AMER. ZOOL., 38:406-409 (1998)
Teaching Zoology
Review of Textbooks of Comparative Vertebrate Anatomy1
HERBERT I. ROSENBERG 2 AND ANTHONY P. RUSSELL
Department of Biological Sciences, The University of Calgary, Calgary, Alberta TIN 1N4, Canada
This is a review of recent texts that may be used in
a course in comparative vertebrate anatomy for undergraduates usually majoring in an organismic biology
program. Books were selected for this review based on
their date of publication; all were published in the
1990s. Reviews are based on our combined experience
of over 50 years as lecturers in a half year course in
comparative anatomy at the University of Calgary;
several of the texts have been used in our classroom.
Reviews appear in reverse chronological order, with
the most recent release appearing first.
Our review is written with a standard in mind, one
that developed over time as we continue to teach comparative anatomy. We are concerned about the degree
of integration of information dealing with structure,
function, embryology, phylogenetics, and paleontology. We look for synthesis of textbook material that
stimulates the student and enhances the learning of important concepts in comparative anatomy. Finally, we
evaluate the depth of the presentation.
We find that the books reviewed all contain three
main parts that we label Introduction, Core, and Support System. The Introduction includes chapters that
discuss the origin, diversity, and phylogenetic relationships of vertebrates; histology of basic tissue types;
and embryology of organ systems.
The Core includes a system by system comparative
analysis of the vertebrate body. The main theme is
evolution of organ systems illustrated by selected examples. Structural similarities and differences are discussed with function in mind. A functional analysis of
each system may be integrated with the descriptive
material or the functional section may stand alone.
The Support System includes diagrams, selected references, glossary, credits, and appendices. A selection
of clear, well labeled diagrams that are tightly integrated with the text is essential. Most students have
never seen the structures being discussed; many have
never heard of the vertebrates chosen to illustrate a
particular point. Full and carefully wordedfigurecaptions increase the educational value of the diagrams.
The selected references at the end of each chapter
should contain a balance of descriptive and functional
papers including those most recently published. It
would be very useful if publishers maintained web
sites for their texts where Selected References sections
could be updated with input from authors.
Instructors will develop different sets of objectives
over the span of their affiliation with a course in comparative anatomy. Some will want a textbook that integrates analysis of structure and function supported
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Invited review.
E-mail: [email protected]
by a wide spectrum of examples. Other instructors may
prefer a separate approach to comparative anatomy and
functional anatomy. Still others may prefer a streamlined overview that softpedals research and empasizes
phylogenetic trends. Each of the five books we have
reviewed fits into one of these three categories. We are
fortunate that we have these different presentations to
choose from.
Vertebrates: Comparative Anatomy, Function, Evolution (second edition). KENNETH V. KARDONG. Wm.
C. Brown/McGraw-Hill, 1998, xviii + 747 pp.,
$70.00, ISBN 0-697-28654-1.
Style and content—Each chapter starts with an outline in point form that indicates the organization and
contents of that chapter. The first five chapters serve
as an introduction to the remainder of the chapters that
each deal with a different organ system; the skeletal
system is covered in three chapters. Chapter 1 is historical and elaborates on paleontological as well as
morphological concepts. Chapter 2 treats the chordates
and their possible origins. Chapter 3 details diversity
of the vertebrates and is the most revised chapter of
the second edition. Kardong has added seven new
cladograms to the second edition that illustrate proposed relationships among different groups of vertebrates. These cladograms are preceded by four pages
of new, useful text in chapter 1 headed "Grades and
Clades." Chapter 4 is an interesting and useful treatment of biomechanics and scaling, but material in this
chapter may better serve the reader if it was integrated
into chapters dealing with the musculoskeletal system.
Chapter 5 consists of a fairly traditional elaboration of
descriptive embryology.
Each of the chapters in the core dealing with an
organ system presents a wealth of anatomical detail,
results of a variety of functional anatomical studies,
and descriptions of major adaptive trends. We question
the placement of the chapter detailing the complex history of the skull as the lead off chapter for the skeletal
system; this appears before students appreciate anything about bones at all. Some of the chapters are not
constructed in parallel fashion and this may distract
the reader. The author should have shortened some
chapters by being less inclusive. One wonders at the
utility of including a table listing those vertebrates with
one ovary; careful editing should have eliminated such
items. On the other hand, the highly motivated undergraduate or graduate student will be pleased with the
wealth of material in Kardong's book that can be used
to jump-start their advanced projects. The author's attempts to highlight investigations developed in Euro-
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COMPARATIVE ANATOMY TEXTBOOKS
pean labs is to be applauded. This is clearly a text that
can be used as a reference throughout the student's
undergraduate program. It is an exciting book featuring
new material that will be appreciated by morphologists.
Given the emphasis on functional anatomical findings, it is fitting that the last chapter of the book offers
four different perspectives on how to analyze the design of vertebrates. This chapter is a fitting coda for
the entire book; most texts in this field end abruptly
and leave the student wondering what comes next.
Diagrams—The author takes a fresh approach and
presents many diagrams derived from the primary literature that are not found in other similar textbooks.
These illustrations form a rich compendium of material
for lecturers and graduate students intent on working
as vertebrate morphologists. However, for some undergraduates using this as a course textbook there may
be an overabundance of diagrams. Would Kardong
consider developing a shorter and a longer version of
his textbook? The second edition includes a full Credits section that acknowledges the sources of illustrations. Interested readers can now track down the original sources of the diagrams and have access to the
full publication. Color is used effectively in the embryology and skull chapters.
Selected references—Chapters terminate with a relatively large number of citations; there is emphasis on
recent articles in the primary literature dealing with
functional anatomy.
Innovations—Boxed essays provide interesting anecdotes and anatomical vignettes that may serve to
stimulate the student. Kardong has added three new
boxed essays to the second edition of his text. New
topics include: a sequel to the hot blooded dinosaurs
controversy, a description of Jon Mallatt's proposal regarding evolution of "the new mouth," and a discussion of castration and its consequences. The latter includes a detailed history of castrati in the entertainment world!
The publishers provide a set of 100 transparency
masters to lecturers; this is a useful bonus usually reserved for introductory level texts. However, the set is
too small and should be expanded to provide lecturers
with greater choice. A laboratory manual that Kardong
has co-authored with Edward Zalisko (Comparative
Vertebrate Anatomy: A Laboratory Dissection Guide)
is now in print (ISBN 0-697-37879-9). While this
manual may accompany any text, WCB/McGraw-Hill
plans to offer a combined package of the second edition of Kardong's text plus lab manual (ISBN 0-69741676-3) for $107.
Comparative Anatomy of the Vertebrates (eighth edition). GEORGE C.
KENT AND LARRY MILLER.
Wm.C. Brown/McGraw-Hill, 1997, xvi + 487
pp., $66.00, ISBN 0-697-24378-8.
Style and content—Each chapter starts with a very
brief introduction and outline. Chapters conclude with
an extensive, point form summary that should serve as
a useful checklist and study guide. The first five chapters form an introduction to the remainder of the book.
407
Chapter 1 describes the vertebrate body plan and vertebrate characteristics. Chapter 2 presents morphological concepts and a very brief historical perspective.
Chapter 3 deals with origins of the vertebrates, chapter
4 covers diversity of the vertebrates, and chapter 5
describes highlights of vertebrate embryology.
The Preface indicates that "This is a textbook of
functional and comparative morphology. . . " In fact,
there is little functional content. The core of the book
emphasizes comparative aspects of vertebrate morphology. This is a straightforward, no frills text that
has been streamlined as a result of multiple revisions.
The authors claim that this redesigned eighth edition
is about 100 pages shorter than previous versions. Promotional materials indicate a discussion of cladistics
has been expanded and distributed throughout the text.
This "discussion" is limited to a single column that is
accompanied by one cladogram (fig. 4.16). Students
proceeding to advanced courses in vertebrate morphology would find little useful information in this
text. However, this text is popular and can be used in
those programs that take a narrow approach to vertebrate morphology.
Diagrams—Illustrations are of uniform style and
contrast is heightened by the subtle use of color; credits or citations of the original are rarely provided.
Selected references—These are relatively few in
number and do not reflect current research trends. The
selections are not extensive enough to get an undergraduate started with a research project.
Analysis of Vertebrate Structure (fourth edition). M l TON HiLDEBRAND. John Wiley and Sons, Inc., 1995,
xiii + 657 pp., $48.00, ISBN 0-471-30823-4.
Style and content—The book is divided into four
parts and starts with an introductory chapter that sets
out basic principles of comparative anatomy and includes a list of general references. Part I, also introductory, is a survey of vertebrate diversity that begins
with a brief discussion of vertebrate relationships and
methods of classification.
Part II is the central core of the text and contains
16 chapters dealing with the systems of the body. This
section starts with a chapter detailing some basics of
vertebrate embryology. The chapters that follow emphasize comparative anatomy; functional material is
reserved for part III. The chapters of part II are streamlined, uncluttered by excessive examples, and make for
easy reading. The author's stated intention is that his
presentation be " . . . sound and solid but not overly
technical. Details that do not serve interpretations are
omitted."
Part III forms a supplementary core that deals with
structural adaptations and incorporates findings of
functional anatomical investigations. This part starts
with three introductory chapters that form an invaluable primer for an undergraduate approach to functional anatomy. The first chapter deals with the characteristics of bio-materials, the second with biomechanics, and the third with growth. The six chapters
that follow deal with different types of locomotion and
feeding.
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H. I. ROSENBERG AND ANTHONY P. RUSSELL
One way to use this text is to assign almost all of
Parts I and II for a one semester course. Portions of
Part III are then integrated back into the course at the
discretion of the lecturer. This proves to work more
smoothly than having to delete bits and pieces of textual material as one advances through the chapters of
a textbook that is overloaded with material.
Diagrams—The book is richly illustrated with a collection of largely original drawings that are well integrated with the text. The rendition of the drawings is,
however, somewhat stylized.
Selected references—Each chapter concludes with a
brief list of citations that are descriptive or functional.
Innovations—Hildebrand's original textbook stood
out as a radical departure from previously published
texts in comparative vertebrate anatomy. The author
made a clear attempt to pique the student's interest and
emphasized functional morphology. The functional
section (Part HI) may well be used as the core of an
advanced course in vertebrate morphology (one that
has comparative anatomy as its prerequisite) that
stresses analytical approaches to morphological adap-
of styles and are derived from many sources. This
tends to distract students. The reader is provided with
the original source of most diagrams; not all citations
are included in the selected reference list. Color is introduced into figures used in chapters dealing with embryology, circulation, and the head skeleton.
Selected references—The authors provide a relatively extensive list with emphasis on recent functional
anatomical investigations that should be useful for the
continuing student.
Innovations—Most of the chapters include a Focus
Essay that details an interesting adaptation or describes
the functional analysis of a particular structure.
Functional Chordate Anatomy. RONALD G. WOLFF.
D.C.Heath and Co., 1991, xx + 840 pp, $71.00,
ISBN 0-669-21895-2.
Style and content—Each chapter starts with a point
form outline, plus an introduction. The chapters are
well written and organized in parallel style; they each
contain sections dealing with development, basic structure, comparative survey of structure and function, and
phylogenetic analysis of vertebrate trends.
The first four chapters provide an introduction to the
Functional Anatomy of the Vertebrates (second edi- text. Chapter 1 provides an historical background plus
tion). WARREN F. WALKER, JR. AND KAREL F. LIEM.
principles of morphological study. Chapter 2 deals
Saunders/Harcourt Brace, 1994, xxi + 788 pp., with evolution of form and function. Chapter 3 deals
$77.00, ISBN 0-03-096846-1.
with chordate characteristics, origins, and diversity
Style and content—Each chapter starts with a precis while chapter 4 covers embryology. The chapters in
and a point form outline. The chapters conclude with the core of the book include recent research with the
a thorough summary. The first four chapters introduce material being considered. The concluding chapter of
the reader to the text that continues as a system by the book integrates anatomy, function, and behavior.
Diagrams—Most of the illustrations are derived
system treatment of the vertebrate body. Chapter 1 introduces anatomical concepts that are comparative and from other sources and they are all fully cited.
Selected references—This section contains entries
functional. A "Focus Essay" on Reconstructing Phylogenies compares traditional and cladistic systematics for material cited within the body of the text as well
and explains how to interpret a cladogram. Cladograms as general references that appear under the heading of
are employed in chapter 3 (Diversity of Vertebrates); General Bibliography. The latter includes a recomsome include characters while others do not. While the mended list of morphological journals, books, and
Preface to this second edition promises "cladograms practical references dealing with anatomical illustraat the end of many chapters. . . " in fact, they are found tion and preparation of reports. These combined citaonly at the end of the chapter dealing with the respi- tions are fairly extensive and form a valuable resource
ratory system. Chapter 2 deals with vertebrate origins for the continuing student.
and chapter 3 presents a detailed treatment of verteInnovations—A great deal of attention has been debrate diversity. Chapter 4 places emphasis on descrip- voted to providing extensive literature citations and
tive embryology but does include material on hom- bibliographic resources. This clearly enhances the valeoboxes.
ue of this text to students that pursue higher level
The first edition of the text has been revitalized. Re- courses and research projects in vertebrate morpholocent functional studies and cladistics have been melded gyto the traditional comparative and early functional apCONCLUSIONS
proach first produced by Walker. In some cases the
meld is a smooth one and in others the poor integration
These books fall into three categories based on the
of old and recent material is quite apparent. Parts of degree of integration of material, synthesis of new perthe chapters dealing with the musculoskeletal system spectives for the reader, and depth of presentation. The
introduce important biomechanical concepts and com- first category consists of texts by Kardong, Walker and
plete chapters are given over to locomotion and feed- Liem, and Wolff. All of these authors integrate strucing. The core of the text is a serious attempt to enrich ture and function and provide a large number of anatraditional, descriptive anatomy with the findings of tomical examples. We have found it necessary to tailor
experimental, functional anatomical studies. Students reading assignments; general requirements to read enare presented with an analysis of vertebrate form and tire chapters may well lead to indigestion. Overfunction in a phylogenetic context.
whelmed students may give up on the author and simDiagrams—Illustrations reflect a very wide variety ply ignore their reading assignments.
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COMPARATIVE ANATOMY TEXTBOOKS
Kardong, having written the most recent text, builds
upon the earlier entries in the field. He stimulates the
reader (textually and visually) and takes a meaningful
approach to a very wide spectrum of topics. Other authors in this category do not venture as far nor as wide.
The difficulty in writing a text for the undergraduate
is to know how to strike a balance between anatomical
detail, central concepts, and matters of interest to professional morphologists. Sometimes Kardong loses his
balance.
The next category is represented by Hildebrand's
text that is different because the entire functional portion has been separated from the descriptive part and
the former has been developed on its own. It is apparent that this book was written with a great deal of
concern for the student reader. Chapters are short and
to the point; extraneous material is kept to a minimum.
The final category is represented by the book of
Kent and Miller. This book lacks the depth and diversity of the others. However, we cannot fail to recognize
that this is the 8th edition and we assume that there is
a market for a presentation of this type.
There are other historically important texts that are
no longer listed in the most recent version of Books
in Print. These include: Romer and Parsons's "The
Vertebrate Body" (6th ed., 1986), Wake's multiauthored "Hyman's Comparative Vertebrate Anatomy"
(3rd ed., 1979), Kluge's multiauthored "Chordate
Structure and Function" (2nd ed., 1977), Weichert and
Presch's "Elements of Chordate Anatomy" (4th ed.,
1975), and Webster and Webster's "Comparative Vertebrate Morphology" (1974). Instructors may wish to
consult library copies for supplementary information.
Over the past decade, comparative vertebrate anatomy has become much less of a purely descriptive
subject and has embraced, and has been embraced by,
a number of complimentary disciplines. The inclusion
of function has been an increasing trend, and is often
reflected in the titles of the books. This form and function approach has been augmented to greater or lesser
degrees by the incorporation of robust systematic
frameworks, the inclusion of developmental information (at both the gross and molecular levels), and the
application of more rigorous interpretation of the fossil
record.
Comparative vertebrate anatomy has a long and venerable history and is at the threshold of another rebirth
as it becomes the focus of intense synthetic investigation. The texts currently available wrestle with this
integration to varying degrees and with different levels
of enthusiasm. The highly motivated student will be
drawn to those books that grasp this synthesis by the
scruff of the neck and try to place it all together. Those
less prepared and less integrated in their thinking,
however, will prefer a more conservative approach.
Thus the relative strengths of the reviewed books cannot be evaluated in a simple manner as, beyond the
basics, they endeavor to do different things. The choice
of a textbook will be made on the depth and integration that is expected and on the philosophy of the instructors). There is no "best" option, although students will certainly find some books more challenging
than others.
ACKNOWLEDGMENTS
We thank Deric Fournier of the University of Calgary Bookstore for his help.
REFERENCES
Kardong, K. V. and E. J. Zalisko. 1998. Comparative
vertebrate anatomy: A laboratory dissection
guide. Wm. C. Brown/McGraw-Hill, 214 pp.
Kluge, A. G. (ed.) 1977. Chordate structure and function. 2nd edition. New York: MacMillan Publ.
Co., x + 628 pp.
Romer, A. S. and T. S. Parsons. 1986. The vertebrate
body. 6th edition, Philadelphia: Saunders, vii +
679 pp.
Wake, M. H. (ed.) 1979. Hyman's comparative vertebrate anatomy. 3rd edition, Chicago: University
of Chicago Press, xi + 788 pp.
Webster, D. and M. Webster. 1974. Comparative vertebrate morphology. New York: Academic Press,
xiv + 517 pp.
Weichert, C. K. and W Presch. 1975. Elements of
chordate anatomy. 4th edition, New York: McGraw-Hill, viii + 526 pp.