certain, purer, more unequivocal examples will be those where physiological processes become exalted as the result of the parasitic irritation. An example in point is a case of volvulus in a Screech Owl {Otus asio asio).{6) Here it is probable that the parasites excited the gut to undue peristaltic action, and that during this process it became twisted. Worms in such passages as the nose and nasal sinuses (I have seen blood- red filarias in the frontal sinuses of an opossum) undoubtedly pro- duce nervous effects by their presence and movements. Those in the subcutaneous tissue (tilariae of wild cats) probably also do so. It is difficult to judge those cases where doubtfully sensitive parts are the ones affected. Probably the intestinal and intraperitoneal worms, and less certainly the generally-migrating ones analogous to Filaria loa, produce no nervous effects mechanically. 3. Production of Hemorrhages. — Hemorrhages large enough to kill suddenly are theoretically possible, since worms occasionally produce aneurysms which may rup- ture; we have seen such an accident in a Paradoxure {Paradoxurus leucomystax). But certainly it is the long- continued, wasteful small hemorrhages that are impor- tant, inducing an anemia often of severe and fatal grade. The hookworms are the shining offenders here, yet we have seen very much the same effect from Acanthostoma in the intestine of monkeys. (E'sophagostomum has also been incriminated at the London Garden in young Rhesus Macaques {Macacus rhesus) {7) where the young forms of the parasite did the damage as they burrowed into the wall of the gut. 4. Opening up Avenues of Infection. — This may be accomplished either by passage of parasites from one position normally containing bacteria to another wliich is susceptible to infection, or by devitalizing a tissue which is ordinarily resistant to infection; i.e., creating a (6) Phila. Zool. Soc. Rep., 1921, p. 31. (7) Proc. Zool. Soc. London, 1919, p. 15. 620 DISEASE IN WILD MAMMALS AND BIRDS locus resistenticB minoris. The intestinal tract is the most common organ concerned, but the illustrations to follow will give variety. Thus, the mature examples of oesophagostoma in young rhesuses just referred to above burrowed into the gut wall and led to both local and gen- eral peritonitis. In one of our **spiroptera" parrots the worm had passed through the proventricular wall and a chronic fibrosis resulted around it. At the autopsy on a Rhesus Macaque Doctor Fox found a localized abscess adjacent to the gut wall, and in it a whipworm was imbedded. Passing from these examples of intestinal worms, I can mention the loss of a valuable Philippine Spotted Deer {Cervus alfredi) as the result of secondary infection of a cysticercus cyst of the lesser omentum which led to a nearby peritonitis. Lung infections are not uncommon. Murray (8) records that forty-four out of eighty-five young rhesus monkeys dying from pneu- monia showed an acarian, and he ascribed the pulmonary irritation to certain crystals in the excreta of the mite. I have studied a case of bronchopneumonia in a prairie dog where great numbers of an arachnid were present. , The reports of the London Zoological Society are replete mth notes of round worm pneumonias of reptiles. These pul- monary cases must result from decreasing of tissue resistance by the presence of the worms, and are easy to understand, much more so than the intestinal infections when one recalls how sensitive lung tissue is to foreign bodies, and that there seems to be no indication that this tissue becomes accustomed to infestation such as may be argued for the gut. j All these citations must convince us that parasites are most important predisposing agents to infection, and that tliis is one of the most sinister phases of animal parasitism. . 5. Destruction of Tissue. — This heading does not refer to the comparatively trivial effects that accompany the more acute inflammations secondary to parasites, (8) Proc. Zool. Soc. London, 1919, p. 15. THE ANIMAL PARASITES 621 albeit certainly the absorption of their disintegrative tissue products has some effect on the economy ; but our ideas of such are so vague as to justify their being disre- garded here. (What I refer to is the more massive destruction such as may occur in the blood, for instance, from the action of protozoa. There is also loss of mucosa in those chronic inf estments of the stomach where we find excessive fibrous tissue overgrowth. The most striking example of tissue destruction we have seen was in the cirrhotic livers of prairie dogs affected by Eepaticola hepatica, where in extreme cases, the amount of function- ating liver substance was reduced to a very smaU fraction of its normal bulk. (9) 6. Toxins. — We have no direct evidence to offer that noxious products of parasites are concerned in producing disease in wild animals. The local effects of such toxins are not distinctive enough — individual enough to toxins or to the animal body — to separate them from the effects of such accompanying factors as bacterial inflamma- tions; nor can we separate the general effects of these toxins from what might have been, for instance, the effects of an accompanying anemia of hemorrhagic or other origin. From a knowledge of what happens in human prototypes though, there is scant doubt that some one of the multitudinous species must be capable of pro- ducing toxins, but just which varieties are concerned cannot be listed by anyone. By analogy we can at most only suspect the hookworms and the dibothriocephalidse. Under this same category of the toxins come the worm- products which are reputed to have a destructive effect upon the digestive enzymes in the gastrointestinal tract of the host, and which would thereby interfere with the proper assimilation of pabulum, resulting in malnutrition. For the same reasons as above indicated for the toxins one is unable to speak for or against these ' * anti- enzymes. ' ' (9) Phila. Zool. Soc. Rep., 1916-1921. 622 DISEASE IN WILD MAMMALS AND BIRDS 7. Preclusion of Nutrition.— This must be a very miimportant phase of the activity of intestinal parasites, when one compares the bulk of food which passes through the bowel and the average number of worms present ; and the same holds good for some interstitial parasites like the adult filariaB. Even in amazingly hea\y infestments of the intestines one will be constrained to dismiss this idea when he compares the bulk of parasites with that of the host, and recalls what the physiologist terms the ''factor of safety" inherent in this tract as elsewhere. But in the case of blood parasites the matter may be different. Here we are concerned with the withdrawal of refined foodstuffs — those which have been worked over and over by subtle internal metabolic processes ; and we are not so sure, especially on recalling the enormous numbers of parasites usual to blood infestments, that there is the capacity on the part of these internal processes to meet increased demands that we count upon for the intestinal functions. It is much more serious to be deprived of the finished product than of the crude because it means the undoing of "digestive" work all along the line, from gut to tissue cell. Furthermore, a blood infestment guarantees that the parasite has been feeding upon and depriving the animal of the precise foodstuffs the cells require, and not by any chance upon, even in part, intestinal substances that were wastes or residues. If we except the blood parasites, then, it seems safe to conclude on the whole that the amount of pabulum used by parasites is unimportant to the animal. Having reviewed the manner in which parasites may conceivably be harmful, it is time to return to the question of the actual exercise of these powers. The older appraisal of parasites in animals, namely that they were rather innocent of disease production, was suggested by and borrowed from the veterinarian, prob- ably being engendered in him by their frequency in what appeared to be normal domestic specimens. Yet it is only THE ANIMAL PARASITES 623 proper to add that one of our former pathologists, and sometime professor of veterinary pathology, Dr. C.^ Y. White, is a medical man and is of much the same opinion. Older writers regarded worms even as "guardian angels" of children. Very recently Schwartz (10) reviews some work in this connection showing that, in vitro, some cestode extracts were inhibitory to certain bacteria {B. anthracis, B. pyocyaneus and B. dysenterice Shiga). This relationship is so different from natural conditions as to need no further comment. At the London Garden the view appears to be dif- ferent. In the 1910 report they charge five deaths against perforation by worms of the stomach and intestines ; in the 1911 report they record giant toads dead from lung infestment; in 1912 ''eighteen cases of enteritis were due to worms"; and in 1917 they mention pneumonia in a toad and perforation of the stomach of a puma. These reports represented evidently the more striking, unequiv- ocal examples of death from parasites which had outspoken anatomical expressions, and omitted those in which the more subtle agencies of parasitic pathogenesis were concerned. Their experience has apparently been much the same as ours. The ideal approach to a decision in reference to the importance of parasites would appear to be a mathe- matical one, something as follows : First, to determine what species infest animals and how commonly, then to decide which ones are pathogenic and tliirdly to estimate the severity of the disease induced; so that finally, by an analysis and comparison of the three results— a com- parison and analysis judicial in the broadest sense— we might hope to come to an opinion. Let us consider the three avenues in order. At the first glance it must be evident that a list of all possible parasitic varieties does not exist and may never be compiled. The most that can be done is to tabulate the findings in scattered labora- (10) Journal of Paras^it., June, 1921, p. 194. 624 DISEASE IN WILD MAMMALS AND BIRDS tories, data usually recorded in terms of the individual observer's studies and often inadequate to give the com- piler all the facts desired. The same remarks apply to the percentage incidence of parasitism. Not to prolong the academic discussion, suffice it to say that very much the same obstacles present in the second avenue — that of pathogenicity of the individual species. Our own data referring to this second heading will be presented later, but after the failure of the first avenue, the second and third lose greatly in value. At best, statistics can be only suggestive. Unless critically and suspiciously interpreted, and with a full appreciation of their limita- tions from a foreknowledge of the way in which they were compiled, they would only delude the reader and offend science, and so we abandon this line of reasoning. At present the best results of the study of patho- genesis by animal parasites will probably be reached by a combination of methods, as follows : 1. Direct. How commonly do we see clinical symp- toms and morbid anatomical changes that are incontro- vertibly due to the parasite ? We restrict ourselves here to a narrow group of inf estments indeed, and think of such diseases as trichosomiasis in prairie dogs and spiroteria- sis in parrots. 2. By comparison with analogous infestments of domestic animals and man — more thoroughly studied and therefore more accurately appraised, in general, as to pathogenicity ; a comparison from the standpoint of dis- ease production rather than natural habits of the para- site. Example, coccidiosis and hookworm disease in foxes and dogs. 3. By inference through deduction. This is the most unsatisfactory consideration of all, and should be well checked up and discounted. Here we would evaluate the known propensities of the parasite first, such as its size, motility, anatomic position in the host and the general pathological traits of the genus and family to which it THE ANIMAL PARASITES 625 belongs, etc., and then compare these verminous properties with those of the host — its size, temperament, physical stamina, etc. This third consideration must necessarily overlap with or be supplementary to the first two. For example, this consideration would have to be resorted to in many cases of ascaris infestment where anatomical changes are generally not demonstrable. Acting on these three considerations, and after twelve years of observation on parasites here in the Garden, a fresh review of our records, and a recent review of the accessible relevant parasitological literature I have come to the conclusion that, considering wild animal collections the world over, there is no justification for an unqualified, definite answer to the question of pathogenic parasitism that will meet all conditions. We lack data on too many species that are not sufficiently represented in collections or indeed not represented at all. It is the liability to infestment of each order or family of beasts that will have to be determined, and, depending on the assortment each garden has on exhibition, will the importance of parasites to the garden as a whole vary. Speaking for the Philadelphia Garden, I have come to the conclusion that on the whole parasitism does play an important part of our animal losses. The financial loss which could be charged against spiroptera alone is in the four figures, to say nothing of the difficulty of replace- ment of rare species. And while touching the financial phase let it be added that scientific work done now, it must be remembered, is not restricted to the present time or place, but is to be measured in dollars and cents with the yard stick applied to the future, and in other places than that where the initial work is done. Even if we cannot answer the question of the matter of importance the world over we can guarantee that it is sufficiently so in the Philadelphia and London Gardens to warrant a rigid supervision for parasitism; and since the other larger collections are probably made up of similar animals, 626 DISEASE IN WILD MAMMALS AND BIRDS albeit in different proportions, we surmise at least that it is likewise so with them. Importance of Parasites in Other Fields. — In addi- tion to their importance to exliibitions, animal parasites of wild animals are important first to man. The animal hosts may serve as porters of infestation, and interfere with attempts at eradication of the disease. The experi- ence of the European with African sleeping sickness attests to this. Not to go farther than immediate examples I wish to note in this connection the occurrence in this Garden of scabies in an orang which was transmitted to a keeper, and of amebic dysentery in monkeys. Leiper(ll) has called attention to a guinea-worm in a leopard. Parasites are important to certain wild animal indus- tries. The ones that have come to my attention are the fur seal {Otoes alaskamis) industry of the Pribiloff Islands and fox-farming in Newfoundland. In both of these instances the hookworm was concerned and entailed losses of thousands of dollars. Lucas, who conducted a United States Government commission to the seal grounds and after whom Stiles named the parasite, has left very full notes of the former disease. I have identi- fied the same infestment in a young California hair seal {Zalophus calif ornianus) which was born and died in this Garden. This indicates that the parasite might perhaps be found farther dowTi the Pacific coast than hitherto suspected. To hunters parasitism of animals must be important, but to an unkno^vn and undoubtedly unimagined extent. The grouse plague of Scotland (12) is an example to point. Who knows but that the disappearance of some of our game animals, particularly birds, was not due more to disease than to the ravages of man I There is at least food for thought here. (11) Proc. Zool. Soc. London, 1910, p. 147. (12) Fantham, Proc. Zool. Soc. London, 1910, p. 672. THE ANIMAL PARASITES 627 OCCUKEENCE OF AnIMAL PaRASITES IN THE WlLD. It would be unbelievable that parasitism did not exist in the wild. It seems proper, however, to record some evidence. Diesing's 8y sterna Hehninthum is replete with ref- erences to Natterer's Brazilian expedition. Nicoll speaks of a German expedition to Spitzbergen in 1898, and a Swedish one to Egypt in 1901, in both of which large numbers of parasitic forms were collected. Nicoll (13) found Trichosoma hepaticum in a hare shot in the wild, and liver-flukes (14) in a kestrel shot on the coast of Scotland. Leiper(15) found nine species of worms in hippopotami during an expedition to Uganda, and (16) states that thirty-seven species of helminths were col- lected on an Antarctic voyage by Surgeon Atkinsons In an investigation of Grouse disease in Scotland, Fantham found many different blood and intestinal parasites. \ Dr. Charles B. Penrose tells me that all of the white- tailed deer he shot in the valley of the Swan River, Mon- tana, were infested with liver-flukes, so much so that the liver was literally riddled by the disease, and yet the deer were fat. The black-tail deer of the same valley were not thus parasitized and were not as fat. In our own Garden we have found many tapeworms in wild cats (17) which had been too recently captured for the worms to have developed in captivity. Such instances might be still further multiplied. A more important consideration is the fate of the parasites thence introduced into our Garden. Do they disappear of themselves? Naturally we can never make sweeping predictions, for future events will depend upon the life history of the individual parasite concerned. But by and large, once introduced it is better to assume the attitude of pessimism, and resign oneself against spon- (13) hoc. cit., 1911, p. 674. (14) hoc. cit., 1915, p. 87. (15) Loc. cit., 1910, p. 233. (16) Loc. cit., 1914, p. 222. (17) Phila. Zool. Soc. Rep., 1912, p. 40. 628 DISEASE IN WILD MAMMALS AND BIRDS taneous disappearance and, what is worse, realize that the parasitism is likely to become indigenous. We have sev- eral pieces of evidence, however, that the infestment may occasionally quite disappear. Thus, I have seen Coccid- ium higeminum spontaneously disappear from a Swift Fox {Canis velox) and Spiroptera incerta from a Macaw as proven at autopsy. Nicoll(18) remarks that certain trematode infestations were heavier in newly arrived animals than in ones long resident in the Garden. This is .conceivable on the basis of individual worms dying out, i.e., fulfilling their life spans without the host becoming reinfested with fresh parasites. Precise information on the subject is supplied by Ackert(19) who found that cestodes disappeared from chickens in six to eight months when the birds were confined, i.e., protected from rein- festment.'i Moreover, it is known that worms can escape during acute infections, the infectious state of the economy producing conditions obnoxious to the parasite. We hear of many instances of their expulsion in human feces and vomitus during malaria and the exanthemata of childhood and know of similar discharge from animals during the death agony. I cite these data largely because they explain the scarcity or absence of parasites at autopsy in animals which were known to have been clinically infested. FREQUENCY OF PARASITISM IN WILD ANIMALS There can be little doubt that mid animals are more frequently infested than man, and furthermore with a larger number of parasites, I have no statistical basis for these opinions — they rest on personal observations of human and animal autopsies, and reports of findings in the tropics and elsewhere. They have therefore but the value of an indi\ddual opinion. I should estimate rather cautiously that ^dld animals are infested at least two or three times as frequently as man and much more heavily. (18) Proc. Zool. Soc. London, 1914, p. 140. (19) Jour. Parasit., June, 1921, Vol. Vll, p. 198. THE ANIMAL PARASITES 629 The first step in the discussion of the incidence of parasites must be that respecting the (host) classes and smaller taxonomic divisions — of course as they have been studied in this Garden. Certain statistical limitations were experienced and can be summarized as follows : Data are not available on a sufficiently large number of animals to justify conclusions as far down as genera and species, except for such commonly and generously exhibited forms as monkeys and parrots. I have there- fore in tabulating and reviewing our records, distributed the animals only as far as families — not into genera and species. The table (24) to follow will be found not to contain every family because to do so would needlessly enlarge it. Accordingly I have followed the policy of only indicating those genera and species showing either frequent or important infestment. I shall refer to those groups later as *' susceptible " groups. If no family is recorded in the table it means that we have had no important numbers of inf estments in it. The ' ' remarks ' ' column shows the individual parasite that has been par- ticularly frequent or otherwise important. If there are no remarks it means that the species of parasites found have been scattering. Results of Review and Tabulations. We now pass to an analysis and discussion of the find- ings brought out in the previously mentioned review of our records and in Table 24. Viewed broadly we find that there is a wide variation in the susceptibility of dif- ferent families to infestment. Those that are susceptible may be located by consulting the table, and each will therefore not be separately culled out and subjected to needless repetition. A few points are however worthy of separate mention. While there is a familial or generic susceptibility within certain orders it is unwise to gen- eralize too broadly. Thus for example the Corvidae have 630 DISEASE IN WILD MAMMALS AND BIRDS Table 24. Incidence of Parasites in Animal Groups. Primates Cercopithecidae Sooty Mangabey Cercocebus fuligi- nosus Rhesus Macaque Macacus rhesus CalUtrichidse Marmosets Cebidae Squirrel Monkeys Other Cebus Monkeys Lemures Carnivora FeUdse American Wild Cat. . . Fehs ruffus Spotted Wild Cat Felis ruflfus texensis Canada Lynx Felis canadensis Lions Fehs leo Ocelot Fehs pardahs Canidse Gray Fox Canis cinereo argen- teus Red Fox Canis vulpes penn- sylvanicus Swift Fox Canis velox Gray Wolf Canis mexicanus Mustehdse American Badger Taxidea taxus Procyonidae Raccoon Procyon lotor Ursidae Bears *538 34 60 43 8 87 86 498 28 5 10 10 15 28 17 5 18 17 42 37 9.4 11.8 10. 16.3 37.5 11.5 7. 16.9 40. 80. 40. 30. 33. 12. 40. 11. 41. 5. 16. Eight had Filaria gracihs. Stomach and intestines, 22; Bronchi, 4; Muscles, 7. Ascarids only. Ascarids in stomach and intestines. Uncinaria. Cestodes. Uncinaria. Uncinaria. Ascarids. Physaloptera. Ascarids. ♦This figure and a number of others in the tables do not correspond with those in other sections of this book because certain injured, decomposed and newly arrived animals were available and accepted for my purposes, but were objectionable for the general medical statistics and therefore excluded. THE ANIMAL PARASITES Table 24 (Continued). 631 I Otariidse Hair Seal Zalophus californi- anus Rodentia Sciuridae Castoridae American Beaver Castor canadensis Hystricidse Canada Porcupine. . . Erythizon dorsatus dorsatus Hyraces Cape Hyrax Procavia capensis Ungulata Equidae Zebras Cervidae Axis Deer. Cervus axis Barasingha Deer Cervus duvanceli Eld's Deer Cervus eldi Fallow Deer Cervus dama Hog Deer Cervus porcinus Japanese Sika Deer. . Cervus sika typicus Red Deer Cervus elaphus Elk Cervus canadensis White tailed Deer . . . Mazama virginiana Mule Deer Mazama hemionus Camelidse Llama Lama glama Camels Suidae 20 198 44 17 47 365 20 14 32 17 44 5. 16. 9. 23. 36. 28. 12. 100. 17. 62. 14. Uncinaria. Scattered through four different genera. In three cases oxyuris and flukes in cecum. Cestodes 8, filaria 11, oxy- uris 9, in peritoneal cav- ity, also intestine. Cestodes in bile ducts. Nematodes, intestine. C. tenuicollis. Echinococcus cysts. Trichocephalus. Echinococcus in lung (2). Four Cyst, tenuicollis. Hydatid cysts. 632 DISEASE IN WILD MAMMALS AND BIRDS Table 24 (Continued). Animal Edentata Armadillos Marsupialia Didelphyidse Common Opossum. . . Didelphys virgini Macropodidae Kangaroos and walla- bies 'I 16 10 175 84 70 12.5 20. 48. Physaloptera, 38; oxyuris, 5; cestodes, 5; nema- todes in lungs, 3; cysts in peritoneal areolar tis- sue, 2; trematodes in ileum, 1. Corvidse Common Crow Corvus brachyrhyn- chos brachjThyn- chos Magpies Jays Pies, choughs, etc... Sturnidae Starlings . Turdidse. 16 35 AVES Canaries. 24 12 19 44. 64. 55. 33. Tropidocerca and occa- sional intestinal cesto- des. Syngamus in crows. Few filaria. Periproventricular filaria, strongylus. There is a striking consis- tency of infestment in the different members of Corvidse both as regards degree of infestment and species of parasite pres- ent. Periproventricular filaria largely. Periproventricular filaria largely. Thrushes and Robins. None in American thrushes, one in a robin. Finches. Not examined closely at autopsy, but there is a scattering of periproventricular fila- ria and intestinal ces- todes through most of the species. Were free from parasites. THE ANIMAL PARASITES Table 24 (Continued). 633 AVES 11 II 3 a 1 1 4 2 50. 30 9 30. 142 2 7. *774 124 16. 24 5 20. 34 2 6. 45 4 9. 453 65 14.3 121 2 1.6 21 6 29. 48 21 44. 86 12 14. 74 13 18. 321 69 21.5 26 9 34. 62 16 26. 164 27 16.5 69 17 10. *201 13 6.7 55 4 7.3 44 1 2.3 29 1 3.4 73 7 9.6 Picarise Picidse Woodpeckers Rhamphastidae Toucans Striges Psittaci Loriidse Lorys Cacatuidse Cockatoos Crested Ground Parra- keet Calopsitta novse-hol- landise PsittacidsB Old World (Totals) . . . Undulated Grass Parrakeet Melopsittacus un- dulatus Pennant's Parrakeet . . Platycercus elegans Rosehill Parrakeet. . . . Platycercus eximius Other old world parra- keets Old world parrots, lovebirds, eclectus . . New World (Totals) . . . . Macaws Conures Amazons Other new world par- rots Accipitres Falconidse Buzzards Eagles Serpentaridse Vultures Miscellaneous * For foot note see page 630 41 Spiroptera largely. Remarkably free of para- sites. 3 spiroptera, 1 hemopro- teus, 1 intestinal worm. 2 spiroptera. 4 spiroptera. 1 spiroptera, 1 coccidium. 6 spiroptera. 20 spiroptera, 1 cestode. 12 spiroptera. 13 spiroptera. 9 spiroptera. 15 spiroptera, 1 hemopro- teus, 1 blood larva. 24 spiroptera, 3 nema- todes. 7 spiroptera. 4 were blood protozoa. 634 DISEASE IN WILD MAMMALS AND BIRDS Table 24 (Continued). AVES Animal •ss II ll 9 a 1 § 1 Remarke Galli.. 299 95 14 70 39 ♦163 *38 *41 *21 ♦22 ♦105 ♦319 48 83 188 36 42 20 3 10 7 14 7 10 3 1 21 28 7 13 8 1 14. 21. 21. 14. 18. 9. 18. 25. 14. 5. 20. 8.8 14. 15.6 4. 2.8 Phasianidae Quail Megapodidae Wild Turkeys Columbse Fulicarise Heterakis. Intestinal cestodes. Coo- cidia twice. Mostly intestinal cestodes, but several spiroptera. Alectorides Gavise. Steganopodes Herodiones Geese Parasites scattering. Few intestinal cestodes. Ducks Struthiones. . * For foot note see page 630. a high percentage in incidence for tropidocerca, 'syn- gamus and periproventricnlar worms, many families of Ungulata harbor echinococcus, and Carnivora are prone to show ascarids. On the other hand, among the copious exceptions to this may be cited the irregular liability to inf estment exhibited by the Galli. Four varieties of these birds are represented but there are missing such impor- tant kinds as curassows, guans, guinea fowl and peafowl. I (Nor do all members of a genus necessarily show the^ same susceptibility, and the heterakis infestment in the pheasants illustrates this matter very well. It was limited almost entirely to two species — Amherst's and Golden, whereas several frequently exhibited species showed none. ) The following table brings this out in more detail : THE ANIMAL PARASITES 635 Table 25. Heterakis in Pheasants. Species Golden Pheasant (Chrysolaphus amherstise) . Amherst's Pheasant (Chrysolaphus pictus). . Silver Pheasant (Gennaeus nycthemerus) . . . . Reeves' Pheasant (Phasianus reevesi) Ringnecked Pheasant (Phasianus torquatus) . Swinhoe 's Pheasant (Gennaeus swinhoii) Enzootics and environment played no part in the above figures. We have had no real heterakis enzootics, for in but two instances did three heterakis deaths occur in a year, and two deaths per year have occurred in but four instances in the past twenty years. During this time there have been sufficient animals on exhibition and sub- jected to autopsy to indicate definitely that the two species named — Amherst's and Golden, must be considered as more susceptible than the other varieties. Nearly all of the heterakis in quail likewise occurred in one species — seven of the ten cases occurred in a total of twenty-three Scaled Quail— but in these birds the infestment appeared in enzootic form and cannot be viewed as indicating a preference for a species. Psittaci are on the whole, not susceptible to worms. It is true that we suffered a serious outbreak of spiro- pteriasis a few years ago, but if we consider this a closed chapter we can accept the above generality as stated. Among 774 parrots autopsied we have encoun- tered but one cestode and three intestinal round worms. , The deer, likewise, are singularly free from intestinal parasites. I gave the detailed records of these animals in Table 24 to emphasize the scarcity of parasites even when fairly numerous specimens had been available for examination. Other interesting features in the table are the out- standing infestments of squirrel monkeys and marmosets among the monkeys, of gastric and intestinal worms in the wild cats, and intestinal worms in the zebras. 636 DISEASE IN WILD MAMMALS AND BIRDS The foregoing has had to do with parasitism from the standpoint of the host. The next phase, that of the individual parasite itself, interests more the strict parasi- tologist than the general zoologist; however, both will see how it may have a very practical value. Table 26. Distribution of Parasitic Cases According to Parasitic Groups. Nematodes Spiroptera Filaridae Ascaris Physaloptera Uncinaria Tropidocerca Heterakis Trichocephalus Syngamus Trichina Hepaticola Other Miscellaneous. Total Nematodes. . . . Cestodes Echinococcus Cysticercus Taenia Miscellaneous Total Cestodes Trematodes Acanthocephalus Protozoa Arthropods Unclassified 183* 145 138 30 28 25 23 22 11 9 2 2 4 165* 9 7 4 3 622 188 22 4 14 6 34 Grand Total. 890 * Not generically diagnosed. Incidence According to Parasitic Groups. Inasmuch as it has been physically impossible to de- termine specifically and classify efficiently the accumu- lations of verminous material from our autopsies I will not be able to tabulate parasitic groups even as closely as I did in the * ' animal host ' ' table. Nevertheless sufficient has been done to illuminate in part certain phases of parasitism and to prevent a summary dismissal of the subject. Reviewing our cross-index I have distributed the data into the following Table 26, the parasites being THE ANIMAL PARASITES 637 listed in the order of their frequency. It may serve only as a panorama of the situation, inasmuch as determinative study of a group amounts to a research in itself, and the multiplicity of them precludes a consistent study of every one. The data are based upon '' cases of parasitism." That is, each and every worm species occurrence has been counted, regardless of whether it was the same species that has been concerned over and over again, or in dif- ferent anatomical positions (of different individual hosts, of course) or whether it was in association with other parasites. Analysis of Table 26. There is a grand total of 890 cases of animal parasit- ism embraced in the above table, which is a sufficiently large number to give representative value to some phases of the analysis. I In the first place nematode worms occur about three times as frequently as all other forms of parasites. In gardens where spiroptera has not figured so largely the proportion might be reduced to about two to one. Ces- todes rank a poor second, trematodes a worse third, and acanthocephali a very bad last. This order agrees with our figures of 1913(20) and with the small series of Nicoll.(21) The latter worker found that the order was not changed when pains were taken to include also such smaller worms as could only be obtained from the host by using sieves, etc. Cestodes were not likely to be over- looked, but very small trematodes and nematodes were easily passed over. ViscEEAL Distribution. As to the individual organs which are most commonly parasitized our records show that with Aves as well as Mammalia the intestines are the parts most commonly affected. The stomach ranks second for both-\-the pro- (20) Proc. Acad. Nat. 8ci. Phila., March, 1913, p. 127. (21) Proc. Zool. Soc. London, 1912, p. 858. 638 DISEASE IN WILD MAMMALS AND BIRDS ventricle rather than the gizzard of birds corresponding, parasitologically speaking, to the stomach of mammals. We have found but one parasitic species in the gizzard of birds, i.e., immature forms of Spiroptera incerta lying under the chitinous lining of the gizzard and only dis- coverable after the lining has been peeled off. The peri- toneum comes third (air sacs of birds) due to the presence of filaridae, and the blood fourth for the same reason." It is to be emphasized that, in our data, identical organs of mammals and birds should be about equally liable to infestment with the possible exception of the lungs. But in view of the small number of cases available there is no justification for speculating about the reason for this last difference, albeit the radical difference in the anatomy of the two classes is very inviting. Now that our spiroptera enzootic has subsided, the order above given will be changed, and in view of like disturbing factors other gardens should not expect the same order to hold invariably for their collection, since their enzootics will depend somewhat on the preponder- ance of animals of one or another family which are likely to compose their exhibits. A single such enzootic may suffice to disarrange the whole fabric, and if two or three are taken into account the order of organ involvement can be quite disrupted. To attempt to construct statisti- cally an '* order of frequency involved " which would stand for every garden would only lead to interminable adjustments on the basis of animals exhibited and of parasitic enzootics, so that I have finally been reduced to a combination of our Garden statistics and the blood- parasitic ones of the London Garden. Doing this I have arranged in Table 27 the frequency of organ involvement as follows and estimated the percentage of animals in- fested. These figures are computed upon a different basis from that of Table 24. They naturally cover all animals and not the *' susceptible " ones as in Table 24. Fig 73.— hugely DISTENDED PROVENTRICLE OF PARROT DYING WITH SPIROPTERIASIS. COMPARE ITS SIZE WITH THAT OF THE HEART WHICH IS ABOVE AND TO THE LEFT, AND THAT OF THE GIZZARD BELOW AND TO THE LEFT. THE ANIMAL PARASITES Table 27. 639 Mammalia Aves per cent. 9.0 3.7 2.3 1.5 1.0 1.0 0.5 Blood per cent. 6.5 Intestines 3.5 Proventricle 1.7 Blood Air sacs 1.3 0.3 0.3 0.4 Total 20.0 Total 14.0 / (The effect of this is at first sight startling in that it places the blood parasites of birds so far in the fore, but it must be at once recalled that the inquiries upon the blood parasites were much more searching — microscopic, than in the case of the other organs. If similar methods were applied to the others their percentage of parasitism might be notably raised — particularly that of the intestines. ) Special Parasitologic Considerations. At this point the statistical considerations of parasit- ism will give way to descriptions of certain specific inf est- ments that have given us more or less concern. The occurrence of single parasitic varieties or of well known species in an isolated host may occasionally be of practical importance, but usually they amount to little more than an academic study, whereas the repeated dis- covery of single parasitic kinds, or inf estment of similar hosts, especially when grouped, raises the matter to a very practical level demanding attention. Such findings being not infrequent in our experience, it has been possi- ble to study our material in a manner designed to show the frequency of various parasites in a certain host, the susceptibility of certain animals to parasites in general and the inf estment of dissimilar hosts by the same para- site. The more important of these now follow. 640 DISEASE IN WILD MAMMALS AND BIRDS Avian Spibopteriasis. ' This disease concerned parrots particularly but tou- cans, pigeons, and such widely separated species of birds as the starling, quail, thicknee and barbet have been occa- sionally affected. To the naked eye the parasite resembles the human hookworm, but differs in location, being a resident of the proventricle where it produces a swelling of the mucosa which interferes with the passage of food. Up to a hundred worms may be present in the one bird, and immature fonns are occasionally found under the chitinous lining of the gizzard. The parasite burrows into the mucous membranes, occasionally pene- trates quite through the wall into the air sacs, and on one occasion induced an adenomatous hyperplasia of the mucous membrane, and an adjacent ' 'peritonitis. ' ' Mucus is sometimes present in the droppings. Death may occur either acutely, or with emaciation. Spiroptera incerta Smith (22) is the common parasitic species of parrots, but I have found at least one other as yet unidentified species in the toucan, and there are proba.bly more. In the eight year period 1906-1913 from 25 to 50 per cent. of our dead parrots showed this parasite every year, the total loss being 113 birds for this period — a most important inf estment. ' We approached the problem by diagnosing and isolat- ing the infested birds through a microscopic examination of droppings, finding that by boiling the droppings in 5 per cent. NaOH solution we clarified them and made ex- amination easier and more certain without at the same time destroying the parasitic ova. The result of the ex- amination of all our parrots was the isolation of 14 per cent, of the parrot population ; and as these died off the diagnosis of inf estment was found confirmed at autopsy in every case. The parrot house was thoroughly reno- vated and no newly arrived parrots were admitted until after quarantining and examining droppings for ova. (22) Proc. Acad. Nat. 8ci. Phila., 1913, p. 133. ' 4^ ^ ' fjv\i-* Fig. 74.— histologic SECTION THROUGH PROVENTRICULAR WALL OF PAR- ROT. SHOWING SECTIONS OF SPIROPTERA IN THE LUMEN AND MUCOSA. THERE IS SOME GLANDULAR HYPERPLASIA (ADENOMATOID) AND NECROSIS OF THE LUMINAL PORTIONS OF THE MUCOSA. Fi<;. 75.— INFLAMMATORY ROUND-CELL INFILTRATION AROUND NERVE TRUNK IN WALL OF PROVENTRICLE. PARROT DEAD WITH SPIROPTERIASIS. THE ANIMAL PARASITES 641 The toucans and other species, being housed elsewhere, were not quarantined. Following this, we were gratified to experience no more spiroptera deaths in parrots for seven years. Then, in 1920 and 1921, a new outbreak occurred in four toucans and several other scattering species, including two parrots; but none of these came from the main parrot house and probably represented a fresh importation. We attempted to cure the isolated verminous birds by medication but were unsuccessful. Likewise attempts at determining the life-cycle of the parasite brought us no farther than that the ova devel- oped larvae in moist sand in six days. Feeding of ova, freshly passed and larvated did not produce infestment in parrots or pigeons. On the whole we can quote our experience with spiroptera as a most satisfactory exam- ple of the value of hygiene and as a result which could never have been accomplished by medication. Hepaticola (Trichosoma)Hepatica in Prairie Dogs. Bancroft (23) and Hall (24) have given us details con- cerning this parasite and the disease it causes. It is threadlike, several inches long, and permeates the livers of the gray rat, white rat and wild hare. (25) We first saw it in the more or less cirrhotic livers of several prairie dogs ; later we observed it in a beaver and the gray rats of the Garden. In the prairie dogs and beaver the liver resembled that of fatty cirrhosis and was so considered on naked eye examination at our first autopsy. We were only set right when we came to the histological examination. It was remarkable how well conditioned some of the prairie dogs were in in the face of very exten- sive liver destruction; but on the other hand some were emaciated and a few of the spontaneously diseased showed at autopsy an enormous ascites. The outstanding features at autopsy were the large size of the liver and its pallor (23) Proc. Roy. Soc. N. So. Wales, Sydney, Vol. 27, pp. 86-90, 1893. (24) Proc. U. 8. Nat. Mus., Wash., D. C, Vol. 50, 1916, p. 31. (25) Proc. Zool. Soc. London, 1911, p. 674. 642 DISEASE IN WILD MAMMALS AND BIRDS and hardness; and fine yellow lines could sometimes be made out twisting over the surface. The disease affects wild rats differently from prairie dogs. In both the spontaneous and experimental disease the infestment was insignificant, amounting to perhaps three or four foci the size of a split pea near the anterior margins of the liver. Diagnosis may be easily confirmed by crushing the yellow infested portions of the liver between glass slides and examining microscopically for ova. "We have seen such a small number of cases of this disease because so few prairie dogs reach the autopsy table, yet there must be some important mortal factor in our prairie dog enclosure, for the Superintendent states that the population there does not increase in spite of the frequent births and additions from dealers. The animals almost invariably die under ground and their bodies are not recovered. In order to test out the origin of the infestment we trapped two of our exhibition specimens, and the liver of both was found infested on surgical examination whereas six newly purchased ones had normal livers. The latter were secured fresh from their native habitat in the West, and their livers were examined through long surgical incisions and found free of infestment. Later we fed the ova (embryophores) from rat livers to these prairie dogs and on destroying them found them infested. We were also successful in transmitting the disease in the oppo- site direction, i.e., from prairie dog liver to white rat. From all this we feel sure that the prairie dog disease in our Garden was transmitted from the rat and that here is another reason for rat extermination in a zoological garden. The adult Hepaticola hepatica of prairie dogs I have not seen in sufficient entirety to compare with the rat spe- cies and therefore cannot affirm that the two are identical species. It is presumably like that of the rat, being thread- nr. 76— OVA OI HtPATICOLA HIPAflCA IN lUFR Ol ''^ARIH • "<- ' ^^^ HAM BIPOI \R OPtMNCb IHhRF IS DI bl RUC I ION OI '-' V^ ' '""^ vo mPMH 1 ^M INHAMMAIORY REACTION OF ChU.ULAR CHARAClhR HLT NO IMPORI^Nl FIBROSIS. r i I ^1 I ^1 I "^1 I ^1 I ^1 1 1 I ®l 1 ^ 'iiiiiiiiiliiiiliiiiliiiiliiiiliiiiliiiiliiiilitiiliiiniiiiltiiiliiiiliiiiliiiil^ Vu.. 77.— r\ti\ ar: A >\iriHi cdiiii) i\ i\ ik \iii:i' \ i k' iiii.i: dicis ok GIRAFFE. Nf)TE MARKKI) TKRIDI CI AL FIBROSIS IN I H K N KI<;HBORHOOI) OF THE PARASITES. THE ANIMAL PARASITES 643 like and most difficult to separate from the liver substance through which it ramifies. At maturity it dies and dis- integrates, leaving the ova distributed more or less in tracts through the liver substance, so that we are limited to a certain period wherein to obtain the mature form. The ova are not passed into the intestine, but re- main in situ, just as in the case of hydatid disease, and therefore diagnosis cannot be achieved by examination of feces. For the disease to be transmitted the host must die and its carcass be eaten or otherwise so disintegrated that the ova are distributed abroad. Another interest- ing observation is the long incubation period of the ova. Confirming Bancroft, we found that the ova only became larvated after they had lain in water at least three months. Hookworms. These important parasites have been taken from several foxes: Gray Fox (Canis cinero argenteus), Arc- tic Fox {Canis lagopus), Swift Fox {Canis velox), Red Fox {Canis vulpes p ennsylv aniens) , a Gray Wolf {Canis mexicanus), divers members of the Felid£e-Eyra {Felis eyra), Jaguarundi {Felis jaguarundi), American Wild Cat {Felis ruff us), Spotted Wild Cat {Felis ruff us texensis), Ocelot {Felis pardalis), from two Giraffes {Giraffa camelopardalis, Giraffa capensis), a Malayan Tapir {Tapirus indicus), and a young California Hair Seal {Zalophus calif ornianus). It has been a most seri- ous infestment in American wild cats {Felis ruffus and Felis ruffus texensis) — animals which generally also har- bor other species of worms. In view of the petechial hemorrhages of the intestines and analogous circumstan- ces in dogs and human beings, it must be conceded that this worm is pathogenic. At this point it is fitting to note the infestment as it affects hair seals. The parasite concerned, Uncinaria lucasi, has long been a scourge among the fur seals 644 DISEASE IN WILD MAMMALS AND BIRDS {Otoes alaskanus) of the Pribiloff Islands. Its punctures are bloodless, being signalized instead by small edem- atous plaques in the intestinal mucosa. The animal we autopsied was a young California Hair Seal born in the Garden, and is singularly the only hair seal in which we have seen it. The natural habitat of the hair seal is the coast of California which means that the range of U. lucasi may extend farther southward than at first sus- pected. We have none of the northern variety. I point out two giraffe cases only because they are unique as to the organ (liver) affected. So far as I know, mature hookworms have never been reported from other organs than the intestines. From the prophylactic standpoint it mil be advisable to have as little moist earth as possible, particularly sandy ground, in and around the enclosures for the above mentioned susceptible animals because it is in such soil that the earlier stages of the life cycle of the parasite are passed. We have never found any of the human hookworm species in our animals, but it must be recognized that transmission is possible to a certain degree. Anchylostoma ceylanicum Lane (26) was found in man, cats, dogs, and a lion; Leiper(27) reports A. duodenale in a dog, and von Linstow(28) states that the latter parasite also occurs in the chimpanzee. Amebic Dysentery in Monkeys. — We recently lost six monkeys in a small outbreak of this disease — four black spider monkeys {Ateles ater), a Pinche marmoset {Leontocehus edipus), and a woolly monkey (Lagothrix lagotricha). Except for non-characteristic looseness of stools, there were no symptoms until the usual terminal lethargy set in. Li\^ng amebae were found in feces. At autopsy only the colon was found to be anatomically (26) Indian Med. Gaz., June, 1913, p. 217. (27) Jour. Trop. Med. Etc., London, 1913, XVI, p. 334. (28) Am. Med. Phila., V. 6 (16), 1903, p. 611. Fig. 78.— microscopic SECTION OF LIVER OF GIRAFFE. SHOWING SECTIONS OF UNCI- NARIA SMITHI IN BILE-DUCT AND MARKED FIBROSIS AROUND THE DUCT. Fig. 79.— colon OF MONKKV OVINC KI.KVATKI) SLOUGHS THE ANIMAL PARASITES 645 affected. It was hugely distended, fully an inch in diameter, and there were numerous confluent ulcers of the mucosa covered by a thick slough. The liver showed no abscesses. In the histological sections amebae were found in the interstices of vital gut tissue just as they are in corresponding human lesions. I have not diagnosed the species yet, but can vouch that it is not Endameha his- tolytica or coli. According to Leidy 's recommendation, grated nutmeg was administered and was followed by an improvement in symptoms. The animals became brighter and the stools firmer, but the amebae were not eradicated. Emetin hypodermically and by mouth had no obvious effects on the disease or the amebae. One monkey thus treated with nutmeg recovered, but died the next year of another affec- tion and disclosed the scars of the old ulcers in the colon. Our experience with this disease, however, is not unique. At Washington, D. C.,(29) eight spider monkeys were affected, and sporadic cases come to light from the West Coast (30), Manila, Khartoum and Ceylon. Prowazek's report concerned a young orang(31). Liver abscesses in addition to the intestinal lesions were found by three different observers. As to the transmissibility of monkey amebiasis to man, reporters are divided. Both sides are probably right, in as much as Endameha histolytica was concerned in some cases and non-human species in others. It is an infestment to be feared, and calls for examination of stools from such newly arrived animals as are known to be susceptible (spider and woolly monkeys, orangs). Parasites of Marmosets and Squirrel, Monkeys. — I give a special place to this subject because Table 24 shows that these monkeys are so commonly infested and because they are so commonly used as household pets. In this (29) Eichhorn and Gallagher, Jour. Inf. Dis., XIX, No. 3, Sept., 1916, p. 395. (30) Macfie, Ann. Trop. Med. and Parasit., 1915, 9, p. 507. (31) Arch. f. Protistenk, Jena, V. 26 (2), 22, July, p. 241. 646 DISEASE IN WILD MAMMALS AND BIRDS connection the questions arising are, first, whether the infestment is a menace to life, and second, whether it is existent outside the Garden or only acquired here. The following lists set forth the parasitic status as sho\vn at autopsy. The figures indicate how long the animal lived in the Garden : Marmosets Squirrel Monkeys Infested Not infested Infested Not infested 1 day 1 day 2 months 6 months 12 months 12 months 12 months 13 months 6-15 days ( 4 animals) 1 month ( 6 animals) 3-5 months ( 9 animals) 6 months ( 2 animals) 7 months ( 2 animals) 8 months ( 1 animal) 9 months ( 2 animals) 10 months ( 1 animal) 12 months ( 1 animal) 14 months ( 1 animal) 15 months ( 1 animal) 17 months ( 1 animal) 18 months ( 2 animals) 20 months ( 1 animal) 21 months ( 1 animal) 2 days 14 days 26 months 3 months 3 months 5 months 14 months 15 months Totals 8 animals 35 animals 3 animals 5 animals Reverting to the questions above raised, the data show that some of the animals were certainly infested on arrival here, and that others probably were; but since these animals were not examined for parasites on arrival in the Garden the duration of infestment remains unknown, and accordingly we are not justified in going farther in our conclusions. In the case of the marmosets, though, if we confine ourselves to the non-parasitized animals, it would appear that the ''acclimatization" period is within the first six months. I have attempted to arrive at a conclusion on this basis, but the average lifetime of the four parasitized marmosets which survived this period is the same as that of the sixteen non-para- sitized survivors, and we do not know at what time the parasitized ones contracted the disease. "^^ ' »ii. 3 ^^ y%^ \^*' Fig. 80. — ARACHNID (PNELiMUN'i ^^l. h lu\l; IN 1,U.\(; OF ADULT MONKEY (MA- CACUS RHESUS). IT OCCUPIES THE CENTRE OF A CYST WHICH IMMEDIATELY UNDERLIES THE PLEURA SEEN AT UPPERMOST PART OF THE ILLUSTRATION. THE ANIMAL PARASITES 647 Cysticeecus tenuicollis. — We have noted this blad- der worm in the Aoudad {Ovis tragelaphus), Red River Hog {Potamochoerus porcus), domesticated Angora Goats and several deer (Cervus alfredi, Capreolus capre- olus, Mazama mexicana, M. hemionus) located with one exception in the peritoneal cavity or membrane. One of the mule deer {Mazama hemionus) exhibited the parasite also in the lung and liver. This parasite is discussed because the very valuable Philippine spotted deer {Cervus alfredi) died from a peritonitis secondary to an infected cyst in the lesser omentum, and because the parasitism {Tcenia marginatum) is contractible from canidae which are also on exhibition in the Garden. It happens that the spotted deer did not become infested from the dogs, but it is quite probable that the goats did, since they passed many times daily in front of the wolf cages, drawing the children's carriages over the walks, and were stabled nearby. We have not discovered any of the other tapeworm cysts in deer which might be trans- mitted to them from the canidae. Camels which are parked directly opposite them have only exhibited echinocoecus cysts, yet we have never found its adult form {Tcenia echinocoecus) or its ova in the canine feces. The danger of fatal disease from C. tenuicollis, even though the infest- ment be present, is remote ; but we feel that it is better, if possible, not to exhibit the canidae adjacent to sus- ceptible animals. Pulmonary Acaeiasis in Monkeys. — We have seen but two instances of this affection in the Philadelphia Garden. The offending parasite in our animals was a new species, Pneumonyssus foxi Weidman ( 32 ) . It occurred sparingly in small cysts under the pleura and was certainly benign in our cases. The importance of the infestment consists in part in that these lesions may be mistaken for tubercles. (32) Jour. Parasit., Sept., 1915, V. 2, pp. 37-45. 648 DISEASE IN WILD MAMMALS AND BIRDS At the London Gardens (33) acariasis was found in forty-four young rhesuses dying of pneumonia, and the observers ascribed the inflammation to irritation of cer- tain doubly refractile crystals which occurred in the excreta of the mite. There are four other recorded instances of such disease in monkeys, all caused by dif- ferent species of parasites. As to pathogenesis of these arachnids, the London experience is most illuminating in that it was young rhesuses that were affected. Our specimens were mature, and nothing was stated to the contrary in the other reported cases from various parts of the world. The parasites are perhaps inhaled from the straw used as bedding, since such vegetable material is a common habitat for mites. If the resultant acute pneumonia is weathered the relics might remain only in the form of the subpleural and parabronchial cysts such as we have seen at the Philadelphia Garden. I am the more willing to accept the possibility that the simian arachnids can induce an acute pneumonia after studying a very definite case of bronchopneumonia in a prairie dog, which was induced by Cytoleichus penrosei Weidman 1916.(34) Periproventriculae FiLARro^ or Birds. — Every year we report a number of cases (up to twenty- three) of these worms, probably several species, coiled under the serosa of the air sacs and most commonly around the proven- tricle. Tentatively we have recognized two forms, a shorter (an inch or so long) and a longer (three to four inches). The latter is most inextricably coiled, but the former may be teased out. Microfilaria occur in the blood of the latter cases, but not in that of the former. The adults have been observed to penetrate from their posi- tion in the air sac serosa into the lumen of the proventricle (goose), to have caused rupture of the inferior cava (33) Proc. Zool. 8oc. Lvndon, 1919, p. 14. (34) Jour. Parasit., Dec, 1916, V. 3, pp. 82-89. Fig. 81.— arachnid (CYTOI.EICHUS PENROSEI) IN A RRONCHOPNEUMONIC FOCUS IN THE LUNG OF A PRAIRIE DOG (CYNOMYS LUDOVICIANUS). Fig. 82. — FILARIAL WORM COILED NEAR PROVENTRICLE OF A FINCH. THE ANIMAL PARASITES 649 (bulbul), to be associated with subserous cysts of the intestine (weaver) and with profound anemia (liothrix). The birds affected are mostly small, inexpensive ones, but the infestment is important because of its frequency and deserves study of the means of transmission. Physaloptera in Opossums and Badgers. — These worms were frequent findings for a period of years and were particularly impressive on account of the large number of parasites present. The stomach often con- tained scores, more or less securely attached to the mucosa by the head. The worms average an inch or two in length and perhaps an eighth of an inch in thickness. P. turgida is the only species we have identified (three examinations). As to pathogenicity we have not observed that definitely constant lesions are induced by the para- sites. In several instances the gastric mucosa has shown the mosaic appearance indicative of chronic gastritis, a condition not necessarily incited by, but certainly aggra- vated by, these worms ; at least significant is the habit of the worm to imbed its head in the gastric mucosa. In one instance the microscope has revealed a most severe fibrosis of the submucosa. The fibrosis was not so much diffuse as it was local or nodular, and in favorable places the ova of physaloptera could be discovered in the centres of the nodules, and thus betrayed the previous presence of the adult worm there. In this individual animal the case against the physaloptera is clinched by direct evi- dence. In other cases we have circumstantial evidence. Whereas it is not a. deeply burrowing parasite, it is still a penetrative one, and this is sufficient to compromise the all important * ' integrity of the mucosa, ' ' It should there- fore be considered pathogenic in all cases, because open to suspicion in several directions — abstraction of tissue juices, irritation by its products or movements and by opening up an avenue for bacterial infection. [ Tropidocerca in Birds. — This is a blood-red nematode of the size of a mustard seed to that of a peppercorn which 42 650 DISEASE IN WILD MAMMALS AND BIRDS inhabits the depths of the proventricidar mucosa. At first sight its spheroidal form suggests that of a fluke, but under the microscope it is found to be a nematode hugely ballooned out by ova, and coiled up into a ball. In spite of its dangerous appearance — being red — it is most likely quite innocuous, for microscopic sections show no sign of inflammation around the worm. Moreover, we know that a Concave Casqued Hornbill / {Dichoceros bicornis) now on exhibition has harbored the worms, as indicated by ova in the droppings, for eight years and yet seems perfectly well. I have made wax reconstructions of three of the worms and find that the coils are not very intricate and that they assume no regular or con- stant arrangement. V Syngamus Trachealis. — Our worst experience with this picturesque parasite was in common crows {Corvus b. brachyrhynchos) . In 1914 and 1915 alone we lost five such birds. Some geese, swans and a pheasant complete the short list of birds affected in addition to the crows. In no case was it a young bird that was affected. Shipley (35) reports this parasite in two grouse at the London Gardens, and Plimmer's tables show that three deaths were directly charged against them in one year(36). ' Extra-intestinal Tapeworms. — This discovery is worthy of record because it is rare for cestodes to appear anywhere save in the intestines. We have observed three instances where they had backed up into the bile duct — twice in the Cape Hyrax {Procaria capensis) and once in a Livingston's Eland {Taurotragus oryx livingstonii). At the London Gardens they were mentioned in the gall- bladder of a wallaby and in Cape Hyraces. Beddard(37) carefully describes four new species of these cestodes from the hyrax. (35) Proc. Zool. Soc. London, 1909, p. 335. (36) Loc. cit., 1912, p. 236. (37) Loc. cit., 1912, p. 576. Fu,. 8?.— PHVSALOPTERA IN STOMACH OK COMMON OPOSSLM (DIDELPHVS VTRGIMANA). THIS IS NOT AN EXCEPTIONAL OE(;REE OK INVOLVEMENT. i^PMPt' l^^M ' ;i: r'v Fir.. 84. — ONE OF THE FIBROUS NODILES IN THE GASTRIC Sl'BMlCOS.\ OF AN OPOSSUM. AN OVUM OF PHYSALOPTERA IS SEEN PRECISELY IN THE MIDDLE OF THIS ILLUSTRATION. THE ANIMAL PARASITES 651 Table 28. Occurrence of Blood Parasites. (Adapted from Plimmer, nine year period) Animals examined-1 2,241 Mammalia-2,924 Aves-6,619 Reptilia-2,( 1. Hemogregarines . 2. Microfilaria 3. Hemoproteus. . 4. Trypanosomes. 5. Plasmodia. 6. Leucocytozoa 7. Intestinal organisms* 8. Toxiplasma 9. Spirochaeta 10. Babesia 11. Haemocystidium . Host Grand Total. Reptilia Mammalia Aves Reptilia Aves Mammalia Aves Reptilia Amphibia Mammalia Aves Reptilia Aves Reptilia Mammalia Aves Reptilia Mammalia Mammalia Reptilia No. Infested 316 33 191 24 140 1 28 4 3 2 39 5 16 16 1 1 1 1 1 1 824 % Infested 11.8 1.1 3. 1. 2.1 0.003 0.4 0.6 0.2 0.5 *The exact taxonomic position could not be stated,— probably an ameba. Summary of Table 28. Mammalia. Aves Reptilia Total. Parasitized 39 415 367 821 Animals examined 2,924 6,619 2,698 12.241 1.5 6.5 — 14.0 6.7 FiLARiASis IN Wild Cats {Felis ruff us) .—This para- site was named Filaria fasciata because it coils in the fascia between the muscles— generally those of the thigh and abdomen. The worms are easily detected on skinning the animal and separating thigh and other muscles. Microfilaria were always present in the blood. The grade of pathogenicity is only conjectural. Peritoneal Filaria in Monkeys.— Thread worms have been encountered eleven times, largely in Cebidae. In 652 DISEASE IN WILD MAMMALS AND BIRDS several instances F. gracilis has been the species identi- fied, always inhabiting the peritoneal cavity, and in one instance also the lung. Microfilaria were always present in the blood. We have never seen lymphangitis or elephantiasis in our filarial cases. Blood Parasites. — I justify this paragraph on the basis of the usefulnes it might have in the clinical direc- tion, for while the taking of blood specimens is not as easy as with man it can still be done wdth some animals. From time to time we have encountered blood parasites in this Garden, but the large numbers occurring in the experience of special searchers in the London Garden and Plimmer 's particular interest in this direction make their data much the more valuable. In one report of 6,430 animals examined he found 7 per cent, infested with blood parasites of one sort or another. I have constructed the foregoing table (28) from his various reports to show which animal classes were affected by the several blood parasites. This table (28) brings out that considering them as a whole and without respect to host, just as the animals come day in and day out to the autopsy table, blood parasites will be met in 6.7 per cent, of all cases. They are seen most commonly in the form of hemogregarines of reptiles (2.5 per cent, of all animals and 12 per cent, of all reptiles) while microfilaria run a close second, being found in 2 per cent, of all animals but much more commonly in birds. Hemoproteus of birds while ranking third, should be emphasized on account of its acknowl- edged blood-destructive properties. The remaining infes- tations were too infrequent to be useful statistically. Turning to individual groups of (blood parasites, micro- filariaB of birds deserve special comment. They occurred four times more often in birds than in other animals, or, put in another way, one out of every twenty-two birds was affected,^ and only one out of every ninety other animals. ( The high figure for birds is significant in rela- Fig. 85. — AD.'VPTATION FROM RECONSTRL'Cl l()\ ol IROPl IK)CERC.\ C THE WORM LAY IN THE WALL OF THE PROVENTRICLE OF A LOUISIANA HER( TRICOLOR RUFICOLLIS). MM. . 1 ijii nil 1 2 iltiiiil. nlilifH ,,1 bl (iiilniiliiimiitliinliiitltiiiHit^ 86.— CESTODES (THREE) PROJECTING FROM THE SEVERED END OK THE DUCT OF A CAPE HYRAX (PROCARIA CAPENSIS). THE ANIMAL PARASITES 653 tion to what we have already said about periproven- tricular filaridae in our Garden, indicating that the same inf estment probably also exists in London. A point brought out by Plimmer is to the effect that, of the several blood parasites, the microfilariae were the least harmful, and that of these the adult forms were the only ones to produce symptoms; yet in one place (38) he records microfilaria as plugging the cerebral capillaries of birds. This is a very important lesion if permanent, and especially so when affecting cerebral capillaries as do the organisms and pigment of malaria. The adult forms were found in one-fourth of the cases where micro- filaria were demonstrated. As to the pathogenicity of these blood parasites in general, it will be unsafe to arrive at a definite conclusion, recalling the pitfalls that I have already outlined in dis- cussing pathogenicity of parasites in general. Keeping in mind the wonderful adaptability on occasion of animals to unfavorable circumstances we must hesitate to declare unqualifiedly the importance of even blood parasites as morbid agents. Where the parasite is known to destroy the blood cells of birds and mammals it is otherwise, but even here experimental work would be necessary to settle the question. The element of ' ' racial ' ' immunity and of phylogeny is the fly in the ointment of our deductions.) Transmission of Animal Parasitism From Wild Ani- mals TO Man. Examples of direct transmission will be only occa- sional, due to the relatively infrequent contacts between the two hosts. Pets threaten the most. Several such examples have been touched upon in the preceding pages and it but remains to gather them into one place. There is one concrete instance in the form of clear-cut simian scabies being transmitted to a keeper in this Garden (39) (38) Proc. Zool. Soc. London, 1910, p. 134. ( 39 ) Weidman ( F. D. ) , " Dermatoses of Monkeys," Arch. Derm, and Syph., Chicago, March, 1923, p. 289. 654 DISEASE IN WILD MAMMALS AND BIRDS and a similar lot fell to the keeper of a wombat at the Paris Garden (40) as well as to the taxidermist who pre- serv'ed its skin. We know that the skin and feathers of our parrots and pigeons harbor mites (41) (plumicoles of Megnin) and, recalling the occasional cases of poultry- men's itch, a transient affection might be conceded from pet parrots and other birds. Pediculi are not as num- erous on monkeys as popularly supposed — ^we see very few at the autopsy table. We have seen Trichinella spira- lis in the polar bear {Ursus maritimus) — an animal whose flesh is edible. The hydatid cysts in the camel appear un- important, but in the livers of deer it is otherwise. Neither of these infestments is dangerous if the meat is sufficiently cooked before eating. Hookworm disease points thus far only to Anchylos- toma duodencde in the chimpanzee and Uncinaria cey- lanicum in the lion and tiger. Both serve as reservoirs of the disease, the ova being discharged by way of the feces. Similarly the Strongyloides intestinalis infest- ment which we have seen in the orang might be trans- ferred to man. Indirectly, Europeans traveling in Africa have made the crucial test that certain ungulates and other wild animals of Africa are the reservoirs of Try- panosoma gamhiense, the parasite of the well known African sleeping-siclmess ; for this example the blood stream of the beast is the reserv^oir and a biting insect the means of transmission. The above examples are cited to emphasize the pos- sibility that parasites of wild animals may have a patho- genic significance for man. They do not exhaust the subject. Many more instances might be cited but the fore- going bring out the important ones which have come to our attention. (40) Railliet Traite de Zool. Med. et Agric. Paris — Asselin et Houzeau, 1895, p. 659. (41) Megnin, Les Parasites Articules, 1895, Masson et Cie, Paris. THE ANIMAL PARASITES 655 Treatment. The recognition of the existence of parasites during the life of an animal, especially those of the sldn and intestinal tract whose discovery is easiest, suggests that some means of combating them should be employed. But we are by now quite satisfied that medicinal and disin- fective therapeutic procedures, while they ha\^e their field of usefulness, are much less to be depended upon for the protection of exhibits than are preventive meas- ures of general hygienic nature. Under the latter head- ing come the prompt removal of excreta, frequent changes of drinking water, routine examinations of feces of cer- tain varieties, autopsy examinations and incineration of autopsy remains — all of which are part of the require- ments of common cleanliness and general disease preven- tion. I wish to amplify the matter of disposal of feces and general cage-police. Our ideas as to what constitutes thoroughness in this work have changed considerably since Fulleborn's recent demonstration that ascarid ova (42) could live in formaldehj^de for four or five years, and the older one of Galli-Valerio (43) that those of Hepaticola hepatica lived one month in 2 per cent, formal- dehyde solution. Evidently the same substances which disinfect do not invariably disinf est ; and if the occasion should arise for the most exacting control in tliis respect, a special investigation of the susceptibility of the indi- ^4dual ova in question would have to be undertaken. In addition to these general measures we have put up certain special safeguards against parasites. Thus, each specimen of the large Carnivora (lions, tigers, leop- ards, etc.), has received routinely a dose of santonin every month over a period of several years. We have no figures on which to base comparison with previous periods, but an examination of feces of all the inmates (42) Quoted by Jensen (V.), Hospitalstidende, Copenhagen, 1922, 65, No. 28. p. 457. (43) Centr. f. Bakt. u. Parasitk., (etc.), Jena 1— Abt. V. 35 (1), 5, 1903, orig. p. 89. 656 DISEASE IN WILD MAMMALS AND BIRDS of the Camivora house in 1916(44) showed that less than one-third of the animals were infested, and of these all save the jaguars showed either small numbers of ova in the feces or relatively non-pathogenic forms. The jaguars had been badly infested for many years with dibothriocephalus. Prior to this examination we had been under the impression that nearly every one of the felidae ordinarily was infested and if this impression was well founded, due credit must be given, in company with general hygienic precautions, to the routine santo- nin dosages. It goes without saying that where animals are detected at autopsy with unequivocal transmissible and dangerous parasites (coccidia, amebae, etc.), the con- tacts are isolated, examined and if necessary treated for the affection or even sacrificed. To continue the preventive measures, it would be most desirable to examine at least the blood and feces of all newly arrived animals, but at present this is not practi- cable on account of the labor involved in the laboratory and in collecting the material, and because all animals do not stand the restraint involved when blood specimens are being taken. At present we are limiting special ex- aminations to the droppings of newly arrived parrots and toucans for Spiroptera incerta and to the feces of certain monkeys for amebae. Further preventive measures will depend on the nature of individual infestments as they crop up. Food inspection, screening, sulphur dips, etc., are but a few examples of what might be found necessary hygienically after investigating or establishing the life cycle of our numerous parasitic groups. However we cannot forbear to emphasize again the value of the blast lamp and of paint in the hygiene of animal enclosures — ^means we believe to be much more potent and quite as practicable as chemical disinfectants. (44) Phila. Zool. Soc. Rep., 1917, p. 36. Fig. 87. — TRICHINELLA SPIRALIS IN MUSCLES OF POLAR BEAR (URSUS MARI- TIMUS). THIS WAS AN OLD INFESTMENT. AS INDICATED BY THE THICK AND HYA- LOID CHARACTER OF THE CAPSULE. THE ANIMAL PARASITES 657 Turning now to the active curative side of the sub- ject, what medical means we have against parasites ap- pertain for the most part to the intestinal ones. The treatment of tapeworms is very hazy and unsatisfactory — areca nut is perhaps more useful in animals than any one other drug. For round worms santonin is most to be depended on although turpentine is useful against the round worm of the Equidae. The dosage of santonin per month has been— for large bears, ten grains ; for lions, tigers, large pumas, six grains; for jaguars, leopards, hyenas, four grains ; for wild cats, etc., two grains. The dose of areca nut recommended for Carnivora is two grains per pound of body weight. Since ung-ulates do not stand areca nut well, iron sulphate may be used. For animals the size of a horse the dosage is two drams, and to this one or two grains of arsenic trioxide may be added. On the basis of very carefully controlled experiments on dogs. Hall recommends carbon tetrachloride for hook- worms in these animals — 0.3 mils per kilo of body weight, without purging. Its efficacy has been confirmed lately but we have not had the occasion to test it. Prom time to time we have broached other lines of medication against worms which may be worth while re- lating if for nothing more than to illustrate the uncertain ways of our vermifuges when applied to wild animals. I can speak first of thymol as employed on parrots parasitized by Spiroptera incerta. The first thing that impressed us was the large dosage which birds could en- dure. The lethal dose for pigeons was four grains, sus- pended in mucilage of acacia. After we had established that certain parrots withstood fourteen grains in muci- lage, we administered on one occasion twelve grains and on another sixteen grains, suspended in glycerin. The drug is reputed to be absorbed when exhibited in the latter vehicle and we hoped to get a certain anthelmintic effect on the parasites from the blood side as well as from the lumen of the gut. The bird itself, a very heavily 658 DISEASE IN WILD MAMMALS AND BIRDS infested cockatoo, showed no ill effects and passed two dead female spiroptera and enormous numbers of ova. But thereafter it passed even greater numbers of ova than before (we estimated 182,000 per day for this bird over a five day period and 288,000 on a single subsequent day), and was obviously unimproved by the treatment. The explanation of failure was clear, for the worms can retire into the protecting mucus or mucous membrane lining the proventricle until the thymol has passed by, and even though paralysed may not be flushed out. In a later test on a toucan which died twenty minutes after thymol administration we found at the autopsy that worms deeply imbedded in the proventricle were translucent and motionless from the effects of the thymol-glycerin mixture, i.e., saturated with the medi- cament and apparently dead. Twenty minutes later they were placed in normal salt solution in the incu- bator, and next morning were found actively motile. Thymol evidently does not kill — it only stupefies, and in the absence of means for flusliing the parasites out, as we do in human hookworm cases, this class of vermifuge will have to be abandoned in work against tliis parasite. Not with any serious hope of success, but feeling that arsenic was the most promising drug available for parenteral use, we tried atoxyl hypodermically and arsphenamine intravenously but without success. The only positive results were to emphasize the tolerance of some lower animals to arsenic. Thus in preliminaiy work pigeons received sixty drops of Fowler's solution by mouth without embarrassment, but five minims killed a pigeon when administered hypodermically. The organic arsenical, arsphenamine, w^as withstood intravenously by pigeons in six times the proportional human dosage. One of our drug trials was instructive in that it worked quite a different effect from that in man, besides being most amusing. In earlier diagnostic work on spiroptera we tested the practicability of examining the THE ANIMAL PARASITES 659 vomitus for the worms, hoping thereby to get a greater concentration of ova, which would facilitate the micro- scopic examination. Hypodermic injections of apomor- phine (0.1 grain) into an amazon did not induce vomiting from the gizzard as hoped — only a regurgitation from the crop, but it did cause some dizziness and most ludicrous talking and laughter. To illustrate further the difficulties of animal medi- cation I quote our experience with four red howling monkeys {Alonatta seniculus). One of these died of intes- tinal obstruction from large ascarids — the case which has been already cited. Ova were found in the stools of the remaining three, and one of the monkeys was treated twice mth santonin. It died in thirty hours after the second dose — not of santonin poisoning, for none of the clinical symptoms were present, but most likely from ab- sorption of toxic substances originating in the decom- posing ascarids wliich crowded the gut. It profits not to destroy these parasites, then, unless we feel assured that they may thereafter be removed immediately. If, for the sake of brevity, I were asked to state in a single sentence the practical status of animal parasitic disease in this Zoological Garden I would put it thus: Since there are various animal parasitic diseases con- tinuously present here of which we know, and since fresh ones are from time to time cropping out, and since these are on the whole of economic importance, it behooves us to continue and extend our efforts against an issue extant — somewhat through therapeutic means, but far more through clinical laboratory examinations, careful au- topsy searches, and by rigid general hygienic measures such as cage-police, new quarters, isolation, or if neces- sary, destruction of the exhibit. PROPf N. C, . INDEX Abortion, 305 Abscess of liver, 231 of lung, 155 Acariasis, lungs, 647 of monkeys, 647 Actinomycosis, 138, 568 in deer, 368, 568 tapirs, 568 treatment, 570 Adenoma, 474 Adrenal body, 336 Alimentary tract, 166 Amblyopia, 403 Amoebae, 606, 644 dysentery from, 644 Amyloid, liver, 227 spleen, 128 Anatomy of labor, 290 Anchylostomum, see hookworms Anemia, 87 primary, 98 secondary, 88 Aneurysms, 65, 80 Animal Parasitism, hygiene, 656 prevention, 656 treatment, 655 Animal Parasites, 614 disappearance of, 627 frequency, 628 of groups, 633 in blood, 652 incidence, 628-636 modes of action, 617 occurrence in wild, 627 transmission animals to man, 653 visceral distribution, 637 Angina pectoris, 49 Aorta, 72 fatty deposits in, 71 Arteries, 66 Arteriosclerosis, 71 Arteritis, 70 Arthritis, 347 gouty, 347, 411 Ascending nephritis, 276 Aspergillosis, 558 Aspergillus, varieties, 558 Ataxia, 375 Atrophy, acute of liver, 228 Autopsy Ust, 47 Avian spiropteriasis, 172, 640 Bacterial flora, 418 Basal cell carcinoma, 475 Beriberi, 439 Bihary tract, 225 calculi, 238 Birth canal, 287, 296 comparative anatomy, 287 et seq obstructions to, 306 Blackhead, 206 Bladder, gaU, 224, 238, 239 urinary, 286 Blood, diseases of, 83 Blood formation in birds, 98 Blood vessels, 66 Bone marrow, 83, 109, 111 Bones, diseases of, 343 effects of trauma, 343 tumors of, 368 Botryomycosis, 564, 602 Botuhsm, 604 Brain, 385 tuberculosis of, 378 tumors of, 384 weight of, 385 references to, 387 Breast, 312 Bronchi, 141 Bronchiectasis, 144 Cage palsy, 349 CalcuU, bihary, 238 renal, 282 Carcinoma, 476 basal cell, 475 Cataract, 403 Cecum, 211 Cestodes. 637 Cholangitis, 239, 256 Cholecystitis, 239 Choledochitis, 239 Cholehthiasis, 238 Chondroma, 472 Cloaca, 211 Coccidiosis, 606 Cirrhosis of hver, 232 Comparative anatomy of uterus, 287 of pelvis, 297-303 Conjunctivitis, 402 Constipation, 209 Constitutional diseases, 410 Convulsions, 373 Cornea, 403 Coronary arteries, 49 Cowper's gland, 313 661 662 INDEX Cretinism, 320. 331 Cvsticercus tenuicollis, 647 Cystitis, 286 Cytoleichus penrosei, 647 Deficiency diseases, 438-443 Degenerations of kidney, 269 of liver, 228 Diabetes, 412 Diet, carnivorous, 452 herbivorous, 452 grain, 455 seed, 454 soft, 453 omnivorous, 402 relation to disease, 415 alimentary tract, 417 Dilatation of heart, 54 Diphtheria, 600 Dislocations, 345 Distemper, 599 Diverticula of intestine, 219 Diverticulitis, 219 Dysentery, amoebic in monkevs, 644 Dystocia, 292 Ear. 409 Echinococcus, 647 Emphysema, 161 Encephalomyelitis, 380 Endocarditis, 52 Endometritis, 305 Endothehoma, 165, 474 Enteritis, 177 in Aves, 202, 205 Mammalia, 185 Enterohepatitis, 605 EpitheUoma, 475 Esophagus, 169 Exophthalmic goitre, 320, 323, 329 Eye, 402 tuberculosis of, 402 Fallopian tubes, 305 Fat infiltrations of kidney, 268 hver, 226 metabolism, 445 Fibroma, 472 Filaria, fasciata, 651 gracilis in monkeys, 651 in blood, 652 fascia, 651 muscles, 651 wildcats, 651 periproventricular, 648 peritoneum, 651 Food, 415 definition, 415 in relation to alimentary tract, 417 Food, disease, 422 poisoning, 457 Fowl cholera, 598 plague, 598 typhoid, 598 Fractures, 344 Gall stones, 238 Gas bacillus infection, 602 Gastritis. 204 GastroenterocoUtis in Ungulata, 194 in MarsupiaUa, 198 Gangrene of lung, 155 Giraffe, hookworm in, 644 Gout, 53, 410 Heart, dilatation of, 54 hypertrophy of, 54 diseases of, 48 effects of, 55 effect of strain, 55-59 weight of, 63 relative vulnerabihty of, 61 Hemorrhagic septicemia, 598 Hemoglobinuric fever, 603 Hemorrhoids, 218 Hepaticola hepatica, 641 Hepatitis, 228 Hernia, 216 Heterakis in avian ceca, 606 Hookworms, 643, 654 in giraffe, 644 Hypernephroma, 339, 341, 342, 475 Hypertrophic periosteitis, 346 Hyperthyroidism, 320 Hypertrophy of heart, 54 in Aves, 60 Hypothyroidism, 320 Ileus, 213, 261 Infantihsm, 433 Infiltrations of kidney, 268 liver, 226 Inorganic salts in diet, 427 Intestinal obstruction, 212 tract, 177 inflammation of, 181 mechanical obstruction of , 212, 617 relation to food, 422 Intestines, diverticula, 219 tumors of, 220 IridocycUtis, 402 Kangaroo disease, 570 bacteriology, 576, 586 course of attack, 573 pathology, 575 prevention, 572 treatment, 591 r INDEX 663 \ Kidney, 263 abscess, 268, 278 calculi, 282 degenerations of, 269 hemorrhages, 271 hypertrophy of, 267 infiltrations of, 268 tumors, 284 weight of, 265 Labor from a comparative stand- point, 290 obstructions to, 306 Laryngitis, 139 Larynx, 138 Leontiasis ossium, 359, 472 Leucemia, 104 in birds, 108 lymphatic, 105 myeloid, 109 Leucocytes, 84-86 Limberneck of ducks, 604 Lipoma, 472 Liver, 222 abscess, 231 acute atrophy, 228 amyloid, 227 cirrhosis, 232 degenerations, 228 fatty changes, 226 infiltration, 226 inflammation, 228 chronic, 232 necrosis in, 230 tumors, 240 Lungs, 146 abscess, 155 congestion, 148 gangrene, 155 infarct, 160 tumors of, 162 Lymphadenitis, 117 Lymphatic leucemia, 105 tissue, 114 hyperplasia of, 115 in pharyngeal wall, 115, 138 Lymph nodes, 114 tuberculosis of, 121 tumors of, 122 Lymphomatosis, 118 Malnutrition, 424 Mammary gland, 312 Marmosets, parasites of, 645 Marrow of bone, 83, 109, 111 Meningitis, 376 Metabolism, carbohydrate, 443 fat. 445 inorganic, 427 protein, 447 Miliary tubercle, avian, 512 bovine, 510 human, 511 monkey, 511 Miscarriage, 305 Molluscum contagiosum, 601 Monckeberg sclerosis, 74, 76 Monkey's temperature, 520-528 Moon blindness, 405 Muscles, 370 Mycosis, 137, 558 of esophagus, 168 histology of, 561 hygiene, 563 incidence, 562 of lung, 562 method of action, 560 pharynx, 168, 564 types of, 560 Myelitis, 350, 381 Myeloma, 111 Myocarditis, 52 Myocardium, 49, 50, 65 Myxcedema, 320, 331 Necrosis, liver, 230 spleen, 130 Nematodes, 636 Neoplasms, 462 incidence of, 463, 468 embryonic origin, 471 in captivity, 469 in the wild, 462, 476 metastasis, 471 visceral origin, 477 Nephritis, 271 ascending, 276 effects of, 280 histology of, 279 toxic, 275 Nervous system, 372 Nocardia macropodidarum, 585 Nocardiosis, 570 Obesity, 446 Ophthalmia, periodic, 405 Osteitis, 346 Osteitis deformans, 359, 431 Osteoma, 368 Osteomalacia, 349 Ovary, cysts, 307 Pachymeningitis, externa, 331, 377 Paget's disease, 359, 431 Pancreas, 244 degenerations, 250 tumors, 259 Pancreatitis, 250 Parasites, see animal parasites, 614 Parovarian cyst, 307 664 INDEX Pasteurelloses, 597 Pearl disease, 491, 501, 505 Pellagra, 441 Pelvis, comparative anatomy, 297-303 Penis, 313 Pericarditis, 53 Pericardium, position of effusion in, 54 Perio&teitis, hypertrophic, 346 Periproventricular worms, 648 Perisplenitis, 131 Peritoneum, 260 tumors, 262 Peritonitis, 260 Pharyngitis, 168 Pharynx, 168 Phimosis, 313 Physaloptera turgida, 649 in opossums, 649 Plants, poisonous, 459 Pleura, 163 Pleuritis, 164 Pneumonia, 149 broncho, 152 fibrinous, 151, 153 in Aves, 153 origins of, 154 lobar, 151 Pneumonokoniosis, 159 Pneumonyssus foxi, 647 Poisonous plants, 459 PoliomyeUtis, 380 Prostate gland, 313 enlargements of, 314 tuberculosis of, 315 tumors of, 314 Proventricle, 171 worms in, 172, 640 Psittacosis, 208, 597 Pyelonephritis, 277 Quail disease, 608 Rabies, 602 Rachitis, 349, 429 Rectum, prolapse of, 218 Reproductive organs, female, 287 male, 317 Respiratory tract, 134 Rhinitis, 135 Rickets, 349, 429 Renal calculi, 282 Salpingitis fallopii, 305 Santonin, 657 Sarcoma, 471, 474 Scurvy, 440 Seminal Vesicles, 315 Sinusitis, 135 Skeleton, 343 Spinal cord, 373 Spiroptera incerta, 638, 640 detection, 640 eradication, 640 in parrots, 172, 208, 640 Spiropteriasis, 172, 640 Spleen, 114, 122 amyloid, 128 congestions, 125 enlargements, 124 hemorrhage, 125 inflammation, 126 in anemia, 130 in hepatic cirrhosis, 130 necrosis, 130 size, 124 tuberculosis of, 132 Squirrel monkeys, parasites of, 645 Starvation, 425 Stomach, 174 tumors of, 176 ulcers of, 175 Streptothricosis, 567 Suprarenal body, 336 Syngamus trachealis, 140, 650 Taenia echinococcus, 647 Tape worms, 637 in Uver, 650 Temperature of monkeys, 520-528 Testes, 313 tumors of, 313 Tetanus, 602 Thrombosis, 69 Thymol, 657 Thymus, 120, 336 Thyroid body, 316 atrophy of, 330 hyperplasia of, 325 size of, 318 tumors of, 333 Tonsils, 115, 138 Trachea, 140 Tropidocerca contorta, 649 Tubercle bacillus, types of, 513 Tuberculin test on monkeys, 518 other animals, 549 dose, 529 effect on kidneys, 548 eye, 546 reaction, 530 skin, 546 Tuberculoma, 505 Tuberculosis of brain, 378 avian characters, 503, 512 Carnivora, 498 control, 514-548 diagnosis of, 514 discovery during life, 514 distribution in birds, 504 91 INDEX 665 Tuberculosis of eye, 402 gelatinous, 504 histology, 510 hygiene, 516 in Aves, 503 in MammaUa, 492 in Primates, 492 in various avian orders 506-510 incidence, 489 intestinal in birds, 505 Lemures, 495 lymph nodes, 121,494 nonsusceptible animals, 490 ordinate characters, 492 frequency, 489 pathological type, 490 Proboscidea. 502 Rodentia, 499 routes of infection, 485 Tuberculosis, sanitation of cages, 516 susceptible animals, 490, 515-517 Ungulata, 500 visceral distribution, 491 Tumors, see neoplasms Ulcer, gastric, 175 Uncinaria, 643 Uremia, 281 Urethra, 315 Uterus, comparative anatomy, 287 inflammation, 305 tumors of, 308 Vitamins, 438 Waterfowl epizootic, 604 Zoological list, 43 A LIST OF THE PUBLICATIONS FROM THE LABORATORY OF COMPARATIVE PATHOLOGY OF THE PHILADELPHIA ZOOLOGICAL SOCIETY 1909—1923 1. Results of Tuberculin Tests in Monkeys at the Philadelphia Zoological Garden, by C. Y. White, M.D. and Herbert Fox, M.D. The Archives of Internal Medicine, December, 1909, Vol. 4, pp. 517-527, Chicago, Illinois. 2. Note on the Occurrence of a Ciliate {Opalinopsis nucleolobata, n.s.) in the Liver of a Mammal (Canis latrans), by Allen J. Smith, M.D. and Herbert Fox, M.D. University of Pennsylvania Medical Bulletin, February, 1909, Philadel- phia, Pennsylvania. 3. The Tuberculin Test in Monkeys: with Notes on the Temper- ature of Mammals, by Arthiur Erwin Brown, D.Sc. C.M.Z.S., Sec. Zool. Soc, Phila. Proceedings of the Zoo- logical Society of London, 1909, pp. 81-90. 4. Observations on the Occurence of Neoplasms in Wild Animals, by C. Y. White, M.D. and Herbert Fox, M.D. Proceedings of the Pathological Society of Philadelphia, February, 19 10. 5. Observations on the Comparative Anatomy of the Female Genitalia, by Edward A. Schtimann, M.D. American Journal of Obstetrics and Diseases of Women and Children, Vol. LXIV, No.. 4, 191 1, New York. 6. Observations Upon Neoplasms in Wild Animals in the Phila- delphia Zoological Garden, by Herbert Fox, M.D. The Journal of Pathology and Bacteriology, Vol. XVII. (191 2), pp. 217-231. England. 7. A Study of Metazoan Parasites Found in the Philadelphia Zoological Garden, by Fred D. Weidman, M.D. Proceed- ings of the Academy of Natural Sciences of Philadelphia , March. 19 13, pp. 126 to 151, Philadelphia, Penna. 8. The Pathology of the Thyroid Gland in Wild Animals, by Herbert Fox, M.D. Journal of Comparative Pathology and Therapeutics, Vol. 27, p. 23. Edinburgh, Scotland. A LIST OF PUBLICATIONS 9. The Mechanism of Labor From the Standpoint of Comparative Anatomy, With a Report of Cases of Dystocia in Wild Animals, by Edward A. Schumann, M.D. American Jour- nal of Obstetrics and Diseases of Women and Children, Vol. LXIX, No. 3, 19 1 4, New York. 10. Cirrhosis of the Liver in Wild Animals, by Herbert Fox, M.D. New York Medical Journal, December 19, 19 14. 11. The Dynamics of the Female Pelvis; Its Evolution and Architecture with Respect to Function, by Edward A. Schumann, M.D. American Journal of Obstetrics and Dis- eases of Women and Children, Vol. LXXI, No. i, 1915, New York. 12. Pneumonyssus foxi, Nov. Sp. An Arachnid Parasitic in the Lung of a Monkey (Macacus rhesus), by Fred D. Weidman, M.D. Journal of Parasitology, September, 19 15, Vol. II, pp. 27-45, Urbana, Illinois. 13. Cytoleichus penrosei, A New Arachnid Parasite Found in the Diseased Lungs of a Prairie Dog, {Cynomys ludovicianus) . Journal of Parasitology, December, 1916, Vol. Ill, pp. 82-89, Urbana, Illinois. Fred D. Weidman, M.D. 14. A Method of Obtaining Duplicate Reconstructions from the One Series of Wax Plates, by Fred D. Weidman, M.D. New York Medical Journal, March 3, 191 7, New York. 15. Papers: Read at the Meeting of the Pathological Society at the Philadelphia Zoological Garden. Pancreatitis in Wild Animals, by Herbert Fox, M.D. Report of an Enzootic of Parasitic Proven tricular Worms {Spiroptera incerta, Smith) of Parrots, with Control of Same, by Fred D. Weidman, M.D, Coccidium bigeminum. Stiles, in Swift Foxes (habitat Western U S.), by Fred D. Weidman, M.D. Distribution of Uncinaria Among the Lower Animals, by Fred D. Weidman, M.D. An Arachnoid {Pneumotuber macaci, Landois and Hoepke ?) Parasitic in the Lungs of a Monkey {Macacus rhesus), by Fred D. Weidman, M.D. A Note Upon the Lesions of the Female Genitalia in Wild Animals, by Edward A. Shumann, M.D. A LIST OF PUBLICATIONS Amblyopia in a Young Monkey (Macacus nemestrinus) , by H. M. Langdon, M.D. and W. B. Cadawalder, M.D. Remarks on Examinations of a Series of Brains, by W. B. Cadawalder, M.D. Journal of Comparative Pathology and Therapeutics, December, 1915, Vol. XXVIII. Part 4, pp. 298-336, Edinburgh, Scotland. 16. Reversionary Pseudobile Canaliculi Formation in the Cirrhotic Liver of a Vulpine Phalanger, by Fred D. Weidman, M.D. New York Medical Journal, March 9, 19 18, New York. 17. A Contribution to the Anatomy and Embryology of Cladorchis (Stichorchis) Subtriquestrus, Rudolphi, 18 14 (Fischoeder, 1901), by Fred D. Weidman, M.D. Parasitology, Vol. X, No. 2, January 22, 191 8, Cambridge University Press, London, England. 18. Nutritive and Blood Changes in Rats on Cancer-Inhibiting and Cancer-Stimulating Diets, by E. P. Corson-White, M.D. Pennsylvania Medical Journal, March, 19 19, Vol. XXII, p. 348, Athens, Penna. 19. Pemphigus in an Orang Utan Infested with Strongyloides (intestinalis ?) and Dying from Advanced Tuberculosis, by Fred D. Weidman, M.D. Journal of Cutaneous Diseases, March, 1919, Vol. XXXVII, pp. 169-173, Chicago, 111. 20. Arteriosclerosis in Wild Animals, by Herbert Fox, M.D. American Journal of Medical Sciences, June, 1920, No. 6, Vol. CLIX, p. 821, Philadelphia, Penna. 21. Osteomalacia in Wild Animals, by E. P. Corson-White, M.D. Archives of Internal Medicine, November, 1922, Vol. 30, pp. 620-628, Chicago, Illinois. 22. Osteitis Deformans in Monkeys, by E. P. Corson-White, M.D. Archives of Internal Medicine, December, 1922, Vol. 30, pp. 790-796, Chicago, Illinois. 23. Certain Dermatoses of Monkeys and an Ape, by Fred D. Weidman, M.D. Archives of Dermatology and Sy philology, March, 1923, Vol. 7, pp. 289-302, Chicago, Illinois. 24. Acute Papular and Desquamative Exanthem in an Orang Utan, by Herbert Fox, M.D., and Fred D. Weidman, M.D. Archives of Dermatology and Syphilology, April, 1923, Vol. 7, pp. 462-464, Chicago, Illinois. (C-^ N. MANCHESTER. '^■^ INDIANA,