Miller and Evans'
Anatomy of the Dog
FIFTH EDITION

John W. Hermanson, PhD
Associate Professor
Department of Biomedical Sciences
College of Veterinary Medicine
Cornell University
Ithaca, New York

Alexander de Lahunta, DVM, PhD
James Law Professor of Anatomy, Emeritus
Department of Biomedical Sciences
College of Veterinary Medicine
Cornell University
Ithaca, New York

Howard E. Evans, PhD
Professor of Veterinary and Comparative Anatomy, Emeritus
Department of Biomedical Sciences
College of Veterinary Medicine
Cornell University
Ithaca, New York

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Table of Contents
Cover image
Title Page
Copyright
Dedication
Special Dedication
About the Authors
Preface
Anatomical Terms
Planes of the Body
Movement
Literature
Acknowledgments
Contributors
Former Contributors in Past Editions
1 The Dog and Its Relatives
The Order Carnivora

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The Family Canidae
Breeds of Dogs
Bibliography
2 Prenatal Development
Early Development

Length of Gestation
Prenatal Periods
Oocyte–Embryo
Embryo
Age Determination
Fetus
Bibliography
3 The Integument
Epidermis
Dermis
Structure of the Dermis and Changes with Age
Pigmentation
Nasal Skin
Digital Pads
Hairy Skin
Muscles of the Skin
Glands of the Skin
Blood Supply to the Skin
Nerve Supply to the Skin
Skin Grafting
Claw

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Bibliography
4 The Skeleton
General
Axial Skeleton

Appendicular Skeleton
Bibliography
5 Arthrology
General
Ligaments and Joints of the Skull
Ligaments and Joints of the Vertebral Column
Ligaments and Joints of the Ribs and Sternum
Ligaments and Joints of the Thoracic Limb
Ligaments and Joints of the Pelvic Limb
Bibliography
6 The Muscular System
Introduction
Skeletal Muscles
Muscle Description
Muscles of the Head
Muscles of the Neck
Muscles of the Dorsum
Muscles of the Thoracic Wall (Musculi Thoracis)
Muscles of the Abdominal Wall (Musculi Abdominis)
Muscles of the Tail (Musculi Caudae)
Fasciae of the Trunk and Tail
Muscles of the Thoracic Limb

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Muscles of the Pelvic Limb
Bibliography
7 The Digestive Apparatus and Abdomen

Oral Cavity
Pharynx
The Alimentary Canal
Liver
Pancreas
Bibliography
8 The Respiratory System
External Nose
Nasal Cavity
Larynx
Trachea
Bronchi
Thoracic Cavity and Pleurae
Lungs
Bibliography
9 The Urogenital System
Urinary Organs
Reproductive Organs
Embryologic Characteristics of the Urogenital System
Bibliography
10 The Endocrine System
General Features of the Endocrine Glands

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The Hypophysis
Thyroid Gland
Parathyroid Glands

Pineal Gland
Adrenal Gland
Pars Endocrina Pancreatis
Enteroendocrine Cells
Endocrine Tissues of the Ovary
Fetal Membrane Endocrine Tissues
Endocrine Tissues of the Testis
Endocrine Cells of the Kidney
Bibliography
11 The Heart and Arteries
Pericardium and Heart
Pulmonary Arteries and Veins
Systemic Arteries
Bibliography
12 The Veins
General Considerations
Cranial Vena Cava
Veins of the Thoracic Limb
Caudal Vena Cava
Portal Vein
Veins of the Pelvic Limb
Veins of the Central Nervous System
Bibliography

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13 The Lymphatic System
General Considerations

Ontogenesis of the Lymphatic System
Lymph Drainage
Lymph Vessels
Innervation of Lymph Vessels and Lymph Nodes
Lymphoid Tissue
Lymph Nodes
Hemal Nodes
Lymph Nodules
Regional Anatomy of the Lymphatic System
Bibliography
14 Introduction to the Nervous System
General
Functional Components of Nerves
Reflexes
Bibliography
15 The Autonomic Nervous System
General Visceral Efferent System
Enteric Nervous System
Bibliography
16 The Spinal Cord and Meninges
The Spinal Cord
Morphologic Features of the Spinal Cord
Spinal Cord Segments

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Segmental Relationships to Vertebrae
Gray Matter of the Spinal Cord

White Matter of the Spinal Cord
Spinal Reflexes
Transverse Sections of the Spinal Cord
Meninges, Brain Ventricles, and Cerebrospinal Fluid
Bibliography
17 The Spinal Nerves
Initial or Primary Branches of a Typical Spinal Nerve
General Features of Spinal Nerves
Cervical Nerves
Nerves to the Diaphragm
Brachial Plexus
Thoracic Nerves
Lumbar Nerves
Sacral Nerves
Caudal Nerves
Bibliography
18 The Brain
The Brainstem
The Subthalamus
The Cerebrum
Basal Nuclei
The Cerebellum
Brain Atlas
Bibliography

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19 Cranial Nerves

Olfactory Nerve (Cranial Nerve I)
Optic Nerve (Cranial Nerve II)
Oculomotor Nerve (Cranial Nerve III)
Trochlear Nerve (Cranial Nerve IV)
Trigeminal Nerve (Cranial Nerve V)
Abducent Nerve (Cranial Nerve VI)
Facial Nerve (Cranial Nerve VII)
Vestibulocochlear Nerve (Cranial Nerve VIII)
Glossopharyngeal Nerve (Cranial Nerve IX)
Vagus Nerve (Cranial Nerve X)
Accessory Nerve (Cranial Nerve XI)
Hypoglossal Nerve (Cranial Nerve XII)
Cutaneous Innervation of the Head by Noncranial Nerves
Bibliography
20 The Ear
The Internal Ear
The Middle Ear
The External Ear
Bibliography
21 The Eye
Development
The Eyeball
The Eye as an Optical Device
Orbit
Eyelids

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Lacrimal Apparatus
Muscles
Innervation
Vasculature
Comparative Ophthalmology
Bibliography
Index

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Copyright
3251 Riverport Lane
St. Louis, Missouri 63043
MILLER AND EVANS' ANATOMY OF THE DOG, FIFTH EDITION  ISBN: 9780-323-54601-0
Copyright © 2020, Elsevier Inc. All rights reserved.
Previous editions copyrighted 2013, 1993, 1979 and 1964.
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Dedication
Sandy de Lahunta and Howard Evans collectively were on the anatomy faculty
from 1950 to 2005 at Cornell University. This represents a remarkable span of
time during which they mentored a multitude of students and worked together as

colleagues. With this fifth edition, we recognize the contribution of Professor Evans
in the title: Miller and Evans' Anatomy of the Dog.

Cornell Graduation, 2005

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Special Dedication

Malcolm E. Miller, BS, DVM, MS, PhD (1909–1960). Dr. Malcolm E. Miller
was born on a farm in Durrell, Pennsylvania; studied for two years at
Pennsylvania State University; and then earned his BS and DVM (1934), MS
(1936), and PhD (1940) degrees from Cornell University. He was appointed
Instructor in 1935 and, at the time of his death, was Professor and Head of
the Department of Anatomy and Secretary of the New York State
Veterinary College at Cornell University. His zest for life, devotion to his
family, and his enjoyment of teaching and anatomical research sustained
his spirit through several brain operations which provided only temporary
relief from epileptic attacks.
This volume was envisioned by Dr. Miller in 1944 as a comprehensive
treatise documenting the morphology of the dog. His efforts were aided
considerably by the encouragement of Dean W.A. Hagan and the
appointment of a medical illustrator in 1946. Preliminary work with the

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help of his wife, Mary, resulted in the 1947 publication by Edwards
Brothers Press of Miller's Guide to the Dissection of the Dog, which now
appears as Guide to the Dissection of the Dog, eighth edition (2017) by Evans
and de Lahunta.

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About the Authors
Howard Evans was an undergraduate student in Entomology at Cornell University
when he was called to active duty in the Army during World War II in 1943. Upon
completion of 3 years of service as a Second Lieutenant and graduation from Cornell
in absentia, he returned as a graduate teaching assistant in Comparative Anatomy.
He received a PhD in 1950, with a thesis on the anatomy of Cyprinid fishes. That
same year he was hired by Dr. Malcolm Miller as an Assistant Professor in the
Veterinary College. He taught Veterinary Anatomy for 36 years until his retirement
in 1986. During his tenure, he taught Dog Anatomy, Horse and Cow Anatomy, and
the Anatomy of Fish and Birds. As an emeritus professor, he has taught a course on
Natural History to veterinary students at Cornell for 12 years. For 2 or 3 weeks each
year since 2002, he has taught Fish and Bird Anatomy and Natural History at St.
George's University Veterinary College in Grenada, West Indies.
He was a co-author of the first and second editions of Miller's Anatomy of the Dog
and the sole author of the third edition. The fourth edition with Dr. de Lahunta was
the first to have all illustrations in color. He is co-author with Dr. de Lahunta of
Guide to the Dissection of the Dog, which is in its eighth edition. Evans has written
chapters in other texts on tropical fish anatomy, bird anatomy, ferret anatomy, and
woodchuck anatomy. His research has concerned fetal development of Beagle dogs,
cyclopia in sheep, and the replacement of teeth in fishes. His sabbaticals were spent
at the Veterinary College in Davis, California, learning surgical techniques for fetal

studies of dog development; the University of Hawaii, teaching Comparative
Anatomy; the Medical School of the University of Pennsylvania, studying fetal
development of sheep; and the Marine Station of the University of Georgia, studying
the anatomy of the Spotted Sea trout.
His interest in natural history led to giving courses to veterinary students at
Cornell and at St. George's University in Grenada, West Indies. He and his wife Erica
have led many Cornell University trips to Africa, Hawaii, New Guinea, and
Antarctica. He has continued to enjoy his interactions with students and sharing the
anatomical collections in his office almost daily since his retirement in 1986.
Alexander de Lahunta (“Sandy” to his colleagues, or “Dr. D.” as he was known to
his students) received his DVM from the College of Veterinary Medicine at Cornell

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University in 1958. After 2 years of mixed practice in Concord, New Hampshire, he
returned to the Department of Anatomy at Cornell where he obtained his PhD. For
the next 42 years he taught gross anatomy of the dog, veterinary neuroanatomy and
clinical neurology, applied anatomy, embryology, and neuropathology.
He established a consultation service in clinical neurology in the Teaching
Hospital and was a founding member of the Neurology Specialty of the American
College of Veterinary Internal Medicine (ACVIM).
He was chairperson of the newly developed Department of Clinical Sciences. He
received numerous awards for his teaching and the Robert W. Kirk Distinguished
Service Award of the ACVIM. He retired in 2005 and has been active in textbook
writing since that time.
In addition to this text, he has authored or co-authored a fourth edition of
Veterinary Neuroanatomy and Clinical Neurology, an eighth edition of Miller's Guide to
the Dissection of the Dog, The Embryology of Domestic Animals: Developmental

Mechanisms and Malformations, Veterinary Neuropathology, and Applied Anatomy.
John Hermanson was an undergraduate at the University of Massachusetts,
Amherst, where he first became interested in comparative vertebrate anatomy as a
student of the late David Klingener. He received an MS from Northern Arizona
University in 1978, and a PhD from the University of Florida in 1983. It was at
Florida where he first engaged in teaching veterinary anatomy at the College of
Veterinary Medicine in Gainesville. After postdoctoral studies at Emory University
(neuroscience) and at the University of Pittsburgh School of Medicine (muscle
regeneration), he began his career at Cornell University in 1987. He has taught
anatomy of “whatever is in the barnyard” since that time, including Horse Anatomy,
Ruminant Anatomy, Bird Anatomy, and comparative anatomy that caters to lab
animal, zoo animal, and wildlife conservation interests. His research has spanned bat
biology, equine biomechanics, and paleontology.

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Preface
The first edition of this text was based on an unfinished manuscript with
illustrations that Professor Miller had been working on for many years prior to his
death in 1960. At the request of his wife, Mary Miller (Ewing), the manuscript was
completed by Howard E. Evans and George C. Christensen. The first edition of
Anatomy of the Dog by Miller, Christensen, and Evans appeared in 1964. “Mac” Miller
had supervised the preparation and completion of almost all of the illustrations by
Pat Barrow and Marion Newson that appeared in the first edition.
The second edition, entitled Miller's Anatomy of the Dog, by Evans and Christensen,
was published in 1979. It incorporated the most recent nomenclature of Nomina
Anatomica Veterinaria and had new chapters on fetal development, the endocrine
system, the spinal cord, and the eye. There were also many new drawings by Marion

Newson, Lewis Sadler, and William Hamilton.
The third edition of Miller's Anatomy of the Dog, by Evans, updated the literature,
added new material, and incorporated nomenclatorial changes that appeared in
Nomina Anatomica Veterinaria 1983. Many figures were modified structurally or
relabeled, and several were replaced. There were chapters by eight new contributors,
and the introductory chapter was expanded to include the phylogenetic
relationships of canids to other carnivores and the history of domestication of the
dog. The chapter on muscles was augmented to include current histochemical and
electrophysiologic evidence of muscle function. The material on the nervous system
was amplified by new chapters on the brain and cranial nerves.
Miller's Anatomy of the Dog attempts to meet the varied needs of anatomists,
veterinary students, clinicians, and experimentalists. Throughout the text, the intent
is to describe and illustrate the specific morphology of the dog, with reference to
older literature. Although there are many similarities between species, it is often
surprising how different anatomical specifics can be. What is a functional structure
in one may be only a vestige or absent in another, and care is required when
extrapolating to other species.
The fourth edition was the first fully colorized version, and it required removal of
all former labels (which were made with LeRoy lettering stencils, something
unknown to new generations of students) and dotted lead-lines. Some of these
illustrations appeared in the eighth edition of Guide to the Dissection of the Dog (2017,

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Elsevier/Saunders) by Evans and de Lahunta.
This fifth edition includes many new radiographs and computed tomography (CT)
scans that were kindly provided by Dr. Peter Scrivani, supplementing a number of
radiographs from earlier editions that were contributed by Dr. Vic Rendano. New

digital imaging technology allows an ever-improving insight into the structure of the
dog's body. Michael Simmons provided new illustrative interpretations of a number
of images of the lymphatic system, replacing images that had grown darker with
each subsequent printing. Illustrations were added or updated in Chapter 21.
Nomenclatural revisions were included as appropriate, and we attempted to retain
some synonyms where their use would clarify discrepancies with human medical
anatomy texts and other veterinary anatomy texts. New research was referenced
throughout the text to enhance our interpretations of anatomic structures.
We appreciated correspondence received and offer our thanks to the many people
who pointed out errors or suggested improvements. The illustrations have been
particularly well received, and requests for their reuse in books and journals stand as
a tribute to the dissectors and illustrators whose combined efforts produced them.

Anatomical Terms
The terms used for structures of the body are numerous, and, in the course of
medical history, about 50,000 names have been given to some 5000 structures. This
has led to considerable ambiguity.
The history of anatomical terminology shows gradual regional changes from the
Arabic to the Greek of Hippocrates, Aristotle, and Galen to the Latin of Vesalius,
Fallopius, Eustachius, Fabricius, and Malphighi, when the center of medical
education shifted to Italy. Some Arabic terms, such as “saphena” and “nucha,”
remain, as do several Greek terms, some with Latin endings. Each country often
used different endings for Latin terms, and later vernacular terms made the medical
vocabulary unusable internationally. (For example, the hypophysis, or pituitary
gland, had at least 30 names in Greek, Latin, German, French, and English.)
For any meaningful communication, it is necessary that anatomical terms be clear
and precise. With this in mind, international anatomical nomenclature committees
sponsored by various anatomical societies have published Nomina for humans
(Nomina Anatomica [NA], 1989 and Terminologia Anatomica [TA], 1998), and Nomina
for domestic animals (Nomina Anatomica Veterinaria [NAV], 2005), and (Nomina

Anatomica Avium [NAA], 1993) to promote international communication and
facilitate learning. There are lists of Nomina for gross, histologic, and embryologic
terms.
Several anatomists, including Burt Green Wilder, MD, Professor of Physiology,
Vertebrate Zoology, and Neurology at Cornell and Secretary of the Committee on
Anatomical Nomenclature of the Association of American Anatomists, tried
(between 1880 and 1890) to standardize anatomical nomenclature, but their results

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lacked international agreement. In 1887, the German Anatomical Society undertook
the task and assembled an international committee that worked for 6 years before
issuing a final list in 1895. This Basel Nomina Anatomica (BNA) for the human
included about 5000 terms from approximately 30,000 proposed by the
subcommittee (O'Rahilly, 1989, “Anatomical terminology, then and now.” Acta Anat.
134:291–300). This BNA formed the basis for subsequent revisions in Birmingham
(BR) in 1933, Jena (JNA) in 1936, and Paris in 1955. The latter was published as the
first edition of the NA. In 1977, the fourth edition of the NA included Nomina
Histologica (NH) and Nomina Embryologica (NE). The current NA for the human is in
its sixth edition. In 1989, the International Federation of Associations of Anatomists
created a new committee to write Terminologica Anatomica (TA), which was
published in 1998 by Thieme. Both the NA and the TA are only for human anatomy.
A committee on veterinary anatomical nomenclature was established in 1895, at
the Sixth International Veterinary Congress in Bern because the BNA was not
applicable to domestic animals. At the next Veterinary Congress in Baden-Baden in
1899, a nomenclature for domestic animals was approved but not printed or
distributed internationally, although the terms were used in several textbooks. In
1923, the American Veterinary Medical Association published the Nomina Anatomica

Veterinaria (NAV) based on the BNA, but it was not widely known or used. In 1957,
the International Association of Veterinary Anatomists established a nomenclature
committee that incorporated the earlier unpublished lists of the American
Association of Veterinary Anatomists with terms in current use. After several
preliminary lists and many meetings in different countries, the first edition of the
internationally approved NAV was published by the World Association of
Veterinary Anatomists in 1968. This Nomina is currently in a sixth edition (2017) and
is available for free on the worldwide web.
Although anatomical structures are quite stable, our understanding and
interpretation of what we see will continue to require changes at all levels: gross,
microscopic, and ultrastructural. Formal procedures exist for making changes in
anatomical terms, and the International Committee for Veterinary Anatomical
Nomenclature of the World Association of Veterinary Anatomists welcomes
suggestions and help.
The terminology used in this text follows NAV 2005, with subsequent
recommendations (NAV 2017). We also consulted Constantinescu and Schaller's
Illustrated Veterinary Anatomical Nomenclature (2012) as we considered terminology
used in this edition. The following constraints serve as guidelines in the work of
anatomical nomenclature committees:
1. Each anatomical concept should be designated by a single term. Synonyms
have been used in rare exceptions, usually as transitional terms, but in some
cases both terms may be used such as: peroneus = fibularis.
2. Each term should be in Latin (Greek remains in some terms: ischiadic [G.] =

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sciatic [L.]; splen [G.] = lien [L.]).
3. Each term should be as short and simple as possible.

4. The terms should be easy to remember and should have instructive and
descriptive value.
5. Structures that are closely related topographically should have similar
names (e.g., femur: femoral artery, vein, and nerve).
6. Differentiating adjectives should generally be opposites (major/minor;
superficial/deep).
7. Terms derived from proper names (eponyms) should not be used because
the choice of the eponym has varied by country and was not descriptive of
the structure (e.g., Eustachian tube = auditory tube; canal of Schlemm =
scleral venous sinus; foramen of Monro = interventricular foramen).
Directional terms as applied to quadrupeds are different from those applied to
humans. The anatomical position of a standing dog is with four paws on the
supporting surface and the abdomen ventral. For a human, the standing position is
with the forelimbs hanging by the side, palms held forward: the palms and the
abdomen are thus considered to be anterior. For the dog, the terms cranial and caudal
apply to the neck, trunk, and tail as well as to the limbs as far distally as the end of
the antebrachium and crus. The terms for the forepaw or manus are dorsal and
palmar; those for the hindpaw or pes are dorsal and plantar. On the head the terms
rostral, caudal, dorsal, and ventral are preferred. Only in a few locations, such as the
jaws, eye, and inner ear, are such terms as anterior, posterior, superior, and inferior
used. Medialis and lateralis apply to the whole body except on the digits, where axialis
and abaxialis refer to the sides of the digit toward the axis of the limb or away from
the axis of the limb, respectively. The axis of the limb passes between the third and
the fourth digits.

Planes of the Body
The planes of the body are formed by any two points that can be connected by a
straight line.
Median Plane: Divides the head, body, or limb longitudinally into equal right
and left halves.

Sagittal Plane: Passes through the head, body, or limb parallel to the median
plane.
Transverse Plane: Cuts across the head, body, or limb at a right angle to its
long axis or across the long axis of an organ or a part.
Dorsal Plane: Runs at right angles to the median and transverse planes and
divides the body or head into dorsal and ventral portions (see Fig. PF1.1).

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FIG. PF1.1 (From Guide to the Dissection of the Dog, by Evans and de Lahunta,
Elsevier, 2017.)

Movement
Parts of the body can move relative to one another primarily because of muscular
action on bones articulated with each other by joints. Flexion is the movement of one
bone on another so that the angle between them is reduced; thus, the limb, digit, or
vertebral column is bent, folded, retracted, or arched. Extension is the lengthening of
a part by increasing the angle between bones, straightening the limb, digit, or
vertebral column. Extension beyond 180 degrees is overextension (sometimes referred
to as dorsiflexion).
Abduction is the moving of a part away from the median plane; adduction is the
moving of a part toward the median plane. Rotation is the movement of a part
around its long axis (action of the radius when using a screwdriver). Supination
(lying on the back = supine) is lateral rotation of the paw so that the palmar or
plantar surface faces medially or dorsally. Pronation (lying on the belly = prone) is
medial rotation so that the palmar or plantar surface of the paw faces ventrally.
On radiographs the view is described in relation to the direction of penetration by
the x-ray: from the point of entrance to the point of exit before striking the film. A

radiograph of the carpus in the standing position with the film under the palmar
surface of the paw would be a dorsopalmar view.
In the text that follows, structures are generally designated by their anglicized
terms in common use unless none exists. Each term, when introduced for the first
time, is followed by its Latin equivalent.

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Literature
Much anatomical information is published in several thousand scientific journals, of
which about 100 frequently contain anatomical articles in one of several languages.
Language differences and the accessibility of periodicals are still considerable
barriers to the dissemination of anatomical information, although abstracting
services and electronic retrieval systems have eased the burden of keeping current.
Literature on anatomy of the dog is not always easy to categorize by system, and,
as a result, there are many old as well as recent monographs and books that are not
cited by abstracting systems or in the chapters that follow. Some are little known: for
example, the extensively illustrated doctoral thesis of Madeleine A. Hamon “Atlas
de la Tete du Chien: Coupes Series-Radioanatomie-Tomographies” (1977) presented
at the University Paul Sabatier of Toulouse. This study includes serial transverse,
sagittal, and horizontal sections and has a bibliography of 997 references relating to
structures of the head of the dog. Current interest in multiplanar imaging, such as
CT scans, ultrasonography (sonograms), and nuclear magnetic resonance (NMR)
scans make such works invaluable. A book that compares body sections of the dog
with scans is the Atlas of Correlative Imaging Anatomy of the Normal Dog: Ultrasound
and Computed Tomography, by Feeney, Fletcher, and Hardy (Saunders, 1991). For
radiographic anatomy, the most detailed source is the Atlas of Radiographic Anatomy
of the Dog and Cat, by Schebitz, Wilkens, and Waibl (Elsevier, 2011). Thrall's recently

published seventh edition of Textbook of Veterinary Diagnostic Radiology (Saunders,
2018) is also highly useful.
Dated but still useful and authoritative anatomical information on the dog can be
found in the out-of-print Handbuch der Vergleichenden Anatomie der Haustiere, by
Ellenberger and Baum (1943). Dissection guides for the dog include Atlas der
Anatomie des Hundes, by Budras (Hannover: Schluter, 2010); Canine Anatomy: A
Systemic Study, by Adams (Ames, Iowa State University Press, 2004); Dog and Cat
Dissection Guide, by Susan and Chris Pasquini (Pilot Point Texas, 2009); A Regional
Approach to the Dissection of the Dog by M.S.A. Kumar (Linus Learning, 2017); and
Guide to the Dissection of the Dog, by Evans and de Lahunta (Elsevier 2017).

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