Preface

This is a textbook on shape analysis for biologists, covering both its basic theory and
its practice. We think that a textbook is needed because the field has changed tremen-
dously in recent years, primarily due to rapid developments in the mathematical theory of
shape. Most of the work done during the past decade has focused on the mathematical
foundations of methods, or on extensions of theory to ever more mathematically complex
situations, and so most treatments of the subject have been written by and for mathe-
maticians. Biologists reading those treatments may find the subject abstract, even obscure
and mysterious, when the reality is that conducting a biological shape analysis requires
no more background in mathematics and statistics than most biologists acquire in their
undergraduate training. The discouragement caused by reading the often highly technical
works in the field is unfortunate, because the tools of shape analysis have great utility
in biology. Furthermore, a major achievement of geometric morphometrics is the ability
to draw pictures of morphological transformations; we can literally see one morphology
transforming into another, which should make geometric morphometrics intuitively acces-
sible to morphologists. Accordingly, we have written this primer emphasizing applications
to biological questions and illustration of results, and we have written it presuming that
the reader’s background consists only of a basic course in statistics and some familiarity
with elementary geometry and algebra.
We wrote this for largely selfish reasons: we teach morphometrics to advanced under-
graduate and graduate students, and we needed a textbook. Like many biologists, our
students ask sophisticated biological questions and require methods that can answer those
questions, but have little (if any) experience with matrix algebra, non-Euclidean geome-
try or multivariate statistics. Also, like many others who are learning new methods, our
students want to apply them (often immediately on learning them), not just to learn their
rationales. Accordingly, we have emphasized the biological questions answered by various
methods and provided examples of applications to both simple and complex biological
questions. We also make software available electronically to conduct all the analyses
explained in the book, and incorporate the manuals within each chapter. This combi-
nation of explanation of methods, examples, illustration of results, software and manuals
will allow students to begin applying the methods as they are learning them.
We strongly encourage all students (and faculty) to begin collecting data as soon as
possible – ideally while reading Chapter 2. Although some datasets are contained within
the software packages so that you can practice analyzing them, it is best to practice on your
own data. They are more familiar and far more interesting to you than any we can provide.
Consequently, you will learn the material most quickly and thoroughly by applying it to
your own organisms, in context of your own biological questions.
We are very grateful to our students, both those in our regular courses at the University
of Michigan and the State University of New York, Buffalo and Canisius College, and
those who participated in our workshops at the University of California, Berkeley and
x PREFACE

the University of Massachusetts, Amherst. They all improved the text considerably. We
are also grateful to others who read earlier versions of this text and pointed out errors and
ambiguities, and also asked probing questions that sometimes required us to rethink as well
as rewrite. We especially thank Barbara L. Lundrigan, Ian Dworkin, and Rebecca German
and her laboratory group. We also thank Jason Mezey and Markus Bastir, who provided
prepublication versions of the methods for comparing subspaces (Chapter 12) and for
evaluating competing hypothesis of morphological integration (Chapter 11), respectively.
Finally, we are especially grateful to Dr Charles R. Crumly of Academic Press, who worked
with us patiently, but not too patiently.

M.L.Z., D.L.S., H.D.S., W.L.F.

April 2004