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British Library Cataloguing in Publication Data Avian medicine 1.Avian medicine I. Tully, Thomas N. II. Lawton, Martin P. C. III. Dorrestein, Gerry M. 636.5'089
Library of Congress Cataloguing in Publication Data Avian medicine/edited by Thomas N. Tully, Jnr., Martin Lawton, Gerry M. Dorrestein. p.cm. Includes bibliographical references and index. ISBN 0 7506 3598 3 1. Avian medicine. I. Tully, Thomas N. II. Lawton, Martin P. C. III. Dorrestein, G. M. SF994.A93 2000 636.5'089–dc21 00–036020
ISBN 0 7506 3598 3
Composition by Genesis Typesetting, Laser Quay, Rochester, Kent Printed and bound by MPG Books Ltd, Bodmin, Cornwall Acknowledgements
The completion of an international text is a We are also grateful to our colleagues in our substantial achievement. Only a cohesive practices who supported us during this team can accomplish what one person can- endeavour. Drs. Mark Mitchell and J. Jill not. We the editors want to first thank the Heatley at LSU, Drs. Lynne Stoakes, Stephen authors, without whom there would be no Divers and Julie Austin in the UK and Dr. Avian Medicine handbook. The idea of an Marein van der Hage in Utrecht were all instru- international text to call upon the expertise mental in the completion of this text. The edi- of a global avian community was encour- tors would also like to thank Marion Jowett for aged by our publisher. Wanting this expertise her belief in this project from the beginning. and having them commit to this project Of course this work could never have come with their busy schedules were two different to fruition without the constant nurturing, matters. Fortunately, we were able to secure commitment and work of our publisher, But- their commitment and feel the veterinary terworth-Heinemann. There are a few special community as a whole will benefit from this people we would like to thank for their help work. and efforts, Mary Seager, Senior Commission- There was a large support staff which was ing Editor, for being the captain and running an invaluable asset in formatting the original this ship, and Angela Davies, Medical Desk manuscript. Joseph A. Smith, whose hard Editor, and Caroline Savage, Editorial Assis- work and dedication will serve him well in his tant, for making the impossible task of putting chosen profession, veterinary medicine, led the original manuscript into a publishable text this support staff team. Others who made possible. significant contributions to this project Finally, a heartfelt thank you to our wives, included Harry M. Cowgill, photographer; Susie Tully, Lynne Stoakes and Marein van der Michael L. Broussard, graphic artist; Sam Hage and families who helped us survive the Moran and the staff at the Greater Baton ups and downs of editing a multi-author Rouge Zoo and Joshua R. Pinkston. medical text. Foreword
‘The night when I came home I was much troubled to hear that my poor Canary Bird, that I have kept these three or four years, is dead’ Samuel Pepys (1665)
Of all animals, those in the Class Aves – the In 1903, Arthur G. Butler produced his Hints birds – have most fascinated the human race. on Cage-Birds (British and Foreign) and in the Their almost ubiquitous distribution, the Preface declared that ‘Aviculturists in the tendency of many species to live in villages, present century ought to do better work than towns and cities and the ease with which their predecessors....’. Butler went on to say some can be tamed or trained: these and other ‘Now that men who have laboured and features have led to birds occupying a very suffered are recording the results of their special place in the lives and traditions of work..., their successors may confidently look various cultures. for brighter and more prosperous times, start- Interest in the biology and health of birds is ing where the pioneers left off they may not new. Two thousand years ago, Pliny, continue to add to the sum of knowledge, Aristotle and others studied birds, taught and may hope eventually to bring the science others and published their observations. In of birdkeeping to something approaching the 17th century European anatomists carried perfection’. out painstaking dissections of many species Nearly a hundred years later, it cannot be and were able to produce scholarly descrip- claimed that the ‘science of birdkeeping’ has tions that have, in a number of cases, with- yet approached perfection, but the advances stood the test of time and are still valid today. have been enormous. Over the past two A century later John Hunter, the great sur- decades our knowledge and understanding of geon, anatomist and champion of the veteri- bird diseases in particular has progressed nary profession, explored the links between extraordinarily and much of the credit for this structure and function in animals and is owed to the ‘pioneers’, amongst them the amongst many other great discoveries, descri- editors and many of the contributors of bed the air sac system and studied factors that this book, who have turned avian medicine influenced the healing of fractures in avian into a bona fide, state-of-the-art, scientific species. discipline. Medical care for birds is also not new. As It gives me particular pleasure in this long ago as 1486 advice on the diagnosis and Foreword to note and to commend the book’s treatment of falcons was given in the Boke of St international orientation. The Editors are from Albans written by Dame Juliana Berners – a three different countries and the authors of reminder of the key role in avian science chapters span even more. This spread reflects played by women, even 500 years ago! The the global importance of avian medicine, in popularity of the sport of falconry over the terms not only of treating individual sick succeeding three centuries ensured that pub- birds but also of promoting the health of wild lications on raptors continued to appear. (free-living) avian populations, particularly There was interest in other types of bird – see those that are under pressure for other rea- the quotation from Pepys at the beginning of sons. The part that the veterinary profession this Foreword – but it was only in the 19th can play in conservation is now widely century that serious attention began to be paid recognized and a Resolution at last year’s to the diseases of those kept in cage and (1999) meeting of the European Committee of aviary. the Association of Avian Veterinarians urged x Foreword international bodies and policy makers to treat birds. Those of us who care about – or for recognize that role and to incorporate veter- – the class Aves owe them a debt of gratitude inarians in their programmes. In my own for all they have done to promote a better work overseas I have witnessed the contribu- understanding of these animals and their tions that can be made by those with a sound requirements. This book will serve as a lasting knowledge of aviculture and pathology, medi- legacy of their work, and at the same time, cine and surgery. Thus, for example, the will encourage others to contribute to the survival and recovery of the Mauritius Kestrel challenging and exciting field of avian (Falco punctatus) owes much to the collabora- medicine. tion in the 1970s between veterinarians and biologists who, often unfunded, worked to Professor John E. Cooper save the species from extinction. This study DTVM, FRCPath, FIBiol, Dip ECAMS, FRCVS has been repeated on a number of occasions Consultant Veterinary Pathologist since then and augurs well for the future. Durrell Wildlife Conservation Trust Tom Tully, Martin Lawton and Gerry Dor- (Jersey Zoo) restein are familiar names to all who keep or February 2000 Contributors
Alan Abrey BVSc Patricia Macwhirter BVSc (Hons), MA, Consultant/Director, Zoological Collections, FACVSc Durban, South Africa Principal, Highbury Veterinary Clinic, Burwood, Victoria, Australia Janette Ackermann DVM, MS Chief, Veterinary Services, American Wildlife Foundation, Molalla, Oregon, USA Glenn H. Olsen DVM, MS, PhD Veterinary Medical Officer, US Department of Interior, US65 Patuxent Wildlife Research Brian H. Coles BVSc, FRCVS, Dipl ECAMS Center, Laurel, Maryland, USA 4 Dorfold Way, Upton, Chester, Cheshire, UK Patrick T. Redig DVM, PhD Gerry M. DorresteinDVM PhD Dipl Vet Associate Professor and Director, The Raptor Pathology Center, University of Minnesota, St. Paul, Associate Professor in Avian, Exotic Animal Minnesota, USA and Wildlife Pathology, Utrecht University, Utrecht, The Netherlands Ian Robinson BVSc, Cert. SAP, MRCVS Peter de Herdt DVM, DVSc RSPCA Norfolk Wildlife Hospital, Station Professor of Veterinary Medicine, University Road, East Winch, Kings Lynn, Norfolk, UK of Gent Director of Clinic for Poultry and Special Andrew Routh BVSc, MRCVS Animal Diseases, Merelbeke, Belgium RSPCA Norfolk Wildlife Hospital, Station Road, East Winch, Kings Lynn, Norfolk, UK Luc Devriese DVM Senior Researcher in Veterinary Bacteriology, University of Gent, Merelbeke, Belgium Stephanie Sanderson MA, VetMB, MRCVS RSPCA Norfolk Wildlife Hospital, Station Nigel H. Harcourt-Brown BVSc, FRCVS, Road, East Winch, Kings Lynn, Norfolk, UK Dipl ECAMS 30 Crab Lane, Bilton, Harrogate, North Yorkshire, UK Katja Trinkaus DVM, FTA Staff member, Clinic for Avian and Reptile Don J. Harris DVM Diseases, Institute for Avian and Reptile Avian and Exotic Animal Medical Center, Diseases, Justus-Liebig-University of Miami, Florida, USA Giessen, Giessen, Germany
Maria–Elisabeth Krautwald-Junghanns Thomas N. Tully, Jnr DVM MS Dipl ABVP PD Dr Med Vet, Dr Med Vet Habil, (avian) Dipl ECAMS FTA Associate Professor, Louisiana State Professor, University of Leipzig, Leipzig, University School of Veterinary Medicine, Germany Baton Rouge, Louisiana, USA Martin P.C. Lawton BVetMed CertVOphthal CertLAS CBiol MIBiol DZOOMed FRCVS Amy B. Worell DVM, Dipl ABVP (avian) RCUS Specialist in Exotic Animal Medicine, Director, All Pets Medical Centre, West Hills, Exotic Animal Centre, Romford, Essex, UK California, USA Introduction
There have been a number of surveys per- over the last 20 years avian medicine and formed by veterinary research groups concern- surgery has made its way into mainstream ing the number of avian patients seen per week academia and veterinary continuing educa- in the average small animal practice. In those tion programmes. practices that see avian and exotic patients, on To add to the existing knowledge base of average two to three bird cases are seen each proceedings and journal articles, a number of week. Most of the veterinary surveys show reference avian texts have been published, as that the vast majority of avian owners, approx- well as periodicals that keep practising veter- imately 80 per cent, do not seek veterinary care. inarians up to date with the latest trends in There is therefore great potential for veter- avian medicine. This information is especially inarians to offer health care for these untreated useful to the high volume avian veterinarians. birds and maintain them as healthy pets for However, what about the majority of veter- their owners. To garner this population of inarians, who see only a few birds a week and potential avian clients, there must be com- enjoy the variety of patients but do not want petent caring avian veterinarians. to specialize? The general public has a basic conception Avian Medicine has been developed to give that veterinarians do not know how to treat the average small animal practitioner a com- birds, or that a sick bird is a dead bird. This plete information source for the basics of avian public perception is slowly changing in the medicine and surgery. It is a multi-author text, face of significant medical advances in the which utilizes the international expertise of the field of avian veterinary medicine made over avian veterinary community. By incorporating the last 20 years. These advances, and the the knowledge of authors from Europe, Africa, expertise of avian specialists, have been Australia and North America, different tech- acknowledged by the aviculture community niques that have been successful in treating the and by lay avian publications such as Bird Talk average avian patient are covered. magazine. These groups of bird owners are The first five chapters cover the basic just the tip of the iceberg; we are looking to medical information needed to run a primary treat and care for all birds, and this can only care avian practice. Again, the focus of the text occur when the general public is aware of the is on introductory level material and the ability and availability of avian veterinarians average companion animal practice. If you see to care for their pets. between one and five birds a week, this text is Private practitioners founded the primary for you. avian veterinary organization, the Association The later chapters are species-specific. If a of Avian Veterinarians (AAV), in 1980. The practice sees avian patients, it is not unusual majority of AAV members have always been to treat any one of these species. The informa- clinical veterinarians. These practice owners tion in these chapters will allow the veter- saw an unfulfilled need in the veterinary inarian a comfort zone of knowledge in order community, and took up the challenge of to evaluate, treat and/or refer. meeting the avian pet owners’ demands for We hope the book reaches the groups of adequate health care for their pets by gaining veterinarians and allied technical staff who knowledge and experience and sharing it with need it the most; also veterinary students, like-minded colleagues. Since the veterinary veterinary technicians and the general veter- schools and colleges at that time were woe- inary practitioner. fully inadequate in teaching companion avian Thomas N. Tully, Jnr. medicine, veterinarians had to learn through Martin P.C. Lawton conferences and wet-labs. Slowly but steadily Gerry M. Dorrestein
1 Basic anatomy, physiology and nutrition
Patricia Macwhirter
Introduction
In companion animal practice, veterinarians opportunity to observe birds in their natural traditionally treat dogs and cats – species that habitat. By thinking beyond the examination are significant because of the bonds they form room, veterinarians can gain insight into with their human owners. Avian practice medical and behavioural problems of indi- shares the focus on the human–animal bond, vidual patients as well as playing a role in but in addition avian practitioners routinely trying to preserve avian biodiversity for treat cage and aviary birds, zoo birds, wild future generations to experience and enjoy. birds, racing and fancy pigeons, ducks, rap- tors, ratites and poultry. A fundamental chal- lenge for the avian practitioner is to be able to offer quality patient care across this wide Feathers range of species as well as to understand the needs and expectations of their owners. There Feathers are unique to birds, and evolved is an added dimension to this discipline, from reptilian scales. They are arranged in because many of the species that practitioners feather tracts called pterylae. The unfeathered treat are currently threatened or endangered. areas between the pterylae are called apteria Sadly, in some cases avicultural pressure has (Figure 1.1). contributed to the demise of species in the New feathers develop from the epidermal wild. However, there is a strong movement collar at the base of the feather follicle, and among caring and responsible avian veter- grow in an upward and outward manner. inarians to develop global links with bird This development process is sensitive, and owners and conservation groups to try and nutritional deficiencies, stress or exogenous redress habitat damage and ensure the sur- cortisone can cause horizontal fault lines to vival of wild as well as pet bird populations. occur on the emerging feather (Plate 1). Because of the diversity of species encoun- The central stalk or shaft of the feather is tered, avian medicine is best mastered by composed of the calamus or quill, which is learning about one family of birds in some present below skin level, and the rachis, above detail and then drawing comparisons (McLel- the skin. In developing feathers there is a land, 1991; Rubels et al., 1992; Smith and nutrient artery that traverses the centre of the Smith, 1992). In this chapter, avian anatomy, shaft and is surrounded in the calamus by physiology and nutrition are reviewed using feather pulp. The feather pulp and artery the psittacine species as a basic model; how- regress as the feather matures, but keratinized ever, where appropriate, there is comparison pulp caps remain as horizontal bars across the with other avian species likely to be encoun- lumen of the shaft (Figure 1.2). If an emerging tered in veterinary practice. When studying or feather is cut or damaged, it may bleed treating birds, it is important to consider how profusely from the nutrient artery (a so-called each species has evolved and take every blood feather). 2 Avian Medicine
Emerging laterally at 45° to the rachis are the barbs, and emerging from the barbs at 45° are barbules. These interlock with each other by a system of hooks (or barbicels), which enables waterproofing. Preening restores interlocking if the barbules become dislodged. The vane is the portion of the feather extend- ing either side of the rachis, and may be plumulaceous (soft and fluffy) or pennaceous (closely interlocked), depending on the indi- vidual type of feather (Figure 1.3). Contour feathers are those that cover the bird’s body. They have well-developed shafts with lumulaceous and pennaceous compon- ents to the vane. Covert feathers are small contour feathers present on the wings and tail. Flight feathers, or remiges, are generally asymmetric in form to enable flight, and the vane is pennaceous. The primary remiges are those that emerge from the periosteum of the metacarpus, while the secondary remiges are those that emerge from the periosteum of the ulna. Primary remiges are numbered from the carpus distally, while secondary remiges are numbered from the carpus proxi- Figure 1.1 The unfeathered apteria may provide skin exposure without removing feathers to provide a site for mally. The tail feathers (or retrices) are struc- subcutaneous fluid administration or, as shown here, for turally similar to remiges, but are symmet- intradermal skin testing. rical. They are numbered from the centre laterally.
(b)
(a)
Figure 1.3 A typical avian feather showing Figure 1.2 Line drawing of a developing feather. (a) plumaceous and (b) pennaceous vane structure. Basic anatomy, physiology and nutrition 3
Birds have other feathers adapted for special barb. These usually lie near feather tracts to aid purposes. Plume (or down) feathers have a in insulation. Filoplumes are hair-like, with a rachis shorter than the longest barb, and non- long rachis and short barbs at the end. Bristles interlocking barbules. These provide the have a stiff rachis with few or no barbs at the undercoat. Powder downs are specialized end, and function like whiskers. After feathers down feathers located anterior to the hips, (hypopnea) are small remnant feather struc- which produce a keratin material from the tures attached to the shaft at the superior surface cells of barb-forming tissue. This umbilicus, the position where the calamus performs a dry lubrication function. Powder emerges from the skin. downs grow continuously, and are often the first feathers to show abnormalities if a bird is infected with circovirus. Powder down feath- Integument ers are most prominent in cockatoos, cockatiels and African grey parrots, so owners prone to Bird skin is thinner and more delicate than feather allergies should avoid these species. that of mammals. In most cases the subcuta- Overgrowth of powder down feathers can neous layer is insufficient to suture surgically. occur if a bird is prevented from preening – for The only epithelial glands are the uropygial example, if it is collared or suffering from beak glands and holocrine glands of the external damage. Semiplume feathers have fluffy ear canal, but the keratinocytes produce vanes, but the rachis is longer than the longest lipid, essentially making the entire skin an
(a)
(b)
(1) (2) (3) Figure 1.4 Line drawings of the uropygial gland (preen gland) representing various avian species. (a) Dorsal view of the gland on a White Leghorn chicken. (b) Details of papilla: (1) delicate type; (2) compact type; (3) unique passerine type. 4 Avian Medicine oil-producing holocrine gland. Because they position. In passerines, digit I points anteriorly lack sweat glands, hyperthermic birds dis- while digits II–IV point in the posterior sipate heat by increasing respiration and direction. holding the wings away from the body. Cold A bird’s feet and legs are covered with scales birds ruffle their feathers and crouch to retain – raised areas of highly keratinized epidermis heat. Normal body temperature for birds is separated by a fold of less keratinized skin. higher than for mammals (generally 40–44°C). Claws have a strongly keratinized dorsal plate Bilobed uropygial glands are found dorsally and a softer ventral plate that grows more at the base of the tail on most psittacine species, slowly, thus producing the curved claws for except for Amazon parrots and some Colum- perching. Vitamin A deficiency, e.g. from an all biformes (Figure 1.4). Each lobe is drained by a seed diet, can produce loss of the plantar single duct, which empties into a lone papilla. footprint as well as hyperkeratosis and over- The gland secretes a lipoid sebaceous material, grown claws. Scaly mite (Cnematocoptes sp.) which is spread over feathers during preening. can also induce hyperkeratosis and disrupt Impaction (which can be treated by manual normal scale formation. expression) or neoplasia are occasionally encountered. Uropygial glands are not essen- tial, and can be removed surgically. A brood patch forms on birds incubating Senses eggs. The dermis on the chest becomes thick- Sight ened and vascular, and feathers are lost. This structure should not be mistaken as Birds possess great visual acuity. Most birds’ pathological. eyes are very large, the globe is asymmetrical, and there are 10–18 ossicles (eyebones) that overlap to form a ring encircling the sclera. The sphincter and dilator muscles of the pupil Legs, claws and feet are striated; therefore, unlike mammals, atro- pine has no effect. In psittacines, digits II and III point anteriorly A soft lens allows for rapid accommodation. while digits I and IV point in a posterior The retina is thick and has no direct blood
Figure 1.5 Cross-section of the avian globe showing the pecten as it projects into the posterior chamber. Basic anatomy, physiology and nutrition 5 vessels but many cones, which allow for The digestive system excellent colour vision, while the fovea is well developed and, in some avian species, mul- Beak tiple, allowing for good visual acuity. The pecten, which projects into the vitrous body The beak consists of the bones of the upper from the optic nerve and consists of capillaries and lower jaws and their keratinized sheaths, and extravascular pigmented stromal cells, the rhamphotheca (Figure 1.6). In psittacines, provides nutrients to the avascular retina the jaws are connected to the skull by kinetic (Figure 1.5). In some birds, small regular joints. The upper jaw is called the rhinotheca torsional movements of the eye sweep the and the lower jaw is the gnathotheca. These pecten through the vitreous (Myer, 1986). are composed of modified epidermis, with Birds’ dorsal and ventral eyelids do not cells of the stratum corneum containing free blink, and there are semiplumes rather than calcium phosphate and hydroxylapatite crys- eyelashes. There are no meibomian glands, tals as well as abundant keratin. Beak tissue and the third eyelid blinks in a nasal to extends outwards towards the surface, over temporal direction. Eye movements are lim- much of the beak, extending to the edges ited, but this is compensated for by greater and tip. In cases of chronic nasal discharge, head mobility and by the ability of both eyes vertical defects in the rhinotheca leading to move independently. from the nares to the tomia (the cutting edge of the beak) are occasionally seen. Hearing Oropharynx Birds have no pinna. Instead, specialized contour feathers (ear coverts) cover the Birds have no soft palate. The opening to meatus. The tympanic membrane projects the nasal passageway, the choana, is a slit outwards rather than inwards, and the col- umella is the only bony ossicle (compared with three ossicles in mammals). The cochlear duct is short when compared to that of mammals. Because of these differences, birds have good discrimination of pitch (frequency) but are less sensitive to higher and lower tones than humans. Temporal resolution is about 10 times faster than in humans.
Taste
Taste buds are present on the roof and floor of the oropharynx, but not on the tongue. Par- rots, budgerigars, hummingbirds and nectar feeders actively select sugar solutions. Insec- tivorous and granivorous species may not be particular in selecting food for taste.
Smell
The sense of smell in psittacines and passer- ines is, in general, poorly developed. In other species – for example, kiwis, vultures and albatrosses – the sense of smell may be well Figure 1.6 The large beak of a green winged macaw developed. (Ara chloroptera). 6 Avian Medicine
Figure 1.7 The choanal slit is the oral nasal interface located in the dorsal aspect of the oral cavity.
between the palatine folds of the roof of the Stomachs mouth (Figure 1.7). Caudally directed papillae are abundant in this region. The infundibular The cranial proventriculus (or glandular cleft, which opens to the auditory tube, is stomach) is thin walled and is lined with situated immediately caudal to the choana. mucus-secreting columnar epithelial cells and Parrots have intrinsic muscles in the tongue, oxynticipeptic cells, which secrete hydrochlo- while other birds have only extrinsic muscles. ric acid and pepsinogen. The caudal ven- Salivary glands produce mucus rather than triculus or gizzard lies ventrally on the left watery secretion, and are abundant on the side of the abdomen, with the pylorus joining walls of the oropharynx. Vitamin A deficiency the duodenum just to the right of the midline. may produce hyperkeratosis of these glands. The proventricular–ventricular junction is a common site for inflammation associated with megabacteria infections in budgerigars. Oesophagus and crop The anatomy of the stomachs relates to the bird’s diet. In granivorous birds, glands in the The oesophagus goes down the right side of thick, biconvex, muscular gizzard wall secrete the neck – opposite to its situation in mam- a hard, proteinaceous, cuticular lining, the mals. The crop is a dilatation of the oesopha- koilin. This helps to pulverize food in pre- gus in which food is stored and softened prior paration for proteolysis in the intestinal tract. to passage into the proventriculus. No diges- The koilin is usually yellow or green due to tion takes place in the crop. Food enters the bile pigment regurgitation, and the gizzard crop on the right and exits caudally on the generally contains grit. In finches, gizzard midline, where the oesophagus extends to the worms (Acuaria sp.) are sometimes found proventriculus. Enlarged thyroid glands or beneath the koilin lining. lesions due to Trichomonas spp. can cause Birds that consume predominately soft food obstructions in this region and induce vomit- or nectar, such as the lorikeets, have poorly ing, crop dilatation or respiratory distress due developed muscular gizzards. On the other to pressure on the trachea. hand, birds consuming large prey items com- A full crop can be viewed under the skin at pared to their body size (for example pen- the base of the neck in nestlings without guins, which feed on whole fish) may have feathers, or in adults that have lost feathers. enormous stomachs relative to their body size. Alarmed owners will sometimes present birds, As a normal digestive process, the raptors not realizing that this structure is normal. regurgitate castings composed of bones and Basic anatomy, physiology and nutrition 7
developed capillary system instead. Many species, particularly poultry, ducks and quail, have large paired caeca, but these are absent or vestigial in psittacines, passerines and Colum- biformes. Caeca are the site of blackhead (histomoniasis) infection in gallinaceous birds. The rectum is usually short and straight. The liver consists of two lobes joined cranially. The gall bladder is usually absent in psittacines and ostriches, but is present in many other species. It receives bile from the hepatocystic duct from the right lobe only. The vitelline diverticulum (Merkel’s diver- ticulum) is the remnant of the yolk duct, and is located opposite the cranial mesenteric artery. In altricial birds the yolk sac is gen- erally resorbed by 4 days, while in precocial Figure 1.8 Regurgitated casts produced in the proventriculus of raptor species. species it may persist for up to 10 days. fur of their prey species from their gizzards Cloaca and vent (Figure 1.8). The cloaca consists of: Intestines the cranial coprodeum, which receives fae- ces from the rectum The duodenum, jejunum and ileum are located the middle urodeum, which receives the on the right side of the abdomen. Folds and oviduct or ductus deferens and the ureters villi do not contain lacteals, but have a well- on the dorsolateral surface
Figure 1.9 Cross-section of the cloaca. 8 Avian Medicine