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Principles of Systematic Zoology
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Principles of Systematic Zoology

Ernst Mayr
Alexander Agassiz Professor of Zoology, Harvard University
Director, Museum of Comparative Zoology
Member, International Commission on Zoological Nomenclature

McGraw-Hill Book Company

New York St. Louis San Francisco Toronto London Sydney
Principles of Systematic Zoology

Copyright © 1969 by McGraw-Hill, Inc. All Rights Reserved. Printed in

the United States of America. No part of this publication may be reproduced,
stored in a retrieval system, or transmitted, in any form or by any means,
electronic, mechanical, photocopying, recording, or otherwise, without the
prior written permission of the publisher.

Library of Congress Catalog Card Number 68-54937

41143

1234567890 MAMM 7543210698

 Dedicated to mv frienas

£. Gorton Linsley
and
Robert L. Usinger

without whose constant encouragement

and inspiration this work could not have
been undertaken nor completed-
Preface

Systematics has had a remarkable renaissance during the last generation.

The reasons for this are diverse. Taxonomists played a leading role in the
new synthesis of evolutionary theory, and they have demonstrated that the
study of organic diversity, the main concern of systematics, is a major integral
branch of biology. Systematics has also been very important in initiating the
entire field of population biology, including population genetics. The recog¬
nition of the importance of taxonomy has been helped by the realization
that there are two great scientific methods, the experimental method and the
comparative method (based on observations). Observational data are rather
meaningless unless they are classified prior to comparison. Understanding
this methodological necessity has created a new interest in methods and
theory of classification in all comparative sciences. This interest has been
enhanced by the coming of the computer and concurrent endeavors to
automate classifying through electronic data processing. All this is super¬
imposed on the increasing need for the applied aspects of taxonomy, such as
the correct identification and classification of species in agriculture, public
health, ecology, genetics, and behavioral biology.
In 1953 when Methods and Principles of Systematic Zoology was published,
no other text on the systematic method was available except Hennig’s
specialized German volume. The current vigor of systematics is indicated
by the number of recently published texts in this area (see Chap. 1), by the
flourishing of the journal Systematic Zoology since its founding in 1952 and of
similar periodicals in England and Germany, and by the annually increasing
number of articles in journals throughout the world dealing with the methods

vu
viii Preface

and principles of systematics. The only way to cope with this massive accumu¬
lation of new information and new theory has been to prepare a radical
revision of the 1953 volume.
Two of the three coauthors of this earlier volume (E. Gorton Linsley
and Robert L. Usinger) were unfortunately too heavily committed by profes¬
sional duties to participate in the time-consuming task of a revision. By
mutual agreement Ernst Mayr, therefore, assumed full responsibility. In
the course of the work it became apparent that more was needed than a
mere revision. The outcome was a new work with a new title. Nevertheless
it is a direct descendant of the 1953 book, and much that was valuable in
the old volume is incorporated in the new one. A first draft of the manuscript
was, therefore, critically read by the former coauthors. I deeply appreciate
the help and encouragement I have had from my two friends, who continue
to maintain an undiminished interest in this volume.
It has been a frequent complaint in the past that the training of the
young taxonomist is too empirical, consisting too much in merely condi¬
tioning him to carry out the operations of experienced taxonomists. Such
criticism is legitimate when this practical training is not supplemented by a
study of the theory. Yet, at the same time, taxonomy is a subject which is so
operational that it could not possibly be learned merely by reading a book.
The main objective of this volume, therefore, is to serve as a guide and com¬
panion for those learning the subject and perhaps even more so for those
who are teaching it. However, it cannot take the place of a laboratory in
which the procedures of classification are actually demonstrated. The prob¬
lems of taxonomy are different in every group of animals, and every teacher
will want to use material and illustrations from the zoological groups with
which he himself is most familiar. This is the reason why this volume does
not cite more examples to illustrate basic principles and methods. The teacher
himself will know best what examples will be most instructive in a given case.
The problems of taxonomy are far too diverse to provide, as the beginner
might love to have it, a set of recipes which can be applied to every situation.
Actually, a clear presentation of the theory will be more helpful in the carry¬
ing out of practical tasks than such a recipe book.
It is for this reason that the theoretical aspects of taxonomy have re¬
ceived so much attention in this work. The object is not so much to make a
contribution to theoretical biology as to establish a sound foundation for
practical operations. It is important for the taxonomist at every stage of his
work to understand exactly the meaning of such terms as species, taxon,
category, classification, and type. The clearer these terms, and the concepts
underlying them, are understood, the greater will be the agreement among
taxonomists and the less time they will waste on sterile controversy.
The 1953 book, while presenting some original ideas of the authors,
concentrated on presenting a balanced digest of the published literature of
 Preface IX

taxonomy. The new work goes somewhat further in offering original mate¬
rial, particularly on the theory of taxonomy (Chap. 4), on the procedures of
classification (Chap. 10), and on the theory of nomenclature (Chap. 13).
It is hoped that this will make the volume important even for the experienced
worker without interfering with its utility for the beginner. As far as the
theory of nomenclature and the procedure of classification are concerned,
there are no other comprehensive treatments available in the literature. To
add to the usefulness of the volume the main text of the Rules of Zoological
Nomenclature is incorporated in Chap. 12.
In order to compensate for the considerable expansion of the volume
through the addition of new material, an effort was made to shorten the
presentation of subjects that are well covered in recent publications. The
new Code of Zoological Nomenclature, for instance, gives so much detail on
the formation of scientific names (in Appendix D) that it seemed advisable
to omit this topic from the new edition. Likewise, the revised edition of
Quantitative Zoology by Simpson, Roe, and Lewontin (1960) covers the aspects
of statistics relevant to taxonomy so excellently that it was possible to shorten
our own account drastically. The recent publication of several books devoted
to scientific illustration and the techniques of drawing (Chap. 11) permitted
shortening the section devoted to this subject.
Finally, the author’s recent publication of a wide-ranging volume on
evolution, Animal Species and Evolution (1963), made it unnecessary to cover
the same subject matter in the present volume, as important as an under¬
standing of the evolutionary processes is for the taxonomist. The exhaustive
bibliography of that volume permits convenient access to the primary evolu¬
tionary literature, and the volume is therefore—specifically for this purpose—
frequently quoted in the present work.
No author can ever adequately thank all those who by their construc¬
tive criticism have contributed to the improvement of the manuscript. The
present work is one more example of such helpfulness. A first draft of the
entire manuscript was read by E. G. Linsley, R. L. Usinger, Michael
Ghiselin, Donn E. Rosen, W. F. Blair, and Richard D. Alexander. Of the
next version Herbert Levi, K. Boss, and J. Lawrence read most chapters.
Chapters 8 and 9 were read by S. Gould and R. A. Reyment and Chaps.
12 and 13 by J. Corliss, Eugene Eisenmann, W. J. Follett, R. P. Higgins,
Myra Keen, Alfred and Helen Loeblich, Hobart M. Smith, and Ellis
Yochelson. Every one of them, but particularly R. D. Alexander, made
valuable suggestions. Each found a number of errors and inconsistencies in
my treatment which I was able to correct. Those faults which still remain
are entirely my own responsibility.
John Lawrence undertook the great labor of compiling the information
on the number of known species presented in Table 1-1, much of it secured
from the leading specialists in the respective groups.
X Preface

I am greatly indebted to Susan Martin, Sabine Wespi, and Sally Loth

for the careful typing of three consecutive versions of the manuscript and to
William Jolly as well as to Vojislav Jovanovic for literature search, help
with the bibliography, and other editorial assistance, which greatly lightened
my own burden. More than anyone else it was my wife who has assisted me
in numerous ways and has truly made the completion of this volume possible.

Ernst Mayr
Contents

Preface vii
Chapter 1 The Science of Taxonomy 1

Part I. Principles of Zoological Classification

Chapter 2 The Species Category 23

Chapter 3 The Polytypic Species, Population Systematics
and Infraspecific Categories 32
Chapter 4 Theories of Biological Classification and Their History 54
Chapter 5 The Hierarchy of Categories and the Higher Taxa 87

Part II. Methods of Zoological Classification

Chapter 6 Taxonomic Collections and the Process of Identification 101

Chapter 7 Taxonomic Characters 121
Chapter 8 The Qualitative and Quantitative Analysis of Variation 144
Chapter 9 Taxonomic Decisions on the Species Level 181
Chapter 10 The Procedure of Classifying 198
Chapter 11 Taxonomic Publication 258

Part III. Principles and Application of Zoological Nomenclature

Chapter 12 The Rules of Zoological Nomenclature 297

Chapter 13 Interpretation of the Rules of Nomenclature 334
Bibliography 381
Glossary 397
Index 415

xi
Principles of Systematic Zoology
Chapter i The Science of Taxonomy

T ......
-M. he amount of diversity in the living world is staggering.
About 1 million species of animals and half a million species of plants have
already been described, and estimates on the number of still undescribed
living species range from 3 to 10 million. An estimate of half a billion for the
extinct species is consistent with the known facts. Each species may exist
in numerous different forms (sexes, age classes, seasonal forms, morphs,
and other phena). It would be impossible to deal with this enormous di¬
versity if it were not ordered and classified. Systematic zoology endeavors
to order the rich diversity of the animal world and to develop methods
and principles to make this task possible.

1.1 TAXONOMY AND SYSTEMATICS

The study of organic diversity has changed its objectives and enlarged
its scope in the course of history, as happens in any branch of science.
A detailed history will be presented in Chap. 4, but some of it must be
anticipated here in order to make clear subtle changes in the meanings
of commonly used terms (e.g., systematics).
The ancients looked for a natural order (kosmos) which would ex¬
plain the bewildering diversity of phenomena. They attempted to discover
the true “nature” of things and approached the classification of inanimate
1
2 The Science of Taxonomy

objects and living beings by the procedures of logic. The major purpose
of a classification was to serve as an identification key, and the philosophy
of the early taxonomists was well suited for the utilitarian purposes of
taxonomy. The objective shifted and the interests of the taxonomist broad¬
ened when, after 1859, organic diversity was interpreted as the result of
evolutionary divergence. No longer was he interested merely in producing
identification keys; he now interpreted groups of organisms as descendants
of common ancestors, and inevitably he became interested in the pathways
and causations responsible for evolutionary changes. Also, since this force
deals with the living organism, the taxonomist became increasingly a student
of living organisms, particularly in the field. These field studies, in turn,
showed that behavior and ecology often supply far more important taxo¬
nomic characters of species than the morphological differences of preserved
specimens. Imperceptibly a new branch of biology began to emerge, the
study of the diversity of organisms.
The ultimate result of these developments has been the recognition
that the universe of the taxonomist is far greater than was previously en¬
visioned. This had an effect on the definition of the terms taxonomy and
systematics. Until quite recently these terms were generally considered to
be synonymous. Now it has become advantageous to restrict the term tax¬
onomy to its conventional meaning, but to define the term systematics
more broadly as the study of organic diversity.
The term taxonomy is derived from the Greek words taxis, arrange¬
ment, and nomos, law, and was first proposed, in its French form by de
Candolle (1813) for the theory of plant classification. Analogous to astron¬
omy, agronomy, economy, etc., it is correctly formed and need not be
amended (Mayr, 1966). It agrees best with current thinking to define
it as follows: Taxonomy is the theory and practice of classifying organisms.
The term systematics stems from the latinized Greek word systema,
as applied to the systems of classification developed by the early naturalists,
notably Linnaeus (Systema naturae, 1st ed., 1735). We follow Simpson’s
(1961) modern redefinition of this term: “Systematics is the scientific study
of the kinds and diversity of organisms and of any and all relationships
among them ” or more simply, systematics is the science of the diversity
of organisms. The word “relationship” is not used in a narrow phylogenetic
sense, but is broadly conceived to include all biological relationships among
organisms. This explains why such a broad area of common interest has
developed between systematics, evolutionary biology, ecology, and behavioral
biology.
Place of Systematics in Biology. Systematics is unique among the bio¬
logical sciences in its dominant concern with diversity. In all subdivisions
of functional biology the main concern is with basic processes and mecha¬
nisms shared by all or most organisms. Hence the reductionist tendency
at the cellular and molecular levels—the endeavor to reduce everything
 The Science of Taxonomy 3

to common denominators, i.e., to penetrate to the universal building stones.

If all biologists were to share this objective, biology would become very
one-sided. It is the student of systematics who helps to restore the balance
by his interest in and insistence on uniqueness, whole organisms, and sys¬
tems. One of the major preoccupations of systematics is to determine, by
comparison, what the unique properties of every species and higher taxon
are. Another is to determine what properties certain taxa have in common
with each other, and what the biological causes for the differences or shared
characters are. Finally, it concerns itself with variation within taxa. In
all these concerns systematics holds a unique and indispensable position
among the biological sciences. Classification makes the organic diversity
accessible to the other biological disciplines. Without it most of them would
be unable to give meaning to their findings.
Systematics deals with populations, species, and higher taxa. No other
branch of biology occupies itself in a similar manner with this level of
integration in the organic world. It not only supplies urgently needed infor¬
mation about these levels but, more important, it cultivates a way of think¬
ing, a way of approaching biological problems which is tremendously impor¬
tant for the balance and w’ell-being of biology as a whole (Mayr, 1968).
As in every generation there is also at present a questioning of the
basic objectives of taxonomy. Have recent developments in other branches
of biology made taxonomy superfluous? Is it futile to search for a classifica¬
tion based on evolution? Is the method of binominal nomenclature in con¬
flict with its function in information retrieval? Although these questions
are sometimes posed in a manner to produce more heat than light, the
taxonomist would ignore his obligations if he did not face them squarely.
Stimulating—if not provocative—reading is provided by Michener (1963),
Ehrlich (1964), Constance (1964), and Rollins (1964). The impression
the taxonomist receives from a discriminating reading of the current biologi¬
cal literature is that there was never a greater need for a strong science
of systematics than there is at present. There is a need for someone to
stress the diversity of the living world, the most truly biological quality
of organisms, in the farce of current reductionist tendencies. However legiti¬
mate the study is of that which all organisms have in common (much
of it being the physics and chemistry of organisms), it is equally legitimate
to study the unique characteristics of taxa at all levels down to the species.
And this is precisely what the taxonomist is doing.

1.2 TERMS AND DEFINITIONS

In systematic zoology, as in all branches of science, the danger of

misunderstandings is greatly reduced by the precise definition of terms.
Terms that are frequently used in this volume, such as species, type, poly¬
typic, etc., will be carefully defined in the relevant chapters and in the
4 The Science of Taxonomy

glossary. At this point some terms will be considered that relate to all
the chapters.
The terms taxonomy and systematics were defined above. This leaves
the term classification, which partly overlaps with taxonomy. The word
is used with two different meanings. Most commonly it designates the prod¬
uct of the activity of the taxonomist, the classification of the primates
or of the bees. But it is also used as a term for the activity of classifying:
“Zoological classification is the ordering of animals into groups (or sets)
on the basis of their relationships” (Simpson, 1961). In this sense classifica¬
tion coincides largely with what is sometimes designated as beta taxonomy.
Both usages are so well established and so easily distinguished by their
context that it would seem futile and impractical to try to restrict the
term classification to only one of the two meanings. A certain amount
of overlap between the terms systematics, taxonomy, and classification is
perhaps unavoidable and not necessarily harmful.
The process of classification is totally different from that of identif ca¬
tion. In classification we undertake the ordering of populations and groups
of populations at all levels by inductive procedures; in identification we
place individuals by deductive procedures into previously established classes
(taxa). (See 4.3.1 for the theory and 6.3 for the practices of identification.)
Zoological nomenclature is the application of distinctive names to
each of the groups recognized in the zoological classification. The rules
governing it and the interpretation of these rules will be presented in Chaps.
12 and 13
The most important aspects of classification are the grouping and
ranking of organisms (Chap. 10). Precision and unambiguity of terms are
of the utmost importance in these operations. Simpson (1961, pp. 16-21;
1963) gives excellent analyses of the relevant terms.
In the earlier taxonomic literature there was frequent confusion be¬
tween the zoological objects which groups of populations represent and
their rank in the hierarchy of taxonomic categories. These are two very
different phenomena. We have somewhat analogous situations in our daily
affairs: Fred Smith is a concrete person, but “captain” (or “associate profes¬
sor” ) is his rank in a hierarchy of levels.
Taxon. The words bluebirds, thrushes, songbirds, or vertebrates refer
to groups of organisms. These are the concrete objects of zoological classi¬
fication. Any such group of populations is called a taxon if the zoologist
considers it sufficiently distinct to be worthy of being formally assigned
to a definite category in the hierarchic classification. As Simpson defines
it, “A taxon is a group of real organisms recognized as a formal unit
at any level of a hierarchic classification.” The same thought can also be
expressed as follow's: A taxon is a taxonomic group of any rank that is
sufficiently distinct to be worthy of being assigned to a definite category.
 The Science of Taxonomy 5

This definition calls attention to the fact that the delimitation of a taxon
against other taxa of the same rank is often subject to the judgment of
the taxonomist.
Two aspects must be stressed. A taxon always refers to concrete
zoological objects. Thus the species is not a taxon, but a given species
such as the robin (Turdus migratorius) is. Secondly, the taxon must be
formally recognized by the taxonomist. Within any large genus, for instance,
groupings of species can be recognized. They are taxa only if and when
they are formally distinguished, as, for instance, by being recognized as
separate subgenera. Likewise, demes and geographical isolates become taxa
only when formally recognized as subspecies.
We speak of higher taxa, such as thrushes, birds, or vertebrates, and
lower taxa, such as bluebirds or robins. Taxa of species rank are what
the taxonomist ordinarily classifies. Yet there is a great deal of variation
within most taxa, as will be discussed in later chapters. The recognition
of what belongs to a given taxon of species rank is often the most difficult
step in classification owing to sexual dimorphism, seasonal changes, age
variation, and genetic polymorphism.
Phenon. The first step in classification is the separation of reasonably
uniform samples and their assortment into taxa at the species level. There
is no generally accepted technical term for a phenotypically reasonably
uniform sample, but it may be designated as a phenon, a term introduced
by Camp and Gilly (1943) for phenotypically homogeneous samples at
the species level. Males and females often belong to different phena, while
in the case of sibling species it is possible that several species belong to
a single phenon. The term morphospecies has sometimes been applied con¬
fusingly to what is here designated as a phenon. Recognition of a technical
term for the phenotypically uniform sample greatly facilitates the description
of the taxonomic procedure. Its recognition is of particular importance
in the procedures of computer taxonomy. Sokal and Sneath (1963) use
the term phenon in a very different sense. Chapters 8 and 9 deal with
the taxonomic treatment of phena.
A category designates rank or level in a hierarchic classification. It
is a class, the members of which are all the taxa assigned a given rank.
For instance, the species category is a class the members of which are the
species taxa.
A full understanding of the meaning of category depends on an under¬
standing of hierarchical classification, which will be discussed in Chap. 5.
Such terms as species, genus, family, and order designate categories. A
category, thus, is an abstract term, a class name, while the taxa placed
in these categories are concrete zoological objects. Until the word taxon
was introduced into the literature, the term category was often confusingly
used both for group and for rank.
6 The Science of Taxonomy

Ghiselin (1966a) suggests that confusion will be further reduced by

exercising more precision in the use of the verbs associated with the terms
category and taxon. Since we define wrords, we may also legitimately define
categories such as the species, or the genus. Taxa, howrever, are things,
and we can only describe (or delimit) but not define things. When this
distinction is clearly kept in mind, many of the arguments about “defining
species” are automatically eliminated.
For a fuller discussion of the theory of classification and the meaning
of the stated terms see Gregg (1954), Beckner (1959), Cain (1958, 1962),
Simpson (1961, 1963), and Buck and Hull (1966).

1.3 THE CONTRIBUTION OF SYSTEMATICS TO BIOLOGY

A consideration of the contribution of systematics to other branches

of biology and to mankind as a whole adds to an appreciation of its scope.
Leaders in many fields of biology have acknowledged their total de¬
pendence on taxonomy. Elton (1947) made this statement regarding
ecology:

The extent to which progress in ecology depends upon accurate identification,

and upon the existence of a sound systematic groundwork for all groups
of animals, cannot be too much impressed upon the beginner in ecology.
This is the essential basis of the whole thing; without it the ecologist is
helpless, and the whole of his work may be rendered useless.

No scientific ecological survey can be carried out without the most

painstaking identification of all the species of ecological significance. A
similar dependence on taxonomy is true for other areas of science. The
entire geological chronology and stratigraphy hinges on the correct identi¬
fication of fossil key species. Even the experimental biologist has learned
to appreciate the necessity for sound taxonomy. There are many genera
with two, three, or more very similar species. Such species often differ
more conspicuously in their physiological traits or in their cytology than
in their external morphological characters. Every biologist will recall exam¬
ples in which two workers came to very different conclusions concerning
the physiological properties of a certain “species” because, in fact, one
of them had been working with species a and the other with species b.
Comparative biochemistry is vitally interested in sound classification. The
evolution of molecules, an increasingly important area of research in mo¬
lecular biology, can be understood only against the background of a sound
classification. It is solely in consultation wfith the taxonomist that the bio¬
chemist can determine what organisms might supply the key to important
steps in the evolution of molecules.
 The Science of Taxonomy 7

One of the greatest assets of a sound classification is its predictive

value. It permits extrapolation from known to previously unstudied charac¬
ters. An analysis of a few species strategically scattered through the natural
system may provide us with much of the needed information on the dis¬
tribution of a new enzyme, hormone, or metabolic pathway. Many animals
cannot be kept in the laboratory, and others will not reproduce in captivity.
Again, a sound system will permit all sorts of inferences from the genetically
well-known types. (The validity of these inferences, of course, is limited
by the fact that every species is a unique system.) The systematist can
fill many gaps in our knowledge that are inaccessible to the specialists
in the experimental branches of biology. There has been increasingly close
collaboration in recent years among taxonomists and workers in immu¬
nology, in comparative biochemistry, in comparative physiology, and in
the study of animal behavior.
It may be proper to single out some specific areas to which taxonomy
has made noteworthy contributions.
1.3.1 Applied Biology. The contribution of taxonomy to the applied
sciences has been both direct and indirect. It relates to medicine, public
health, agriculture, conservation, management of natural resources, etc.
Taxonomic breakthroughs have often supplied the key to the solution of
previously perplexing problems in economic entomology. The famous case
of the epidemiology of malaria is a good example. The supposed vector
in Europe, the malaria mosquito Anopheles maculipennis Meigen, was re¬
ported throughout the continent, and yet malaria was restricted to local
districts. Large amounts of money were wasted because no one understood
the connection between the distribution of the mosquito and of malaria.
Careful taxonomic studies, summarized by Hackett (1937) and Bates
(1940), finally provided the key to the situation. The maculipennis complex
was found to consist of several sibling species with different habitat prefer¬
ences and breeding habits, only some of which are responsible for the trans¬
mission of malaria in a given area. This new information enabled control
measures to be directed to the exact spots where they would be most
effective.
With biological control of insect pests again receiving increased atten¬
tion, the determination of the exact country of origin of insect pests and
of their total fauna of parasites and parasitoids is restored to the great
importance it had prior to the short period when at least some applied
entomologists thought that they could completely control insects with
pesticides.
Pemberton (1941) cites an outstanding instance of the value of insect
collections, assembled for taxonomic study, in the solution of a biological
control problem. Some twenty years ago the fern weevil, Syagrius fulvitarsis
Pascoe, became very destructive to Sadleria ferns in a forest reserve on
8 The Science of Taxonomy

the island of Hawaii, and control measures became necessary. Entomological

literature failed to reveal its occurrence anywhere outside Hawaii except
in greenhouses in Australia and Ireland. These records, of course, gave
no clue as to the country of origin. However, while engaged on other
problems in Australia in 1921, Pemberton had the opportunity of examining
an old private insect collection at Sydney, and among the beetle specimens
was a single Syagrius fulvitarsis bearing the date of collection, 1857, and
the name of the locality in Australia from which it was obtained. This
provided the key to the solution of the problem, for a search of the forest
areas indicated on the label revealed a small population of the beetles
and, better still, a braconid parasite attacking the larvae. Collections were
made immediately for shipment to Hawaii, and the establishment of the
parasite was quickly followed by satisfactory control of the pest. The data
borne on a label attached to a single insect specimen in 1857, in Australia,
thus contributed directly to the successful biological control of the pest
in Hawaii sixty-five years later.
Clausen (1942) and Sabrosky (1955) have presented in detail some
of the achievements of taxonomy in biological control as well as reports
on very expensive failures due to incorrect identifications.
1.3.2 Systematics and Theoretical Biology. The service functions
of taxonomy are often stressed to such a degree that the important contribu¬
tions of systematics to the conceptual structure of biology are overlooked.
Population thinking, for instance, has come into biology through taxonomy
(Mayr, 1963), and indeed one of the two roots of population genetics
is taxonomy (Chap. 3). The problem of the multiplication of species was
solved by taxonomists, and they have made the greatest contributions to
our understanding of the structure of species and of the evolutionary role
of peripheral populations. It was taxonomists who continued to uphold
the importance of natural selection at the time when the early Mendelians
thought that mutation had eliminated the role of natural selection as an
evolutionary factor. Taxonomists like H. W. Bates and F. Muller made
significant contributions to the understanding of mimicry and related evolu¬
tionary phenomena. Taxonomists and naturalists in close contact with tax¬
onomy were instrumental in the development of ethology and the study
of the phylogeny of behavior. Taxonomists have consistently played an
important role in counteracting the reductionist tendencies dominant in
so much of functional biology. They have contributed thereby to a healthy
balance in biological science. (See also Simpson, 1962a, on the status of
taxonomy.)
The Role of Taxonomy. The multiple role of taxonomy in biology
can be summarized as follows:

1. It works out for us a vivid picture of the existing organic diversity

of our earth and is the only science that does so.
 The Science of Taxonomy 9

2. It provides much of the information permitting a reconstruction of the

phylogeny of life.
3. It reveals numerous interesting evolutionary phenomena and thus makes
them available for causal study by other branches of biology.
4. It supplies, almost exclusively, the information needed for entire branches
of biology (e.g. biogeography).
5. It supplies classifications which are of great heuristic and explanatory
value in most branches of biology, e.g., evolutionary biochemistry, im¬
munology, ecology, genetics, ethology, historical geology.
6. It is indispensable in the study of economically or medically important
organisms.
7. In the hands of its foremost exponents it makes important conceptual
contributions (such as population thinking), not otherwise so easily acces¬
sible to experimental biologists. Thus it contributes significantly to a
broadening of biology and to a better balance within biological science
as a whole.

1.4 THE TASK OF THE TAXONOMIST

There is much uncertainty in the minds of some zoologists as to the

function of the taxonomist. Some laboratory workers and ecologists limit
his role to service. He should content himself with identifying material
and devising keys. Beyond that he should keep his collections in good order,
describe new species, and have every specimen properly labeled. According
to this view, systematics is the mere pigeonholing of specimens.
In actuality systematics is one of the major subdivisions of biology,
broader in base than genetics or biochemistry. It includes not only the
service functions of identifying and classifying but the comparative study
of all aspects of organisms, as well as interpretation of the role of lower
and higher taxa in the economy of nature and in evolutionary history.
It is a synthesis of many kinds of knowledge, theory, and method, applied
to all aspects of classification. The ultimate task of the systematist is not
only to describe the diversity of the living world but also to contribute
to its understanding.
The modern taxonomist is far more than the caretaker of a collection.
He is a well-trained field naturalist who studies the ecology and the behavior
of his species in their native environment. Most younger systematists have
had a thorough training in various branches of biology, including genetics.
This experience in both field and laboratory gives them an excellent back¬
ground for more fundamental studies.
The practice of classifying consists of two principal procedures, each
in turn consisting of two separate steps (see also Chap. 10).
1.4.1 The Discrimination of Entities. The systematises first task is
to sort that portion of the diversity of individuals which he encounters
into easily recognizable and internally homogeneous groups, and to find
10 The Science of Taxonomy

constant differences between such groups. Each such aggregate is a phenon

(see 1.2). A phenon is not necessarily a population in the biological sense
but may also be either a biased sample from a population (males, juveniles,
morphs, etc.) or else (in the case of sibling species) a mixture of several
populations, and finally, in the case of geographically heterogeneous mate¬
rial, possibly a mixture of several subspecies.
The second step is the assignment of phena to species, the lowest
taxon routinely used in classification (Chap. 9). In most zoological groups
instances are known in which several phena within a single biological species
have been described and named as species. In certain groups the specialists
have not succeeded completely even today in matching the sexes (e.g. mutil-
lid wasps), the castes (e.g. ants), or the generations (e.g. trematodes).
For some groups we have separate identification schemes for larvae and
for adults. These should not be called biological classifications because we
as biologists classify species, while larvae and adults are merely different
phenotypic expressions of the same genotypes.
As soon as the basic units are discriminated, it becomes necessary
to supply identifying symbols, “names,” to facilitate communication among
zoologists. Universality and stability are among the two most important
qualities of such names, as in all other communication systems (Chap.
12).
The Inventory of Species. Even this first task of the taxonomist, the
discrimination and description of species, becomes the more formidable
the greater the increase in our knowledge. Linnaeus in 1758 recorded 4,162
species of animals; Mobius in 1898 listed 415,600 Recent species. At present
the figure presumably exceeds 1 million (Table 1-1).
It must be remembered that all estimates or counts contain two sources
of error, which fortunately tend to cancel one another out. First, a lesser
or major proportion of the existing species is still undescribed in all groups
of animals. On the other hand, in the more poorly known groups the
so-called number of species is the number of recorded names. This includes
many synonyms and the names of geographic races still listed as full species.
Estimates of the total number of living species (including nematodes,
mites, and protozoans) range as high as 5 or even 10 million. The number
of fossil species has been estimated to be 50 or 100 times as high (Cailleux,
1954). Thus only about one in 5,000 fossil species has so far been described.
Of the one single group of protozoans, the Foraminifera, 28,000 species
had been described by 1964. 685,000 species of insects were counted by
Sabrosky in 1953, and allowing for an annual increment of about 6,000
species, the figure would now be around 750,000.
It is of considerable significance that some strikingly new types are
still being discovered. This includes the only surviving coelacanth, Lati-
meria, discovered in 1938; the primitive mollusk, Neopilina, described in
 The Science of Taxonomy 11

Table 1-1. Number of Species in Major Animal Taxa

ANIMALIA (Recent). 1,071,000

Protozoa. 28,350
Sarcomastigophora. 17,650
Mastigophora. 6,000
Opalinata. 200
Sarcodina. 11,450
Sporozoa. 3,600
Cnidospora. 1,100
Ciliophora. 6,000
Mesozoa. 50
PORIFERA. 4,800
COELENTERATA (CnIDARIa). 5,300
Ctenophora. 80
Platyhelminthes. 12,700
Turbellaria. 3,000
Trematoda. 6,300
Cestoda. 3,400
Gnathostomulida. 45
Entoprocta (Kamptozoa). 75
Nemertinea. 800
Aschelminthes (Nemathelminthes). 12,500
Gastrotricha. 170
Rotatoria. 1,500
Nematoda. 10,000
Nematomorpha. 230
Kinorhyncha. 100
Acanthocephala. 500
Priapulida. 8
Mollusca. 107,250
Polyplacophora (Loricata). 1,000
Aplacophora (Solenogastres). 150
Monoplacophora. 3
Gastropoda. 80,000
Scaphopoda. 350
Bivalvia (Lamellibranchia). 25,000
Cephalopoda. 750
SlPUNCULIDA. 250
Echiurida. 150
Annelida. 8,500
Onychophora. 70
Tardigrada. 350
Pentastomida (Linguatulida). 65
Arthropoda. 838,000
Chelicerata. 57,500
Merostomata (Xiphosura). 4
12 The Science of Taxonomy

Arachnida. 57,000
Pantopoda (Pycnogonida). 500
Mandibulata. 780,500
Crustacea. 20,000
Chilopoda. 2,800
Diplopoda. 7,200
Pauropoda. 380
Symphyla. 120
Insecta. 750,000
Lophophorata (Tentaculata). 3,750
Phoronidea. 18
Bryozoa. 3,500
Brachiopoda. 230
Hemichordata (Branchiotremata). 80
Echinodermata. 6,000
Echinozoa. 1,750
Holothuroidea. 900
Echinoidea. 850
Crinozoa. 650
Asterozoa. 3,600
Somasteroidea. 1
Asteroidea. 1,700
Ophiuroidea. 1,900
POGONOPHORA. 100
Chaetognatha. 50
Chordata. 43,000
Tunicata. 1,300
Cephalochordata. 25
Vertebra ta. 41,700
Agnatha. 50
Chondrichthyes. 550
Osteichthyes. 20,000
Amphibia. 2,500
Reptilia. 6,300
Aves. 8,600
Mammalia. 3,700

Total about 1,071,000

1956; the phylum Pogonophora, with over 100 species mostly described
since 1950; the Cephalocarida, discovered in 1955; and the Gnathosto-
mulida, discovered in 1956. Several of these discoveries have had a strong
impact on our concepts of phylogeny.
The degree to which the inventory taking has been completed differs
from group to group. In birds 99 percent of all living species have surely
been described, and in mammals and reptiles it may well be more than
 The Science of Taxonomy 13

90 percent. Yet there are poorly studied groups of arthropods, protozoans,

and marine invertebrates in which less than 10 percent of the species existing
in the world have so far been described. These are merely estimates, as
no one really knows.
Only a few non-taxonomists appreciate how poorly most animal
groups are known taxonomically. A striking illustration of this is presented
by Remane’s work on the microscopic marine fauna of the Kieler Bucht,
an area previously considered to be well known. By thorough search and
with the application of new methods, Remane found 300 new species in
ten years, including representatives of 15 new families. Sabrosky (1950)
has pointed out how poorly much of the North American insect fauna
is known. Many so-called “common species” actually represent whole com¬
plexes of previously overlooked good species. This is the situation in the
temperate zone. In the tropics the discrimination of species is even farther
behind; often in a generic revision every species is found to be new.
To complete this part of the task of the taxonomist will require the
labors of several more generations. Considering the limited number of spe¬
cialists, we cannot expect, for instance, to settle all the problems of mite
taxonomy in the next thirty years. We must take it for granted that a
large part of the mite fauna of the world will remain unsampled, unnamed,
and unclassified for decades to come. The same is probably true for the
majority of the kinds of animals. It is regrettable that literally thousands

Fig. 1-1. Number of new species of Drosophila described in stated periods. Almost as many (516)
new species were described in the last 18 years as in the preceding 163 years (566 species). [Data from
M. R. Wheeler ]
14 The Science of Taxonomy

of species will become extinct in the next generation in all parts of the
world owing to habitat destruction before they were ever collected and
scientifically described.
1.4.2 Grouping and Ranking. The recognition and accurate descrip¬
tion of species is only the first step in classification. Should the taxonomist
stop here, he would soon be confronted with a chaotic accumulation of
species names. The second task, then, of the taxonomist is to put the species
in order. He must group them into smaller and larger arrays of related
species, taxa, and higher taxa, and must place these in a hierarchy of
categories. In other words, he must make a classification (see Chaps. 4
and 10).
Information Storage. There is perhaps no other branch of science
in which information storage and retrieval is as formidable and as crucially
important a task as in taxonomy. It is impossible to prepare a reliable
revision or monograph of a higher taxon if one does not first compile
a listing of the described species. Mutatis mutandis this is true for any
operation in beta and gamma taxonomy. To make matters worse there
are few fields in which the literature is more scattered. The more species
are described and the more people work in taxonomy, the worse the problem
gets.
Hardly anyone doubts that computerized information storage is the
inevitable solution. It is probably correct to say that every taxonomist spends
more time on searching and extracting the literature than on original work.
This is so traditional and taken so much for granted that it is virtually
never pointed out how appallingly inefficient this research method is. Coding
all taxonomic information for computer storage will be very expensive,
but in the long run perhaps considerably less so than the uneconomic waste
of time now required of the working taxonomist. Whatever new kinds
of information become important to the taxonomist—behavioral, cytological,
biochemical, ecological—he will soon be totally overwhelmed by it if new
ways for storing and retrieving it are not soon found.
1.4.3 Biological Systematics. The making of a classification is not
the end of the taxonomist’s concern. Consistent with Simpson’s definition
of systematics as “the scientific study of the kinds and diversity of organisms
and of any and all relationships among them,” the systematist studies all
aspects of living organisms. He is interested in more than description;
as a scientist he is concerned with meanings and the study of causation.
Species formation, the factors of evolution, the structure of natural popula¬
tions, biogeography—all these are of concern to the taxonomist, and he
has made important contributions to all these branches of biological science.
Indeed it may be claimed that they all are part of systematics. The entire
broad field of comparative zoology is included in his concern. Activities
in this area are sometimes referred to as gamma taxonomy (see next
paragraph).
 The Science of Taxonomy 15

Stages in Classification. The classification of a given taxonomic group

goes through various stages of maturation. In the first stage, often called
alpha taxonomy, emphasis is on the description of new species and their
preliminary arrangement in comprehensive genera. In beta taxonomy rela¬
tionships are worked out more carefully on the species level and on that
of the higher categories; emphasis is placed on the development of a sound
classification. At the level of gamma taxonomy much attention is paid
to intraspecific variation, to various sorts of evolutionary studies, and to
a causal interpretation of organic diversity. The three levels overlap and
intergrade, but the trend from alpha to gamma is unmistakable in the
taxonomic history of any group. Though there is need for more work on
the alpha and beta levels even in the taxonomically best-known groups,
it is the ultimate endeavor of the biologically minded taxonomist to achieve
an understanding of the group in which he specializes.
The reason for gamma taxonomy is the fact that causal explanation
is part of any science, and systematic zoology is no exception. If systematics
were satisfied with merely pigeonholing the diversity of the living world,
it would be a technology but not a science.
1.4.4 Strategy of Taxonomic Research. Like most other scientists,
the taxonomist must frequently make the decision what to do next. Should
he concentrate on work in alpha taxonomy, describe new species, and pre¬
pare catalogs of the available names? Should he concentrate on beta tax¬
onomy and prepare a basic revision or monograph, based on an examination
of all included species and preferably all type-specimens, and culminating
in a well-balanced, carefully reasoned reclassification? Or finally, should
he devote himself to some aspects of gamma taxonomy, such as making
a detailed study of individual and geographical variation in a single species,
based on numerous large population samples, or studying the behavioral
or chemical characteristics of a set of species, etc.? The answer to these
questions is twofold. First, depending on the level of maturity reached
in the group of which he is a specialist, either one or the other of these
levels will permit the most productive work. Secondly, most specialists avoid
mental fatigue by alternating different approaches. When one gets tired
of the tedious compilation of data necessary for the preparation of a revi¬
sion, it is relaxing to do a study of geographic variation in a single species
with rich material, or of living material, or of certain distributional aspects.
Field systematics offers countless challenges which are particularly stimulat¬
ing when alternated with museum research.
Finally, as pointed out by Gosline (1965), the geographic location
of a taxonomist will considerably influence his research strategy. Some prob¬
lems are most advantageously pursued in a small institution, others in a
large national museum. The specialist working in Washington, London,
or Leningrad, in spite of his many other advantages, will be unable to
settle certain questions, particularly those dealing with life cycles, sexual
16 The Science of Taxonomy

dimorphism, seasonal occurrence, niche occupation, concealing or warning

coloration, relative frequency, etc., that are easily accessible to the specialist
in Trinidad, Belem, Honolulu, Lae, Kuala Lumpur, or Entebbe. To devote
one’s life to the compilation of a nomenclator of generic names, when
one happens to be at a tropical station with a poor library, is about as
misguided a strategy as can be imagined (and yet this has actually
happened).
Some nonsystematists have proposed in recent years to abandon most
of taxonomy as “old-fashioned,” “an exhausted mine,” or “useless,” and
to concentrate instead on some specialized area, such as comparative protein
chemistry, the taxonomy of behavioral traits, or the principles of geographic
variation. Advocates of such specialized approaches forget a number of
things: (1) that the various approaches are not mutually exclusive, (2)
that in many groups there is still abundant need for the most basic alpha
taxonomy, and (3) that man is one of the most polymorphic species. Some
workers are interested in using electronic computers, others in w'atching
behavior; some are interested in insects, others in fish; some like to work
with books, others with test tubes, some with preserved specimens, others
with living material. Human polymorphism cannot be controlled by regi¬
mentation. The breadth of systematics has room for students with the most
diversified interests and talents.
A good biological education will reveal to the student where the prob¬
lems are that are most exciting to him. His natural inclination will lead
him to select one area or another. And it will be only for the good of
biological science if not too many students of a given year class jump
on the same bandwragon.
Questions of strategy also arise in connection with publication. These
are dealt with in Chap. 11.

1.5 SYSTEMATICS AS A PROFESSION

1.5.1 Opportunities and Difficulties. Positions for professional tax¬

onomists exist in museums, at universities, and in various agencies of the
government. In the United States there are perhaps at present more tax¬
onomists in teaching positions in various colleges and universities than in
government or museum positions. A well-trained taxonomist is particularly
well qualified to teach a general course in zoology or biology, because
he has the broad background in zoology, morphology, physiology, genetics,
and ecology which some specialists, for lack of time, are unable to acquire.
At the present time (1968) there are numerous unfilled positions for tax¬
onomists, particularly for specialists in the Arachnida and marine inverte¬
brates. Indeed, the professional opportunities in taxonomy are far greater
 The Science of Taxonomy 17

than is generally realized. There are, however, drawbacks. The situation

in applied taxonomy often changes rapidly. When the importance of Fora-
minifera for the stratigraphic determination of oil-bearing geological forma¬
tions was discovered, there was such a demand for micropaleontologists
that it took decades to fill it. With geophysical methods now dominating
oil exploration, the demand for micropaleontologists has decreased sharply.
During the height of chemical insect control, the demand for well-trained
insect taxonomists lagged. With the renewed recognition of the importance
of biological control, there is a greatly increased need for qualified insect
taxonomists. The recent stepping-up of oceanographic research revealed
an extraordinary shortage of marine zoologists. New positions for specialists
in marine invertebrates were soon created in numerous museums and uni¬
versities. No one can predict where the next need will arise. It is conceivable,
for instance, that institutes in molecular biology may begin to employ tax¬
onomists for consultation on the numerous evolutionary and taxonomic
problems which comparative biochemistry encounters at every step. As in
all fields of biology, there is always room for superior workers, regardless
of temporary fluctuations in the employment situation.
To the same extent to which die field of taxonomy as a whole has
become increasingly ^diversified, the need for different kinds of taxonomists
has sharply increased. As Rozeboom (1962) has said rightly: “The person
trained to program data for analysis with electronic computers is not likely
to be willing to spend three or five years in the jungles of Africa collecting
ecological data on Anopheles gambiae. I question also whether he would
be willing to spend an equal amount of time searching for minute mor¬
phological characters suitable for some kind of measurement.” Nor is the
person who is painstaking enough to compile a checklist or nomenclator
likely to be equally interested in comparative behavior and biochemical
studies, or vice versa. There are some taxonomists who do their best work
when undertaking population analyses, while others are virtual geniuses
in the weighting of characters preparatory to construction of a classification
of higher taxa. All these talents are needed, and someone can usually be
found to fill each niche, thanks to the extraordinary variation in man’s
talents.
The number of pure research positions is as limited in taxonomy
as in most branches of science. Most taxonomists earn their living as teach¬
ers, curators, members of identification services, or in other branches of
applied biology. Another group of taxonomists are “amateurs,” that is,
they earn their living as civil servants, businessmen, lawyers, or doctors,
and conduct their taxonomic research as a hobby. The role of the amateur
in systematics has been changing over the years. When collecting and “nam¬
ing” was believed to be the essence of taxonomy, almost any untrained
person could become a specialist in a group of which knowledge was still
18 The Science of Taxonomy

at the level of alpha taxonomy. The stamp-collector type of amateur has

been losing ground as the quality of taxonomic research has been improving.
However, when field systematics became an increasingly important branch
of taxonomy, a new niche opened up for the amateur naturalist. He supplies
information on behavior and ecology which may be very important tax-
onomically. The work of the better amateurs is sometimes of the same
high level of quality as that of the good professional. Considering the enor¬
mous size of the taxonomic task still to be done, taxonomy would make
slow progress if it were not for the dedicated amateur.
One of the most interesting current developments is the blurring of
the borderline between “taxonomists” and “other biologists.” Population
geneticists such as Sturtevant, Dobzhansky, Carson, and Wheeler have been
most active in elucidating the classification of the genus Drosophila and
in the describing of new species. Herpetologists have discovered new sibling
species of frogs by analyzing their vocalization. Entomologists have done
the same for cicadas and grasshoppers. Ecologists have sometimes taken
up taxonomic studies in order to give a new dimension to their ecological
research. Virtually all taxonomists have a broad interest in biogeography,
and most major contributions to this branch of biology were made by
taxonomists. Now that “taxonomic character” means not only aspects of
morphology, but every kind of manifestation of the genotype, there is con¬
tinuous transgression of the borders between taxonomy, physiology, behavior,
genetics, biochemistry, and so forth. This active interchange is evident from
recent symposia (Leone, 1964; Handler, 1965; Bryson and Vogel, 1965).
What this means is that there is room in taxonomy for biologists of the most
diverse interests. Even the mathematically inclined student can make con¬
tributions by applying computer methods to taxonomy.
1.5.2 The Training of the Taxonomist. Regular courses in methods
and principles of taxonomy are a comparatively new phenomenon. Formerly
the young taxonomist learned as apprentice to a master. Perhaps this is
in part responsible for the highly uneven quality of taxonomic work done
in the past. A competence of such breadth is demanded from the profes¬
sional taxonomist of today that the method of apprenticeship is no longer
sufficient. One expects from the well-trained taxonomist a broad knowledge
of zoology, a thorough knowledge of the comparative morphology of the
group in which he is specializing, and an understanding of genetics and
evolutionary biology. He must be versed in statistical methods and, if possi¬
ble, in the use of computers. The discussion of taxonomic characters in
Chap. 7 indicates some of the other areas of biology in which the taxonomist
should be knowledgeable. One might add, incidentally, that a broad knowl¬
edge of the classification of animals should be not merely useful, but an
indispensable part of the equipment of every biologist, regardless of field
of specialization.
 The Science of Taxonomy 19

1.5.3 Professional Organizations. Systematic zoologists of most coun¬

tries have their own professional organizations. Some of these are devoted
to systematics as a whole, others are specialized societies, like the American
Ornithologists’ Union, the Entomological Society of America, or the
Lepidopterists’ Society. Blackwelder (1967) gives a listing of such societies.
All aspects of animal taxonomy are represented in the Society of
Systematic Zoology, the only broad systematics society in this country and
publisher of Systematic Zoology. A comparable organization in Great Britain
is the Systematics Association, publisher of symposia.
As far as general systematics is concerned, much relevant material
is also published in the journals Evolution, American Naturalist, and
Ecology; in Great Britain in the Journal of the Linnaean Society of Lon¬
don; and in Germany in the Zeitschrift fur zoologische Systematik und
Evolutionsforschung (from 1963 onward).
1.5.4 The Future of Systematics. Judging from the developments
of recent decades, the systematist has every reason to be optimistic about
the future of his field. Simpson (1945) put it very well when he said
that systematics “is at the same time the most elementary and most inclusive
part of zoology, most elementary because animals cannot be discussed or
treated in a scientific way until some taxonomy has been achieved, and
most inclusive because [systematics] in its various branches gathers together,
utilizes, summarizes, and implements, everything that is known about ani¬
mals, whether morphological, physiological, psychological, or ecological.”
There will always be a need for the ordering and classifying activities
of the taxonomist, even if society should, unexpectedly, stress increasingly
the applied aspects of science. There will always be a need, and as it
seems to us, a growing need, to study organic diversity and its meaning,
for there is no other subject that teaches us more about the world we
live in than systematics. And is it not equally important to reveal the
unknown life on our own planet as it is to map additional specks on the
sky?
This age of leisure and automation offers unparalleled opportunities
for the cultivation of avocational interests. There are few hobbies as reward¬
ing as natural history, and the greatest joy always comes from deep knowl¬
edge and true understanding. We agree with Crowson (1958) that the
pursuit of taxonomy not only gives great pleasure to its devotees, but that
it conveys a reverence for the wonders of living nature which should be
part of the outlook of every truly human being.
 I
Part Principles of Zoological
Classification
ChapterThe Species Category

T ,
M hp twenty or more categories which the taxonomist uses
.
in classification are of unequal value and of different significance. They fall
quite naturally into three groups:

1. The species category (Chap. 2)

2. Categories for distinguishable populations within species (= infraspecific
categories, Chap. 3)
3. Categories for higher taxa, that is, for groupings of species (collective
categories = higher categories, Chap. 5)

The species, in a number of different ways, occupies a unique position

in the taxonomic hierarchy.

2.1 THE SPECIES PROBLEM

It seems to be one of man’s most elementary urges to want to identify

things and name them. Even the most primitive native peoples have names
for kinds of birds, fishes, flowers, or trees. If only individuals existed, and
the diversity of nature were continuous, it would be difficult to sort them
into groups and distinguish “kinds.” Fortunately, at least in the animal
world, the diversity of nature is discontinuous, consisting in any local fauna
23
 4 Principles of Zoological Classification

of more or less well-defined “kinds” of animals which we call species.

Around New York City, for instance, there are about 150 “kinds” of breed¬
ing birds. These are the species of the taxonomist. Primitive natives in
the mountains of New Guinea will distinguish the same kinds of organisms
as, quite independently, does the specialist in the big national museums
(Diamond, 1966).
The concept of species seems so absurdly simple that it always comes
as something of a shock to a beginning taxonomist to learn how voluminous
and seemingly endless the debate about the species problem has been. In
zoology there is now fair agreement on the species concept, although
heterodox views are still vigorously defended. For recent summaries see
Mayr (1957a, 1963) and Simpson (1961). For statements of the botanical
point of view see Heslop-Harrison (1963) and Love (1964).
The species problem has been made to appear more difficult than
it is by a confusion of the concepts underlying the terms phenon, taxon,
and category (see 1.2). The working taxonomist sorts specimens (individ¬
uals) into phena and decides which of these are members of populations
each of which belongs to a single taxon of the species category. To be
able to undertake the ranking of taxa, the taxonomist must have a clear
conception of the category species. If he defines it (as a morphospecies)
in such a way that it coincides with the phenon, he may facilitate his
task of sorting specimens, but his activity will result in species that are
biologically, and hence scientifically, meaningless. The objective of a scien¬
tifically sound concept of the species category is to facilitate the assembling
of phena into meaningful taxa on the species level. A short survey of the
history of species concepts will show how different the species taxa are
which one is forced to distinguish if one adopts different species concepts.

2.2 SPECIES CONCEPTS

Taxonomic literature reports innumerable species concepts (Mayr,

19576; Heslop-Harrison, 1963). By their philosophical basis, all these con¬
cepts fall into three groups. The first two have mainly historical significance,
but are still upheld by a few contemporary authors.
2.2.1 Typological Species Concept. According to this concept the
observed diversity of the universe reflects the existence of a limited number
of underlying “universals” or types (eidos of Plato). Individuals do not
stand in any special relation to each other, being merely expressions of
the same type. Variation is the result of imperfect manifestations of the
idea implicit in each species. This species concept, going back to the philoso¬
phies of Plato and Aristotle, was the species concept of Linnaeus and his
followers (Cain, 1958). Since this philosophical tradition is sometimes re¬
ferred to as essentialism, the typological definition is also sometimes called
 The Species Category 25

the essentialist species definition. (For a discussion of essentialism see

4.3.2.) Various attempts at a purely numerical or mathematical species
definition (e.g. Ginsburg, 1938) are logical equivalents of this species con¬
cept. It must be emphasized that there is a complete difference between
basing one’s species concept on morphology and using morphological evi¬
dence as inference for the application of a biological species concept (Simp¬
son, 1961).
Two practical reasons exist for the now quite universal rejection of
the typological species concept: (1) Individuals are frequently found in
nature that are clearly conspecific with other individuals in spite of striking
differences in structure owing to sexual dimorphism, age differences, poly¬
morphism, and other forms of individual variation. Although often de¬
scribed originally as different species they are deprived of their species
status, regardless of the degree of morphological difference, as soon as they
are found to be members of the same breeding population. Different phena
that belong to a single population can not be considered different species.
(2) Sibling species differ hardly at all morphologically, yet are good biologi¬
cal species. Degree of difference is not the decisive criterion in the ranking
of taxa as species.
Its own adherents abandon the typological species concept whenever
they discover that they have named as a separate species something that
is nothing but a conspecific phenon. At present the typological species con¬
cept is still defended by some writers adhering to Thomistic philosophy.
2.2.2 Nominalistic Species Concept. The nominalists (Occam and
his followers) deny the existence of “real” universals (4.3.3). For them
only individuals exist, while species are man-made abstractions. (When
they have to deal with a species, they treat it as an individual on a higher
plane.) The nominalistic species concept was popular in France in the
eighteenth century (Buffon in his early writings, Robinet, Lamarck) and
has adherents to the present day. Bessey (1908) expressed this viewpoint
particularly well: “Nature produces individuals and nothing more . . . spe¬
cies have no actual existence in nature. They are mental concepts and
nothing more . . . species have been invented in order that we may refer
to great numbers of individuals collectively.”
Any naturalist, whether a primitive native or a trained population
geneticist, knows that this is simply not true. Species of animals are not
human constructs, nor are they types in the sense of Plato and Aristotle,
but they are something for which there is no equivalent in the realm of
inanimate objects.
2.2.3 The Biological Species Concept. In the late eighteenth century
it began to be realized that neither of the medieval species concepts dis¬
cussed in the two preceding sections was applicable to biological species.
An entirely new species concept began to emerge after about 1750. It
26 Principles of Zoological Classification

is augured by statements made by Buffon (in his later writings), Merrem,

Voigt, Walsh (1864) and many other naturalists and taxonomists of the
nineteenth century. K. Jordan (1905), however, was the first who clearly
formulated the concept in all of its consequences. It combines elements
of the typological and nominalistic concepts by stating that species have
independent reality and are typified by the statistics of populations of indi¬
viduals. It differs from both by stressing the populational aspect and genetic
cohesion of the species, and by pointing out that it receives its reality
from the historically evolved, shared information content of its gene pool.
As a result, the members of a species form (1) a reproductive com¬
munity. The individuals of a species of animals recognize each other as
potential mates and seek each other for the purpose of reproduction. A
multitude of devices ensures intraspecific reproduction in all organisms.
The species is also (2) an ecological unit which, regardless of the individuals
composing it interacts as a unit with other species with which it shares
the environment. The species, finally, is (3) a genetic unit consisting of
a large, intercommunicating gene pool, whereas the individual is merely
a temporary vessel holding a small portion of the contents of the gene
pool for a short period of time. These three properties raise the species
above the typological interpretation of a “class of objects” (Mayr, 1963,
p. 21). The species definition which results from this theoretical species
concept is:

Species are groups of interbreeding natural populations that are re-

productively isolated from other such groups.

The development of the biological concept of the species is one of

the earliest manifestations of the emancipation of biology from an inappro¬
priate philosophy based on the phenomena of inanimate nature. This species
concept is called biological not because it deals with biological taxa, but
because the definition is biological. It utilizes criteria that are meaningless
as far as the inanimate world is concerned.
When encountering difficulties (see also 2.5) it is important to focus
on the basic biological meaning of the species: A species is a protected
gene pool. It is a Mendelian population which has its own devices (called
isolating mechanisms) which protect it against harmful gene flow from
other gene pools. Genes of the same gene pool form harmonious combina¬
tions because they have become coadapted by natural selection. Mixing
the genes of two different species leads to a high frequency of disharmonious
gene combinations; mechanisms that prevent this are therefore favored
by selection. This makes it quite clear that the word species in biology
is a relational term: A is a species in relation to B and C because it is
reproductively isolated from them. It has its primary significance with re¬
spect to sympatric and synchronic populations, and these are precisely the
situations where the application of the concept faces the fewest difficulties
 The Species Category 27

(“the nondimensional species”). The more distant two populations are

in space and time, the more difficult it becomes to test their species status
in relation to each other, but the more irrelevant biologically this also
becomes.
The biological species concept also solves the paradox caused by the
conflict between the fixity of the species of the naturalist and the fluidity
of the species of the evolutionist. It was this conflict which made Linnaeus
deny evolution and Darwin the reality of species (Mayr, 1957fr). The bio¬
logical species combines the discreteness of the local species at a given
time with an evolutionary potential for continuing change.
The unique position of species in the hierarchy of taxonomic categories
has been pointed out by many authors. Taxa of the species category can
be delimited against each other by operationally defined criteria (“inter¬
breeding versus noninterbreeding of populations”). It is the only taxonomic
category for which the boundaries between the taxa at that level are defined
objectively.
Logicians (e.g. Gregg, 1954; Buck and Hull, 1966) do not fully under¬
stand that the terminology of “class” and “member” fails to bring out
the complete difference between, on one hand, the relation of individuals
to the species and, on the other hand, of species taxa to higher taxa. The
statement that something is “a member of a class” has an entirely different
meaning for an individual which through its genotype is a member of
a species, and a species taxon which is included in a higher taxon. A
category is not a class in the same sense as a higher taxon, but a designation
of rank. Logicians do not appreciate that the “higher” and “lower” rank of
taxa is a relative, not an absolute property. One can compare taxa in
a single phyletic line, but one cannot say that the genus is the same thing
in birds, ammonites, bivalve mollusks, protozoans, and weevils, and that
they are all equal. Again, the species is an exception because (at least
in sexual species) the species is an equivalent phenomenon in all groups
of animals.
Intraspecific categories designate groupings of populations within spe¬
cies. Normally, however, the species is the lowest category used in routine
taxonomy. The higher categories are groupings of species. In view of this
key position of the species and the fact that in nature one encounters
individuals and phena, the assigning of individuals and phena to species
taxa is one of the key problems of taxonomy.

2.3 FROM PHENON TO TAXON TO CATEGORY

A failure to understand the meaning of these three terms and their

theoretical foundation has led taxonomists into much confusion. It has
been the cause of most attacks on the biological species concept. When
28 Principles of Zoological Classification

an author says, “As a paleontologist I cannot employ the biological species

concept because I cannot test the reproductive isolation of fossils,” he reveals
his lack of understanding. What the taxonomist observes directly are indi¬
viduals which he sorts into phena. On the basis of certain biological concepts
and information, such as an awareness of the possibility of sexual dimor¬
phism, growth, alternation of generations, nongenetic modifications of the
phenotype, etc., he assigns the phena to populations, which in turn he
classifies into taxa. The ranking of a taxon as subspecies, species, or genus
by a taxonomist is based on inferences drawn from the available data.
This methodology of basing inferences on evidence and its justification
has been perceptively discussed by Simpson (1961, pp. 68-69) :

“Evolutionary classification uses, for the most part, concepts and definitions
for which the data are not directly observable. This is not a feature peculiar
to taxonomy. It is shared in greater or less degree by most of the inductive
sciences. . . . Here it is necessary again to emphasize the distinction between
definition and the evidence that the definition is met. We propose to define
taxonomic categories in evolutionary and to the largest extent phylogenetic
terms, but to use evidence that is almost entirely non-phylogenetic when
taken as individual observations. In spite of considerable confusion about
this distinction, even among some taxonomists, it is really not particularly
difficult or esoteric. The well-known example of monozygotic (“identical”)
twins is explanatory and is something more than an analogy. We define
such twins as two individuals developed from one zygote. No one has ever
seen this occur in humans, but we recognize when the definition is met
by evidence of similarities sufficient to sustain the inference. The individuals
in question are not twins because they are similar but, quite the contrary,
are similar because they are twins. Precisely so, individuals do not belong
in the same taxon because they are similar, but they are similar because
they belong to the same taxon. (Linnaeus was quite right when he said
that the genus makes the characters, not vice versa, even though he did
not know what makes the genus.) That statement is a central element
in evolutionary taxonomy, and the alternative clearly distinguishes it from
non-evolutionary taxonomy. Another way to put the matter is to say that
categories are defined in phylogenetic terms but that taxa are defined by
somatic relationships that result from phylogeny and are evidence that the
categorical definition is met.

The reproductive isolation of a biological species, the protection of

its collective gene pool against pollution by genes from other species, results
in a discontinuity not only of the genotype of the species, but also of
its morphology and other aspects of the phenotype produced by this geno¬
type. This is the fact on which taxonomic practice is based. Reproductive
isolation cannot, of course, be directly observed in samples of preserved
specimens. However, it can be inferred on the basis of various types of
evidence, as for instance the presence of a discontinuity, a bridgeless gap,
 The Species Category 29

between two correlated character complexes. In living species, of course,

such inferences can be tested by observation and experiment.
The crucial difference between the reasoning of the typologist and
the adherent of the biological species concept is as follows: The typologist
says, “There is a clear-cut morphological difference between samples a
and b, therefore they are, by definition, two morphospecies, that is, two
species.” Any list of synonymies will quickly reveal how often this philosophy
has led to the description of phena as species. The biological taxonomist
asks, “Is the morphological difference between samples a and b of the
kind one would expect to find between two reproductively isolated popula¬
tions, that is, between two biological species?” In other words, he uses
the amount and kind of morphological difference only as an indication
of reproductive isolation, only as evidence to draw an inference. This is
a legitimate and reliable technique. Where the typologist would recognize
phena as (morpho-) species, the biologist will draw the right inferences
from largely morphological evidence, and his species are usually confirmed
by subsequent researches. When competent taxonomic work based on mor¬
phological evidence is re-examined in the light of the findings of behavior
or biochemistry, it is usually confirmed in its entirety.
It is not always realized that the classification of phena is based on
entirely different evidence than the classification of species. The classifica¬
tion of species is based on weighted similarity, evaluating all sorts of com¬
parative data, be they morphological, physiological, behavioral or what
not. The classification of phena is based on their relation to the gene pool
of the population to which they belong. Ultimately this can be established
only by breeding behavior. This in turn can be either observed in nature
or studied experimentally. It does not matter whether one deals with strik¬
ingly different sexes in birds, insects, or marine invertebrates, or with larval
forms, or alternating generations of parasites; breeding alone (or the piecing
together of growth stages) will establish what phena together form a popula¬
tion. The experienced taxonomist knows what variation to expect within
a biological species. No computer method has yet been found that would
empirically assign phena to species. The taxonomist does this rapidly and
with a high degree of precision on the basis of his accumulated knowledge
of the biology of the species concerned. In this taxonomic operation the
classical methods still reign supreme “because they are enormously faster
than the numerical methods” (Michener, 1963).

2.4 SPECIES NAMES

The scientific name (binomen) of a species consists of two words,

the generic name and the specific name. The rules concerning the coining
of species names and all other aspects of the nomenclature of such names
30 Principles of Zoological Classification

are discussed in Chap. 13 (particularly 13.31, 13.40, and 13.43), where

cross references to the appropriate articles of the Code (Chap. 12) are
also provided.
Species names, to become available, must be accompanied by a de¬
scription. Advice concerning the preparation of descriptions is given in
Chap. 11.

2.5 DIFFICULTIES IN THE APPLICATION OF THE

BIOLOGICAL SPECIES CONCEPT

The fact that difficulties sometimes arise when the biological species
concept is applied to natural taxa does not mean that the concept as such
is invalid. This has been shown by Simpson (1961, p. 150) and Mayr
(1963, pp. 21-22).
Many generally accepted concepts face similar difficulties when they
have to be applied in a particular situation or to a specific sample. The
concept tree, for instance, is not invalidated by the existence of spreading
junipers, dwarf willows, giant cacti, and strangler figs. One must make
a clear distinction between a concept and its application to a particular
case.
The more ordinary problems of taxonomic discrimination at the spe¬
cies level, in particular the criteria for ranking a taxon as a species rather
than a subspecies, will be dealt with in Chap. 9B.2.
The three most serious difficulties in the application of the biological
species concept, discussed in this chapter, are those caused by the lack
of pertinent information, those caused by uniparental reproduction, and
those caused by evolutionary intermediacy.
2.5.1 Insufficient Information. Individual variation in all of its forms
often raises doubt as to whether a certain morphotype is a separate species
or only a phenon within a variable population. Sexual dimorphism, age
differences, polymorphism, and other such types of variation can be un¬
masked as individual variation through a study of life histories and through
population analysis. This is dealt with more fully in Chap. 8A and 8B.
The neontologist who normally works with preserved material is confronted
by the same problems as the paleontologist, who likewise must assign phena
(morphotypes) to species.
2.5.2 Uniparental Reproduction. In many organisms systems of re¬
production are found that are not based on the principle of an obligatory
recombination of genetic material between individuals prior to the forma¬
tion of a new individual. Self-fertilization, parthenogenesis, pseudogamy,
and vegetative reproduction (budding or fission) are some of these forms
of uniparental reproduction.
 The Species Category 31

A population, as defined in evolutionary biology, is an interbreeding

group. By definition, therefore, an asexual biological population is a contra¬
diction, even though the word population has other usages in which a
combination with asexual is not contradictory. The biological species con¬
cept based on the presence or absence of interbreeding between populations
is therefore inappropriate for uniparentally reproducing organisms.
How to solve this dilemma has been discussed by Simpson (1961,
pp. 161-163) and by Mayr (1963, pp. 27-29). Attempts to define agamo-
species or asexual species, with or without using the word population, have
not been particularly successful. Fortunately, there are usually well-defined
morphological discontinuities among kinds of uniparentally reproducing
organisms. These discontinuities are apparently produced by natural selec¬
tion among the various mutants which occur in the asexual clones. It is
customary to utilize the existence of such discontinuities, and the amount
of morphological difference among them, to delimit species among uni¬
parentally reproducing types.
Species recognition among asexual organisms is based not merely on
analogy but also on the fact that each of the morphological entities, sepa¬
rated by a gap from other similar entities, seems to occupy an ecological
niche of its own; it plays its own evolutionary role. In groups like the
bdelloid rotifers, all of which reproduce by obligatory parthenogenesis, there
is evidence for a definite biological meaning to the recognized morphological
species. The treatment of asexual entities that do not qualify as species
is discussed in Chap. 3 (3.3.8).
Examples are known in which a form that is as distinct as a good
species reproduces strictly parthenogenetically, and no biparental species
is known from which it might have branched off. Nomenclatural recognition
is justified in such cases. Whenever several reproductively isolated chromo¬
some types occur within such a “species,” as in various crustaceans (e.g.
Artemia salina Linnaeus) (White, 1954), it may be convenient to distin¬
guish them nomenclaturally. Although conventionally referred to as races,
reproductively isolated chromosomal populations are more logically desig¬
nated (micro-) species.
2.5.3 Evolutionary Intermediacy. The species, as manifested by a
reproductive gap between populations, exists in full classical distinctness
only in a local fauna. As soon as the dimensions of space (longitude and
latitude) and time are added, the stage is set for incipient speciation. Popu¬
lations will be found under these circumstances which are in the process
of becoming separate species and have acquired some but not yet all of
the attributes of distinct species. In particular, the acquisition of morpho¬
logical distinctness is not always closely correlated with the acquisition of
reproductive isolation. The various difficulties for the taxonomist which
may result from evolutionary intermediacy may be tabulated as follows
(see also Mayr, 1957a, p. 375) :
 32 Principles of Zoological Classification

1. Acquisition of reproductive isolation without equivalent morphological

change. Reproductively isolated species without (or with very slight) morphologi¬
cal difference are called sibling species. Their taxonomic treatment will be dis¬
cussed in Chap. 9A.2.
Acquisition of strong morphological difference without reproductive isola¬
tion. A number of genera of animals and plants are known in which morphologi¬
cally very different populations interbreed at random wherever they come in con¬
tact. The typological solution of calling every morphologically distinct population
a species is clearly inappropriate in such situations. Conversely, there are genera in
which the isolating mechanisms between any two species may break down occa¬
sionally. To consider such species conspecific would be going to the opposite ex¬
treme. No generalized solution is possible where morphological divergence and
acquisition of reproductive isolation do not coincide. The only recommendation to
the specialist is that he delimit his species in such a way that they form biologically
meaningful, natural entities. The difficulty posed by the rapid morphological diver¬
gence of populations without acquisition of reproductive isolation is well-illustrated
by the West Indian snail genus Cerion (see Fig. 2-1).
3. The occasional breakdown of isolating mechanisms (hybridization). Re¬
productive isolation may break down occasionally even between good species.
Most frequently this will lead only to the production of occasional hybrids that are
either sterile or of lowered viability, and this will not cause any taxonomic difficulty.
More rarely, there is a complete local breakdown of isolation resulting in the pro¬
duction of extensive hybrid swarms and more or less complete introgression
(Mayr, 1963, pp. 110-135).
_ Hybrid individuals are sometimes described as species before their hybrid
nature is discovered. Such names lose their validity as soon as the hybridism is
established (see 13.21). Only populations are recognized as taxa, and hybrid in¬
dividuals are not populations.
Taxonomically more difficult are situations where new populations are
formed as a result of hybridization. We can recognize several types of natural
populations that owe their origin to hybridization. The taxonomic treatment of
secondary intergradation, the result of the fusion of previously isolated populations,
is discussed in Chap. 9B.2b. Two other kinds of hybridism concern us here.
3a. Sympatric hybridization. In all instances in which the two parental
species maintain their genetic integrity over a more or less wide area in which
they occur together, it is advisable to uphold their species status even though in a
portion of their ranges there is a breakdown of the isolation. The example of the
two Mexican towhees (Pipilo erythrophthalmus and P. ocai) is an excellent
illustration of this situation (Sibley, 1954) (Fig. 2-2). No taxonomic recognition
is given to hybrid populations that result from such a local breakdown of reproduc¬
tive isolation. The only possible exception is such a complete breakdown of the
isolation that the two parental species fuse into a single new species. The taxonomic
literature records a number of instances that have been interpreted in this manner,
but we are not aware of a single thorough analysis that would have established
this unequivocally.
3b. Amphiploidy. Hybridism in plants may lead to the instantaneous pro¬
duction of an allopolyploid, an individual that combines the chromosome sets of
two parental species. Such hybrids may give rise by uniparental reproduction to a
 The Species Category 33

Cerion

Banes

Fig. 2-1. Irregular distribution of populations of the halophilous land snail Cerion in eastern
Cuba. Numbers refer to different races or species. Where two populations come in contact (with
the exception of lepida) they hybridize (//), regardless of difference. In other cases contact is
prevented by a barrier (B). In = isolated population (from Mayr, 1963).

new population that is reproductively isolated from both parents and behaves like
a new species if it is able to reproduce and, by occupying a new ecological niche,
is able to compete with other species (including the parents). It is doubtful that
such speciation by amphiploidy has ever been incontrovertibly established in sex¬
ually reproducing animals. Polyploidy among parthenogenetically reproducing
insects, annelids, and turbellarians is not infrequent, but it is not certain that any
of these parthenogenetic lines owes its origin to hybridization.
4. Semispecies. Geographical isolates occasionally have an intermediate
status between subspecies and species. On the basis of some criteria they would be
considered species; on the basis of others, they would not. It is usually more con-
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 The Species Category 35

venient for the taxonomist (see 9B.2) to attach such doubtful populations to the
species with which they are most nearly allied. Circular overlaps and other
borderline cases (Mayr, 1963, pp. 496-512) are other instances of evolutionary
intermediacy that will have to be decided from case to case on the basis of con¬
venience and degree of evolutionary intermediacy.

2.6 SPECIAL SITUATIONS

2.6.1 Paleontology. The paleontologist has to cope with two special

kinds of difficulties in species delimitation in addition to those which the
neontologist encounters.
These difficulties have been discussed frequently and with considerable
detail in the paleontological literature. We refer to Imbrie (1957), Sylves-
ter-Bradley (1956, 1958), Simpson (1960; 1961, pp. 152-155 and 163-171),
and various articles in paleontological journals. As these authors point out,
the difficulties encountered by the paleontologist are often exaggerated.
Many paleontologists have confused phenon and species, taxon and cate¬
gory, and evidence and inference. A clear understanding of these terms
and the underlying concepts removes much of the difficulty.
Evolutionary Continuity. Species are evolving systems, and the vertical
delimitation of species in the time dimension should in theory be impossible.
Unbroken sequences of fossil populations are, however, extremely rare.
Where they exist, the named morphospecies are usually so similar that
they are better not recognized at all or, at most, ranked as subspecies.
In most fossil sequences there are convenient breaks between horizons
to permit a nonarbitrary delimitation of species. This is true even for Brink-
mann’s (1929) remarkably complete sequence of Jurassic ammonites (Kos-
moceras). Since much speciation occurs in peripheral isolates, the discovery
of strata with intermediate populations (incipient species) is highly im¬
probable and will occur only rarely. Range fluctuations contribute to the
appearance of breaks even in cases of speciation owing to uncomplicated
phyletic evolution in a single vertical column (cf. Fig. 3-8). An evolutionary
species may be absent from a given locality for shorter or longer periods,
and when it reoccurs, it may have changed sufficiently to be classified
as a different species.
Amount of Evidence Limited. Taxonomic evidence supplied by behav¬
ior, chromosomes, properties of proteins and other chemical constituents, and
by other attributes of living populations, is not available to the paleontolo¬
gist. However, most classification of living species is also made without
access to such information, and when it finally becomes available it usually
confirms the existing classification. As Imbrie (1957) and others have
pointed out, the paleontologist has far more information available than
36 Principles of Zoological Classification

that of mere morphology. The variation of his samples, the associated

faunas, the paleoecology, the geography, horizon and time level, etc., con¬
tribute abundant information that facilitates inferences on the delimitation
of taxa and their categorical ranking. The analysis, as in much neontological
work, often has to begin with a separation of the samples into phena,
but a consideration of all the collateral evidence usually allows an unequivo¬
cal assignment of these phena to species or subspecies, as was well demon¬
strated by Sylvester-Bradley (1958) for the two chronological species Ostrea
knorri knorri and O.k. lotliaringica (Table 2-1).

Table 2-1. Frequency of 12 Phena in Two Subspecies of Ostrea

knorri (from Sylvester-Bradley, 1958)

12 Phena Schonmatt (knorri) Geisingen (lotharingica)

D + B + M 49 0
A + G 41 5
K + J + E 10 5
F + H 3 14
C + L 0 45
Total 103 69

2.6.2 Parasites. Populations of parasites on different hosts are often

slightly different. There are three possibilities for interpreting these morpho¬
logical differences.

1. The differences are caused by nongenetic modification resulting from the

different physiological environments of the different hosts. Many trematodes may
mature in a large number of possible hosts and show great differences. Specimens
of the liver fluke Fasciola hepatica from a cow, a rabbit, and a guinea pig mani¬
fest differences far greater than those usually employed to distinguish species
(Stunkard, 1957). It is possible that large numbers of nominal species in certain
genera of trematodes such as Hymenolepis may be shown eventually to be nothing
but host-induced, nongenetic modifications.
2. The differences are indicative of subspecific rank. The differences between
host populations of Mallophaga, although constant, are often so slight that they
are best treated as subspecies (Clay, 1958). Host separation in this case corre¬
sponds to geographic isolation in free-living species, and the amount of gene flow
between Mallophaga occurring on different host species is presumably very slight.
3. The differences are indicative of rank as full species. Numerous cases
have been described in the parasitological literature where exceedingly similar
parasites could not be transferred from one host to another. There is no oppor¬
tunity for gene interchange in the case of species without intermediate host. In
such a case, even though the morphological difference is comparatively minor, one
must assume that the genetic barrier has reached species level.
Chapter The Polytypic Species
O Population Systematics
and Infraspecific Categories

A
-A- -A_t a given locality a species of animal is usually sepa¬
rated from other sympatric species by a complete gap. This is the species of
the local naturalist, the species of Ray and Linnaeus. It may also be called
the nondimensional species because it lacks the dimensions of space and time.
Combining the properties of a species and of a single local population,
the nondimensional species can usually be delimited unequivocally.
Every species consists, however, of numerous local populations, and
some of these are visibly different from each other. If a taxonomist finds
a population which he considers sufficiently distinct from the population
of the original type locality of the species (or of other previously named
subspecies), he calls it a new subspecies (see below). Species that contain
two or more subspecies are called polytypic species. Species that are not
subdivided into subspecies are called monotypic species. Recognition of
the fact that many species, particularly widely distributed species, are poly¬
typic was one of the most important developments in taxonomy. For a
full treatment of this development and of various aspects of the polytypic
species, see Mayr (1963, chap. 12).
37
38 Principles of Zoological Classification

3.1 IMPORTANCE OF POLYTYPIC SPECIES

The greatest benefit derived from the recognition of polytypic species

is that in well-known groups of animals such as birds, mammals, butterflies,
or snails it has led to a considerable simplification of the classification.
The reclassification into polytypic species of geographically representative
forms that had originally been separately described as monotypic species
led to a great clarification of the system. This reorganization of classification
on the species level is virtually completed in birds. It is in full swing for
mammals and some groups of insects and land mollusks, but has hardly
begun in most other groups of animals. The 19,000 monotypic species of birds
listed in 1910 (together with numerous species discovered since then) have
now been reduced to about 8,600 species. A similar simplification has been
reported for many other groups of vertebrates and invertebrates. Of much
greater significance is the restoration to the species category of a definite
biological meaning and homogeneity. Awarding species rank to every local
population, no matter how slight its difference, completely destroyed the
biological significance of the species category.
The task of assembling local populations into polytypic species, or,
more broadly, of sorting large numbers of “nominal species” and “varieties”
into polytypic species, reveals many taxonomically and biologically interesting
situations (Mayr, 1963, p. 343). The consistent application of the polytypic
species concept to all groups of animals is one of the major tasks of
taxonomy.
3.1.1 Difficulties. When establishing polytypic species the taxonomist
encounters two difficulties. Polytypic species are composed of allopatric
or allochronic populations that differ from each other. However, all popula¬
tions of sexually reproducing organisms differ slightly from each other,
and certain standards must be met before subspecies can be recognized
(9B.1). The other difficulty is that, occasionally, closely related species
with similar ecological requirements replace each other geographically and
yet are full species and not subspecies. How to choose between these two
alternatives will be discussed in 9B.2.

3.2 OCCURRENCE OF POLYTYPIC SPECIES IN

THE ANIMAL KINGDOM

The frequency of polytypic species differs from animal group to animal

group. They are most frequent where species tend to form geographic iso¬
lates. In most well-studied groups between 40 and 80 percent of the species
are polytypic, but some highly specialized species, particularly host-plant
 The Polytypic Species Population Systematics and Infraspecific Categories 39

Fig. 3-1. The distribution of 35 subspecies of the kangaroo rat Dipodomys ordii Woodhouse, an
example of a range map of a polytypic species (after Setzer). Numbers designate the ranges of the
various subspecies.
40 Principles of Zoological Classification

Fig. 3—2. Type localities (1-15) of fifteen subspecies of the scorpion Mesobuthus eupeus in the
middle east (from Vachon, 1958).

specific insects, do not readily form polytypic species. Polytypic species are
also scarce or absent in groups with slight species differences (e.g., groups
of sibling species).
A number of other technical terms, such as Formenkreis (Lorenz,
Kleinschmidt) and Rassenkreis (Rensch), have been applied to polytypic
species but have not become established.
The polytypic species is, in a sense, the lowest of the higher categories.
Being multidimensional it lacks the simplicity and objectivity of the nondi-
mensional species. Most of the difficulties of delimiting species of animals
concern situations where there is doubt as to whether two allopatric popula¬
tions belong to the same polytypic species or not. Among birds, even such
borderline cases are in the minority. The claim that such difficulties are
more frequent in other groups of animals still awaits verification.
3.2.1 Nomenclatural Problems. A polytypic species is often a com¬
pound of several “species” originally proposed as monotypic. It differs thus
from the Linnaean species by no longer being the lowest category (which
is now the subspecies) and by being a collective category. What scientific
name should be given to this new collective taxon, and who should be
the author? When Linnaeus named the white wagtail Motacilla alba, to
cite one example, he had in mind the Swedish population with the diagnos¬
tic characters described by him. The M. alba of Linnaeus is now called
The Polytypic Species Population Systematics and Infraspecific Categories 41

the nominate subspecies M. alba alba Linnaeus. When M. alba of Linnaeus

was combined with eight or more other taxa, all originally described as
separate species (lugubris Temminck, dukhunensis Sykes, baicalensis Swin-
hoe, leucopsis Gould, personata Gould, hodgsoni Blyth, ocularis Swinhoe,
lugens Kittlitz, etc.), the newly-formed polytypic species was something
very different from the original M. alba of Linnaeus. If we still associate
the name Linnaeus with the new polytypic species M. alba, it is to indicate
Linnaeus as the author of the name alba, and not of the drastically recon¬
stituted taxon to which we now attach the name.

3.3 INFRASPECIFIC CATEGORIES AND TERMS

3.3.1 The Variety. This, as varietas, was the only subdivision of

the species recognized by Linnaeus. It designated any deviation from the
type of the species. As a consequence the varieties of the early taxonomist
were a heterogeneous potpourri of individual variants (see Chap. 8) and
various kinds of races. As a result of this confusion the term variety came
into discredit and is hardly used any longer by animal taxonomists. For
a further discussion see Simpson (1961, p. 177) and Mayr (1963, p. 346).
3.3.2 The Subspecies. The term subspecies when it came into general
usage during the nineteenth century was a replacement for the term variety
in its meaning of “geographic race.” It was considered a taxonomic unit
like the morphological species, but at a lower taxonomic level. Many early
authors used the term subspecies indiscriminately, almost like the term
variety, for distinguishable entities that were not as distinct as species. Ant
specialists, for instance, employed the term for sibling species and for indi¬
vidual variants. When an author reports several subspecies of one species
from the same locality, it strongly indicates a wrong usage of the term.
Subspecies are normally allopatric and allochronic, exceptions occurring,
however, in migratory species and in parasites with sympatric host subspe¬
cies. A purely morphological definition of the subspecies, as attempted by
typologists, often results in a sympatry of entities, thus defined. In view
of the many misuses of the term it must be emphasized that the subspecies
is a category quite different from the species. No nonarbitrary criterion
is available to define the category subspecies. Nor is the subspecies a unit
of evolution, except where it happens to coincide with a geographical or
other genetic isolate.
The subspecies may be defined as follows: A subspecies is an aggregate
of phenotypically similar populations of a species, inhabiting a geographic
subdivision of the range of a species, and differing taxonomically from
other populations of the species.
42 Principles of Zoological Classification

The reasons for the particular wording of the definition are as follows:

1. A subspecies may consist of many local populations all of which, though

very similar, are slightly different from each other genetically and pheno-
typically; a subspecies is therefore a collective category.
2. Every local population is slightly different from every other local popula¬
tion, and the presence of these differences can be established through
sufficiently sensitive measurements and statistics. It would be absurd and
would lead to nomenclatural chaos if each such population were given
the formal trinominal name that is customary for subspecies. Therefore,
subspecies are to be named only if they differ “taxonomically,” that
is, by sufficient diagnostic morphological characters (see 9B.1).
3. Even when it is possible to assign populations to subspecies, an assignment
on the basis of phenotype alone is not necessarily possible for every
individual, owing to an overlap of the ranges of variation of neighboring
populations.

The term overlap is often misused. The breeding ranges of two species
may overlap geographically but not those of two subspecies of the same
species. If two discrete breeding populations coexist at the same locality,
they are full species (except in the rare cases of “circular overlap”). Where
two subspecies meet, intermediate or hybrid populations may occur which
combine the characters of both subspecies. It would be misleading in such
a case to say that the two subspecies overlap in this area, since the species
is represented in this area only by a single population, no matter how
variable.
Difficulties in the Application of the Subspecies Category. Recognition
of the polytypic species requires the use of the subspecies category, with
all the concomitant benefits described above. However, various aspects of
geographic variation cause difficulties. Indeed, the subspecies has been mis¬
used in many ways. Some authors applied the term to individual variants
and sibling species, many authors named insignificant local populations
as subspecies, and finally some authors considered the subspecies as a unit
of evolution rather than as an arbitrary device to facilitate intraspecific
classification. As a result the practice of describing subspecies was criticized
by numerous authors, most cogently by Wilson and Brown (1953) (see
also Inger, 1961). They pointed out four aspects of the subspecies which
reduce its usefulness:

1. The tendency of different characters to show independent trends of geo¬

graphic variation
2. The independent reoccurrence of similar or phenotypically indistinguish¬
able populations in geographically separated areas (“polytopic subspe¬
cies”—see Sec. 9B.1.4)
3. The occurrence of microgeographic races within formally recognized
subspecies
 The Polytypic Species Population Systematics and Infraspecific Categories 43

4. The arbitrariness of the degree of distinction considered by different

specialists as justifying subspecific separation of slightly differentiated
local populations

Numerous articles on the pros and cons of the subspecies question

can be found in volumes 3 to 5 of Systematic Zoology (1954-1956). For
general reviews of the subspecies question see also Simpson (1961, pp.
171-176) and Mayr (1963, pp. 347-350).
Recent arguments have, we hope, led to a more critical attitude to¬
ward subspecies. It has also shown that sensible use of the category sub¬
species is still a convenient device for classifying population samples in
geographically variable species.
Practical problems which the population taxonomist frequently faces
will be dealt with in Chap. 9. They concern particularly the following
questions: How different must a population be in order to justify subspecific
recognition? How shall one treat intermediate populations? How does one
delimit subspecies against adjacent subspecies? Shall one recognize polytopic
subspecies for indistinguishable but geographically separated populations?
When should geographical isolates be called species, and when subspecies?
The Nomenclature of Subspecies. A trinomen is used to designate
a subspecies taxon (Arts. 5 and 45 of the Code). For instance, the British
Red Deer is called Cervus elaphus scoticus, while the continental Red
Deer, originally named by Linnaeus binominally, becomes the nominate
subspecies Cervus elaphus elaphus. For further nomenclatural aspects see
13.40 and 13.41; see also 13.25 (combined and divided taxa), 13.36 (sub¬
ordinate taxa), 13.46 (homonymy among subspecies), and 13.57 (type-
localities) .
3.3.3 Temporal Subspecies. In paleontology slightly different popu¬
lations separated in time are increasingly often assigned to the subspecies
category. It does not seem advisable to make a terminological distinction
between geographical and temporal subspecies because it is usually impossi¬
ble, when different subspecies of a fossil species are found at different
localities, to determine whether or not they are precisely contemporary.
Even when there is a sequence of subspecies at a single locality, it need
not necessarily be purely temporal. Subspecies found in succeeding strata
may actually be geographic races that replaced each other owing to climatic
changes (Fig. 3-3).
The paleontologist, when applying the subspecies category, faces cer¬
tain difficulties not encountered by the neontologist. There may be a differ¬
ential deposition of various age classes and sexes in different horizons, as
well as the occurrence of nongenetic habitat forms. Yet treating fossils
as the remains of formerly existing populations usually leads to a deeper
analysis as well as to a better understanding of relationships and of the
meaning of evolutionary trends. As with living species, it must be kept
44 Principles of Zoological Classification

Fig. 3—3. (a) Geography and time in subspeciation and speciation. Diagram illustrating how geo¬
graphical fragmentation of successive populations (the numbered rectangles) may accompany
vertical differentiation of a phyletic line. The populations rarely remain in one locality for long but
migrate. Some migrants become isolated from the parent stock by barriers, becoming ultimately
differentiated into geographical races. The faunal succession in any locality (A or B) is never absolutely
continuous, even though gaps may be obscure. The gaps may be produced by migrations, by deposi-
tional hiatus, and by local extermination. (b) A population becomes divided by a barrier causing
partial isolation with limited gene flow for a time—the subspecies stage in speciation. After sufficient
genetic differentiation has been reached, interbreeding ceases, gene flow is stopped, and the two
branches become separate species (Newell, 1947).

in mind at all times that the subspecies is merely a classificatory device.

For further discussions on the subspecies in paleontology see Newell (1947,
1956a), Sylvester-Bradley (1951, 1956), and Simpson (1961, pp. 175-176).
3.3.4 Race. A race that is not formally designated as a subspecies
is not recognized in the taxonomic hierarchy. However, the terms subspecies
and geographical race are frequently used interchangeably by taxonomists
of mammals, birds, and insects. Other taxonomists apply the word race
to local populations within subspecies.
The nature of ecological races among animals is still controversial
(Mayr, 1963, p. 355). Since no two localities are exactly identical with
respect to their environment, every subspecies is at least theoretically also
an ecological race. However some populations differ in their ecological
requirements without acquiring taxonomically significant differences. More
important from the taxonomic as well as the evolutionary point of view
are host races among parasites and species-specific plant feeders. If gene
flow between populations on different hosts is drastically reduced, such
host races are the equivalent of geographic races in free-living animals.
Also, such host races often develop subspecific characters.
3.3.5 Cline. This term was coined by J. S. Huxley (1939) for a
character gradient. It is not a taxonomic category. A single population
may belong to as many different dines as it has characters. A cline is
 The Polytypic Species Population Systematics and Infraspecific Categories 45

formed by a series of contiguous populations in which a given character

changes gradually. At right angles to the cline are the lines of equal expres¬
sion of the character (points of identical phenotype) ; such a line is called
an isophene. For instance, if in the range of a species of butterfly the per¬
centage of white specimens varies from north to south, the corresponding
isophenes may be indicated on a map (Fig. 3-4).
Any character, be it a morphological, physiological, or other geneti¬
cally determined character, may vary clinally. Clines may be smooth, or

Fig. 3-4. Cline in the darkness of the upper side at different Fennoscandian localities of Pieris
napi females first generation. Isophenes of various darkness values indicated on the map (from
Petersen, 1949).
46 Principles of Zoological Classification

they may be “step clines” with rather sudden changes of values. Clines
do not receive nomenclatural recognition. Indeed, when the geographic
variation of a species is clinal, it is usually inadvisable to recognize subspe¬
cies, except possibly for the two opposite ends of the dine when they are
very different or separated by a pronounced step. For a further discussion
of clines see Simpson (1961, p. 178) and Mayr (1963, pp. 361-366).
3.3.6 Infrasubspecific Categories. The subspecies is the lowest tax¬
onomic category recognized in the Code of Nomenclature (Art. 45c) (see
13.41).
In the days when the subspecies was still defined typologically, many
proposals were made to subdivide heterogeneous subspecies into still smaller,
hopefully uniform taxa, and terms were proposed for such taxa, e.g. natio.
Now that it is being realized that every local population is different from
every other one, even if they live only a few miles apart or less, and that
these populations are not sharply separated from each other (except where
separated by barriers), there is no longer any excuse for a formal recognition
of innumerable local subdivisions of subspecies. The term deme adopted
by zoologists for the evolutionary unit corresponding to a local population
(Mayr, 1963, p. 137) is not the name of a taxonomic category.
3.3.7 Intrapopulation Variants. Taxa are populations, and popula¬
tions are the material of classification. Phena, composed of intrapopulation
variants, have no taxonomic status and deserve no formal recognition in
nomenclature. They can be referred to by a vernacular terminology, e.g.
“albinos,” as discussed in 13.41. The morphs found in polymorphic popula¬
tions are such variants.
3.3.8 Asexual Entities. Uniparental reproduction through partheno¬
genesis, vegetative budding, or fission is not infrequent among the lower
invertebrates, with parthenogenesis occurring even among insects and lower
vertebrates up to reptiles. Since interbreeding is the ultimate test of con-
specificity in animals, and since this criterion is available only in sexually
reproducing organisms, determination of categorical rank is difficult for
taxa of uniparentally reproducing organisms. How should the taxonomist
treat clones, pure lines, biotypes, and so-called “strains” or “stocks” of
such organisms?
Parthenogenesis is usually only a temporary condition in animals. In
aphids, cladocerans, rotifers, and various others, females of many species
are parthenogenetic during part of the year but return to sexual reproduc¬
tion when environmental conditions change. Nomenclatural recognition is
not given to such temporary clones. They terminate sooner or later, either
by extinction or by returning to the common gene pool of the parental
sexually reproducing species.
In the case of permanently uniparentally reproducing lines, the species
category is applied on the basis of degree of morphological difference. Mor-
 The Polytypic Species Population Systematics and Infraspecific Categories 47

phological difference between clones can be used as an indication of the

underlying genetic difference and this in turn for an inference on probable
species status. To group uniparentally reproducing organisms into distinct
taxa is often not difficult. The ranking of these taxa, that is their assignment
to a definite category, is almost invariably somewhat arbitrary. Nature,
unfortunately, is not always as tidy as the taxonomist would wish.
3.3.9 Neutral Terms. It is very convenient in taxonomic work to
have some terms that can be given informally to phena, particularly in
incompletely analyzed cases. These are the so-called “neutral terms.” The
ones that are most frequently used in taxonomy are form, for a single
unit, and group or complex for a number of units. We may speak of
a “form” when we do not know whether the phenon in question is a
full species or a subspecies, or whether it is a subspecies or an individual
variant. Seasonal and polymorphic variants are often referred to as forms.
The term is also used in the plural when two unequal units are referred
to. For instance, when describing attributes common to a species and a
subspecies of another species, one may refer jointly to the species and the
subspecies as “these two forms.”
The term group is more commonly applied to an assemblage of
closely related taxa which one does not want to place in a separate category.
In the large genus Drosophila, for example, numerous species groups are
recognized, such as the melanogaster group, the virilis group, the obscura
group, and so forth. A species group is a group of closely related and
presumably recently evolved species. The use of the species group in formal
taxonomy has been spreading in recent years, because it reduces the need
for the recognition of subgenera. In large polytypic species the term group
is also applied to subspecies groups. The common Palearctic jay, Garrulus
glandarius, has a total of 28 subspecies which can be arranged in seven
subspecies groups, the garrulus group, the bispecularis group, and others.
The term group is used more rarely for aggregates of genera and other
higher categories. The word complex is sometimes used synonymously with
the word group.
Terms like section, series, and division, are generally used for the
higher categories. Their use is not standardized, however, and in different
branches of systematic zoology they may be used above or below the family,
the order, or the class.

3.4 POPULATION TAXONOMY

The increasing preoccupation with the description of new subspecies

and the establishment of polytypic species, beginning in the second half
of the nineteenth century and continuing into the present, resulted in a
48 Principles of Zoological Classification

subtle change of emphasis and outlook. The emerging understanding of

the species taxon as a geographically variable aggregate of populations
accelerated the replacement of the typological species concept, and its tax¬
onomic equivalent, the morphospecies, by the biological species concept.
The taxonomist was no longer satisfied to separate his collections into types
and duplicates. He began to sample the species at many localities, and
tried to assemble large series from every locality. This type of study was
initiated in the second half of the nineteenth century almost simultaneously
by ornithologists, entomologists, and malacologists.
Although the study of populations reached its dominant position in
systematics only within recent generations, its roots go back to the pre-
Darwinian period. A short history is found in Mayr, 1963, chaps. 11 and
12.
Populations are variable, and consequently the description, measure¬
ment, and evaluation of variation has become one of the principal preoccu¬
pations of the taxonomist who studies taxa in the lower categories. A typolo-
gist needed only one or two “typical” specimens of a species; when he
had more, he disposed of them as “duplicates.” The modern taxonomist
attempts to collect large series at many localities throughout the range
of a species. Subsequently he evaluates this material with the methods
of population analysis and statistics (see Chap. 8c).
The work of population taxonomy not only led to a simplification
of taxonomy through the introduction of polytypic species, but it also led
to a new approach in the study of evolution. Systematics has made many
important conceptual contributions, and one of its greatest achievements
is to have assisted in the introduction of the population concept into biology
(1.3.2).

3.5 POPULATION STRUCTURE

The study of the population structure of species shows that the conven¬
tional division of species into subspecies is a very inadequate and sometimes
even misleading representation of the actual phenomena. A species does
not consist of a number of “little species” called subspecies. Rather, a
species consists of innumerable local populations or demes which stand
in a certain relationship to each other. When species are studied strictly
from the viewpoint of their population structure, it is found that they
can best be described in terms of three major population phenomena (Mayr,
1963, chap. 13) :
3.5.1 The Population Continuum. A large part of the range of
many species, particularly the central part, is occupied by a series of essen¬
tially contiguous populations. Even when there are minor breaks in distribu-
 The Polytypic Species Population Systeniatics and Infraspecific Categories 49

tion, owing to the unsuitability of the habitat, such breaks are bridged
by steady dispersal, resulting in copious gene exchange among populations.
Variation in such a population continuum is essentially clinal. Terminal
populations at the opposite ends of a continuum may be very different
phenotypically and may deserve subspecific recognition (3.3.5).
3.5.2 The Geographical Isolate. This term designates all geographi¬
cally isolated populations, or groups of populations, which have only limited
or no gene exchange with other populations of the species. Any insular
population is normally such an isolate, and isolates are therefore particularly
common near the periphery of the species range. Isolates are frequently
of sufficient difference to merit subspecies rank. The biological importance
of the geographical isolate is that every isolate, regardless of its taxonomic
rank, is an incipient species; it is an important unit of evolution.
3.5.3 The Zone of Secondary Intergradation. Whenever a geographi¬
cal isolate reestablishes contact with the main body of the species, the
two will interbreed, if the isolate has not yet acquired an effective set
of isolating mechanisms. Depending on the degree of genetic and phenotypic
difference achieved by the previously isolated populations, a more or less
well-defined hybrid belt or zone of secondary intergradation will develop.
Fusion lines between ex-isolates can be found in many species.

Fig. 3-5. Pattern of geographic variation in the bumble bee Bombus agrorum. There is little geo¬
graphic variation in the continuous range of the nominate subspecies agrorum, while each periph¬
erally isolated population (nos. 1-24) is distinct and mostly recognized as a separate subspecies
(after Reinig).
50 Principles of Zoological Classification

Fig. 3-6. Peripheral isolates at the ends of various lines of expansion in the polytypic bird species
Dicrurus hottentottus. The figures indicate the ranges of the nine forms, the tails of which are
shown in the insert. The tails of 4 and 6 are typical for most populations of the species; the tails of
the peripheral forms 1-3, 5, and 7-9 are aberrant and specialized in various directions (from Mayr and
Vaurie, 1948).

There are only a few groups of animals in which the population

structure of species is sufficiently well known to permit such a detailed
analysis. This has been done by Keast (1961) for all the species in a
number of families of Australian birds. The study of the population struc¬
ture of species does not replace classical taxonomy but is a superimposed
refinement of the classical methods. It is possible only in groups where
taxonomic analysis and population sampling has reached a degree of ma¬
turity that permits such detailed analysis.

3.6 THE NEW SYSTEMATICS

The approach of the population taxonomist differs rather drastically

from the simple pigeonholing of classical Linnaean taxonomy. To emphasize
the difference, Huxley (1940) introduced the term new systematics for
the newer approach, but its roots actually go back to the first half of
the nineteenth century, some traces of the new systematics being found
in the writings of taxonomists as far back as 125 years ago. Every generation
 The Polytypic Species Population Systematics and Infraspecific Categories 51

l'ig. 3-7. Secondary hybrid belts among tree runners (Neositta) of Australia. The arrows indi¬
cate expansion from post-Pleistocene aridity refuges. Wherever former isolates have met, they
have formed hybrid belts (indicated by hatching). R, subspecies with red wing bar; W, subspecies
with white wing bar (from Mayr, 1963).

has its own new systematics, and what we consider as new systematics
in the year 1968 may indeed be very old systematics 50 years hence. Some
writers have placed the new systematics in opposition to alpha taxonomy.
This is a misleading antithesis. Mayr (1964c) wrote:

What then is the new systematics? Perhaps it is best described as a viewpoint,

an attitude, a general philosophy. It started primarily as a rebellion against
the nominalistic-typological and thoroughly non-biological approach of cer¬
tain, alas all too many, taxonomists of the preceding period.
The worker in the new systematics considers himself a biologist rather
than a filing clerk. This has a number of well-defined consequences in
his attitude toward his material and toward various techniques:
1. He is conscious at all times that he is classifying organisms, not the
remains of organisms or merely names.
2. As a consequence, he places considerable emphasis on so-called biological
characteristics, that is on non-morphological information derived from
behavior, physiology, biochemistry, ecology, and so forth.
3. He appreciates that all organisms occur in nature as members of popula¬
tions and that specimens cannot be understood and properly classified
unless they are treated as samples of natural populations.
4. As a consequence, he attempts to collect statistically adequate samples,
in the case of variable species often amounting to hundreds and thousands
52 Principles of Zoological Classification

of specimens, in order to be able to undertake a study of individual

and geographic variation with the help of the best biometric and statistical
tools.
If we would attempt to describe the current model of the new systematics,
we would see at once that every single item is merely the continuation
of a trend which in most cases had started more than one hundred years
ago. Let me mention some of these points:
1. The utilization of an ever-increasing number of kinds of characters and
a continued depreciation of key characters and of single character classi¬
fications in contrast to the typological approach.
2. A ready acceptance of new tools and techniques, such as
(a) the visual analysis (by sonagrams, etc.) of sounds in insects (cicadas,
orthopterans), frogs, and birds;
(b) the analysis of courtship displays and other behavior;
(c) the utilization of biochemical characters, particularly those yielded
by various methods of protein analysis; and
(d) the utilization of computers, to reduce the danger of subjectivity in
character evaluation.
3. A further clarification of concepts, for instance,
(a) a clearer separation of taxon from category;
(b) the recognition of the subspecies as a category, not an evolutionary
unit; and
(c) a clearer understanding of the causes for similarities and differences
between taxa.
It is evident that the new systematics is neither a special technique nor
a special method, but rather, as I have said, a viewpoint or attitude which
can be applied at every taxonomic level.

For a somewhat different statement on the characteristic trends in

modern taxonomy see Simpson (1961, pp. 63-66).

3.7 THE SUPERSPECIES

Allopatric populations are often so distinct from each other that there
is little doubt about their having reached species level. Rensch (1929)
proposed for such groups of allopatric species the German term Artenkreis.
Since the literal translation “circle of species” was frequently misunderstood,
Mayr (1931) introduced the term superspecies as a convenient international
equivalent.

A superspecies is a monophyletic group of closely related and largely

or entirely allopatric species.

When the ranges of the component species are plotted on a map,

the superspecies usually presents the picture of a polytypic species. Yet,
 The Polytypic Species Population Systematics and Infraspecific Categories 53

Fig. 3-8. A superspecies of paradise magpies (Astrapia) in the mountains of New Guinea. Some
hybridization has been recorded in the zone of contact between mayeri and stephaniae (from
Mayr, 1963).

there are three kinds of evidence to indicate that the component species
have attained reproductive isolation. Either these species, although com¬
pletely isolated from each other, are morphologically as different as are
sympatric species in the respective genus, or they are in some areas in
geographical contact (parapatry) without interbreeding, or there is actually
a slight distributional overlap. Superspecies are not given nomenclatural
recognition, but are listed as such in monographs and catalogs by an appro¬
priate use of headings or symbols. They are important chiefly in zoogeo-
graphical and speciation studies. See Mayr (1963, pp. 499-501) for exam¬
ples of superspecies and further discussion (also Cain, 1955, and Amadon,
1967).
The component species of a superspecies were originally designated
semispecies. However, various authors have suggested a broadening of the
term semispecies to include not only members of superspecies but all border¬
line cases in speciation (Mayr, 1963, p. 501).
Chapter yt Theories of Biological
TT Classification
and Their History

4.1 INTRODUCTION

If the taxonomist were satisfied merely with the describing and naming
of species, he would be left with total chaos, considering the several million
species of animals in existence. To convert this chaos into order is the
task of classification. The ability to classify components of the environment
did not originate with man. Indeed, classifying of objects of the environment
occurs well below the human level, although not codified in language.
Animals prove by their reactions that they classify objects of their environ¬
ment as “food” or “non-food,” as “competitors” or “potential mates,” as
“enemies” or “prey.” Man has classified things by his use of generic or
collective terms ever since he began to communicate by speech. Classification
by man of animals encountered in his environment is as old as man himself,
even though early classifications may have been broad and vague as indi¬
cated by terms like animals, bugs, and worms. A good classification makes
an enormous amount of diverse information readily and conveniently avail¬
able to biologists and, indeed, to anyone dealing with organisms. It gives
meaning by association. It should therefore be the objective of the tax¬
onomist to produce a system which has high predictive value and will
allow maximum information retrieval.
The meaning and the principles of biological classification, as well
54
 Theories of Biological Classification and Their History 55

as all the associated difficulties, have been excellently discussed by Simpson

(1961) and by numerous other authors—for instance, Michener (1957),
Beckner (1959), Cain and Harrison (1960), Cain (1962), Gisin (1964),
and Mayr (1965a, 1965c). The present chapter is devoted to a discussion
of the history and theory of classification. The application of this theory,
the procedure of constructing a classification, will be dealt with in Chap. 10.
To define classification without being circular and without including
terms that bias the definition in favor of a particular philosophy of classi¬
fication is difficult. Most zoologists would perhaps be willing to accept
a tentative definition, such as:

Zoological classification is the ordering of animals into groups on the

basis of their similarity and relationship (in the broadest, not necessarily
biological, meaning of the latter word). The two terms, similarity and
relationship, used in this definition, are the reason for controversies that
have raged for hundreds of years.

4.2 HISTORY OF THEORIES OF CLASSIFICATION

The history of taxonomy goes back to the beginnings of mankind.

Since that time we can distinguish a number of periods or levels of increas¬
ing knowledge and understanding. Simpson (1961) in his second chapter
gives a valuable survey of the history of taxonomy and the development
of modern theory. His treatment goes much deeper into the philosophical
aspects of the subject than the present short one, and it should be read
in conjunction. Simpson also cites bibliographic references to other papers
on the history of taxonomy, such as his own earlier one (Simpson, 1959)
and, for example, Cain (1958fif) and Hopwood (in Sprague, 1950).
We shall distinguish six historical periods, a division which is ad¬
mittedly somewhat arbitrary. They are not sharply separated from each
other, and sometimes several trends are concurrent during a single period.
Progress in the classification of various animal groups (and in the study
of animals from different regions) has been uneven. Activity in some groups
of arthropods and other invertebrates still concentrates on identification
and description, while the methods of population systematics have been
prominent in some better-known groups, for instance birds, mammals, and
butterflies.
An authoritative history of taxonomy has not yet been written; the
treatments in the standard histories of zoology and botany are superficial
and purely descriptive. Indeed, it remained impossible to attempt a pene¬
trating history as long as the roots of taxonomic method and principle
in essentialism (4.3.2), in Aristotelian logic, and in nominalism (4.3.3)
56 Principles of Zoological Classification

were not understood.1 The following short treatment attempts to incorporate

some of the recent findings.
4.2.1 First Period: The Study of Local Faunas. Natives of even
the most primitive tribes are often excellent naturalists, and attach specific
names to the more prominent plants and animals of their home country.
Seashore tribes usually have names for all species of shoreline fishes and
for all edible or poisonous inshore invertebrates. Schemes to classify the
distinguished entities are usually rudimentary (Conklin, 1962), but binomi¬
nal nomenclatures are found in native tribes of Asia and America—they
are not the sole invention of Linnaeus.
Several early Greek scholars, notably Hippocrates (460-377 b.c.),

enumerated kinds of animals, but there is no indication of a useful classi¬

fication in the remaining fragments of their work. There is no doubt that
Aristotle (384—322 b.c.) was the father of biological classification. He lived
for some years on the island of Lesbos, where he seems to have devoted
himself almost entirely to the study of zoology, in particular to the study
of marine organisms. He not only studied comparative morphology but
also paid much attention to embryology, habits, and ecology. Emphasizing
that all attributes must be taken into consideration, he said, “Animals may
be characterized according to their way of living, their actions, their habits,
and their bodily parts.” He referred to such major groups of animals as
birds, fishes, whales, and insects, distinguishing among the last both man-
dibulate and haustellate types and winged and wingless conditions; he
used certain terms for lesser groups, such as Coleoptera and Diptera, which
persist today. He established numerous collective categories, or genera, using
as differentiating characters blooded or bloodless, two-footed or four-footed,
hairy or feathered, with or without an outer shell, and so forth. All this
was a tremendous advance over anything that had previously existed, and
it completely dominated animal classification for the next 2,000 years. Nev¬
ertheless, Aristotle did not supply an orderly, fully consistent classification
of animals.
Aristotle, however, was not only the father of biological taxonomy,
he was also the proponent of a well-rounded philosophy (including meta¬
physics) and the founder of logic, and his impact on taxonomy by means
of these was far greater than through the actual classification he proposed.
Among other things it was he who chiefly suggested to naturalists and
philosophers the idea of arranging all animals in a single graded scala
naturae according to their degree of “perfection” (Lovejoy, 1936, pp.
58-59). This led to the endeavor to classify animals into those that are
“lower” and those that are “higher,” an endeavor translated after 1859
into evolutionary terms.
1 Consult the glossary and index for a definition of terms left unexplained in this historical survey.
 Theories of Biological Classification and Their History 57

Likewise, it was through Aristotle and his followers (including Lin¬

naeus) that “typological” or essentialist thinking became entrenched in
taxonomy (4.3.2). Eventually typological thinking was replaced by popula¬
tion thinking (1.3.2 and 3.4). Typology should not be confused with the
type method of modern nomenclature (13.48).
The energetic revival of natural history after the Dark Ages, together
with the vigorous worldwide explorations and discoveries since the fifteenth
century, led to a precipitous increase in the number of known kinds of
animals and plants. Yet the great encyclopedias of Gesner (1551-1558)
and Aldrovandi (1599) were either alphabetical or followed Aristotle’s
rudimentary classification. Of all the pre-Linnaean authors the one who
arrived at the most natural higher classification was John Ray (1627-1705).
On the whole, throughout this period the botanists were far ahead of the
zoologists and were the first to look for new methods and new principles.
These endeavors reached their climax in the next period.
4.2.2 Second Period: Linnaeus and His Contemporaries. The great
Swedish naturalist Linnaeus (1707-1778) exerted such an important influ¬
ence on the entire subsequent development that he has been called the
father of taxonomy. The binominal method of nomenclature (13.11, 13.12)

CAROLI LINN/EI
Equitis De Stella I'olari,
Archiatri Reg ii, Mm. A Botan. Profess. Upsal.
Acad. Upsal. Holme:;>■ Pftkofol. Berol. Imper.
Lond. Mosspei.. Tolos. Florent. Soc.

SYSTEMA
NATUR7E Per.

REGNA TRIA NATURAL,

Secundum

CLASSES, ORDINES,
G ENERA, SPECIES,
Cum
CHAR ACTE RI HUS , DIFFER ENTIIS ,
S7 NONFMIS , LOCIS.

Tom us I.
Editio Decjma, Reformata.

Cum Pritilegit S:* R:<t M:tit Svta*.

HOLMIJE,
Impensis Direct. LAURENTII SALVII,
J7 ft-

Fig. 4-1. Carolus Linnaeus (1707-1778) and title page of the foundation work in zoological nomen¬
clature.
58 Principles of Zoological Classification

was for the first time consistently applied by him to animals in the tenth
edition of his Systema naturae (1758). Linnaeus himself considered this
a rather inconsequential achievement, and was far prouder of his consistent
application of the Aristotelian system of logic to classification. The world
as we see it was to him the work of the Lord, and classification was the
presentation of the plan of creation. The words “nature” and “natural,”
as in natural system, had a highly specific technical meaning to the Aris¬
totelians (Cain, 1958). They referred to the operational manifestations
of the “essence” of a being (in terms of the Thomistic philosophy) (4.3.2).
Great innovator that Linnaeus was in his binominal method and in his
extraordinarily useful catalogs of names and diagnoses, in his basic philoso¬
phy he was looking backward to the scholastic philosophy of the Middle
Ages and to Aristotle. Yet the conveniences of a rigid hierarchy of categories
and an unambiguous binominal nomenclature proved themselves so great
that succeeding generations had little trouble getting rid of the Linnaean
philosophy and yet retaining the best of his method. In more recent years
even the value of binominalism has been questioned (13.13, 13.14).
Most early classifications were quite frankly identification schemes,
based on single characters. They were usually presented as simple dichoto¬
mous keys, using single key characters. These often related to broad or
special adaptations, as when Pliny classified animals into those of land,
water, and air. Water birds with webbed feet were classed together, and
so were all wading birds with long legs. Superimposed on this pragmatic
approach were various taxonomic theories (4.3.2) culminating in the Sys¬
tema naturae of Linnaeus. The more perceptive naturalists soon realized
that these identification schemes often resulted in very heterogeneous group¬
ings, that is, in “artificial classifications.” The botanist Hieronymus Bock
was the first (in 1546) to endeavor explicitly to place together those plants
which resembled one another in external characters, or, as he put it, those
which Nature seemed to have joined together by similarity of form.
Even these guiding principles led to improvement only slowly. Early
classifications were a curious mixture of “natural” and artificial groups.
Plants, for instance, might be divided into mosses, ferns, grasses, herbs,
shrubs, and trees. The former three correspond to natural taxa, the latter
three do not. A division of animals into Vermes, Insecta, and Vertebrata
was equally mixed because Vermes is an artificial conglomerate of worm¬
shaped animals. Continuing improvement resulted from a study of an in¬
creasing number of characteristics which led to the delimitation of groups
that had many characters in common, not merely a key character. Linnaeus
himself adopted this empirical approach for the groups he was most familiar
with, for instance the insects, producing a classification that is still largely
acceptable. His classifications of other groups, like birds, amphibians, and
lower invertebrates, were very poor.
 Theories of Biological Classification and Their History 59

The Aristotelian philosophy of Linnaeus was heavily attacked by con¬

temporary naturalists with empirical or nominalistic tendencies, particularly
by M. Adanson.
4.2.3 Third Period: The Empirical Approach. The hundred years
between the tenth edition of the Systerna naturae and the publication of
Darwin’s Origin of Species was a period of subtle but steady transition.
Deductive, a priori principles were increasingly rejected, and taxonomists
more and more delimited taxa empirically, on the basis of the totality
of characters, not just a few “essential” ones. The term “natural system”
lost the Linnaean meaning of a system based on the inherent “natures”
of organisms (as defined in scholastic philosophy) ; “natural” came to sig¬
nify unbiased by a priori considerations, and based on a consideration of
the totality of characteristics. Eventually what the Aristotelians had consid¬
ered as “natural” came to be considered arbitrary and artificial.
Lamarck (1744-1829), who lived during this period, had no visible
influence on these developments except for some purely practical contribu¬
tions he made to the classification of the invertebrates. Cuvier (1769-1832)
was far more influential (Coleman, 1964). His taxonomic theory was an
extraordinary mixture of antiquated concepts (such as his a priori weighting
of characters on the basis of their physiological importance) and some
very sound practical taxonomy. His insistence on the total independence
of four major types (embranchements) of animals—vertebrates, mollusks,
arthropods, and radiates—was the death knell of the scala naturae which
had still dominated Lamarck’s classification. Now that the invertebrates,
the “bloodless” animals, were broken up, it became easy to continue the
dismemberment of unnatural groupings, and this was indeed a major pre¬
occupation of zoologists during the first half of the nineteenth century.
A steady and enormous increase in the number of known animals
characterized this period. Voyages all over the globe acquainted zoologists
with the animals of Africa, Australia, and the Americas. The local natural¬
ists were being replaced by specialists who studied birds, reptiles, mollusks,
or insects, or indeed only one particular group of insects, be it butterflies,
beetles, or ants.
The systematic work of this period confirmed the conviction of the
early naturalists that the endless variety of organic life is organized into
natural groups. There are bluebirds (Sialia), which are one of the subdivi¬
sions of the thrushes (Turdidae), which together with many other similar
subdivisions form the songbirds (Oscines), one of the subdivisions of the
birds, belonging to the vertebrates, and so on. The empirical taxonomists
of this period did a magnificent job in developing a “natural system,”
in the new meaning of this term. However, disappointed by the philosophi¬
cal speculations of the Aristotelians, of the nominalists, and of Lamarck,
and also kept far too busy by the avalanche of new species descending
60 Principles of Zoological Classification

upon the museums, they did not try to give meaning to the orderliness
of nature discovered by them.
The best empiricists were not satisfied merely to search for “characters
in common,” they also established all the major methods and principles
of classification that are still recognized as leading most efficiently to the
establishment of sound taxa. In particular, they developed principles of
a posteriori weighting of characters, for instance through the study of their
correlation with other characters (10.4). Even though Ray and others before
him had stressed the fact that some characters correlate better with natural
groups than others, it is only in this post-Linnaean period, and in part
as a conscious rebellion against Aristotelian apriorism, that the system of
empirical a posteriori weighting developed. The empiricists also undertook
the evaluation of gaps between taxa, and they supported hierarchical ar¬
rangements of categories (Chap. 5) on the basis of degree of similarity.
4.2.4 Fourth Period: Darwin and Phylogeny. Prior to 1859 the tax¬
onomist had to choose between two alternatives to explain the naturalness
of the system. He could side with the nominalists and claim that natural

ON

THE ORIGIN OF SPECIES

BY MEANS OF NATURAL SELECTION,

OR THE

PRESERVATION OF FAVOURED RACES IN THE STRUGGLE

FOR LIFE.

By CHARLES DARWIN, M.A.,

FELLOW OF TUB ROYAL, GEOLOGICAL, I.IXNAIAN, ETC., SOCIETIES;
AUTHOR OF ‘ JOURNAL OF RESEARCHES DURING II. M. S. DEAGLE’S VOYAGE
ROUND THE WORLD.'

LONDON:

JOHN MURRAY, ALBEMARLE STREET.

1859.

Fig. 4-2. Charles Robert Darwin (1809-1882) and title page of the foundation work in evolutionary
biology.
 Theories of Biological Classification and Their History 61

groups do not exist and that taxa are merely the arbitrary products of
the ordering human mind. This conclusion was so clearly contradicted by
the empirically found naturalness of most taxa that it had hardly any
adherents by 1859. The alternative was to believe that the order of nature
was due to the plan of the Creator, and that each taxon consisted of
variants of an underlying type, all of them containing, however, the essence
of this type. It is Darwin’s everlasting merit to have proposed a third
alternative in the Origin of Species.
When Charles Darwin (1809—1882) in 1831 joined the Beagle as
a naturalist, he still accepted the creationist dogma. During this voyage,
however, he encountered so many phenomena of distribution, variation,
structure, and adaptation that were quite improbable on a creationist inter¬
pretation that he adopted the evolutionary interpretation. All at once the
enigma of the natural system was solved. “Natural” groups exist because
the members of such a group had descended from a common ancestor.
Fortunately, accepting evolution did not necessitate any change in the tax¬
onomic technique. No longer did the taxonomist have to “make” taxa,
evolution had done this for him. All he needed to do was to discover
these groups.
It is not surprising in the least that the adoption of the evolutionary
theory had virtually no impact on the established classifications. All it did
was to give intellectual justification to what had already been standard
practice among the best empirical taxonomists. What the evolutionary the¬
ory supplied was the explanation for the fact that variation in nature is
not continuous but consists, in Darwin’s words, of “groups within groups.”
But Darwin did more than provide the theoretical basis for a natural
system. He also gave some clear practical rules on how to avoid the circular
reasoning of Linnaeus, Cuvier, and other predecessors. It is here that his
many years of concentrated work on the classification of barnacles repaid
him abundantly. Far more clearly than most of his successors, Darwin
realized that two processes occur during phylogeny—branching and subse¬
quent divergence. Accordingly Darwin stressed (1859, p. 420) that the
separation of taxa must be based on branching (“propinquity of descent”),
but that in the ranking of these taxa into various categories due considera¬
tion must be given “to the different degrees of modification which they
have undergone.”
Darwin made yet one other fundamental contribution to taxonomic
theory. He, like the empiricists, rejected both the a priori weighting of
taxonomic characters, as practiced by Linnaeus and Cuvier, as well as
the disapproval of all weighting; he proposed instead (1859, pp. 425-426)
a number of empirical rules on how to discover taxonomically useful charac¬
ters, that is, how to undertake a posteriori weighting. They include the
constant presence of the character in related forms, “especially those having
62 Principles of Zoological Classification

very different habits of life,” and particularly a constant association of

several characters (“we know that such correlated or aggregated characters
have a special value in classification”). Simpson (1959, 1961) and Cain
(1959<z) have given us lucid interpretations of Darwin’s contribution to
taxonomic theory.
The natural system was studied in the generations following Darwin
primarily as important evidence in favor of the evolutionary theory. It
is only rather recently that the theory of biological classification has been
studied as a branch of methodology and philosophy of science as such.
Only then was it recognized that the a priori principles that are useful
in the classification of inanimate objects are largely inapplicable, if not
decidedly misleading, when applied to organisms with an evolved informa¬
tion content (4.3.6).
The empirical taxonomists were greatly encouraged to learn that Dar¬
win and his evolution theory gave meaning to their classifying activities
and continued with increased vigor. One aspect in particular now came
to the fore, namely, the search for missing links between seemingly uncon¬
nected taxa and the reconstruction of “primitive ancestors.” Ernst Haeckel’s
(1834-1919) phylogenetic trees and speculations greatly stimulated this
type of activity. The search for facts to substantiate these phylogenetic
trees and to improve their design dominated biology during the second
half of the nineteenth century and led to a boom in the fields of comparative
systematics, comparative morphology, and comparative embryology. Even
though the results were far less permanent than was then believed, this
concern with phylogeny resulted in much sound research and in an interest
in aberrant groups of organisms that might have otherwise been totally
ignored. More importantly, these descriptive studies laid the foundation
for the functional and experimental branches of biology whose flowering
started in the 1870s and 1880s.
The period of the discovery of major new types of animals was essen¬
tially over well before the end of the nineteenth century. By then the
need to prove the fact of evolution had ceased to exist. Taxonomy no
longer was an exciting bandwagon, and taxonomists were forced to concen¬
trate on the necessary though tedious job of describing, diagnosing, and
classifying the seemingly endless number of species. A minority of the species
describers were dilettantes who brought discredit to the field by the creation
of numerous synonyms and by an excessive splitting of families and genera.
Others concentrated on the unearthing of long-forgotten synonyms, thus
arousing the ire of general biologists, who complained quite rightly that
this defeated the basic objective of nomenclature as an information retrieval
system. There is little question that taxonomy fell into some disrepute during
the latter part of the nineteenth and the early twentieth century. The
fact that the taxonomists sided against the immensely popular and powerful
 Taf. I

I , FpIiI : p in n q < /!)Stdtnmc)

D, Feld: pxya (.’) Stamme)
HI, Feld: p s t (j (1 Stamm) II ;t d i x
.deltca may l'iefa, Salic dt r roiiiitiunis
unioersalat Ccncalogic tlar .
On>;inismmuii[

l ig. 4-3. The phylogeny of living beings as conceived by Haeckel (1866) and expressed in a formal
treelike diagram.

63
64 Principles of Zoological Classification

early Mendelians with their antiselectionist and saltationist interpretation

of evolution aggravated the situation. However, a turn for the better began
to develop in the 1920s.
4.2.5 Fifth Period: Population Systematics. For the sake of conven¬
ience, taxonomists had continued to treat species, in a typological manner,
as invariant units until long after the invalidity of the typological dogma
had become apparent. Yet, whenever population samples from different
portions of the geographic range of a species were compared, smaller or
greater differences were found. This resulted eventually in the replacement
of the typologically defined species by the polytypic species composed of
different populations in the dimensions of space and time. The study and
comparison of intraspecific populations became the objective of population
systematics (3.4). The history of this development, beginning in the first
half of the nineteenth century and reaching its climax in the 1930s and
1940s, has been described in detail by Mayr (1942, 1963). Replacement
of typological thinking by population thinking has had important conse¬
quences in many areas of taxonomy. Considering taxa as populations or
aggregates of populations greatly facilitated the study of variation and the
definition of lower taxa and categories. Labeled by J. S. Huxley (1940)
as the “new systematics it led to a reevaluation of the species concept
and to a more biological approach in taxonomy. The population systematist
understands that all organisms occur in nature as members of populations
and that specimens cannot be understood and properly classified unless
they are treated as samples of natural populations.
The same period saw two additional aspects come to the fore. One
is what might be called the biological approach to taxonomy. The taxono¬
mist moved more and more from the museum into the field, he increasingly
supplemented the morphological characters with characteristics of the living
animal, such as behavior, voice, ecological requirements, physiology, and
biochemistry. Taxonomy truly became biological taxonomy. The other de¬
velopment was the introduction of the experiment into taxonomy. Although
this has been far more characteristic for botany than for zoology, neverthe¬
less the experimental analysis of isolating mechanisms—particularly in verte¬
brates, in Drosophila, and in protozoans—and the application of other
experimental methods have been very helpful. At no time, however, did
a separate field, experimental taxonomy, develop, because there was little
resistance against the use of experimental methods.
Population systematics is not an alternative to classical taxonomy,
but only an extension of it. In those groups in which the inventory-taking
of species is still in full progress, and where too few local populations
have been sampled, one cannot easily apply the methods of population
systematics. Being centered on the population level, the new systematics
naturally had little impact on the theory of classification at the level of
the higher taxa. The population thinking of the new systematics was one
 Theories of Biological Classification and Their History 65

of the major sources of the new science of population genetics, which in

turn influenced the further development of population systematics. Together
they greatly helped to clarify our thinking about evolution on the species
level and were instrumental in producing the great synthesis in evolutionary
biology.
4.2.6 Sixth Period: Current Trends. The current period is character¬
ized by three developments. One is renewed examination of the whole
theory of taxonomy, as indicated by the publications of Hennig (1950,
1966), Remane (1952), Gregg (1954), Bloch (1956), Cain (1958ff), Beck-
ner (1959), Simpson (1961), Gunther (1962), and Mayr (19656). Articles
published in Systematic Zoology and in the Publications (nos. 1-6) of the
Systematics Association are indicative of the same interest. The use of elec¬
tronic computers and associated endeavors to revive a nominalistic approach
to taxonomy (Sokal and Sneath, 1963) form the second significant develop¬
ment of this period. The third is the vigorous introduction of biochemical
techniques and, more importantly, a growing realization among molecular
biologists of the importance of understanding the phylogeny of organisms
as a basis for the understanding of the evolution of macromolecules. Com¬
parative ethology likewise has stimulated a deeper interest in taxonomy.

4.3 THEORIES OF CLASSIFICATION

The time since about 1930 has been a period of unprecedented activity
in systematics. After an early and almost total preoccupation with popula¬
tion systematics a concern began to develop over the meaning of higher
taxa and the hierarchy of categories, and indeed over the theory of classifica¬
tion itself. Previously, when asked to explain their theory of classification,
taxonomists had been singularly inarticulate. They would have said that
they wanted to establish “natural groups” or that they wanted to group
together species agreeing in their “essential properties” or showing “natural
affinity.” Such vague and seemingly noncommittal phrases are actually the
expression of succinct philosophies or taxonomic theories. The publications
listed in 4.2.6, among others, have helped to disentangle the manifold,
often strangely interwoven lines of thought. It is not yet possible to write
a balanced history of taxonomic theory, but it seems that from the beginning
there have been only five theories of classification. Either pure or mixed
they seem to supply the theoretical foundation of the work of all practicing
taxonomists:

1. Essentialism (Aristotle to Linnaeus)

2. Nominalism
3. Empiricism
4. Cladism
5. Evolutionary classification
66 Principles of Zoological Classification

The first three are pre-Darwinian, the latter two post-Darwinian.

The emergence of these theories is closely correlated with the general history
of taxonomy (4.2).
Much of the recent literature is devoted to criticism. Curiously, tax¬
onomic theory is being attacked for aspects that are common to all scientific
theory and that are taken for granted when found in other sciences, for
instance in physics. The provisional nature of theories is one of them. The
fact that most findings of science cannot be observed directly but must
be inferred is another. The necessity for unproven models and working
hypotheses is a third. Evolutionary taxonomy is sometimes accused of circu¬
lar reasoning, but without foundation (4.4). Actually, the theoretical foun¬
dations of taxonomic science are far more solid, secure, and elaborate than
realized by most nontaxonomists.
Before these theories can be discussed, the place of identification in
the conceptual framework of taxonomy must be clearly understood.
4.3.1 Identification. Many classifications attempt to be both identi¬
fication schemes and classifications. This, as we now know, creates conflicts.
The procedure of identification is based on deductive reasoning. One starts
out with a given set of taxa (classes) and attempts to fit the investigated
specimen into one of them. If one succeeds, one has identified it. Identifica¬
tion deals with individuals. The procedure of classification is inductive.
Unlike identification, which deals with a few characters (ideally a single
one) that throw a given specimen into one or the other line of the key,
classification deals with and evaluates a multitude of characters, ideally
all of them. Classification deals with populations and aggregates of popula¬
tions. Much of the development of taxonomy has been toward clearer and
clearer separation of these two entirely different operations. He who tries
to include classification and identification in a single operation is bound
to become confused and thwarted in both endeavors. For details on the
procedure of identification see 7.3.
4.3.2 Essentialism (Aristotle’s Natural System). The dominant the¬
ory of classification for many centuries was based on Aristotelian logic.
It was adopted and elaborated by the Thomists and later by Linnaeus;
its major tenets are still defended by some taxonomists, e.g., Thompson
(1952, 1962), Borgmeier (1957), Blackwelder and Boyden (1952).
The philosopher Karl Popper (1950, p. 34) describes this approach
in these words:

I use the name methodological essentialism to characterize the view, held

by Plato and many of his followers, that it is the task of pure knowledge
or “science” to discover and to describe the true nature of things, i.e.
their hidden reality or essence . . . [all essentialists, including Aristotle]
agreed with Plato in determining the task of pure knowledge as the discovery
of the hidden nature or form or essence of things. All these methodological
 Theories of Biological Classification and Their History 67

essentialists also agreed with Plato in holding that these essences may be
discovered and discussed with the help of intellectual intuition; that every
essence has a name proper to it, the name after which the sensible thing
is called; and that it may be described in words. A description of the
essence of a thing they called a “definition.”

This philosophy, when applied to the classification of organic diversity,

attempts to assign the variability of nature to a fixed number of basic
types at various levels. It postulates that all members of a taxon reflect
the same essential nature, or in other words that they conform to the
same type. This is why the essentialist ideology is also referred to as typology.
Variation, consequently, is considered by the typologist as trivial and irrele¬
vant. The constancy of taxa and the sharpness of the gaps separating them
tend to be exaggerated by him. The fatal flaw of essentialism is that there
is no way of determining what the essential properties of an organism
are and why these and no other properties of an organism are essential.
Simpson (1961) and Hull (1965) have adequately refuted the theoretical
arguments of the essentialist school. It also encounters numerous purely
practical difficulties. The first is the polythetic nature of most taxa (4.7).
Virtually every higher taxon contains certain species which, on the basis
of the total evidence, are clearly members of the taxon even though they
lack some of the “essential natures” of that taxon. A second reason is
that it does not distinguish between “characters in common” resulting from
descent and those resulting from convergence. Indeed its entire emphasis
on “characters in common” is misleading (Simpson, 1961). Finally the
essentialist approach is singularly defenseless against conflicting evidence
from different groups of correlated characters (Crowson, 1965). Essentialists
attempt to combine classification and a system of logic in a single scheme.
Everything has to be classifiable into a and non-a, b and non-6. The essential
characters in this theory of classification have two great virtues: they are
exclusive (“winged” versus “nonwinged,” “six legs” versus “four legs”),
and they serve as diagnostic (= key) characters. Indeed many early “classi¬
fications” were published in the form of dichotomous keys, and there are
still taxonomists who find it difficult to distinguish between a classification
and an identification key.
Classification continues to attract the attention of logicians right to
the present—see Gregg (1954), Beckner (1959), Simpson (1961), or Buck
and Hull (1966). Perhaps these attempts have helped to make the language
of the taxonomist more precise, but they do not appear to have added
to the theory of classification.
The ideal of essentialist classification was the discovery (rather than
establishment!) of the Natural System. Their natural system was nothing
more or less than the plan of creation. Among the empiricists and evolution¬
ists the term natural system acquired very different meanings, yet the con-
68 Principles of Zoological Classification

cept is so permeated with essentialist-creationist ideology that its use in¬

variably evokes a misconception among nontaxonomists. Since there is no
such thing as the Natural System, the term is best not used at all.
One curious by-product of the search for order in the natural system
was the attempt to express the order in numbers, attempts that can be
traced all the way back to the Pythagorean school. Linnaeus had a great
fondness for numbers, and for very specific numbers at that (for instance,
six). Vigors, Swainson, Oken, Kaup, and others attempted to devise systems
based on the numbers three, four, or five. All these misguided plans to
discover in nature a numerical blueprint collapsed in 1859 (Stresemann,
1950).
4.3.3 Nominalism. According to this philosophy only individuals ex¬
ist. All groupings, all classes, all universals are artifacts of the human mind.
This is equally true for species and for higher taxa. According to this
philosophy (Gilmour, 1940) :
The process of classification is as follows: The classifier experiences a vast
number of sense data which he clips together into classes . . . thus a class
of blue things may be made for sense data exhibiting a certain range of
color, and so on . . . the important point to emphasize is that the construc¬
tion of these classes is an activity of reason, and hence, provided they
are based on experienced data, such classes can be manipulated at will
to serve the purpose of the classifier. . . . The classification of animals
and plants ... is essentially similar in principle to the classification of
inanimate objects.
Thus there are no such things as birds or snakes, but only names
invented by man and attached to groups of individuals considered by him
to be similar. Bessey’s statement on the species category (2.2.2) describes
the nominalist theory particularly well.
This philosophy ignores the fact that there is indeed a difference
in principle between classifying inanimate objects (including human arti¬
facts) and organisms (Darwin, 1859, p. 411). It ignores the fact that
groups of organisms, related by descent, possess a unity by the shared portion
of their DNA heritage, a causation for shared characteristics for which
there is no equivalent among inanimate objects. Birds are not an arbitrary
aggregate of organisms resulting merely from “an activity of reason,” as
claimed by the nominalists, but they are a natural group because of the
common heritage they share.
The basic fallacy of the nominalists is their misinterpretation of the
causal relation between similarity and relationship. As Simpson (1961)
has emphasized correctly, members of a taxon are similar because they
share in a common heritage; they do not, as the nominalist would have
it, belong to the taxon because they are similar. It is exactly as with identical
twins; two brothers are not identical twins because they are similar, but
 Theories of Biological Classification and Their History 69

they are similar because they are both derived from a single zygote, that
is, because they are identical twins (2.3). The fatal weakness of nominalist
thinking, when applied to the classification of organisms, is the reversal
of an existing causal relation between “similarity and affinity.” *
The numerical pheneticists (Sokal and Sneath, 1963) have, in princi¬
ple, adopted the nominalist philosophy. A purely phenetic approach, an
approach that “makes taxa” on the basis of the degree of observed similarity,
usually leads to a classification similar to one based on the evolutionary
approach. The reason is that, by and large, two organisms will be the
more similar the more closely related they are by descent (10.2). Neverthe¬
less, the phenetic approach is exposed to the risk of reaching unsound
classifications, because in giving equal weight to all characters it does not
allow for mosaic evolution, special adaptation, convergence, parallelism,
developmental and genetic homeostasis, and other evolutionary, genetic,
and developmental phenomena that disturb the expected close correlation
between phenetic similarity and phylogeny. Worst of all, the theoretical
basis of its nominalistic approach is unsound.
For recent critiques of the philosophy of numerical taxonomy see
Mayr (19656), Simpson (1961),Gisin (1964), and Rollins (1965).
A direct consequence of the assumption that natural groupings do
not exist, but that all “species” or “classes” are the product of the human
mind, is the postulate that definitions should be “operational.” This might
be a legitimate request for arbitrary classes of inanimate objects, and it
is therefore not surprising that a physicist (Bridgman) was the original
proponent of the operational approach. It works best for the definition
of units of measurements but breaks down, even in physics, for more com¬
plex concepts. Operational definitions are certainly not only inapplicable
but altogether inappropriate for evolved phenomena. A species or the species
concept is not made nor tested by my operations. The fact that it is possible
to call delimitations of higher taxa “operational” when they are based
on calculated degrees of difference of a set of arbitrarily selected characters
proves the inappropriateness of operationalism when applied to the products
of evolution. Birds, bats, and other higher taxa are not made by the arbitrary
operations of the taxonomist, rather they are the products of evolution.
Operationalism is an altogether invalid approach in most areas of evolution¬
ary biology, when based on the phenetic method.
4.3.4 Empiricism. According to this approach to taxonomy (4.2.3),
there is no need for a theory of classification. Provided enough characters
are intelligently evaluated, a natural system (the meaning of “natural”
being very different from the Aristotelian one) will emerge automatically.
Even though the working taxonomist usually proceeds on the basis of these
empiricist principles, he feels that the resulting classifications would be
biologically meaningless if not supplied with a theoretical foundation. This
70 Principles of Zoological Classification

was provided by Darwin. Two new theories of classification were proposed

after 1858, cladism (4.3.5) and Darwin’s own evolutionary taxonomy
(4.3.6), at present adopted by the majority of animal taxonomists.
4.3.5 Cladism. This term refers to a taxonomic theory' by which
organisms are ranked and classified exclusively according to “recency of
common descent.” Categorical status according to this theory depends on
the position of the branching points on the phylogenetic tree. Hennig (1950,
1966), the most consistent proponent of this thesis, and others have desig¬
nated themselves misleadingly as the phylogenetic school, and this has con¬
fused all arguments since 1950. The heated controversy of that period,
allegedly concerned with the defense or criticism of “phylogenetic classifica¬
tion,” actually deals with the validity of basing classification exclusively
on the position of branching points (Mayr, 19656). Since the splitting
of branches is only one of several phylogenetic processes, it will avoid misun¬
derstandings to refer to this viewpoint as cladism, according to the terminol¬
ogy of Rensch (1947) and Cain and Harrison (1960). Users of the recent
literature are warned to look out for the misleading use of the term phy-
logeny by the cladists. Cladism has also been designated the genealogical
approach (Gisin, 1964).
The basic fallacy of cladism is to overlook the fact that “relationship”
in the evolutionary sense is determined by both processes of phylogeny,
namely, branching and subsequent divergence (Darwin, 1859, p. 420).

The argument of the cladist fails to recognize that the term relationship
has two distinct meanings, genetic relationship and genealogical relationship.
The two happen to coincide for all practical purposes as long as we deal
with close relatives. ... In phylogeny, where thousands and millions of
generations are involved, thousands and millions of occasions for changes
in gene frequencies owing to mutation, recombination, and selection, it
is no longer legitimate to express relationship in terms of genealogy. The
amount of genetic similarity now becomes the dominant consideration for
a biologist ... if one of the lines is exposed to severe selection pressures
and as a result diverges dramatically from its genealogically nearest relatives,
it may become genetically so different that it would be a biological absurdity
to continue calling them near relatives. Even though the crocodilians are
cladistically nearest to the birds (both having descended from the pseu-
dosuchians) [Fig. 4-4], the crocodilians are still closer to many of the
other reptiles, as far as the total gene composition is concerned, than they
are to the birds, which have so drastically altered their genetic composition
[as a result of their adaptation to life in the air]. (Mayr, 19656, p. 79.)

To rank taxa according to branching points is nearly always misleading.

It might necessitate, for instance, the inclusion of the African apes (Pan)
in the family Hominidae and their exclusion from the family Pongidae.
Their exclusive concern with branching causes cladists to ignore and
 Theories of Biological Classification and Their History 71

Fig 4-4. Inferred phylogeny of the reptilian branch of the vertebrates. The branching point between
birds and crocodilians (xx) is much more recent than that between crocodilians and other surviving
classes of reptilians.
A = mammals, B = turtles, C = ichthyosaurs, D = plesiosaurs, E = rhynchocephalians,
F = lizards, G = snakes, H = ornithischian dinosaurs, I = ptcrosaurians, K = crocodilians,
L = birds, M = saurischian dinosaurs; -+- = extinct.

even deny the existence or relevance of different rates of evolutionary

change in different phyletic lines. Indeed, Hennig ignores rate of evolution
so completely that he asserts (1966, p. 235): “Decisive is the fact that
processes of species cleavage are the characteristic feature of evolution;
they are the only positively demonstrable historical processes that take place
in supra-individual organism groups in nature.” This elimination from con¬
sideration of different evolutionary rates would be defensible only if evolu¬
tionary rates were the same in all lines. By equating genealogical distance
with genetic distance, Hennig does indeed make this assumption.
This, in turn, leads him to postulate that one can determine the
categorical rank of a taxon by fixing the branching point (from its sister
taxon) in the geological time scale. The taxa in the hierarchy “are subordi¬
nated to one another according to the temporal distance between their
origins and the present; the sequence of subordination corresponds to the
‘recency of common ancestry’” [of the taxa] (p. 83). . . . “In the phy-
72 Principles of Zoological Classification

logenetic [= cladistic] system the absolute rank order cannot be indepen¬

dent of the age of the group since . . . the subordination of groups is
by definition set by their relative age of origin” (p. 160). Consistently
with these principles Hennig decides that the position of the branching
point on the geological time scale determines categorical rank. Taxa that
originate from a split in the Precambrian are to be ranked as phyla; be¬
tween Cambrian and Devonian as classes; between Mississippian and Per¬
mian as orders; between Triassic and Lower Cretaceous as families; be¬
tween upper Cretaceous and Oligocene as tribes; and in the Miocene as
genera (his fig. 58, p. 186). “Then the mammals would have to be called
an order . . . the Marsupialia and Placentalia would have to be down¬
graded to families, and the ‘orders’ of the Placentalia would be tribes”
(p. 187). The absurdity of the proposal is self-evident.
Throckmorton (1965) shows that the invalidity of the assumptions
of the cladists is singularly well demonstrated by evolution in the family
Drosophilidae. “Most of the diversification in this family has occurred by
divergence from a single lineage that was itself changing slowly in time”
(p. 233). Lines diverging from the same point diverge and diversify to
different degrees. The end points of many side lines are formed by different
genera. “Those that have diverged in their external and traditionally diag¬
nostic features are classified in other genera. Where these same features
have remained unchanged and in spite of other changes, the forms are
classified as Drosophila” (p. 233). “In most instances in Drosophila closely
related species are complex mosaics of the characteristics of their nearest

ADAPTIVE AND STRUC T UR A L-FUNC T ION AL ZONES

Fig. 4-5. Recency of descent versus degree of adaptive divergence. Dendrogram of probable affinities
of recent hominoids in relationship to their radiation into adaptive-structural-functional zones. The two
major occupied adaptive zones are bordered by solid lines. Pongid radiation into sub- and sub-subzones
is schematically suggested by broken and dotted lines (from Simpson, 1963).
 Theories of Biological Classification and Their History 73

tripunctata

Fig. 4-6. Phylogeny of Drosophila and related genera. Many of these genera are specialized end points
of certain species groups of Drosophila (from Throckmorton, 1965).

relatives. They show individually very little that is unique to themselves.

They show instead unique combinations of the characters found among
other close relatives” (p. 227).
As Sharov (1965) points out, cladists make the further assumption
(which in most instances is not true) that a parental taxon expires when
it gives rise, by splitting, to two daughter taxa (= sister groups of Hennig).
In reality, this is what rarely happens in phylogeny. A new group almost
invariably buds off from a parental taxon which continues to exist with’
very little change, sometimes for more than 100 million years (Fig. 4-7).
A new group in these cases is the “sister group” of the parental taxon.
The fallacy of the cladistic approach consists not only in the equation
74 Principles of Zoological Classification

A B D C E

Fig. 4-7. Tetrapod phylogeny. New classes originate by

the branching off of daughter lines while the parental
line continues, essentially unaffected. A = Crossopterygia,
B = Amphibia, C = Reptilia, D = Mammalia, E =
Aves. B is a daughter line of A, C of B, and D and E of C.

of genealogical and genetic distance but also in the ignoring of mosaic

evolution (Chap. 10) and in the fact that a classification is an information
retrieval system and not just an intellectual exercise. It places highly dis¬
similar organisms (like crocodilians and birds) in a single taxon while separat¬
ing extremely similar ones of slowly evolving lines simply because the branch¬
ing points which separate them occurred early on the geological time scale.
The ranking of taxa on the basis of the cladistic method is quite
meaningless. The method is difficult to apply in all groups with a poor
fossil record. In such groups the branching points are reconstructed on
the basis of evaluated similarity, and thus the method itself is negated.
Cladists tend to forget that the ordering of the diversity of nature
is the foremost task of classification. To use only one of the two processes
leading to the diversification of groups—namely, branching—as evidence
in the ordering procedure is self-defeating. The evolutionary taxonomist
agrees with the cladist in assuming that, usually, the more recently their
phyletic lines separated, the more similar two taxa are. However, the evolu¬
tionary taxonomist also gives due weight in his classifications to any unequal
divergence of the descendant lines. By deliberately ignoring these differences,
the cladist is often forced to recognize taxa of very unequal value.
Cladists sometimes claim that their theory of classification is the only
 Theories of Biological Classification and Their History 75

one that can lead to the establishment of monophyletic taxa. This claim
is based on a misinterpretation of the concept monophyly.
Monophyly. Once we accept the basic principle of biological classifica¬
tion, that organisms are to be classified according to the information content
of their genetic program, it is evident that monophyly must be required
of all taxa. This is therefore one of the postulates of evolutionary taxonomy.
Artificial taxa, containing the descendants of different ancestors, would
be unable to fulfill the demands one places on a scientific theory (see
4.5), owing to the heterogeneity of the included genetic programs.
The issue of monophyly has been clouded by various confusions. Some
authors have referred to a “polyphyly” of a taxon when only a polyphyly
of the diagnostic character of the taxon was involved, the taxon itself being
monophyletic. It sometimes happens that a certain grade of morphological
change is reached independently in several lines derived from a single ances¬
tral group. The group showing this grade of development is of course
monophyletic. As always in evolution, one must distinguish between what
happens to the phenotype and what happens to the genotype. The diagnostic
mammalian structure of the jaw-ear region, for instance, is believed to
have evolved several times from ancestral therapsid reptiles, which had
the needed genetic program to predispose them toward evolving the mam¬
malian grade when exposed to the same selection pressures. This is not
polyphyly, because the genotype permitting these parallel evolutionary
changes goes back to the same ancestral program. We classify taxa (= geno¬
types) and not characters ( = phenotypes). The usual phrasing of the prin¬
ciple of monophyly (“a taxon is monophyletic if its members are descendants
of a common ancestor”) is too vague to be helpful in more complicated
cases, such as that of the mammals. Simpson (1961) has therefore given
a more concrete definition: “Monophyly is the derivation of a taxon through
one or more lineages, from one immediately ancestral taxon of the same
or lower rank.” The class Mammalia is monophyletic because all mam¬
malian lines were derived from the immediately ancestral taxon of therapsid
reptiles. Most cases of alleged polyphyly reported in the literature do not
stand up under critical analysis.
The other misconception about monophyly is to consider it not only
a retrospective but also a prospective principle. “The species included in
each higher taxon must be derivable from a common stem species [= retro¬
spective postulate], and no species having arisen from this stem species
can be placed outside this taxon [= prospective postulate]” (Hennig, 1966,
p. 71, also pp. 72-73). The latter postulate, of course, is completely contra¬
dicted by common sense and is in opposition to the phenomena of evolution¬
ary divergence. If a descendant group, such as the birds among the archo-
saurian reptiles, evolves more rapidly than the other collateral lines, it
not only can but it must be ranked in a higher category than its sister
76 Principles of Zoological Classification

groups. This does not violate the principle of monophyly, retrospectively

defined. The taxon Aves is monophyletic, and so are the taxa Crocodilia
and Reptilia.
The concept of monophyly is important only at the level of the higher
categories. Taxa that are still crossable, like subspecies, may produce hybrids
which by definition would not be monophyletic. Yet the zoologist is not
concerned with monophyly at the species level.
4.3.6 Evolutionary Classification. Like the empirical schemes of clas¬
sification, evolutionary classification is based on the simple fact that readily
delimitable groups of species, such as birds, penguins, bats, beetles, and
the like, occur in nature. Evolutionary taxonomy differs from empiricism
by demanding an explanation for the existence of such groupings and by
using the answer to this question for the improvement of classification.
The answer, of course, was given by Darwin (1859, p. 413): “I believe
that something more is included [in our classification, than mere resem¬
blance]; and that propinquity of descent,—the only known cause of the
similarity of organic beings,—is the bond, hidden as it is by various degrees
of modification, which is partially revealed to us by our classifications.”
By finding the reason for the existence of natural groups, Darwin changed
the whole basis of classification. The taxonomist no longer “makes” taxa,
he becomes a “discoverer” of groups made by evolution. Now he classifies
not characters but organisms, and characters are downgraded to serve
merely as evidence for something known by the biologist to have indepen¬
dent existence in nature (Simpson, 1961). The consequences of this differ¬
ence in approach are by no means fully understood by all taxonomists.
This is the reason why we may define taxa polythetically (4.7) and are
not concerned when a species lacks some of the diagnostic characters of
the higher taxon to which it belongs or when an individual lacks the diag¬
nostic characters of its species. Stenzel (1963) has perceptively called atten¬
tion to such cases.
A failure to understand the fact that we classify taxa and not charac¬
ters has been the reason for most oversplitting of genera and higher taxa.
The fact that a genus of spiders differs in eye structure from all other
genera of its family is not sufficient cause for the creation of a new subfamily
or family, if the genus in question agrees with the remaining genera in
all other characters. Mutatis mutandis this principle is true for all kinds
of organisms and all taxonomic levels. Classifying taxa means looking at
the totality of characters as a single integrated ensemble, not at single,
disconnected characters in an atomistic manner.
The difference between the logician—who applies the same rules to
the classification of organisms and of artifacts—and the evolutionary tax¬
onomist is now clear. First of all, the biologist classifies populations, not
individuals or phena. The lower taxa are not arbitrary aggregates, but repro-
 Theories of Biological Classification and Their History 77

ductive communities tied together by courtship responses and separated

from other similar units not by arbitrary decisions of the classifier but
by isolating mechanisms encoded in the genetic program of the organism
(2.2.3). The higher taxa, likewise, are characterized by the joint possession
of components of an ancestral genetic program.
Organisms have another unique property which distinguishes them
from inanimate objects: they have a phenotype and a genotype (7.2).
The classification of an inanimate object is completed once its “phenotype”
has been classified. When we classify organisms, classification by phenotype
is only the first step. As the second step we attempt to infer the genotype,
the evolved genetic program, which has a far greater explanatory and
predictive value than the phenotype. The phenotype is susceptible to all
sorts of irrelevant similarities, and it is only the analysis of the inferred
genotype which permits us to determine what similarities in the phenotype
are due to convergence and what others are an expression of the ancestral
genotype. The various kinds of similarities, and how to evaluate them when
classifying taxa, are treated in Chap. 10.
The biological significance of discontinuities between taxa reveals an¬
other distinction between organisms and inanimate objects (Inger, 1958).
Adaptive radiation, extinction, unequal rates of evolution, and many other
purely biological phenomena discussed in the evolutionary literature (Simp¬
son, 1953; Rensch, 1960; Mayr, 1963) are responsible for the observed
inequalities in the gaps between species and groups of species. The fact
that the observed clustering of species can be made the basis for many
alternate classifications does not contradict the fact that species in nature
are clustered and that there are real gaps between taxa at any given time
level. Nor is this conclusion weakened by the fact that “missing links”
must have existed at one time and that throughout the animal system
a few species and species groups exist which connect major higher taxa.
How to deal with clusters and gaps is explained in Chap. 10.

4.4 CLASSIFICATION AND PHYLOGENY

Classifications and the reconstruction of phytogenies are derived from

the same basic evidence, a comparison of more closely or more distantly
related species and an evaluation of similarities (or differences) in indi¬
vidual characters. One might call this method comparative character analy¬
sis. Its heuristic value has long been known among biologists and has led
not only to the flourishing science of comparative anatomy but also to
comparative physiology and ethology.
The raw data permit (1) the reconstruction of phylogenies and (2)
the establishment of classifications. Yet, neither is “phylogeny based on
78 Principles of Zoological Classification

classification” nor “classification based on phylogeny.” Both are based on

a study of “natural groups” found in nature, groups having character combi¬
nations one would expect in the descendants of a common ancestor. Both
sciences are based on the same comparisons of organisms and their charac¬
teristics and on a careful evaluation of the established similarities and differ¬
ences. This method is not circular (Hull, 1967).
The importance of phylogeny for classification is that a meaningful
interpretation and evaluation of the characters of evolving and evolved
organisms is not possible without carefully considering their probable evolu¬
tion. What are primitive (ancestral) characters? Which characters form
a single character complex that changed in response to a single adaptive
shift? What similarities are the result of convergence (including the inde¬
pendent loss of structures) ? We must ask these searching questions concern¬
ing characters because we use characters merely as evidence for the deter¬
mination of genetic relationships among taxa. An answer cannot be given
until several phylogenetic models have been tested. Classification then is
not based on phylogeny, but phylogenetic considerations are important in
the weighting of characters that are used in the construction of classifica¬
tions. The practice of such weighting is discussed in Chap. 10.
Many misleading statements have been made about evolutionary tax¬
onomy. For instance, it is not true that it is “the sole aim of evolutionary
classification to reflect as accurately as possible the facts of evolution.”
Actually the most important aim of evolutionary classification is exactly
the same as of all genuine classifications (in contrast to identification
schemes) : to combine maximal information content with maximal ease
of retrieval of this information. The evolutionist believes that a classification
consistent with our reconstruction of phylogeny has a better chance of
meeting these objectives than any other method of classification. Taxa de¬
limited in such a way as to coincide with phylogenetic groups (lineages)
are apt not only to share the greatest number of joint attributes, but at
the same time to have an explanatory basis for their existence. Since classi¬
fications are not based on established phylogenies, the objection that phy¬
logeny is still largely unknown is irrelevant. What we do base our classifica¬
tion on is the universal fact that all organisms are the product of evolution.
This permits us to set up models of inferred phylogeny, to test them against
various alternate schemes of classification, and to undertake a taxonomic
weighting of characters. There is no excuse for abandoning the evolutionary
approach to classification because no man-made, hierarchical system of
categories is capable of expressing precisely all the known or inferred facts
of evolution. In spite of all its admitted inadequacies, the evolutionary
approach produces a sounder basis for the classification of organisms than
any other.
Evolutionism, as a philosophical basis for classification, is a valid ap-
 Theories of Biological Classification and Their History 79

proach only if and when natural groups of organisms are the result of
divergent evolution. If reticulate evolution were common owing to the
frequent fusion of previously separated evolutionary lines, or if convergence
were frequently so complete as to lead to groupings that could not be
unmasked as having a polyphyletic origin, then the claims of evolutionism
as a proper theoretical basis for biological classification would indeed be
questionable. No one has so far presented convincing evidence that these
two potential difficulties are at all frequent in the evolution of animals.
Unnatural taxa in animals almost invariably result from the use or availabil¬
ity of too few characters, and particularly from reliance on a few poorly
chosen key characters.

4.5 THE OBJECTIVES OF A CLASSIFICATION

There has been much argument about the objectives of a classification.

Some taxonomists claim that the only purpose of a classification is to create
a reliable, easy-to-use filing system for the bewildering diversity of nature.
All the early “identification systems” (see 4.3.1) had this as their objective,
and even some modern authors contend that this is the only purpose for
which a classification is constructed. This claim is mistaken. As important
as this function of classification is, it is not the only one. To reduce the
taxonomist to a filing clerk is to misunderstand his role. This would be
even more true if the filing clerk-taxonomist were asked to file the items
by superficial resemblance rather than on the basis of a thorough under¬
standing of the contents. It would mean not only reducing taxonomy to
a service function for other branches of biology, but also causing it to
do this service badly.
4.5.1 Scientific Theory. For the scientist-taxonomist the most impor¬
tant meaning of a classification is that it is a scientific theory, with all
the qualities of a scientific theory. First of all, it has an explanatory value,
elucidating the reasons for the joint attributes of taxa, for the gaps separat¬
ing taxa, and for the hierarchy of categories. It is precisely this explanatory
property of evolutionary classifications which led to their rapid and almost
universal acceptance after 1859. It is sometimes argued that the descriptive
and the explanatory aspects of classifications should be neatly separated.
This is impossible. A good classification of organisms is automatically
explanatory.
The second property of a good classification is that it has a high
predictive value. The common genetic program characteristic for the mem¬
bers of a natural taxon guarantees with a high probability that all the
members of this taxon share certain characteristics. If I identify an individ¬
ual as a thrush, I can make precise statements concerning its skeleton,
80 Principles of Zoological Classification

heart, physiology, and reproduction without ever testing it. A good classifica¬
tion predicts future experiences for the taxonomist. Indeed, one can test
the soundness of a classification by the ease with which it can accommodate
the findings derived from new characters and newly found species (Mayr,
19656).
In turn the closeness of correlation between characters and classifica¬
tion permits conclusions on the genetic basis and biological significance
of characters. Characters controlled by one or few genes are usually irregu¬
larly distributed or limited to lower taxa. Characters that are shared by
most species of a higher taxon are believed to be the expression of a complex
genotype, particularly when the given character is not directly correlated
with utilization of the adaptive zone of the taxon. Like any scientific theory
a classification has a strong heuristic aspect. An evolutionary classification
stimulates efforts to recognize homologous structures and to test the con¬
cordance of various types of characters (e.g. gross morphological, cytologi-
cal, biochemical, and behavioral). The advantage of this approach in pro¬
tozoology has been well described by Corliss (1962a).
Finally, like any theory, a classification is provisional. The discovery
of new species and the availability of new character complexes is likely
to lead to a modification of the theory, that is, to an improvement of
the classification. Single-character identification schemes inevitably lead to
artificial groupings which have to be abandoned sooner or later. Yet even
the most sophisticated multi-character approach is provisional and subject
to future improvement. The demand for “final classifications” for the con¬
venience of computer programmers can rarely if ever be fulfilled. In this
respect classification is not different from any other theory. All scientific
theories are provisional, subject to continuous testing, and rejected when
found to be wanting.
Recognizing a classification as a scientific theory also answers the
questions as to how important it is to have a classification, and whether
it makes any difference what kind of classification one adopts. Our discus¬
sions must have made it clear that the predictive worth of a classification
depends on the genetic homogeneity of the recognized groupings. Arbitrary
groupings have a very low predictive capacity. Consequently, it is indeed
important to have an “evolutionary” classification, that is, one based on
monophyletic groups that share much of their phenotype and genotype.
The question, Are all aspects of classification equally important? is
also sometimes asked. Surely classification at the species level is of first
importance because the results are of the most immediate concern to fellow
biologists in physiology, ecology, and behavior. The classification of the
classes and phyla, on the other hand, is of importance only for those who
ask phylogenetic questions, whether these concern macromolecules or
organ systems.
 Theories of Biological Classification and Their History 81

4.6 INFERENCES ON THE DELIMITATION OF TAXA

4.6.1 Inferences on the Shared Genetic Program. With few excep¬

tions taxonomists agree that higher taxa (smaller or larger groupings of
species) are the result of evolutionary divergence (descent with modifica¬
tion). The genetic program of the founder species of each evolutionary
line gives a certain amount of genetic unity to its descendants. The genetic
programs of the descendants will usually resemble each other more than
the genetic programs of other phyletic lines not derived from this founder.
If we knew the entire genotype of each organism, it would be possible
to undertake a grouping of species that would accurately reflect their
“natural affinity.” Unfortunately, all attempts to determine the genotype
of higher taxa directly have encountered insurmountable difficulties, among
which three are preeminent:

1. Genetic analysis through crossbreeding is difficult, if not impossible, above

the level of the species.
2. Correlation between observable phenotypic difference and genetic differ¬
ence is rarely complete and often not even close. Intraspecific morphs
are often far more different from each other than sibling species.
3. The characters of higher taxa are highly polygenic, but not even a guess
is possible as to the number of genes involved.

Finally, the basic biochemical building stones of organisms are largely

the same throughout the animal kingdom. Comparative biochemistry does
of course permit an indirect genetic approach, because it can infer how
many mutational steps separate the macromolecule of, let us say, a mammal
from the homologous macromolecule of a lower invertebrate or microorga¬
nism (7.4.9). In the case of biochemical similarities, a particularly careful
distinction must be made between similarities resulting from common de¬
scent and convergent similarities resulting from a response to similar selec¬
tion pressures. Molecules are far simpler structures than anatomical struc¬
tures or behavior patterns, and they may permit convergent evolution in
nonrelated groups up to the point of essential identity. The independent
evolution of a powerul nerve poison (Tetrodotoxin) in certain newts (e.g.
Taricha) and fishes (particularly puffers) is an outstanding example. In
principle, the most promising biochemical approach to the classification
of the higher taxa is that of DNA matching (7.4.9), but the technical
difficulties have not yet been overcome.
Recognition of the fact that characters are often the highly polygenic
product of a complex genotype has resulted in a better understanding of
two kinds of difficulties, the independent acquisition of a new character
in parallel lines and the independent loss of characters in related lines
82 Principles of Zoological Classification

(10.2). Phenomena of parallelism always reveal a hidden genetic potential

derived from a common ancestor.
4.6.2 Inferences from the Phenotype. Even though the direct genetic
approach fails, there are a number of indirect methods which permit us
to infer the genetic program of a taxon from a study of taxonomic charac¬
ters. This process of inference is highly controversial, to a large extent
owing to confused terminologies and unclear thinking. Simpson (1961, pp.
67-106) has clarified this complex problem in the most admirable manner,
and we adopt his analysis almost without reservations.
A classification, as stated above (4.5.1), is a scientific theory. The
members of a higher taxon are those species that show the greatest affinity
with each other. Affinity, however, is not synonymous with superficial simi¬
larity, as is sometimes claimed. Nor is affinity, owing to the dual nature
of phylogeny, synonymous with propinquity of descent, as believed by the
cladists. How to determine affinity is the crucial problem in classification.
Phylogeny cannot be directly observed. It is something that happened
in the past and must be reconstructed, it must be inferred from the available
evidence. All scientific knowledge is in part based on inferences. As Simpson
said (1961, p. 68), “This [dependence on inference] is not a feature pe¬
culiar to taxonomy. It is shared in greater or less degree by most of the
inductive sciences.” The Watson-Crick DNA model, the Krebs cycle, or
indeed most findings of molecular biology cannot be observed directly. They
are inferred from certain observations and are subsequently tested against
further observations or experiments. It is therefore completely legitimate
to define taxonomic categories in evolutionary (largely phylogenetic) terms,
but to use evidence (comparative character analysis) that, as such, is almost
entirely nonphylogenetic. Categories are defined in phylogenetic terms, but
taxa are described by characteristics “that result from phylogeny and are
evidence that the categorical definition is met” (Simpson, loc. cit.).
The evidence, used in the construction of classifications, consists of
taxonomic characters (Chap. 7). Characters, however, differ in their phy-
letic information contents and must be evaluated accordingly. This proce¬
dure, called weighting, is treated in Chap. 10 (10.2).

4.7 THE POLYTHETIC CHARACTERIZATION OF TAXA

In classical taxonomy taxa were usually described by citing characters

that were absolutely diagnostic for the given taxon. Linnaeus constantly
revised his differentiae when the characters of newly discovered species
showed that a character of a previously known taxon was no longer exclusive
to that taxon. Higher taxa were characterized in terms of “characters in
common.” Sometimes a single character is fully diagnostic in this procedure,
sometimes a set of characters. The combination of egg-laying, a flat bill,
 Theories of Biological Classification and Their History 83

and certain anatomical characters is both sufficient and necessary for mem¬
bership in the taxon Platypus. Beckner (1959, pp. 14-31) and Simpson
(1961, pp. 42-43) have discussed such uniquely characterized taxa under
the name monotypic, but since this term is used in taxonomy with a very
different meaning, Sneath (1962, p. 291) has proposed the term monothetic
for this concept.
After 1859 a new definition was adopted for what we now call a
taxon. When the definition of the logicians—“individuals sharing common
characters”—was replaced by “members of a group having descended from
a common ancestor,” a monothetic characterization of a taxon was no
longer necessary. Actually, Adanson (in 1763) and other members of the
empirical school had decided long before that a member of a taxon did
not need to possess all the characters of the taxon and that such a deviant
component of the taxon (e.g. species in a genus) did not need to be ex¬
cluded and placed in a separate taxon. Taxa characterized by a set of
characters of which each member has a majority are called polythetic taxa.
For further details on this concept (called polytypic) see Beckner (loc.
cit.) and Simpson {loc. cit.).
A higher taxon is polythetic if it satisfies the following three
conditions:

1. Each species possesses a large (but unspecified) number of the total

number of properties of the taxon.
2. Each property is possessed by a large number of the species.
3. No property is possessed by every species of the aggregate but is missing
in the species of all other taxa.

Consequently, no single feature is essential for membership in a poly-

thetically defined taxon nor is any feature sufficient for such membership.
Almost every monograph dealing with suprageneric taxa refers to
characters that have some diagnostic value in conjunction with a couple
of other characters but that are sufficiently irregular in distribution not
to be reliably diagnostic by themselves. It is this incomplete correlation
between characters and taxa which makes the construction of diagnostic
keys so difficult. The fleas (Siphonaptera) supply abundant illustrations
of suprageneric taxa that can be reliably defined only by combinations
of characters, each one of which may also occur outside the given taxon
or may occasionally be absent in a member of the taxon (Holland, 1964).

4.8 INFERRING RELATIONSHIP

Evolutionary classification demands the delimitation of taxa consisting

of closest relatives. The evidence for relationship (10.1) consists primarily
of weighted similarity. The interpretation of similarity is made difficult
84 Principles of Zoological Classification

by a number of practical and theoretical problems which the taxonomist

must clearly understand in order to avoid mistakes.
The polythetic diagnosis of taxa brings out the well-known fact that
related taxa often overlap in some of their characteristics. This, together
with convergence, is indeed the most formidable difficulty faced in routine
taxonomy. It is one of the consequences of mosaic evolution that the charac¬
ter progressions of several character trends in a group are realized by the
descendants independently and haphazardly. The living representatives of
different phyletic lines derived from the same common ancestor may have
a similar ratio of primitive and advanced characters, but a different assort¬
ment in each case (Table 4-1). The correct assortment of taxa under
these circumstances depends on the finding of additional characters which
will help in the weighting of the previously used ones. The methods of
Maslin (1952), Wagner (1962), Hennig (1950, 1966), Wilson (1965),
and Camin and Sokal (1965) are attempts to arrive at the construction
of sound taxa in the face of widely overlapping character distributions
(see Chap. 10).
4.8.1 Homology. The occurrence of convergence and various other
kinds of similarity not resulting from common descent demonstrates that
in the construction of classifications one must distinguish several kinds of
similarities. Only similarities between homologous characters are of tax¬
onomic importance. The term homology, like the terms species and classi¬
fication, antedates evolutionary biology, but like these other terms it has
acquired since 1859 a new, more precise, and biologically more significant
meaning. Some workers have adopted different terminologies for structural
and for nonstructural similarities, but with the ever greater utilization of
nonmorphological characters in taxonomy, such a distinction would seem
impractical and misleading [see discussions by Remane (1952, pp. 31-103),

Table 4-1. Mosaic Evolution. Each of the six taxa shows a different
assortment of primitive and advanced characters of features A-G.

Feature
Taxon
A B C D E F G

I 1 5 3 2 4 1 6
II 3 6 1 4 1 3 3
III 6 4 1 5 2 2 5
IV 2 2 6 1 4 5 3
V 5 1 4 6 2 3 1
VI 1 2 5 6 3 5 1

1 = primitive, 6 = very advanced character.

 Theories of Biological Classification and Their History 85

Table 4-2. Difference between Homology and Analogy

Derived from equivalent feature in common ancestor

Appearance
Yes No

Similar Homologous Analogous

Not similar Homologous

Simpson (1961, pp. 77-93), and Wickler (1961)]. With Bock (1963) we
prefer to go back to Owen’s two terms, but to define them as follows:
Homologous features (or states of the features) in two or more orga¬
nisms are those that can be traced back to the same feature (or state)
in the common ancestor of these organisms.
Analogous features (or states of the features) in two or more orga¬
nisms are those that are similar but cannot be traced back to the same
feature (or state) in the common ancestor of these organisms.
In the case of homology, similarity is not part of the definition because
homologous structures are by no means necessarily similar (e.g. ear ossicles
of mammals and the corresponding jaw bones in the lower vertebrates).
Similarity must be referred to in the definition of analogy because non-
homologous features that are not similar are not considered analogous
(Table 4-2).
Having an unambiguous definition of homologous permits us to pro¬
ceed in a similar way as in the application of the concepts biological species
or biological classification. We must now seek the evidence that two features
which we compare meet or do not meet our definition. Lists of criteria
exist which help us in making the right decision; they will be discussed
in Chap. 10.
4.8.2 Phylogenetic Laws. One of the reasons why the phyletic ap¬
proach to classification came into discredit has been the reliance of some
taxonomists on so-called phylogenetic laws and principles. Many of these
are not only unreliable but totally false. Among so-called phylogenetic laws
that should be rejected are the following:

1. Simple is always ancestral to complex.

2. Ontogeny (larval or embryonic stages) recapitulates phylogeny.
3. The “type” evolves harmoniously, and consequently all structures and
organ systems evolve at equal rates.
4. There are goal-directed, teleological, evolutionary trends (orthogenesis).
5. New types of organisms originate by saltation.

Rensch (1947), Remane (1952, pp. 164-301), and Simpson (1953, 1961)
have analyzed some of these so-called phylogenetic laws as well as additional
86 Principles of Zoological Classification

ones not here enumerated. Hennig (1950, 1966) has attempted to develop
new phyletic laws on the basis of cladism.

SUMMARY

A classification based on phyletic weighting has numerous advantages.

It is the only known system that has a sound theoretical basis, it has greater
predictive value than other kinds of classifications, it stimulates a character-
by-character comparison of organisms believed to be phylogenetically re¬
lated, and it encourages the study of additional characters and character
systems in order to improve the soundness of the classification and hence
its information content and predictive value. Finally, it leads to the dis¬
covery of interesting evolutionary problems. Systems based on phyletic
weighting thus not only have scientific advantages but are actually best
able to answer the demands of the practice by having a greater total infor¬
mation content than artificial systems.
Chapter^ The Hierarchy of Categories
and the Higher Tax a

E. ach major group of animals can be subdivided into

smaller and smaller subgroups. Within the vertebrates we can distinguish
subgroups such as birds and mammals. Within the mammals, carnivores and
rodents. Within the carnivores, those that are doglike, those that are catlike,
and so forth. As Darwin said (1859, p. 411), “All organic beings are found
to resemble each other in descending degrees, so that they can be classed in
groups under groups,” and if one wants to construct a classification of these
species, “this classification is evidently not arbitrary like the grouping of the
stars in constellations.” The task of classification then is the delimitation of
these groups and their arrangement in an orderly sequence. For this en¬
deavor it is of the utmost importance that the student has a clear under¬
standing of the meaning of the terms taxon and category (1.2).

5.1 HIGHER TAX A

The groups of species found in nature are higher taxa. Cats, car¬
nivores, mammals, and vertebrates are higher taxa of different rank. The
first step in classification, discussed in Chap. 4, is to determine which species
show similarities indicating that they belong to one group, and furthermore
to determine the delimitation of these groups, that is, the location of the
discontinuities between neighboring groups.
87
88 Principles of Zoological Classification

A higher taxon is an aggregate of related species separated from

others by a discontinuity. The ideal situation implied by this definition
is not always met. For instance, in monotypic taxa the “aggregate of species”
consists of a single species (because there are no close relatives), and the
discontinuity, the gap separating one higher taxon from the next, varies
greatly in extent. It is sometimes almost entirely bridged by intermediate
species. See Chap. 10 and Simpson (1961, chap. 6). The definition fails
to give any help in determining the rank of the taxon, because it is equally
valid for higher taxa of all levels, from genus and family up to the phylum
(see 5.4, 5.5, 5.6).
The Meaning of Higher Taxa. Most well-defined higher taxa, particu¬
larly at the genus and family level, occupy a well-defined niche or adaptive
zone. They owe their origin to the invasion of this zone by a founder
species and to the subsequent active and adaptive radiation which usually
follows a successful adaptive shift (Simpson, 1953, 19596, 1961; Mayr,
1960, 1963, chap. 19). It helps in the delimitation of higher taxa and
in their ranking to be aware of their evolutionary origin. In recent years
taxonomists have given much attention to the problem of the origin of
higher taxa (for instance, Schaeffer and Hecht, 1965). It is advisable for
two reasons to stress the ecological significance of higher taxa (5.3.2, 5.4.3,
10.4). Since the members of such a taxon are the descendants of the founder
species that invaded the new adaptive zone, they clearly qualify as a mono-
phyletic group. Secondly, since all the species occupy the same adaptive
zone, in spite of some secondary radiation, they usually show a considerable
amount of structural unity. Keeping this in mind may well prevent unneces¬
sary “splitting” (10.3). Species or groups of species that are not separated
by a well-defined discontinuity also fail, in most cases, to occupy a well-de¬
fined niche or adaptive zone.
As a heritage from the days when classification was considered synony¬
mous with identification, there is an erroneous concept of the higher taxon,
or rather the members of a higher taxon, as the carriers of an identifying
character. A taxon is in fact a group of relatives, and whether or not
they have the same “characters in common” is irrelevant. Many taxa are
based on a combination of characters, and frequently not a single one
of these characters is present in all members of the taxon, yet such a
taxon may have a sound “polythetic” basis (4.7).
A group of related species consists of species descended from a com¬
mon ancestor. Sound grouping of species is the indispensable foundation
of a sound classification. In order to compensate for the possibility of the
misleading effect of mosaic evolution, parallelism, and convergence, a care¬
ful phyletic weighting of numerous characters must be undertaken (10.4).
The classifications of bees (Michener, 1944), Saturnid moths (Michener,
1952), and butterflies (Ehrlich, 1958) are exemplary analyses of this sort.
 The Hierarchy of Categories and the Higher Taxa 89

Where a fossil history is available, as in the case of mammals, its evaluation

often leads to a better understanding of the higher taxa (Simpson, 19596).
The practical aspects of the procedure of delimiting taxa and ranking
them are treated in Chap. 10.

5.2 THE LINNAEAN HIERARCHY

The taxa of animals and plants, according to their comprehensiveness,

are ranked in a hierarchy of categories. Within the animal kingdom the
highest regularly used category is the phylum, and the lowest the species.
Linnaeus, the first taxonomist to establish a definite hierarchy of taxonomic
categories, recognized within the animal kingdom only five: classis, or do,
genus, species, and varietas. Two additional categories were soon generally
adopted when the number of known animals grew—making finer divisions
necessary: the family (between genus and order) and the phylum (between
class and kingdom). The varietas, used by Linnaeus as an optional category
for various types of infraspecific variants, was eventually discarded or re¬
placed by the subspecies (3.3).
The remaining categories form the basic taxonomic hierarchy of ani¬
mals. Any given species belongs thus to seven obligatory categories, as
follows:

Wolf Honey Bee

Kingdom Animalia Animalia

Phylum Chordata Arthropoda
Class Mammalia Insecta
Order Carnivora Hymenoptera
Family Canidae Apidae
Genus Canis Apis
Species lupus mellifera

The basic five higher categories (genus, family, order, class, and phy¬
lum) permit the placing of a species of animals with a fair degree of
accuracy. However, as the number of known species increased, and with
it our knowledge of the degrees of relationship of these species, the need
arose for a more precise indication of the taxonomic position of species.
This was accomplished by splitting the original seven basic categories and
inserting additional ones among them. Most of these are formed by combin¬
ing the original category names with the prefixes super or sub. Thus there
are superorders and suborders, superfamilies and subfamilies, etc. The most
frequently used additional new category name is perhaps the term tribe
for a category between genus and family. Vertebrate paleontologists also
routinely use the category cohort between order and class. Some authors
90 Principles of Zoological Classification

use terms for additional subdivisions, such as cladus, legio, and sectio. Some
use infraclass below the subclass, and infraorder below the suborder.
The generally accepted categories are the following:

Kingdom
Phylum
Subphylum
Superclass
Class
Subclass
Cohort
Superorder
Order
Suborder
Superfamily (-oidea)
Family (-idae)
Subfamily (-inae)
Tribe [-ini)
Genus
Subgenus
Species
Subspecies

Indicated in parentheses are the standardized endings for the names

of tribes, subfamilies, families, and superfamilies (13.33). Standardized end¬
ings for the categories above the family group have not yet been adopted
in zoology.
The Linnaean hierarchy, with its need for arbitrary ranking, has often
been attacked as an unscientific system of classification. Alternate methods,
such as numerical schemes, have been proposed but have not found favor
among taxonomists, primarily for two reasons. Assigning definite numerical
values to taxa demands a far greater knowledge of the relationships of
taxa than can be inferred from the available evidence. Secondly, an assign¬
ment of such values would freeze the system into a finality which would
preclude any further improvements. It is the very subjectivity of the Lin¬
naean hierarchy which gives it the flexibility required by the incompleteness
of our knowledge of relationships. It permits the proposal of alternate
models of relationship and gives different authors an opportunity to test
which particular balance between splitting and lumping permits the presen¬
tation of a maximal amount of information. Like any other scientific theory
it will forever be provisional (4.5). For the logical structure of the hierarchy,
see Buck and Hull (1966).
 The Hierarchy of Categories and the Higher Taxa 91

5.3 THE HIGHER CATEGORIES

Definition: A higher category is a class into which are placed all

the taxa that rank at the same level in a hierarchic classification. The
category selected for a given taxon indicates its rank in the hierarchy.
As explained in Chap. 1, taxa are based on zoological realities, categories
are based on concepts. In that respect there is no difference between the
category species and the higher categories from the genus up. In many
other respects there is a great deal of difference between the concept of
the species and the concepts of the higher categories.
The category species is “self-operationally” defined by the testing of
isolating mechanisms in nature while nonarbitrary definitions for the supra-
specific categories are not available. The species category signifies singu¬
larity, distinctness, and difference, wrhile the higher categories have the
function of grouping and ordering by not emphasizing differences between
species, rather by emphasizing affinities among groups of species. They
are collective concepts. Even though an operational definition for the higher
categories does not exist, nor for the rank which they signify, they do
have an objective basis because a taxon placed in a higher category (if
correctly delimited) is “natural,” consisting of descendants from a common
ancestor. Higher taxa are often, if not usually, well delimited and separated
from other taxa of the same rank by a pronounced gap.
Finally, comparative data furnish the evidence used for the delimita¬
tion of higher taxa and their ranking into categories, w'hile interbreeding
is the criterion used for ranking at the species level, because the species
is a relational concept (Mayr, 19576) and the higher categories are not.
Preevolutionary taxonomists, including Linnaeus, used higher cate¬
gories, but they wrere unable to denote their significance in the framework
of Aristotelianism, as was particularly evident for the categories above the
genus level. They tried to explain the origin of the categories by deriving
the lower ones from the higher through splitting, a remnant of Thomistic
thinking. It was Darwin who supplied the scientific interpretation (1859,
p. 422) : “The natural system is genealogical in its arrangement, like a
pedigree; but the degrees of modification which the different groups have
undergone have to be expressed by ranking them under different so-called
genera, sub-families, families, sections, orders, classes.” The descendants
of an aberrant species may evolve into a different genus, the genus in
the course of geological history into a different family, and so forth. The
origin of higher categories is thus exactly opposite from that envisioned
by the scholastic philosophers. Higher categorical rank evolves through evo¬
lution, not lower rank through subdivision of higher categories.
Most taxa above the family level are sharply delimited. Mollusks,
92 Principles of Zoological Classification

penguins, beetles, and indeed most higher taxa are separated from their
nearest relatives by a decided gap, far more so than most genera and
families. Nevertheless it remains true that the higher categories in which
we place these taxa are ill-defined. Category means categorical rank, and
no yardstick has yet been found for the nonarbitrary ranking of taxa. There
is hardly a higher taxon that is not ranked higher by some and lower
by other specialists. It is in the arbitrariness of definition that all higher cate¬
gories differ from the species category. The criteria and operations used
during the ranking procedure are discussed in detail in Chap. 10 (10.4).

5.4 THE GENUS

5.4.1 Definition. The genus is the lowest higher category and the
lowest of all categories established strictly by comparative data (Cain,
1956). For the modern taxonomist the genus is no different in concept
from family, order, or other higher categories. For Linnaeus, who based
his theory of classification on the principles of Aristotelian logic, the genus
occupied a very special place (Cain, 1958). This fact would have only
historical interest for us if Linnaeus had not incorporated Aristotelian logic
into the binominal system of nomenclature.
Since there is no operational definition available for any of the higher
categories, one is forced to adopt a pragmatic definition: A genus is a
taxonomic category containing a single species, or a monophyletic group
of species, which is separated from other taxa of the same rank [other
genera] by a decided gap. It is recommended for practical reasons that
the size of the gap be in inverse ratio to the size of the taxon. In other
words, the more species in a species group the smaller the gap needed
to recognize it as a separate taxon, and the smaller the species group the
larger the gap needed to recognize it. One of the functions of the genus,
from Linnaeus’ time on, is to relieve the memory (to facilitate information
retrieval), and the “inverse ratio” recommendation prevents the recognition
of a burdensome number of monotypic genera. To delimit as genera species
groups of optimal size is an operation that requires experience, good judg¬
ment, and common sense. For a discussion of helpful criteria, see Chap.
10 (10.4).
An equivalent, nearly identical, pragmatic definition is applicable to
the categories above the genus—family, order, class, etc.
In order to qualify for a given rank, a taxon must satisfy a number
of conditions (see 10.5). It must be sufficiently different from other taxa
of the same rank; it must be separated by a discontinuity; it should occupy
a distinctive niche or adaptive zone; and in the absence of a marked
discontinuity it should not display too great an internal diversity (hetero-
 The Hierarchy of Categories and the Higher Taxa 93

geneity). Finally it should, if possible, satisfy certain practical requirements,

in consequence of which the recognition of a higher taxon is often a
balanced compromise between the stated qualifications. Only the ideal genus
is well separated by a gap, is of the proper size, is internally homogeneous,
and fills a distinctive adaptive zone.
For the nomenclature of generic names see 13.38, see also 13.31 (gen¬
der) and 13.39 (collective groups).
5.4.2 Generic Characters. Taxonomic characters that prove generic
distinctness do not exist (7.5). Taxonomic literature would have been
spared innumerable generic synonyms if taxonomists had always remem¬
bered Linnaeus’ (1737) dictum: “It is the genus that gives the characters,
and not the characters that make the genus.” This, in a sense, is still
generally valid, even though we have abandoned the Aristotelian logic on
which Linnaeus based his statement. The soundest genera are based on
an overall appreciation and weighting of the various considerations previ¬
ously cited (Michener, 1957).
The species included in a genus usually have many features in com¬
mon, thus facilitating its delimitation. Recognition of a higher taxon is
generally based on the occurrence of correlated character complexes (see
10.4). These may include some rather minute and inconspicuous characters,
but, as Darwin said (1859, p. 417), “The importance, for classification,
of trifling characters mainly depends on their being correlated with several
other characters of more or less importance. The value indeed of an aggre¬
gate of characters is very evident in natural history.” So important has
this principle been considered by taxonomists that it led to much generic
splitting when a taxonomist found a species which lacked one or another
character of the correlated complex. Instead of revising his image of the
genus, he named a new one.
Some genera are clearly natural groups yet cannot be diagnosed un¬
equivocally by a single character. This occurs because every character, even
though diagnostic for the majority of the species, is modified or absent
in at least one or the other species of the genus (4.7). This is true, for
example, of many genera and even families of birds.
5.4.3 Meaning of the Genus. When we assign generic rank to a
group of species, we want to express a number of things that are characteris¬
tic of all the higher categories. A genus taxon is a phylogenetic unit, which
means that the included species are descended from a common ancestor.
Almost invariably it is also true that the genus is an ecological unit consisting
of species adapted for a particular mode of life. The genus niche is obviously
broader than the species niche, but both kinds of niches exist. Lack (1947)
has convincingly shown the adaptive significance of genera for the Gala¬
pagos finches (Figs. 5-1, 5-2).
Like any operation in classification, recognition of a genus taxon cor-
94 Principles of Zoological Classification

Fig. 5-1. Adaptive radiation of Darwin’s finches (Geospizinae) on the Galapagos Islands into a number
of different niches (from Lack, 1947).

responds to the proposing of a scientific theory. Like all scientific theory

it must have explanatory, heuristic, and predictive value (4.5). If there
are several alternate ways of delimiting genera, we must be guided by
the same principles as in the recognition of any scientific theory. “Where
alternatives are available, we stand by the theory or concept that is most
useful—the one that generalizes the most observations, and permits the
most reliable predictions” (Inger, 1958, p. 383). For comments on the
generic concept, see also Rosen and Bailey (1963).

5.5 THE FAMILY

As in the case of the genus and the other higher categories, it is

not possible to give a nonarbitrary definition of the family category. What
a lay person would often designate a “kind of animal” is often a family:
The ladybird beetles (Coccinellidae), the long-horned beetles (Ceramby-
cidae), the woodpeckers (Picidae), the swallows (Hirundinidae), and so
forth. How distinctive a group of genera must be in order to be considered
a family varies from one zoological group to another, for the various reasons
previously indicated. If one wants to give a definition of the family category,
it would be equivalent to that of the genus:
A family is a taxonomic category containing a single genus or a
monophyletic group of genera, which is separated from other families by
a decided gap. It is recommended, as in the case of the genus, that the
size of the gap be in inverse ratio to the size of the family.
 The Hierarchy of Categories and the Higher Taxa 95

Like the genus, but perhaps to an even greater degree, the family
is usually distinguished by certain adaptive characters which fit it for a
particular niche or adaptive zone, e.g. the woodpeckers of the family
Picidae, the leaf beetles of the family Chrysomelidae, etc. In most cases,
families obviously are older than genera and have more often a worldwide
distribution. An entomologist who knows the 422 families of British insects
can go to Africa, or even Australia, and recognize nearly all the same
families occupying similar niches.
Thus the family is a very useful category. The British entomologist
would have to learn only 422 names to place a total of about 4,767 genera
and 20,244 species. It is especially useful to the general zoologist because
each family usually presents a general facies which is recognizable at a

Geospizo
mognirostris

/ioo%\
ANIMAlAfk'
FOOD W-

MM Hi*

Certhidea
olfvacea
Fig. 5-2. Niche occupation, feeding habits, and bill structure in ten species of Geospizinae from
Indefatigable Island (from Bowman, 1961).
96 Principles of Zoological Classification

glance, and all its species occupy a similar niche in their particular com¬
munity, as, for instance, most of the thousands of species of Cerambycidae
(long-horned beetles) in the world.
At any given locality the various families, like the various species,
are generally distinct. Decided gaps between families are the rule rather
than the exception, and little or no difficulty is encountered in “keying
out” families in local faunal works. Unfortunately, the situation becomes
much more complicated when a worldwide study is undertaken. Families
are often found to break up into different distinctive groups on each con¬
tinent, and types bridging the gap with other families are sometimes found.
Relict groups may exist at the family level and defy efforts to attain a
clear-cut classification. Thus, in many insect groups (scale insects, aphids,
water striders, etc.) a choice has had to be made between enlarging the
family concept beyond the limits of local convenience or recognizing con¬
necting exotic types as separate families and using a superfamily category
for the group as a whole. In entomology there appears to be a trend,
not necessarily desirable, in the direction of the second of these choices.
In ornithology a knowledge of the tropical relatives of the Temperate Zone
forms has led to a reduction in the number of families. For instance, a
study of tropical genera has induced many authors to consider the Old
World flycatchers (Muscicapidae), warblers (Sylviidae), and thrushes
(Turdidae) as only subfamilies of a more broadly conceived family,
Muscicapidae.
Linnaeus did not recognize the family as a category, but it is significant
that most of his genera have since been elevated to the rank of families.
From this we may infer that his generic concept was not incompatible
with our modern family concept, the difference between the genus and
family being merely one of degree. With only 312 genera of animals in
1758, Linnaeus had no need for an intermediate category between genus
and order. Flowever, the number of newly discovered animal types increased
so rapidly that the early nineteenth-century naturalists gradually evolved
and universally applied the family concept to designate an intermediate
level between genus and order.
The number of families continues to grow because of the advance
in knowledge of existing animals and the discovery of new types. Thus
by the end of the nineteenth century approximately 1,700 families of animals
were recognized (Perrier, 1893-1932, Traite de Zoologie). That the trend
is continuing is indicated by the fact that Brues, Melander, and Carpenter
(1954) recognize 941 families of insects alone.
For matters relating to the nomenclature of families, see Chap. 12
and 13.30 (stem of family name), 13.33 (family names), 13.35-36 (coordi¬
nated and subordinate taxa), and 13.37 (homonymy).
 The Hierarchy of Categories and the Higher Taxa 97

5.6 ORDERS, CLASSES, AND PHYLA

The higher taxa above the family level are on the whole very well
defined in the Recent fauna, and much less often connected by intermediates
than families or genera. There are two exceptions to this broad statement.
First, there is still doubt as to the significance of taxonomic characters
in certain groups of lower invertebrates, for instance the sponges and the
turbellarians. In some cases there has been a complete reclassification on
the ordinal level, owing to the reevaluation of the weight of certain charac¬
ters. The second reason is that even where there has been complete consen¬
sus as to the delimitation of the taxa, there has been strong disagreement
as to the ranking. Instead of recognizing more suborders and superfamilies,
certain authors have raised almost all taxa in rank, which has resulted
in a great imbalance of the respective portions of the system (see 10.3,
10.4).
The taxa ranked in higher categories represent the main branches
of the phylogenetic tree. They are characterized by a basic structural pattern
laid down early in evolution, the special adaptive significance of which
can now be perceived only dimly, if at all. Superimposed on it are seemingly
endless adaptive modifications resulting from series of adaptive radiations
that have taken place in the classes and phyla. In general, then, taxa
in the higher categories are definable in terms of a basic structural pattern,
but except for certain highly specialized groups, such as the order Siphonap-
tera (fleas), the order Chiroptera (bats), the order Impennes (penguins),
etc., the higher taxa are not primarily or even predominantly distinguished
by ad hoc adaptations. The taxa contained in the higher categories are
in most cases widely distributed in space and time. For the names of higher
taxa, see 13.34.
As in the case of genera and families, there has been a trend toward
increase also in the number of recognized taxa above family rank. According
to recent tabulations there are approximately 25 phyla, 80 classes, and
350 orders of Recent animals.

5.7 PRACTICAL CONSIDERATIONS IN THE

CONSTRUCTION OF A CLASSIFICATION

Biological classification, as stressed in Chap. 4, is based on the fact

that evolution produces groups of more nearly or more distantly related
species. Evolution thus sets severe limits to the number of possible classifica¬
tions. Nevertheless, even where there is complete consensus about the phy-
98 Principles of Zoological Classification

logeny of a group of organisms, it is nearly always possible to translate

this knowledge into a number of alternate classifications. The reason for
this is that three operations of the taxonomist cannot be carried out without
an element of arbitrariness: (1) the delimitation of groups which we for¬
mally recognize as taxa (their “size”), (2) the rank in the hierarchy which
we assign to a given taxon (e.g. tribe, subfamily, family), and (3) the
position in the sequence of taxa. The necessity of having to translate a
multidimensional phylogenetic tree into a linear sequence permits many
alternate arrangements (Chap. 10.7). A taxonomist will be the more suc¬
cessful in his classification the more clearly he realizes that it is the major
function of a classification to be useful. A classification is a communication
system, and the best one is that which combines greatest information content
with greatest ease of information retrieval. For instance, a classification
which attempts to express every possible shade of relationship (by “split¬
ting”) makes information retrieval exceedingly difficult and hence defeats
its own purpose. The taxonomist must remember that the key to a filing
system does not contain all the information contained in the file.
The specific points that need to be taken into consideration in the
making of a useful and stable classification are discussed in Chap. 10.
Part Methods of Zoological
Classification
Chapter s' Taxonomic Collections and
\J the Process of Identification

A
-A-11 classification is based on the comparison of specimens
representing populations and species. We can determine the species-specific
characteristics of a species only by comparing it with other similar species,
preferably its nearest relatives. An adequate comparative collection is there¬
fore as indispensable to the taxonomist as are electron microscopes, Warburg
apparatuses, ultracentrifuges, and similar equipment for the cellular and
molecular biologist.
Collections must either be borrowed from museums, which are re¬
positories of systematic collections, or collected by the specialist himself.
In most cases both sources must be tapped. Borrowed material is usually
insufficient for certain crucial areas and does not give the biological informa¬
tion which is so vital in modern taxonomic research. On the other hand,
it would require many years of effort for a single collector to achieve the
broad geographic scope available in a museum collection accumulated dur¬
ing scores of years, possibly centuries.

6.1 SYSTEMATIC COLLECTIONS

6.1.1 Value of Collections. Museums serve an important role as

centers of documentation. They supply a permanent record of faunas and
floras, particularly where the biota has since been destroyed by natural

101
102 Methods of Zoological Classification

catastrophes or the activities of man. This is particularly true of the localized

biota of streams, lakes, and islands. Museums contain a sampling of many
areas that are more or less inaccessible owing to their remoteness or for
political reasons. Much of the material now preserved in various institutions
can be replaced only at very high cost, or not at all. Other material is
of unique value because it forms the basis of published research. It may
be needed again at a later period for verification of the original data or
for renewed study in the light of more recent knowledge or new techniques.
For a survey of the scientific significance of taxonomic collections, see Bio¬
logical Material (Mayr and Goodwin, 1955).
A taxonomic revision of a given group is possible only if adequate
material of the majority of included species can be assembled. It is therefore
legitimate, indeed necessary, that curators of museums build up their collec¬
tions in groups which they or their associates plan to study in the future.
The working taxonomist knows from experience how many revisions were
started, only to be set aside for lack of adequate material. Currently unused
material in a collection may be dormant, but this does not mean that
it is dead. Collections are reference material, and in that respect they are
just as necessary as books in a library, which likewise are not in continuous
use yet must be available when needed.
6.1.2 Purpose of a Scientific Collection. For the old-style, typologi-
cally oriented taxonomist a collection was an identification collection. When
he obtained additional specimens of species already represented in the col¬
lection, he considered them duplicates to be used for exchanges or to be
given away. According to current thinking, biological classification is the
ordering of populations (Chap. 4). Collecting, then, is the sampling of
populations. Considering the great variability of most natural populations
(Chap. 8), an adequate sample of every population should be collected
and preserved (6.1.6).
But what is “adequate”? There are compulsive collectors, veritable
pack rats, who insist on storing thousands upon thousands of specimens
of each species. The ideal is somewhere between the typological extreme
of preserving only a typical specimen of each species and the hoarding
tendency of the compulsive collector. In making the right decision one
must be guided by various considerations. More material is needed in a
species with strong individual and geographical variation than in a uniform
species. More material is needed for studies of specific and subspecific char¬
acters than of the characters of higher taxa. Birds, for instance, are on
the whole more uniform in their anatomy than in their plumage characters.
It would be illogical for this reason to preserve large series of spirit speci¬
mens of every species, such as are needed for the study of the geographic
variation of plumage characters.
The size of the sample thus depends on the objectives of the research.
 Taxonomic Collections and the Process of Identification 103

The presence or absence of an anatomical character diagnostic for a higher

taxon is in most cases not subject to much variation, and one or two
preserved specimens of a number of genera may be all the working material
needed. However, when a population analysis is attempted, whether con¬
cerned with a study of size, proportions, coloration, or polymorphism, large
samples from numerous localities are needed. In fishes, for instance, where
each population of a species may have different statistics for various meristic
characters (fin rays, vertebrae, etc.), a single specimen per locality would
tell us very little. All phena of a species should, so far as possible, be
represented in the collection.
6.1.3 Collecting and Research. On the whole, taxonomists spend
only a small fraction of their time on the collecting of new material. Like¬
wise, most specialists have caught up reasonably well with material they
themselves collected. There are exceptions. Some big expeditions of the
nineteenth century gathered material that is still not yet fully worked out.
Some taxonomists cannot resist the temptation to use the summer to go
in the field hoping to work out the material during the ensuing winter.
However, those who are teaching at universities are often too busy during
the academic year to continue their taxonomic research. A moratorium
on further collecting is advisable in such cases to prevent too large an
accumulation of unopened packing cases from previous field seasons. Large
oceanographic expeditions also have a tendency to accumulate far more
material than the extremely small number of specialists can work up at
once. However, even such material is generally processed so as to make
it readily available to specialists (see 6.3.1).
6.1.4 Scope of Collections. Only a few large national museums at¬
tempt worldwide coverage in all groups of animals. Most museums are
largely restricted to a geographic area and to certain groups of animals.
It is most important for the staff of a museum to have a clear-cut acquisition
policy. Too broad a coverage inevitably leads to shallowness and a failure
to obtain the depth required for monographic studies. There has been
a steady and wholesome trend away from broad-purpose expeditions and
faunal surveys in favor of intensive collecting of specific families or genera.
The late Admiral H. Lynes, for instance, who was especially interested
in Cisticola, a genus of African warblers with some 40 species, made a
whole series of collecting trips to nearly every corner of Africa. He combined
the collecting of specimens with a detailed study of the ecology, habits,
songs, and nest construction of these birds. The result was that the genus
Cisticola, formerly the despair of the bird taxonomists, is now reasonably
well understood (Lynes, 1930). The work of J. Crane on the fiddler crabs
(Uca) is another example.
A number of considerations are important if a faunal survey is at¬
tempted. It should be directed to a natural geographical area and not
104 Methods of Zoological Classification

be hampered by a blind dependence on political boundaries. Faunal surveys

have had the best results when concentrating on a particular taxonomic
group. The most ambitious single undertaking along these lines in recent
decades was probably the Whitney South Sea Expedition, operating under
the auspices of the American Museum of Natural Flistory in New York.
This expedition visited practically every island in the South Pacific from
the Tuamotus and Marquesas in the east to the Bismarck Archipelago,
Palau, and the Marianas in the west, largely concentrating on bird collect¬
ing. It operated continuously from 1921 to 1934, and its work was continued
by single collectors into 1940. The essentially complete collections made
by this single expedition supplied the material not only for scores of detailed
revisions but also for basic zoogeographic and evolutionary studies, such
as Systematics and the Origin of Species (Mayr, 1942). The value of a
single well-made collection is generally far greater than that of an equivalent
number of specimens in casually made collections.
6.1.5 Where and How to Collect. Any collecting trip must be care¬
fully planned. All possible geographic information must be obtained before¬
hand, including the distribution of vegetation types, altitudes, seasons, means
of public and private transportation, etc. In addition, previous collections
must be carefully analyzed and existing type-localities mapped. A plotting
of collecting stations and, in particular, a mapping of species distributions
will reveal the location of crucial gaps. If the study of geographic variation
is a major objective, the periphery of the range of each species should
be given particular attention. This is where geographical isolates and in¬
cipient new species occur most frequently. If a species shows seasonal varia¬
tion, the collections should be spaced seasonally. The season during which
sexual maturity occurs is relatively short in many animals, so collecting
should be done during that time. This is even more important if recordings
of song and courtship, nest structure, egg proteins, embryos, or other mate¬
rials are needed that can be obtained only during the breeding season.
In the case of allopatric populations, the categorical status of which is
uncertain (species or subspecies), a special effort should be made to collect
in the intervening region to determine whether or not intergradation occurs.
Owing to the rapid recent increase of human populations, resulting in
more intensive agriculture and in drastic deforestation, there are many
areas that are in desperate need of immediate collecting before the localized
faunas become extinct. At the present time this task is far more urgent
than collecting in remote uninhabited areas.
Innumerable techniques for the collecting of different groups of ani¬
mals are described in standard collecting manuals. New techniques are
continually being developed, such as the use of mist nets for bird collecting
and of “black light” (ultraviolet lamps) for insect collecting. Different
kinds of traps, baits, poisons, and so on are well known to specialists, who
generally share such information with beginners. Depending on the tax-
 Taxonomic Collections and the Process of Identification 105

onomic group to be covered, one or the other of the following books (of
which the full title can be found in the terminal bibliography) will be
most useful:
Anderson, 1948 (vertebrates), Anthony, 1945 (mammals), Beer and
Cook, 1958 (ectoparasites), Bianco, 1899 (marine animals), British Mu¬
seum, 1936ff (various groups), Kirby, 1950 (protozoans), Kummel and
Raup, 1960 (fossils), MacFadyen, 1955 (soil arthropods), Oldroyd, 1958
(insects) Peterson, 1934, 1937 (insects), Oman and Cushman, 1946 (in¬
sects), Russell, 1963 (marine), Van Tyne, 1952 (birds), WagstafTe and
Fidler, 1955 (invertebrates). Knudsen (1966) contains many references
to special techniques.
6.1.6 Contents of Collections. The classical image of a systematic
collection is that of preserved specimens, either dried or submersed in a
preservative such as alcohol. Even though such specimens are the indispens¬
able basis of all taxonomic research, the information which one can derive
from dead, preserved specimens is limited. The modern systematist needs
a great deal of additional information, much of which he has to collect
himself while studying the living organism, either in its native environment
or in the laboratory.
As far as possible, the collecting should provide unbiased population
samples. No effort should be made to accumulate large numbers of “aberra¬
tions” or, in sexually dimorphic species, to concentrate entirely on the con¬
spicuous sex. Not only adults should be collected, but adequate samples
of all growth stages (including larvae) and associated parasites. Sampling
should be done in such a way as to provide study material not only for
the species describer but also for the evolutionist.
Collections of specimens may be augmented by collections of all sorts
of recordings. These include films of courtship displays and other aspects
of behavior; recordings of the vocalization of animals (sound libraries,
tapes, sound spectrograms) ; collections or photographs or casts of the work
of animals (nests, galls, spider webs, tracks, etc.). Collections of whole
animals must be supplemented by collections that permit histological, cyto-
logical (chromosomal), and biochemical research (see 7.4).
To obtain such additional material requires, as a minimum, not only
field work, but in many instances prolonged stays at a field station, particu¬
larly in the tropics, subtropics, or the Arctic. Museums increasingly include
aquaria, terraria, insectaries, aviaries, and the like among their facilities
in order to permit study and experimentation with living species. The mu¬
seum of today is fully aware that the study of the diversity of nature
requires a far broader approach than was envisioned by taxonomists of
bygone generations.
6.1.7 Preservation of Specimens. This differs from one taxonomic
group to the next. The basic rule is to preserve specimens in such a way
as to make them least subject to any kind of deterioration, whether through
106 Methods of Zoological Classification

the action of insect pests, mold, oxidation or bleaching by sunlight, drying

out, protein decay, etc. Again we must refer to the appropriate manuals
and handbooks for a description of specific methods (see above under 6.1.5).
Properly preserved specimens of some groups are still in satisfactory condi¬
tion 200 years after being collected. With more and more species becoming
extinct, the problem of “permanent” preservation is raised increasingly
often. Some recently suggested methods, such as embedding in plastics,
are too new to permit predictions. Preservation in alcohol raises problems
with sealing and appropriate containers, as discussed by Storey and Wil-
mowsky (1955) and Levi (1966).
6.1.8 Labeling. A specimen that is not accurately labeled is worthless
for most types of taxonomic research. So important, in fact, is the label
that it is sometimes stated jocularly that the label is more important than
the specimen. Many kinds of information are desirable, but by far the
most important single piece of information is the exact locality of collection.
In forms like certain land snails that may have racially distinct populations
as little as 0.5 km apart, the locality must be stated with great precision.
If the locality is a small community, farm, hill, creek, or other geographical
feature which cannot easily be found on commercial or geodetic (e.g.
United States topographic) maps, its position relative to a well-known
place should be added on the label (“15 mi. NW of Ann Arbor, Mich.”).
The county or district name should be given with all less well known
localities. If the specimen was collected in mountains, the altitude should
always be given, and if in the ocean, the depth. Additional ecological infor¬
mation is valuable or even essential in forms like plant-feeding insects or
host-specific parasites.
Whenever possible, the label should be written in the field at the
time the specimen is prepared. Any replacing of temporary labels with
later permanent ones is a potential source of error. However, this cannot
be avoided with insects when labels are printed for entire lots of specimens.
All essential data should be recorded on the original labels. Data recorded
in a field book are frequently overlooked and may be unavailable if the
collection is divided. The original label should never be replaced by a
museum label. A certain number of mistakes are always made in the trans¬
fer. If a museum label is desired, it should be added to the original label.
Labels attached to specimens preserved in alcohol or formalin must be
corrosion-proof. Any writing must be resistant to fading or washing out.
What data, in addition to locality, are needed depends on the given
group. Most good bird collectors, for instance, record on the label not
only locality, date, and collector’s name, but also the sex (based on autopsy),
the actual size of the gonads, the degree of ossification of the skull (impor¬
tant for age determination), the weight (in grams), and the colors of
the soft parts (Van Tyne, 1952).
 Taxonomic Collections and the Process of Identification 107

The little extra time required to make these records is more than
compensated for by the increased value of the specimens.

6.2 CURATING OF COLLECTIONS

Every taxonomist, sooner or later, is given the responsibility of curating

collections. This requires a great deal of expert knowledge, but also, more
importantly, a clear understanding of the function of collections, of the differ¬
ent kinds of collections needed in various areas of taxonomy, and of various
policies concerning the use of collections. The journal Curator publishes
much information on this subject.
6.2.1 Preparation of Material for Study. Bird and mammal skins
are ready for study as sent from the field by the collector. Mammal skulls
have to be cleaned. Some insects should never be placed in alcohol or
other liquid preservatives; others are useless when dried. Invertebrates that
are preserved in alcohol or formalin are usually ready for study as preserved.
Microscopic slide mounts or slides of part of the organs may have to be
prepared for the smaller forms. Instructions may be found in textbooks
of microscopic technique [Clarck (1961), Francon (1961), Gray (1954,
1958), Jones (1950), Lee (1950), Needham (1958)]. Most insects are
pinned, and the wings are spread if they are taxonomically important (or
beautiful), as in butterflies and moths and some grasshoppers (for refer¬
ences, see 6.1.5). Species can be identified in many groups of insects only
by a study of their genitalia. Microscopic slides or dry or liquid mounts
of the genitalia may have to be prepared. Special techniques are needed
for the preservation of protozoans (Corliss, 1963).
6.2.2 Housing. Research collections should be housed like research
libraries in fireproof buildings that are reasonably dustproof. More and
more museums keep their collections in air-conditioned buildings. Buildings
originally designed for public exhibits are usually quite unsuitable for the
housing of research collections. Rapid changes in temperature and humidity
are detrimental to museum cases and to specimens. Photographs and films
should be stored in air-conditioned rooms. Storage cases for specimens
should be built sufficiently well to be insect-proof and, ideally, also dust-
proof. Various firms now construct steel cases that satisfy all these require¬
ments. It must be remembered that insect-proof museum cases reduce the
labor of curating.
6.2.3 Cataloging. The method of cataloging depends on the group
of animals. With the higher vertebrates, where collections consist of a lim¬
ited number of specimens, it has been traditional to give each specimen
a separate number and to catalog it separately. This very time-consuming
procedure has been challenged, and it has been suggested that the methods
108 Methods of Zoological Classification

of entomology and malacology be adopted for vertebrates (see below).

All the specimens collected at a given locality or district or by one expedition
are entered in the catalog together. This greatly facilitates the subsequent
retrieval of distributional data and the preparation of faunisic analyses.
Cataloging is usually done after the specimens have been identified, at
least as far as the genus. This permits a permanent reference to the contents
of the collection long after it has been broken up and distributed in the
systematic collection or even dispersed to other institutions.
Catalog entries of vertebrates usually contain the following items:

1. Consecutive museum number

2. Original field number
3. Scientific name (or, at least, generic name)
4. Sex
5. Exact locality
6. Date of collecting
7. Name of collector
8. Remarks

In groups where the collections consist of large numbers of specimens,

as, for example, insect collections, where additions of 100,000 specimens
per year are not uncommon, it is customary to catalog accessions by lots,
each lot consisting of a set of specimens from a given locality or region.
Lot numbers, in turn, may refer to collector’s diaries or to other sources
of information on each collection. It is also customary to note whether
a lot was received as a gift or by purchase or exchange; the names of
the collector and donor are always given.
When museums and their collections were small, curators often had
rather elaborate card-filing systems permitting easy retrieval of all sorts
of information such as collecting station, name of the collector, etc. Main¬
tenance of such files is so expensive in clerical as well as curatorial time
that they have been abandoned in most museums where the emphasis is
on research rather than providing information to the public. A properly
organized and well-curated collection is a reference catalog in itself and
permits rapid information retrieval. Experiments are now being made in
some museums to place all information on each specimen on a separate
IBM card. Study of pilot projects will reveal whether or not this plan
is feasible (Squires, 1966). There is great danger that the preparation
of the cards will cost more in staff time than it will eventually repay.
Yet electronic data processing (EDP) is rapidly maturing, and taxonomists
and curators must keep up to date with its advances.
The maintenance of catalogs and card files is very time-consuming
and should never be carried to the point where it interferes with work
on the collections. A list of the accessions is indispensable, however, since
 Taxonomic Collections and the Process of Identification 109

it often allows the recording of additional information on localities which

cannot be entered in full on the specimen labels.
6.2.4 Arrangement of the Collection. As far as possible, the collec¬
tion should be arranged in the same sequence as some generally adopted
classification. The sequence of orders and families is reasonably standardized
in many classes of animals. Unidentified material, whenever it is not to
be worked out as a separate collection, is placed with the family or genus
to which it belongs (see later in this chapter, under identification). The
contents of trays and cases should be clearly indicated on the outside. The
bird collection of the American Museum of Natural History lists on
the tray labels the scientific names not only of all the available species
and subspecies but also of those still lacking in the collection (indicated
as such) (Fig. 6-1). The names on the collection cases and trays thus
serve as a checklist of all the known species and subspecies of birds. Of
course such a system is impossible in most groups, where the collections
are far too incomplete. Where specimens are of highly unequal size (e.g.
fish), storage in a strict taxonomic sequence is very wasteful of space.
Extralarge specimens may have to be stored separately.
6.2.5 Curating of Types. The names of species are based on type-
specimens (see type method, 13.48), which are thus official standards and,
being virtually irreplaceable, must be curated with special care. Whenever

Fig. 6-1. Method of storing study specimens of birds in open trays (collection case in the American
Museum of Natural History).
110 Methods of Zoological Classification

doubt arises as to the zoological identity of a nominal species, only reference

to the type can resolve the doubt.
Many descriptions of classical authors are equally applicable to several
related species. Early entomologists rarely, if ever, referred to the structural
detail of the genitalia, now so indispensable for diagnostic purposes in
most groups of insects. Only reference to the type can establish the basis
for the classical name. The curator is responsible for making every effort
to ensure the safety of these irreplaceable specimens. Types are customarily
deposited in large collections of public or private institutions which have
come to be recognized as standard type repositories.
When conducting an authoritative revision of a given genus, a special¬
ist should be able to see all the existing types. If many of them are in a
single institution, he should travel there for examination. He should obtain
scattered types by mail loan. Modem curators are quite liberal in lending
type specimens to qualified specialists. The number of recorded losses of
shipments is very' small, and if there is a real need for the replacement
of a lost type, a neotype can be designated, as permitted by the Code
under specified circumstances (13.55). Ideally types should be housed in
a separate collection in order to facilitate rapid removal in case of emer¬
gency and to avoid the constant handling in a general-study collection.
They should be clearly labeled in distinctive colors. If not previously cata¬
loged, they should be numbered individually to facilitate referring to them
in the literature and finding them in the collection. Since many types are
those of synonyms, a card index by genera and another by species will
save much time in locating the desired type. It is not economical to include
in type collections all sorts of pseudotypes—that is, specimens that are
not name bearers, in other words not holotypes, lectotypes, neotypes, or
syntypes (13.48).
There is much to recommend the arrangement of type-collections
alphabetically according to the given specific name. A type-collection is
a reference collection (rather than a classification), and ease of reference
should determine what system is adopted. When number systems are
adopted, too many errors are usually committed for them to be practical.
Type-specimens assume such an important role in the taxonomy of
lesser-known groups that many workers believe no one has the right, ethi¬
cally, to retain a type in a private collection after study has been completed.
Specialists sometimes donate their collections to a public institute but retain
them on a permanent loan up to the end of their active research period.
This includes the obligation to send out specimens on loan if such requests
are made by cospecialists.
6.2.6 Exchange of Material. In the days when taxonomists considered
most of their specimens “duplicates,” exchanges were very popular. How¬
ever, selecting material for exchanges and keeping records required by ex-
 Taxonomic Collections and the Process of Identification 111

changes is very time-consuming, and except among private collectors ex¬

changes are not as popular as they used to be. A specialist doing a mono¬
graph on a certain genus or family can always borrow material from other
institutions and return it when he has completed his studies. Exchanges
are most desirable in groups where series of unlimited size can be obtained
and where the exchanges concern areas not readily accessible to the respec¬
tive institutions (e.g. intercontinental exchanges). Exchanges are least de¬
fensible when they lead to the dispersal of biologically important population
samples. Exchanges are sometimes necessary to build up complete identifica¬
tion collections. When an exchange is carried out, generosity is always
the best policy. It is generally not advisable to insist upon exchanging
specimen for specimen, except where institutional policies or other unusual
factors demand it. Many specialists give away excess specimens of a large
series as “open exchanges,” not necessarily expecting any return.
6.2.7 Expendable Material. Improperly preserved or inadequately
labeled specimens are usually valueless. Collections would be better of!
without them. Yet they are much less of a burden on a collection than
would be the task of the curator to find and eliminate them. He must
make sure with every specimen that it is not an unlabeled type or a specimen
which, in one way or another, is unique or of historical value. The most
efficient method for the elimination of useless material is to ask specialists
to pull out such specimens when scrutinizing the material during a revision.
6.2.8 Loans. Modern curators are very generous in lending speci¬
mens to qualified experts. The axiom that systematic collections are the
general property of science, not of a specific institution or curator, is being
acknowledged more and more widely. Every loan, however, involves loss
of time and effort, and the borrower should refund the lender for this
outlay. Research grants now often include an item for such purposes, to
cover not only the costs of postage but also of selecting the specimens,
recording the loan, and getting the material packed for shipment. The
modern curator, being essentially a research worker, must delegate these
tasks to hired clerical help.
The borrower also has well-acknowledged obligations. Requests for
loan of specimens should be as specific as possible, including a statement
of the reason for the request and some indication as to the length of time
for which the material is needed. The beginner may be unable to borrow
certain material except through a loan to his institution, or a well-known
colleague, or his major professor. In such cases any laxity in carrying out
the conditions of the loan reflects not only on himself but also on the
sponsoring individual or institution. If it should develop that the borrower
is unable to complete his studies in the time designated, he should inform
the party or institution which made the loan, without waiting and placing
them in the embarrassing position of having to write and ask about the
112 Methods of Zoological Classification

status of the study. The borrower should not ask for material that he
does not actually need, nor should he ask for material that he could easily
study by traveling to the museum.
If a specialist has agreed to identify a collection provided he receives
certain specimens, he should make sure that the terms of the agreement
are well understood and should return to the lender a list of the specimens
which he has retained. All types and unique specimens must be returned
to the lender in such cases. In the case of anatomical material, it is under¬
stood that dissection, the purpose for which the anatomical specimen was
originally collected, will partially or entirely destroy the specimen. The
borrower, in this case, is under obligation to preserve a pictorial record
of the dissection. To cut down on loans, institutions sometimes make tem¬
porary or permanent transfers of collections. For instance, an inland mu¬
seum, which has a small collection of marine invertebrates not used for
exhibition or instruction may transfer it to a large museum that is active
in marine research; but excessive consolidation of taxonomic collections
would create not only monopolies but also an inherent danger. The concen¬
tration of so much irreplaceable material in a single institution makes it
exceedingly vulnerable to destruction in the event of a catastrophe. Long¬
term loans to leading specialists are in most cases a better solution.

6.3 IDENTIFICATION

Under the word identification a number of very different activities

are usually combined. They all involve the identification of previously un¬
identified material, but for very different purposes. They will be discussed
separately here. [See also 1.2 and 4.3.1 for a definition of identification
(in contrast to classification).]
6.3.1 Sorting of Collections. All material gathered on collecting trips
and expeditions must be sorted and at least tentatively identified before
it can be incorporated into a collection. The first rough sorting of freshly
collected material is often done in the field. An entomologist may keep
specimens of different species from different hosts in separate containers.
Collections made by oceanographic expeditions are often at once roughly
sorted, in part because different kinds of animals may require different
methods of preservation. After the specimens are properly preserved and
labeled, the usual practice is to segregate unstudied material down to orders
and, whenever possible, to families or even genera. Such material is then
available to the specialist, who can undertake the precise identification.
Even such a tentative identification requires skill and experience. When
large collections are involved, the establishment of a special sorting organiza¬
tion is advisable. The Smithsonian Institution in Washington maintains
 Taxonomic Collections and the Process of Identification 113

the Oceanographic Sorting Center (SOSC) charged with the preliminary

sorting of the material gathered by the Indian Ocean Expedition and other
oceanographic expeditions. When identified as to family or genus, the mate¬
rial is shipped to those specialists, all over the world, who have agreed
to study it.
The ease of sorting newly received collections depends on the nature
of the material (larvae or adults, microscopic in size or not) and on the
maturity of the taxonomy of a given group and in a given region. In
birds, for instance, identification at the species level is virtually never a
problem, although subspecific identification may be difficult. In less well
known groups for which no recent revisions, keys, or manuals are available,
definite identification may be possible only down to the level of the family
or at best to that of the genus.
Linnaeus thought that every zoologist should know every genus (only
312 genera of animals were known to him). In this spirit, at a time when
the collections and the number of described species were still small, tax¬
onomists attempted to identify every specimen, even when it belonged to
groups of which they had no special knowledge. It is now realized that
this is a very wasteful approach. When accessions are sorted in a modem
zoological museum, they are identified only to the level (order or family)
where they become available to the specialist. It is far more economical
and important for a taxonomist to devote his time to the preparation of
new monographs and keys than to attempt identification down to the species
level in groups with which he is not familiar.
Unfortunately, identifications by nonspecialists are often erroneous.
Most large collections contain numerous examples of misdeterminations.
The original specimen in a series may have been quite authentic in such
cases, but other specimens were subsequently added without critical analysis
and without determination labels by experts. Such misdeterminations are
more bothersome than leaving the material unidentified until it can be
studied by an expert.
6.3.2 Determination Labels. Sooner or later all material in collec¬
tions is seen by a qualified taxonomist or specialist who is able to identify
it down to the species. Each specimen or each series should be labeled
at the time such identification is made. The determination label should
give the scientific (generic and specific) name and author, and in addition
the name of the determiner and the year in which the identification was
made. With this information on every specimen, the authenticity of the
determination is established, and its dependability may be readily evaluated
at any subsequent date on the basis of progress which may have been
made in the study of the group during the intervening years. In bird and
mammal collections these names are usually written in pencil so that they
can be changed easily if there should be a change of nomenclature.
114 Methods of Zoological Classification

6.3.3 Identification of Individual Specimens. The taxonomist is fre¬

quently called upon to identify a particular specimen or species. If such
identification is highly important for a special research project in applied
biology or in experimental zoology, a taxonomist will gladly make every
effort to identify the species on which this research is based. On the other
hand, it is not the job of the taxonomist to undertake the routine identifica¬
tion of ecological collections or of archeological material. Such identification
work is the responsibility of the ecologist or archeologist who wants his
material identified.
There are, however, federal and state agencies charged with the re¬
sponsibility of identifying economically important animals. These agencies
employ specialists, to each of whom a particular group is assigned. The
research taxonomist, on the other hand, does not have any responsibility
whatsoever for routine identifications. Nothing is more detrimental to the
productivity of a research museum than unreasonable identification de¬
mands by the public.
6.3.4 Process of Identification. Even a rank beginner trying to iden¬
tify a specimen can usually tell that it is a bird, a spider, a grasshopper,
or a butterfly. He can immediately go to keys and manuals specializing
in these zoological groups. The real beginner, however, will have trouble
with all but the most common kinds of animals. When in doubt about
the order to which an animal belongs he should try the simple keys given
in general textbooks and handbooks. Even the advanced student may en¬
counter unusual species or immature or exotic forms which cannot be placed
on sight in the proper family or order. However, modern works are generally
available which provide family and subfamily keys and greatly assist in
this stage of identification.
Driver’s Name That Animal (1950) is a good elementary guide to
the principal groups of animals. In addition to the general keys, a bibliog¬
raphy of the most important works on each group of animals is given.
For example, Brues, Melander, and Carpenter (1954) is cited among the
general works on insects; this book in turn gives keys to the orders, families,
and subfamilies of insects and cites the more important monographs under
each group. For North American freshwater animals, consult Pennak
(1953) or Edmondson (1959); for marine animals, Light (1954) for the
Pacific and Smith (1964) for the Atlantic. There are several series of
very useful field guides now available, such as Putnam’s Nature Field Books,
the Peterson Field Guide series, the H. E. Jaques Pictured Key Nature
Series (William C. Brown Company), the Doubleday Nature Guides Series
and the Golden Field Guides. Other useful publications are those of Pratt
(1935, invertebrates), Baker (1958, mites). Bishop (1952, salamanders),
Stebbins (1954, reptiles and amphibians), Wright and Wright (1949, frogs),
Smith (1946, lizards), Wright and Wright (1957, snakes).
 Taxonomic Collections and the Process of Identification 115

There are many works on the animals of specific regions [e.g., Park,
Allee, and Shelford (1939) for the Chicago region, or Eddy and Elodson
(1955) for the North Central states]. There is an excellent survey of tax¬
onomic works dealing with the British Isles (Smart, 1953). Other regional
works for Europe are Die Tierwelt Deutschlands (Dahl, 1925 et seq.),
Faune de France (1921-1950 and continuing), and Die Tierwelt der Nord-
und Ostsee (Grimpe and Wagler, 1925 et seq.). Unfortunately there are
no comprehensive bibliographies of regional taxonomic treatises available
for the United States.
Identification is vastly more difficult when no convenient keys and
manuals are available. The beginner is advised not to attempt it. If there
is available a monograph or technical revision of recent date (see below
how to find such literature), the specimen is run through the keys; the
description of the appropriate species is checked, character by character;
the specimen is compared with any illustration that may be given; and
the recorded geographical distribution is checked. If all these points agree,
the identification is considered as tentatively made, subject to comparison
with authentic specimens and provided that no additional related species
have subsequently been described. For further details on these steps, see
the following sections. When there is not even a recent monograph or
revision available, no one but a specialist should attempt a determination.
Even he will not waste time trying to identify single specimens except
under the exceptional conditions specified in 6.3.3.

6.4 MATERIAL FOR REVISIONARY OR MONOGRAPHIC WORK

The major preoccupation of the taxonomist, except perhaps of the

specialist in the best-known groups of animals, is the preparation of revi¬
sions. The early part of the process, the determination of the species to
be classified, resembles in some ways the process of identification, yet it
is of a somewhat different nature. The details of this procedure are de¬
scribed in Chaps. 9 and 10.
However, before the actual work of revising can be started, the tax¬
onomist must gather the needed specimens and the literature. Let us assume
he wants to revise a certain tribe of beetles from South America. After
he has completed a first examination of the material available in his own
collection, he will write to the major museums whose scope includes the
Coleoptera of South America and ask for the loan of their material. He
should carefully observe the advice given in 6.2.8. During such correspon¬
dence he may discover that some other taxonomist has also started to revise
this group; he will then have to negotiate with this specialist how to divide
the task. In the case of large collections rich in types, he will have to
116 Methods of Zoological Classification

make arrangements to visit these museums rather than ask for the loan
of the material.
Far more difficult than tracing the available material is usually the
tracing of the literature (Bottle and Wyatt, 1966). The zoologist may start
with some general zoological bibliography such as Smith and Painter, Guide
to the Literature of the Zoological Sciences (7th ed., 1967) and may consult
various annual bibliographies and literature-abstracting services (see below).
When there is no recent monograph or revision, the most recent cata¬
log for the group should be consulted. The catalog (11A.9) will give litera¬
ture citations leading to the descriptions of all species known up to the
time of its completion. Some catalogs furnish even more information, e.g.,
complete bibliographies under each genus and species, lists of synonyms,
and geographical distribution. Taxonomic research is greatly facilitated by
a good catalog, because it brings together the most significant published
references to the group.
6.4.1 Reference to Current Bibliographies. Catalogs are inevitably
out of date soon after they are published. This difficulty may be partially
compensated for by the issuance of supplements. Nevertheless, it is not
at all unusual to find that even the most recent catalog is twenty years
old. In some of the major insect orders no general catalog has been prepared
since 1900, and some groups have never been cataloged from a world
standpoint.
Fortunately, there exists an unusual bibliography of the literature
in systematic zoology, a great reference work entitled The Zoological Record.
It is indispensable for taxonomic work. The Zoological Record has appeared
every year from 1864 up to the present time. Each new scientific name
is given, together with a reference to the place of publication and the
type locality. The names are arranged alphabetically under families, but
a systematic arrangement is followed for families and higher groups. Current
numbers are available separately by purchase or subscription.
The Zoological Record is published by the Zoological Society of Lon¬
don in cooperation with the British Museum (Natural History) and the
Commonwealth Institute of Entomology. The following 20 sections of The
Zoological Record are published separately and may be obtained singly
or as an entire volume each year: (1) Comprehensive Zoology, (2) Pro¬
tozoa, (3) Porifera, (4) Coelenterata, (5) Echinodermata, (6) Vermes,
(7) Brachiopoda, (8) Bryozoa, (9) Mollusca, (10) Crustacea, (11) Tri-
lobita, (12) Arachnida1 and Myriapoda, (12) Insecta, (14) Protochor¬
data,2 (15) Pisces, (16) Amphibia, (17) Reptilia, (18) Aves, (19)

1 To Arachnida are added Merostomata, Pantopoda, Pentastomida, Tardigrada, Myriapoda, and

Onychophora.
2 Protochordata together with Pogonophora, Enteropneusta, Graptolithina, Pterobranchia, and
Phoronidea.
 Taxonomic Collections and the Process of Identification 117

Mammalia, (20) List of New Generic and Subgeneric Names. These

are ordered from The Zoological Society of London, Regent’s Park, London,
N.W. 1, England (except for Insecta, to be ordered from the Common¬
wealth Institute of Entomology, 56 Queens Gate, London, S.W. 7).
The commonest method of using The Zoological Record is to start
with the most recent volume and work back to the date of completion
of the most recent catalog or revision. The particular genus or other group
in question may be located in the table of contents of the section devoted
to the particular class of animals involved. New names, synonymies, distribu¬
tions, and in some instances even biological references are given. If the
citation is not clear because of its abbreviated form, or if the exact title
of the publication is important, reference may be made to the bibliography
of papers arranged according to authors at the beginning of the section.
For special needs, there is an elaborate subject index covering various phases
of morphology, physiology, ecology, and biology.
Some groups of animals have never been cataloged or monographed.
This is especially true of certain groups of insects. In such cases, it is
necessary to work through the entire Zoological Record back to volume 1
(1864).
The best annual review of the taxonomic literature, prior to 1864,
is found in the Berichte iiber die wissenschaftlichen Leistungen in different
branches of zoology, including entomology and helminthology, published
in Wiegmann’s Archiv fur Naturgeschichte (Berlin, 1835 et seq.). Addi¬
tional important bibliographic aids covering this early period of zoology
are Engelmann (1846), Agassiz and Strickland (1848), and the catalog
of scientific papers published by the Royal Society (1800-1863). Sherborn’s
Index Animalium (1758-1800, 1801-1850) gives a complete list of generic
and specific names proposed up to 1850 and Neave’s Nomenclator Zoologi-
cus (1939-1940, 1950, 1966) lists all generic names for the period 1758
to 1955.
The Zoological Record is always one or two years behind, so that
other bibliographies must be consulted for the most recent literature. Bio¬
logical Abstracts (1926 to date) is an important source of recent literature.
Its section on systematic zoology contains abstracts of taxonomic papers,
hence is a valuable source of information for papers which are not imme¬
diately available elsewhere. However, since Biological Abstracts covers the
taxonomic field very incompletely, it is no substitute for the Zoological
Record. There are important foreign abstracting journals which must be
consulted. The Berichte iiber die gesamte Biologie, Abt. A, Berichte iiber
die wissenschaftliche Biologie (1926 to date), and Bulletin signaletique 16,
Biologie et physiologie, Animates (1962), are particularly important. In
addition, Gunther’s reviews of the systematic literature (1956, 1962) are
invaluable.
118 Methods of Zoological Classification

There are numerous zoological bibliographies dealing with zoology

as a whole, or with vertebrate zoology (Wood, 1931), or with special groups,
such as birds, fishes, or other taxonomic subdivisions. The student of tax¬
onomy is advised to familiarize himself with the bibliographic aids that
are available in his special field. Experienced specialists will be glad to
give him advice.
6.4.2 Nominal Species and Zoological Species. Not all names found
in the literature (“nominal species”) represent actually different zoological
species. In many cases individual variants of all sorts were erroneously
described as separate species. Chapters 8 and 9 give instructions on how
to determine what names refer to valid species and what others to intra¬
specific phena.
Equally troublesome are cases where similar (or not even very simi¬
lar!) species masquerade under the same name. The correct determination
of the actual zoological species in the group to be revised is the most
important basic step in taxonomic research. This requires consultation of
original descriptions (or improved redescriptions), a study of authentic
specimens, and, in any authoritative revision or monograph, a study of
the actual type specimens.
6.4.3 Original Description. Although the secondary literature is
often a great help, reference should always be made to original or more
recent authoritative descriptions. Unless this is done, there is a possibility
that the species was misidentified in the secondary literature. Original de¬
scriptions are located by means of catalogs, monographs, the Zoological
Record, or other bibliographical sources, as described above.
Copies of original descriptions may be difficult to find. Even the largest
libraries are not complete, and the average university library will often
be found wanting. This is not so much a reflection on the caliber of libraries
as it is evidence of the extent and diversity of scientific publications through¬
out the world. Although largely confined to a half-dozen languages, taxono¬
mic papers are published in practically every country in the world. This
poses a very real problem for libraries with limited budgets. The situation
is further complicated because priority places a premium on the earlier
works. No taxonomic work published since 1758 becomes “out of date”
if it contains new names, and as a result of limited editions, losses through
the years, and other factors, there are not enough copies available to supply
all biological libraries. To an increasing extent, specialists make photocopies
of those pages of periodicals which they have to consult most frequently.
As long as this is not done commercially, it is not likely to infringe on
copyright (but see Science, vol. 152, pp. 291-292).
The search for original descriptions is often involved. Helpful, if not
essential, to the searcher are: full use of the familiarity with all locally
available scientific libraries; reference to the Union List of Serials to locate
 Taxonomic Collections and the Process of Identification 119

publications in other libraries for interlibrary loan; extensive use of micro¬

film services and other copying devices; and accumulation of reprints, by
purchase or by exchange with other workers, and of photostats (Xerox
copies).
Descriptions are the foundation of taxonomy, since only the printed
word is indestructible. Types may be lost, and the original author is available
for only a brief span of years to pass on “his” species.
The original description should be read several times, first to obtain
a general impression or mental picture of the actual specimen which the
original author had before him. Then, particular characters which the origi¬
nal or subsequent authors considered important should be extracted and
checked against the specimens in question. Finally, any comparative notes
given by the original author should be checked. In many cases such com¬
parative characters are the most useful clues to identification.
Original descriptions are normally the court of last appeal for purposes
of general identification. However, many of them are totally inadequate,
particularly those published prior to 1800. The value of a description is
in direct proportion to the judgment of the author and his ability to select
significant characters and describe them in words, also to the extent and
nature of the material available to him at the time of description. For
these reasons descriptions given in a thorough and authoritative monograph
of recent date are usually more usable than original descriptions.
Illustrations are often as valuable as, or more valuable than, original
descriptions, particularly when there are language difficulties. In popular
groups, such as birds or butterflies, there are many works with colored
plates. Such works are often a great help in the rapid identification of
specimens. Colored plates are not always well reproduced, however, and
there are many opportunities for error if too much dependence is placed
upon them.
If the original description is accompanied by an illustration, the diffi¬
culty occasionally arises that characters of illustration and description are
in conflict. It can sometimes be proved in such cases that the artist did
not have access to the type specimen and used another in the belief that
it agreed with the type. Such discrepancies are not infrequent in the works
of early authors.
6.4.4 Comparison with Authentically Determined Specimens. It is
sometimes impossible to make a satisfactory determination from the litera¬
ture alone. Such a situation exists if the group has been neglected and
the descriptions are poor. Even under ideal conditions identification is
greatly facilitated if “authentic material” is available for comparison—mate¬
rial named by a specialist and preferably compared by him with the
holotype. Paratypes do not always necessarily qualify, because series of syn-
types are sometimes composites of several species.
120 Methods of Zoological Classification

Comparison with specimens is a highly technical task requiring a

considerable background of knowledge of and preparation in the particular
group in question. For this reason preliminary identification made by direct
comparisons with authentic collections, without first studying the literature
and the significant characters of the group, is often valueless.
Reference collections are often accumulated for the express purpose
of facilitating identification. In such cases comparison is made with what¬
ever series of specimens is available, and one must judge whether the speci¬
men in hand falls within the possible range of variation of a given species.
Care should be taken not to rely exclusively on comparison with sup¬
posedly authentic specimens. Even “authoritative” collections may contain
wrong identifications, or may be incomplete. In such cases hasty comparison
made without following the other steps in identification may lead to er¬
roneous conclusions.
6.4.5 Comparison with the Type. Type-specimens are the most au¬
thentic of all but are much too valuable to be used for routine identifica¬
tions. Ideally, in the course of a monographic study of a group all type-speci¬
mens should be reexamined. At this time the significant characters are
usually known and can be checked using the same technique and the same
interpretation of the characters as are applied to the rest of the material.
In work with subspecies it is not always necessary to have type-speci¬
mens for comparison (if there is no question as to the identity of the
species). On the other hand, a series of specimens from the type-locality
(“topotypical specimens”) is desirable to provide information on the char¬
acters and variability of the subspecies.
Chapter Taxonomic Characters

B iological classification consists of the assembling

organisms into groups that are similar as a result of their common descent.
of

One of the main operations of taxonomy is thus the determination of this

special kind of similarity. This, as the history of taxonomy shows, is a difficult
and highly controversial procedure (10.1). The experienced taxonomist
compares two kinds of organisms and, merely by inspection, integrates a
large number of attributes into a “similarity value.” His judgment is based
on years of analysis, comparison, and weighting of so-called taxonomic
characters.

7.1 NATURE OF TAXONOMIC CHARACTERS

Definition. A taxonomic character is any attribute of a member of

a taxon by which it differs or may differ from a member of a different
taxon.
This definition implies a number of important points. A characteristic
by which members of two taxa agree but differ from members of a third
taxon is a taxonomic character (a statement true for the characters of
all higher taxa). To define taxonomic character as “any attribute of an
organism” would not be correct. Features by which individuals of the same

121
122 Methods of Zoological Classification

population differ from each other, such as differences between the sexes
and age classes, are not taxonomic characters. However, when populations
(taxa) differ from each other by the presence or absence of sexual dimor¬
phism or in larval characteristics or in any other manifestation of sex or
age, then such a difference becomes a taxonomic character. Taxonomic
characters are population characteristics. The comparison of populations
and taxa is the standard method for the study of taxonomic characters,
and any attribute qualifies if it is established as different through such
comparison.
“Taxonomic character” means actual or potential taxonomic differ¬
ence, such as “eyes red” versus “eyes white.” The definition of taxonomic
character as that by which taxon A differs from taxon B has been universally
accepted in the taxonomic literature of the last 200 years. The computer
taxonomists, perhaps not fully aware of the previous definition, have sug¬
gested changing this terminology. They would call “eyes” the character,
and “red” or “white” character states. However, “red eyes,” “spiny legs,”
etc. are what the literature has universally called taxonomic characters,
and it is the feature (eyes, legs, etc.) which displays this taxonomic variation
for which a new term needs to be found. To transfer the term charac¬
ter from its traditional usage to a new one is bound to prove confusing.
Taxonomic characters have a double function:

1. They have a diagnostic aspect uniquely specifying a given taxon; an

emphasis on the differentiating properties of taxa is particularly strong
at the level of the lower categories.
2. They function as indicators of relationship; this property makes them
especially useful in the study of the higher taxa.

Not every publication in zoology deals with taxonomic characters.

Work published on species that are expecially suited for experimental re¬
search, such as Canis lupus, Macaca rhesus, Rattus norvegicus, Mus
musculus, Drosophila melanogaster, or Tribolium confusum, deal with gen¬
eral biological phenomena, such as the behavior of genes, the chemistry
of enzymes, the physiology of muscle action, or the nature of learning.
Most of the results of such studies are of wide applicability. They are
valid for all mammals, all terrestrial warm-blooded vertebrates, all meta¬
zoans, or all life. Literature of this type of research is of taxonomic interest
only to the extent that comparison between different taxa reveals differ¬
ences. But even eliminating such literature, there are at least 20,000, but
more likely about 50,000 papers published annually which present evidence
on taxonomic characters and in part discuss them quite fully. It is impossible
in this chapter to present more than the barest outline of this important
topic. Chapters 3 to 5 of Animal Species and Evolution (Mayr, 1963,
pp. 31-109) deal with certain kinds of taxonomic characters, particularly
 Taxonomic Characters 123

characters such as isolating mechanisms and differences in niche utilization,

which permit the sympatry of closely related species.

7.2 CHARACTERS AND CLASSIFICATION

Taxonomic practice reveals that not all taxonomic characters are

equally useful. Some are powerful indicators of relationship, others are
not. The usefulness of a character depends on its information content,
that is, on its correlation with the natural grouping of taxa produced by
evolution. The taxonomist, either consciously or unconsciously, assigns a
certain weight to each character according to its “goodness,” that is, accord¬
ing to its ability to reveal relationship. The problem of weighting will
be dealt with more fully in Chap. 10 (10.4).
Some discussion has taken place in recent years whether to define
taxonomic characters in terms of the phenotype or of the underlying genetic
program. Some authors felt that the latter would be “more scientific.”
There is considerable doubt as to the validity of this claim. The pathway
from the gene to the phenotype is a long one in higher organisms and
the “one gene, one character” hypothesis is now thoroughly discredited.
What the taxonomist deals with, his taxonomic characters, are components
of the phenotype and have almost invariably a highly complex genetic
basis. The genotype of most taxonomic characters is now, and probably
will remain so for a long time, refractory to precise genetic analysis. On
the other hand, a number of recent studies on gene-controlled enzymatic
intrapopulation polymorphism indicate that much of the variation of the
genotype is a poor source for taxonomic characters. It is thus evident that
the direct study of the genetic basis of taxonomic characters is either
technically impossible or else not very helpful.
Unfortunately, it is likewise impossible to take the needed information
directly from the phenotype. Size illustrates the danger of an unevaluated
use of the phenotype particularly well. A large-sized species may differ
from a small-sized species in every measurable character, hence in literally
thousands of phenotypic characters. There is, however, almost total redun¬
dancy, since the size difference might be caused by a single genotypic factor.
The literature abounds in cases where such redundancy was not eliminated
in the phenetic character listing, so that an absurdly distorted classification
resulted. A single adaptation (wing diving or seed cracking in birds; see
below) may lead to an impressive number of changes in the overall pheno¬
type, most of which are redundant as far as information on relationship
is concerned (see also “redundant characters” in 10.4.3).
Until he has a proper understanding of the value of various potential
characters, a taxonomist must consider a large number of characters and
124 Methods of Zoological Classification

compromise between using too many and using too few. Considering too
many characters is uneconomical because it requires too much time: it
introduces redundancy because different sets of characters often lead to
essentially the same answer. Giving equal weight to many characters is
sometimes actually misleading because it may conceal the effects of special
adaptations, parallelism, and convergence. For advice on the taxonomic
weight of single characters see 10.2.3.
Unreliable Characters. Every specialist knows characters which in
his group are “unreliable,” that is, poor indicators of relationship; such
characters are said to have low weight (10.4.3). All highly variable charac¬
ters are in this category. The pattern of branching in arteries of vertebrates
may be different not only in different individuals of the same population
but even in the left and the right side of the body. Differences in arterial
pattern are not nearly as helpful for classification as some authors thought
they were. Wing venation provides important characters for the classification
of insects. However, Sotavalta (1964) showed that there was far more
variation in this character in the tiger moths (Arctiinae) than known
and that the traditional generic arrangement of the family, based on wing
venation, is thoroughly in need of revision. For a further discussion of
variability of characters see 10.4.3.
Any kind of regressive character is usually of low taxonomic weight.
Taxa based on the loss of eyes, wings, toes or other appendages, or certain
teeth, etc., are often unnatural (10.4.3).
In order to get away from unreliable single characters taxonomists
in recent years have been searching for an “overall character” which would
represent a single denominator for the taxonomic position of a taxon. Phe-
netic distance was believed by the pheneticists to be such an indicator
(10.2). The serologist thought that protein interaction was such an overall
measure, while DNA matching is the most recent candidate for such a
measure (7.4.9). All these methods are helpful, but since all have failed
on occasion (sometimes frequently) they cannot be considered to be the
hoped-for panacea.
Dual Function of Characters in Classification and Identification.
Characters serve different functions in different operations of taxonomy,
as discussed in Chap. 4. They are used during the analytical phase to
determine the units of classification (species and other lower taxa) and
during the synthetic phase to help in the correct delimitation and ranking
of higher taxa. The same characters, but usually only a selected small
number of them, are used in a very different operation, namely that of
identification. Key characters are characters that are easily perceived, of
very low variability, usually present in preserved material, and useful as
convenient labels for the taxa distinguished by the process of classification.
Many taxonomic characters, such as chemical, chromosomal, physiological,
 Taxonomic Characters 125

and behavioral ones, may have high value for purposes of classification
but are poor or useless for identification because they are inaccessible in
preserved material or because their determination requires difficult tech¬
niques. Chapter 11 contains a discussion of characters most useful in keys.

7.3 TAXONOMIC CHARACTERS AND ADAPTATION

Formerly taxonomists tended to consider characters as if they were

specially created for their convenience. Now we realize that as phenotypic
manifestations of the genotype taxonomic characters are the product of
a long history of natural selection.
Nature of Adaptation. The taxonomic weight of a character depends
to a large extent on the nature of its adaptive significance. The following
comments might be helpful to those undertaking such evaluation. For a
more detailed discussion, see Chap. 10.

1. Adaptation to the General Environment. A given type of adaptation

may be very widespread, yet the specific mode in which it is expressed may
be of high taxonomic significance. For instance, the pelagic larvae of most marine
invertebrates have ciliary bands, yet the particular configuration of the bands
is characteristic for classes and sometimes even phyla. All pelagic organisms have
floating devices, yet the particular manner in which floating is achieved differs
from taxon to taxon. Pigments, poisons, and other chemical constituents of the
body may by convergence have become similar in appearance or effect, yet a
proper analysis almost invariably shows highly specific correlation with taxonomic
groups.
2. Specific Adaptations. Most adaptations for special niches are far less
revealing taxonomically than they are conspicuous. Typical examples of this are
substrate adaptations, such as white coloration in arctic mammals and birds, sandy
coloration of desert animals, the bright coloration of tropical treetop birds, warning
colorations, and various types of Mullerian mimicry.
Occupation of a special food niche and the correlated adaptations have
a particularly low taxonomic value. The bill in birds and the teeth in cichlid
fish may be very different in closely related species, in response to a shift in
niche occupation. See 10.4.2 and 10.4.3.
3. Isolating Mechanisms. Any species-specific character that serves as an
isolating mechanism may be reinforced by natural selection and become exceed¬
ingly conspicuous. In birds this is true particularly for species without pair forma¬
tion (Mayr, 1942, Sibley, 1957). However, such characters are of low weight
for classification on the generic and higher levels.
4. Competitive Character Divergence. Some differences, particularly in niche
utilization, between closely related sympatric species are a consequence of competi¬
tive exclusion (Mayr, 1963, chap. 4). A process which Brown and Wilson (1956)
called character displacement may lead to a stronger morphological divergence
between such species than would be expected on the basis of relationship.
126 Methods of Zoological Classification

There is always a residue of taxonomic characters of which the func¬

tional and hence selective significance is not evident. To determine whether
such characters have an unknown significance or are incidental phenotypic
by-products of a selected genotype must remain for later research. One
can summarize the relation between adaptation and the value of taxonomic
characters by saying that narrowly adaptive characters generally do not
give as much information on relationship as do characters that owe their
conformation to the constitution of the overall genotype, which selection
has shaped for thousands and millions of generations.

7.4 KINDS OF CHARACTER

Almost any attribute of an organism might be useful as a taxonomic

character, if it differs from the equivalent feature in members of another
taxon. However, proper classifying work is possible only when adequate
material of many species is simultaneously available for comparison. Mu¬
seums provide this opportunity and this is the reason why the taxonomist
prefers characters that can be easily observed in preserved specimens (e.g.
morphological characters).
In each group of organisms, whether they are birds, butterflies, sea
urchins or snails, different taxonomic characters exist. It is part of the
training of the taxonomist to become familiar with the characters that
are most useful in the particular taxon in which he plans to specialize.
Monographs and handbooks usually give detailed descriptions of the charac¬
ters used. In his revision of the North African scorpions, for instance,
Vachon (1952) devotes 27 pages to a detailed description and illustration
of the taxonomic characters of that group.
Morphological Characters. Features of the external morphology vary
according to kind of animals. They range from such superficial features
as plumage and pelage characters of birds and mammals through scale
counts of fishes and reptiles, to the highly conservative and phylogenetically
significant sutures and sclerites of the arthropod body. The internal anatomy
provides an abundant source of taxonomic characters in practically all
groups of higher animals. The extent to which such characters are used
routinely varies from group to group, generally in inverse ratio to the abun¬
dance and usefulness of easily observed external characters. In mammals
the skull (with teeth) is routinely preserved and used in classification,
while reptiles, amphibians, and fish are normally preserved in alcohol and
are always available for dissection. On the whole, aspects of the internal
anatomy more often supply characters for the classification of the higher
taxa than for discrimination at the species level. Fossils consist almost en-
 Taxonomic Characters 127

Table 7-1. Kinds of Taxonomic Characters

1. Morphological characters
a. General external morphology
b. Special structures (e.g., genitalia)
c. Internal morphology (= anatomy)
d. Embryology
e. Karyology (and other cytological differences)
2. Physiological characters
a. Metabolic factors
b. Serological, protein, and other biochemical differences
c. Body secretions
d. Genic sterility factors
3. Ecological characters
a. Habitats and hosts
b. Food
c. Seasonal variations
d. Parasites
e. Host reactions
4. Ethological characters
a. Courtship and other ethological isolating mechanisms
b. Other behavior patterns
5. Geographical characters
a. General biogeographical distribution patterns
b. Sympatric-allopatric relationship of populations

tirely of preserved hard parts—in the case of Mesozoic mammals, for in¬
stance, largely of teeth.
Even in this traditional area great advances have been made in recent
decades. Descriptions have become more detailed and better standardized.
The taking of measurements is being automated (Garn and Helmrich,
1967). In the case of lower invertebrates careful microscopic analysis has
revealed an abundance of characters even in such seemingly nondescript
forms as nematodes. The development of new silver impregnation tech¬
niques has revealed a wealth of characters even among protozoans, particu¬
larly ciliates.
New organs and structures are steadily added to those that show
taxonomically important differences. The spermatozoa of many taxa, for
instance, have a highly peculiar and specific morphology and may serve
as useful indicators of relationship.
7.4.1 Hard Parts (shells, external skeletons, etc.) and the Work
of Animals (mines, tracks, etc.). It would be senseless to worry whether
to consider hard parts, etc., as morphological, physiological, or behavioral
characters. Much of the classification of the invertebrates is based on char¬
acters of exoskeletons and shells. Likewise, among the protozoans tests,
128 Methods of Zoological Classification

shells, thecal plates, cysts, and other hard parts are vital in the classification
of foraminiferans, radiolarians, testaceous rhizopods, flagellates, and others.
The orientation types of calcite crystals in the skeleton of echinoderms
agrees well with their classification in families and orders (Raup, 1962).
Many taxa of dinosaurs have been based on fossil tracks. In the classi¬
fication of gall insects the gall sometimes yields as good a clue to relationship
as do the insects themselves. The form of the mines is an important tax¬
onomic character in mining insects, and it even sheds light on their history
since these mines are sometimes well preserved in fossil leaves. However,
since 1930 it has no longer been permissible to base the name of new
species exclusively “on the work of an animal” [Arts. 13a and 24b (iii) ].

Fig. 7-1. Diagnostic mine patterns caused by six species of leaf miners of the genus Phytomyza on
the leaves of Angelica (50, 51) and Aquilegia (54). The letters a, b, c refer to different species of
Phytomyza on the same host plant (from Hering, 1957).
 Taxonomic Characters 129

Fig. 7-2. Types of spermathecae in the Drosophila repleta group placed on a phylogenetic tree
constructed on the basis of chromosomal evolution (after Throckmorton, 1962).

7.4.2 Coloration. Color pattern and other aspects of coloration are

among the most easily recognized and thus the most convenient characters
in certain groups of animals. Every species of birds can be recognized by
its coloration except for a few genera with sibling species (e.g., Collocalia,
Empidonax). The same is true for certain reef fishes, butterflies, and other
animals. Even where coloration is not completely diagnostic, it often helps
to narrow down drastically the number of species to be considered. In
groups in which subspecies are recognized routinely, such as mammals,
birds, butterflies, or certain wasps, color again plays an important role;
many subspecies are entirely based on coloration.
The quality of color is not easy to describe in words, so that a descrip¬
tion by one author is often misunderstood by his readers. In original revi¬
sionary work, therefore, it is preferable not to rely on descriptions but
to base one’s judgment on the comparison of specimens if at all possible.
There are, however, various ways by which greater precision can be given
to color determination and description, as described in Chap. 10.
7.4.3 Genitalic Structures. For reasons that are not yet fully under¬
stood (Mayr, 1963, p. 103), the genitalia of many animals, particularly
arthropods, not only show a great deal of structural detail but are also
highly species-specific. Being three dimensional structures they have to be
carefully prepared to be strictly comparable (Dreisbach, 1952). In many
groups of insects and spiders genitalic structures are more important for
species diagnosis than is any other character. Yet even here it has been
found either that a single species may have a good deal of variation or
else that two related species have indistinguishable structures. Genitalic
130 Methods of Zoological Classification

(a) talpoides (douglasi)

(6) mazama (tacomensis)

I_i_i_i_i_I_i_i_i_i_I_i_i_i_i_I_i_i_i_i_I—i_i_i_i_I_i_i_i_i_I
0 5 10 15 20 25 30 mm.
Fig. 7-3. Shape of baculum as taxonomic character, (a) Thomomys talpoides; (b) Th. mazama.

structures are most useful among the arthropods. In most vertebrates they
are soft, but the gonopodium of some fishes, the hemipenis of snakes, and
the baculum of mammals supply good taxonomic information.
7.4.4 Other Characters. Morphological characters of adult speci¬
mens are still used more frequently than any others but they are supple¬
mented to an ever-increasing extent by other characters, as listed and dis¬
cussed below (e.g., Blair, 1962; Munroe, 1960). This is particularly true
for “difficult” species, genera, and families where the evidence from mor¬
phology has been equivocal or contradictory. There has been an increasing
utilization of new characters, because (1) morphology reflects only part
of the genotype and may not reflect genetic relationship accurately; (2)
morphology in certain taxa does not supply sufficient characters; and (3)
external morphology is sometimes misleading owing to special adaptations.
The introduction of new kinds of taxonomic characters has been one of
the aspects of the so-called new systematics. The new characters (proteins,
chromosomes, behavior, etc.) supplement but do not displace the use of
morphological characters.
A particularly important reason for the utilization of new characters
is that they serve as check on the conventional morphological characters.
When discrepancy between a morphology-based and a non-morphology-
based classification occurs, still other sets of taxonomic characters must
be used. Fortunately, it turns out that the newer characters usually confirm
the classifications based on the more classical approaches. It seems that
the general morphology is a reflection of a large part of the genotype
 Taxonomic Characters 131

and thus permits, on the whole, reliable conclusions on relationships.

7.4.5 Larval Stages and Embryology. Various immature or larval
stages, the embryology, and sometimes even the eggs may provide taxonomic
information. The various sibling species of the Anopheles rnaculipennis
complex (malaria mosquitoes) were discovered owing to differences in egg
structure. The classification of the Aleyrodidae (whiteflies) is based pri¬
marily on the pupae. Discovery of the leptocephalus-like larval stage of
the fish order Heteromi confirmed the previously suspected relationship
with the eels.
In groups with complete metamorphosis, entirely different sets of char¬
acters evolve in larvae and adults, and conclusions drawn from the charac¬
ters of one stage form a very useful check on conclusions from those of
another. This was shown for frogs (Anura) by Orton (1957) and Inger
(1967). A careful comparison of larval and adult characters of the digger
wasps (Sphecidae) permitted Evans (1964) to show that some features
of the adult structure (emarginate eyes, loss of wing veins) had been over¬
valued. A proper evaluation of larval characters led to an improved arrange¬
ment. Similar findings have been made in other groups of insects (Van
Emden, 1957). Larval and adult characters are visible manifestations of
the same genotype. There can be different identification schemes for
larvae and adults, but there must be only one classification for a given
group of organisms. This must be based on the proper weighting of both
adult and larval characters. It is sometimes the adults and sometimes the
larvae which have acquired specialized adaptations, and only a biological
analysis can lead to the proper evaluation. In groups like the sponges
(Porifera) with extremely few morphological characters, a due considera¬
tion of the embryology has been a great help in classification (Levi, 1956).
7.4.6 Genetic Characters and Sterility. There has been much con¬
fusion as to the meaning of the term genetic characters. Broadly speaking,
all characters, except nongenetic modifications of the phenotype, are genetic
characters. Isolating mechanisms and monogenic characters have sometimes
been designated as genetic characters, but this is an unjustified restriction.
The term genetic characters is actually meaningless.
On the whole, only closely related species can hybridize successfully.
In groups of related forms the presence or absence of cross-sterility is impor¬
tant information, but this information must be employed carefully. Ducks,
for instance, show much cross-fertility not only of species but even of related
genera. Yet two such closely related species as the Wood Duck (Aix sponsa)
and the Mandarin Duck (A. galericulata) are completely sterile owing
to a chromosomal rearrangement. In birds, on the whole, there is much
fertility among closely related species, while in some groups of insects there
is already much sterility when geographically remote populations of the
132 Methods of Zoological Classification

same species are crossed. In other groups of insects congeneric species may
be fully fertile with each other. A careful analysis of the genetic compatibil¬
ity of various species of toads has led to a regrouping of species in the
genus Rufo (Blair, 1963). Indeed, a few exceptions notwithstanding, the
relative degree of fertility in species crosses is, on the whole, a very sensitive
measure of relationship.
7.4.7 Chromosomes. Botanists have made use of the abundant infor¬
mation provided by the chromosome pattern far longer than have zoologists.
Improvements in cytological technique during the past twenty years now
permit chromosomal studies even in so-called difficult groups, such as the
mammals. Birds and lepidopterans with their small chromosomes and high
chromosome numbers create the greatest difficulties. Diptera, particularly
those with giant chromosomes, and Orthoptera are among the groups that
are most suitable for chromosomal studies. White (1954, 1957) has provided
useful summaries, already made partially obsolete by the recent rapid ad¬
vances. The current abstract and review literature provides convenient ac¬
cess to this field. Matthey for mammals and reptiles, S. G. Smith for beetles,
Hughes-Schrader for mantids, White for grasshoppers, Stone, Carson, and
Wasserman for Drosophila, Keyl for chironomids, and Halkka for homop-
terans are among the numerous authors who have advanced the classifica¬
tion of animals, particularly that of insects, through studies of chromosomal
patterns. The study of primate chromosomes is another very active field,
yielding much information on relationships (Chiarelli, 1966).
Chromosomes are particularly useful on two different levels. On the
one hand, they aid in the comparison of closely related species, including
sibling species. These are often far more different chromosomally than in
their external morphology (Mayr, 1963, chap. 3). On the other hand,
chromosomal patterns are of extreme importance in establishing phyletic
lines. Most chromosomal changes are unique events which are then charac¬
teristic for all descendants of the ancestral population in which the new
pattern first became established. Changes in sex determination, in all sorts
of rearrangements of the chromosomes and centromeres, in fusions, fissions
or translocations, in the acquisition of supernumeraries, etc., often give
unequivocal clues to relationship. For instance, the similarity in the sperma¬
togenesis of mallophaga and anoplura (true lice) strongly supports the
belief in a close relationship of these taxa. The neotenic salamanders Nec-
turus and Proteus were believed to have become similar by convergence,
but both genera have a haploid chromosome number of 19, otherwise un¬
known among urodeles, and the homologous chromosomes are very similar
(Kezer et al., 1965).
Polyploidy is rare in animals, as compared with plants, but there
are numerous other ways by which increases or decreases in chromosome
number may occur. The most frequent evolutionary trend, curiously, in
 Taxonomic Characters 133

many groups is from high to low chromosome number through chromosomal

fusion. Even though the chromosomes represent the genetic material, it
is not true that the amount of chromosomal change reflects the amount
of genetic change. Close relatives may show considerable rearrangement;
indeed many species are polymorphic for various types of chromosomal
rearrangements. On the other hand, cases are known in which a consider¬
able degree of genic change is not or only lightly reflected in the chromo¬
somal pattern, as in the Hawaiian Drosophila (Carson, 1967).
7.4.8 Physiological Characters. This group of characters is hard
to define. All structures are the product of growth processes, that is of
physiological processes, and are thus ultimately physiological characters.
Also all physiology is regulated by enzymes and other macromolecules and
thus is not separable from biochemical characters. By “physiological charac¬
ters” one generally means growth constants, temperature tolerances, and
the various processes studied by the comparative physiologist. Species differ¬
ences are abundant in these characters, but since they are not present in
preserved material and usually require special apparatus for their study,
they are rarely used by the taxonomist. Some representative cases are dis¬
cussed by Mayr (1963, pp. 60-65), and fuller treatments are given in
textbooks of comparative physiology (e.g. Prosser and Brown, 1961).
7.4.9 Biochemical Characters. The major molecular inventions were
made by the earliest living organisms. Even the most primitive procaryotes
have on the whole the same kinds of macromolecules and metabolic pro¬
cesses that we find in the highest animals and plants. Still, there is enormous
specificity at every taxonomic level, and this is being increasingly exploited
by the taxonomist.
Serology provided the earliest widely used method of comparing pro¬
teins. This method is based on the principle that the proteins of one orga-

Fig. 7-4. Reaction between the blood sera of six species of bovids with the antisera against three
species (from Moody, 1958).
134 Methods of Zoological Classification

nism will react more strongly with antibodies to the proteins of a closely
related organism than to those of one more distantly related. Unfortunately,
the method encounters various technical difficulties, and though used for
more than forty years it has not contributed as much to a clarification
of otherwise ambiguous cases as had been hoped. A summary of some
of the achievements of this method is found in the Kansas Symposium
(Leone, 1964). As a result of improvements in the techniques there has
recently been a revival of interest in the quantitative study of antigenic
reactions (e.g. Williams. 1964). The study of blood-group genes (“immuno-
genetics”) has shed light on relationships between species of pigeons
(Irwin, 1947, and later papers of his school) and is now used extensively
in the study of primates.
Chemical Taxonomy. Much recent work has been devoted to a tax¬
onomy of specific chemical components and macromolecules. Paper chroma¬
tography has been used by numerous authors to compare the chemical
composition of closely related species, with particular attention to amino
acids and peptides revealed by ninhydrin treatment, and the purines, pyrimi¬
dines, or other substances that either fluoresce or absorb ultraviolet light.
Various methods of electrophoresis reveal the molecular composition of
complex proteins. Sibley (1960) analyzed the egg-white proteins of more
than 100 species of birds and was able not only to add clear-cut indications
of relationship in previously equivocal cases, but also to raise doubts con¬
cerning previous arrangements and to make suggestions as to relationship
of taxa previously considered highly isolated. Throckmorton (1962) did simi¬
lar studies for the species of Drosophila. Paper electrophoresis, which was
used in these earlier studies, has been largely replaced by newer techniques
that permit a much finer resolution, but still newer techniques are con¬
tinually being introduced. These techniques have been used also in the
study of reptiles, amphibians, insects, mollusks, and other animals, and
there is no doubt as to their increasing importance. It is necessary to consult
the current literature for the latest techniques and instrumentation in pro¬
tein, peptide, and amino acid analysis (e.g. Wright, 1966).
Still another approach is to consider a single complex molecule for in¬
stance the hemoglobin, of one species, and compare its amino acid composi¬
tion with that of closely or more distantly related species. Patterns of re¬
placement often indicate whether or not two organisms belong to the same
phyletic line. Two recent symposia (Handler, 1964, and Bryson and Vogel,
1965) present such evidence from this rapidly developing field. Any given
enzyme can be used for these studies, but the results are, of course, subject
to the same shortcomings as any other single-character classification.
The newest and most exciting development is the attempt to go back
to the very basis of relationship, the genetic program itself. Techniques
of DNA matching are now being developed (Hoyer et al., 1964) which
 Taxonomic Characters 135

hold much promise. It is not yet certain that these methods can be made
sensitive enough to add to the existing information, but the study of evolving
molecules superimposed on the background of classical taxonomy is bound
to reveal discrepancies and inaccuracies which will lead to improvements
in classification.
The study of structural characters has revealed that each organ or
organ system may have its own specific rate of evolutionary change (mosaic
evolution, 10.4.5). Much evidence indicates that this principle is equally
valid for chemical characters. A comparison of man (Homo) with the
African apes (Pan) shows, for instance, that there has been little evolution¬
ary divergence in their hemoglobins and serum proteins since they branched
from each other, even though the hominid line has since entered an entirely
new adaptive zone (see Fig. 4-5). When using taxonomic characters to
draw inferences on classification one must always balance the potentially
conflicting information derived from different characters. One must also
understand the subtle differences between evolutionary phenomena at the
molecular and at the organismic level (Mayr, 19646; Simpson, 1964).
7.4.10 Behavior. Behavior is undoubtedly one of the most important
sources of taxonomic characters. Indeed, behavioral characters are often
clearly superior to morphological characters in the study of closely related
species, particularly sibling species (Mayr, 1963). Yet there are two major
technical drawbacks. Behavior cannot be studied in preserved material,
and it is intermittent even in the living animal. Certain types of behavior

SEC 01YDS-
0 1 2 3 4 5 6 7 8 9 10 II 12 13 14
SPECIES

t PRIMITIVE UN-NAMED
» i i

2. CINCTlPENNIS • • »
3 HEBES
SALINUS • • • ► • » ►

4. POTOMACA 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

& FRONTALIS
l i 1 i i i i i i i • i i

S VERSICOLOR HttH tan Iht*

7 VERSICOLOR VAR
OUAORIFULGENS M III III
8 VERSICOLOR, TRIPLE
FLASH (DELAWARE) HI II Ml
9 FAIRCHILDI i i
i • i i

10. LUCICRESCENS

II. T»ec-T0P SPCC'CS.

the same as 1

12. PENSYLVANICA !«■ «•

11 PYRALOMIMUS m
14 AUREOLUCENS »
• »

15. CAERULUCENS

16 TREMULANS MM m
Fig. 7-5. Pattern of light flashes in North American fireflies (Pholuris). Height and length of the
marks indicate intensity and pattern of the flashes (from Barber, 1951).
136 Methods of Zoological Classification

occur only during the breeding season or during part of the 24-hour period.
The comparative study of behavior of related species has become an autono¬
mous discipline, comparative ethology. It has already made major contribu¬
tions to the improvement of classifications of birds, bees, wasps, orthop-
terans, frogs, fishes, and other groups.
The reason for the importance of behavior is obvious. Behavior charac¬
teristics are the most important isolating mechanisms in animals, and new
adaptations are often initiated by changes in behavior. The rapidly expand¬
ing literature on behavioral systematics has, in part, been summarized
in a number of reviews: Mayr (1958), Baerends (1959), Cullen (1959),
Alexander (1962), and Wickler (1961, 1967). Exemplary studies are those
of Evans (1957, 1966) on the digger wasps (Sphecidae), of Sakagami
and Michener (1962) on sweat bees, of Spieth (1952) on the genus Dro¬
sophila, of Johnsgard (1965) on ducks (Anatidae), of Tinbergen (1959)
and Moynihan (1959) on gulls (Laridae), and of Smith (1966) on
Tyr annus.
A great technical advance in the study of behavior has been the
development of accurate sound-recording devices and of further devices,
such as sonagraphs, for the translation of sounds into graphic patterns
(Fig. 7-6). More than 40 species of North American crickets were either
discovered or rediscovered as a result of the careful analysis of their songs
by B. B. Fulton and his followers, (Walker, 1964, Alexander, 1967). The
classification of species in several avian genera (for instance, Myiarchus,
Empidonax, Tyr annus) has been greatly helped by an analysis of sound
recordings. A comparison of the calls of frogs and toads has not only led
to the discovery of previously unrecognized sibling species but has also
shed light on the relationship between previously established species. Impor¬
tant studies of comparative sound analysis in anurans were made by Barrio,
Blair, Bogert, Littlejohn, Main, and Mecham. The acoustics of behavior
of animals has been summarized in a number of recent volumes (Tavolga
and Lanyon, 1960, and Busnel, 1963).
In addition to courtship behavior and acoustic behavior, various other
kinds of behavior elements have taxonomic value. For example, the pattern
of the webbing constructed by various spiders, mites, and caterpillars may
be used at various levels in the classification. The two bee genera Anthidium
and Dianthidium were slow to be recognized on morphological grounds,
yet all known species of the former construct their nests of cottony plant
fibers and those of the latter construct theirs from resinous plant exudations
and sand or small pebbles.
The use of extraneous materials in the construction of nests or of
larval or pupal cases provides characters at various levels in the classification
of caddisworms and bagworms. The manner in which such materials are
attached to the shell is a useful taxonomic character in distinguishing species
 Taxonomic Characters 137

of the Molluscan genus Xenophora. The egg cases of praying mantids

have a species-specific form.
Behavior patterns that are characteristic of higher taxa are far more
rare. Examples are the use of mud in nest building by barn swallows and
crag martins (Hirundo), certain “comfort movements” (scratching, stretch¬
ing, bathing, etc.) in birds (McKinney, 1965), and grooming movements
in insects. Attention up to now has been directed so strongly at the compari-

Fig. 7-6. Graphic representation of the songs of three species of sparrows (Passer domesticus, Passer
montanus, Petronia petronia) (from Thielcke, 1964).
138 Methods of Zoological Classification

son of closely related species that the diagnostic value of behavior patterns
at the level of higher taxa has been relatively neglected.
7.4.11 Ecological Characters. It is now well established that every
species has its own niche in nature, differing from its nearest relatives in
food preference, breeding season, tolerance to various physical factors, resis¬
tance to predators, competitors and pathogens, and in other ecological fac¬
tors. Where two closely related species coexist in the same general habitat,
they avoid fatal competition by these species-specific niche characteristics
(principle of competitive exclusion, Mayr, 1963, chap. 4). A number of
sibling species were discovered as a result of discrepancies in food preference
(“host specificity”)—for example, the apple and blueberry maggot—or
habitat preference (op. cit., chap. 3). Many aspects of the life cycle, such
as life span, fecundity, and length or time of breeding season, may be
different in closely related species (op. cit.).
Niche specificity is quite pronounced even in species such as birds,
mammals, or mollusks that are not particularly substrate-specific. Kohn
(1959) found that every species of the genus Conus in the Hawaiian Islands
differs ecologically from related species. Two sibling species (ebraeus and
chaldaeus) feed on nereid polychaetes, but of 199 ebraeus 136 contained
nereid species a and none species b, while of 106 chaldaeus 5 contained
species a and 98 species b. The larvae of Drosophila mulleri and aldrichi
live simultaneously in the decaying pulp of the fruits of the cactus Opuntia
lindheimeri. The two species are, however, markedly specialized in their
preference for certain yeasts and bacteria (Wagner, 1944).
Niche specialization is even more important for animals that are sub¬
strate-specific.. This includes host-specific plant feeders among insects and
mites as well as host-specific parasites. Unfortunately there is no up-to-date

Table 7-2. Ecological Differences between Sibling Species in the

Anopheles maculipennis Complex

Species Habitat Water type Hibernation Malaria carrier

melanoon Rice fields Fresh water No No

messeae Cool standing Fresh water Yes Almost never
water
maculipennis Cool running Fresh water Yes No
water
atroparvus Cool waters Brackish No Slightly
labranchiae Mostly warm Brackish No Very dangerous
waters
sacharovi Shallow stand¬ Often brackish No Very dangerous
ing water
 Taxonomic Characters 139

summary of the field except the excellent treatment of Dethier (1947),

the emphasis of which is on the sensory aspects of host selection. Many
new species of insects were discovered by comparing populations of the
“same” species occurring on different plant hosts. Some enthusiasts carried
this principle too far and invariably made the occurrence on a different
host an excuse for the description of a new species. In the North American
Cryphalini (bark beetles) alone, 53 of the “species” described by Hopkins
turned out to be synonyms. Downey (1962) and Kohn and Orians (1962)
have given useful summaries of some of the relevant literature. Host speci¬
ficity of external parasites was discussed by Clay (1949), Hopkins (1949),
and Holland (1964).
Ecological differences between subspecies are frequent. Indeed most
widespread species exhibit differences in the ecology of local populations,
particularly of peripherally isolated populations (Mayr, 1963, pp. 312ff,
355).
Again, as with behavioral characters, differences on the species level
have been studied far more extensively than have ecological differences
between higher taxa. Many of these are simply taken for granted. That
whales occupy a different adaptive zone from that of bats is too obvious
to be mentioned. Yet a close study shows that even most genera, when
well-founded, occupy definably different niches or adaptive zones. Lack
(1947) showed for instance that each genus of Galapagos finches is charac¬
terized by its utilization of the environment. Geospiza is a ground finch
(chief food, seeds), Camarhynchus a tree finch (chief food, insects), and
Certhidea a warbler finch (chief food, small insects). For more on this
topic, see 5.3.2 and 10.5.36, also Bowman (1961).
7.4.12 Parasites and Symbionts. In several instances sibling species
were discovered because their parasites were different. Two species of
Octopus in California are hosts to two different species of mesozoan para¬
sites. A new species of termites was discovered because its nests contain
a different set of termitophile staphylinid beetles than those of a previously
known species. Parasites are also important in contributing to our knowledge
of the relationship of higher taxa. Parasites evolve together with their hosts
and are in some cases more conservative than their hosts. Unfortunately,
they shift to new hosts more frequently than is sometimes admitted, and
the evidence on parasites must be evaluated very carefully (Baer, 1957).
For instance, the flamingos (Phoenicopteri) exhibit characteristics which
they share with both storks and geese. Their bird lice (Mallophaga) belong
to the same genera that occur on the geese. At first sight this might suggest
close relationship, but it actually indicates merely a comparatively recent
transfer of the lice from the geese to the flamingos. If the bird lice had
been derived from a common ancestor, we would expect related, but slightly
different, parasites in the two orders of birds. The anatomical evidence
140 Methods of Zoological Classification

indicates intermediacy, but on the whole, closer affinity with the storks
than with the geese. Man {Homo) and the African apes {Pan) share
more external and internal parasites with each other than does Pan with
the orang {Pongo). This strengthens the case for a close relationship be¬
tween Homo and Pan (established on other grounds). However, it might
also be due to cross-infection between sympatric host species, although man
also occurs in the habitat of Pongo.
The fact that intracellular symbionts supply important taxonomic
characters is the discovery of P. Buchner (1966a) and his school. For
instance, the most primitive tribes of the coccids (Steingeliini, etc.) have
no symbionts, but once a coccid taxon has acquired them this symbiont
(with all its highly specific adaptations) will be found in the derived phy-
letic lines of coccids. Repeatedly, unnatural taxa of coccids could be un¬
masked because they had heterogeneous complements of symbionts (see
also Buchner, 19666). The same is true for the symbionts of other groups
of insects. The protozoan faunas in the intestines of termites evolved to¬
gether with their hosts and are potentially useful indicators of relationship
in cases of ambiguity in the termite classification (Kirby, 19506).
7.4.13 Geographical Characters. Geographical characters are among
the most useful tools for clarifying a confused taxonomic picture and for
testing taxonomic hypotheses. Most sound classifications show some correla¬
tion with geographic or associated ecologic features. The taxonomist is
primarily interested in two kinds of geographical characters, (1) general
biogeographic patterns, which are especially useful in the arrangement and
interpretation of higher taxa, and (2) the allopatric-sympatric relationship,
which is most helpful in determining whether or not two populations are
conspecific (see Chap. 9). For a detailed treatment of various aspects of
the relation between taxonomy and geography, see also Systematics Associa¬
tion Publication no. 4 (D. Nichols, 1962), and Simpson (1965).
Studies of the distribution of large numbers of groups of plants and
animals have revealed well-defined geographic patterns. Biogeographers
have divided the world into various realms, regions, provinces, subprovinces,
etc. based upon generalized comparisons of faunas and floras. These are
not rigidly defined, but in general they represent distributional centers which
exist today or have existed in the past. The units of distribution are not
static but may be expanding or retreating, and we thus find it more useful
to refer to them as faunas, floras, or biotas rather than as zones or areas.
A taxonomist should have an understanding of the geological history of
the regions in which such biotas center, as well as a knowledge of the
past relationships of the faunas and floras concerned (Mayr, 1964a). Posses¬
sion of this information will permit a much sounder interpretation of various
higher taxa.
The reason for the great taxonomic value of geographical distribution
 Taxonomic Characters 141

lifc. 7-7. Distribution pattern of groups of species of lugworms (Arenicolidae). Figures indicate
isotherms of ocean surface waters (from Wells, 1963).

is evolution and monophyly. Just as every taxon is descended from a com¬

mon ancestor, so is every colonization across a barrier effected by a founder
species. In other words, there is high probability that related species in
an area are descendants of a common ancestor and that in the majority
of the cases no other species of this group occur disjunctly at a far distant
place. For instance, the mammals of South America are either not related
to those of Africa or, if of common ancestry, have presumably reached
South America by way of North America. The hystricomorph rodents, seem¬
ingly close to the African porcupines, appeared to be an exception, owing
to the absence of porcupines in the early Tertiary period of North America.
A reexamination of these rodents, prompted by this zoogeographical puzzle,
revealed that the porcupines of South America and of Africa are of inde¬
pendent origin (Wood, 1950). Distributional difficulties have shed light
on taxonomic relationships in many other instances. They suggest, for exam-
142 Methods of Zoological Classification

pie, that the New Zealand “thrushes” (Turnagra) are not thrushes but
presumably Pachycephalinae, and the New Zealand “tits” not Paridae but
Malurinae. Both reassignments lead not only to an improved classification
but also to a considerable zoogeographical simplification. Distribution is
thus shown to be an important tool in taxonomic analysis. Its most meaning¬
ful application consists in always asking whether our inferences on the
relationship of taxa are consistent with their pattern of distribution.
7.4.14 The Relative Value of Different Kinds of Characters. Each
kind of character adds to the total information. Very often the taxonomist
is limited to preserved material, in other words essentially to morphological
characters and to such geographical and ecological information as came
with the specimens. Whenever the morphological analysis permits ambiguity,
and this is often the case, every effort should be made to supplement the
basic information with biochemical, behavioral, or other additional informa¬
tion. Different kinds of characters are used in the discrimination of species
than in the determination of relationships among higher taxa. It is only
within the most recent past that an effort has been made to determine
the relative weight of different kinds of characters at different levels of
the taxonomic hierarchy.

7.5 CHARACTERS AND CATEGORICAL RANK

Characters that unequivocally designate categorical rank do not exist.

We cannot say that species A must be placed in a separate genus because
it has a generic character (5.4.2). The degree of morphological difference
between species and genera is much greater in some families than in others.
There are stronger behavioral differences among closely related species in
some groups than in others. The same character that is associated with
generic difference in one family may vary individually in species of another
family. Nor does the total level of difference tell us much. Morphs within
a single population may differ more from each other than do good sibling
species, at least in visible morphological characters. Again and again we
must remember that it is the taxon which gives us the characters, not
the reverse.
Empirically, the experienced taxonomist knows what kind of charac¬
ters have what kind of value in the particular group in which he is a
specialist. Any difference that functions as an isolating mechanism or facili¬
tates competitive exclusion is primarily useful for the discrimination of
species. What particular system of communication is used in isolating
mechanisms (light signals in fireflies, calls in frogs, etc.) may serve as
character of a higher taxon, while a specific isolating mechanism may char¬
acterize a species. On the other hand, characters that indicate occupation
 Taxonomic Characters 143

of a particular adaptive zone or that are by-products of a well-defined,

well-integrated genotype generally help in the discrimination and delimita¬
tion of higher taxa. Recent studies of biochemical characters have clearly
shown that some of them are most useful at the species level, for instance
some immunological methods, while others are not sensitive at that level
but are full of information concerning the higher taxa; still others are
subject to local polymorphism and have no taxonomic value. It is up to
the taxonomist to find out what character will be most useful at a given
categorical level.
Even though broad generalizations cannot yet be made, a differential
significance of characters is gradually becoming apparent within each higher
taxon. Different kinds of characters appear to be most useful at the fol¬
lowing categorical levels:

1. The recognition of subspecies (geographic variation)

2. Distinguishing closely related species, particularly sibling species
3. The grouping of related species into genera
4. Determining the relationship of higher taxa, from families to phyla

CONCLUSIONS

A taxonomic character is any attribute by which a member of a

population may differ from members of other populations of organisms.
This attribute may relate to any feature of the dead or living organism
that is amenable to comparison.

1. Taxonomic characters that are conservative (i.e., that evolve slowly)

are most useful in the recognition of higher taxa; those that change
most rapidly or concern isolating mechanisms are most useful in the
lower taxa.
2. Taxonomic characters that are subject to parallelism, especially those
involving loss or reduction, should be used only with great caution.
3. Taxonomic characters are expressions of the biology of their carriers.
An understanding of this biology is a prerequisite for the proper evalua¬
tion of these characters.
4. The same phenotypic character may vary in value and constancy from
taxon to taxon and even within a single phyletic series. The weight
to be given to such a character depends largely on its constancy in
the given group.

The entire zoological classification is based on the proper evaluation

(weighting) of taxonomic characters. This operation, then, is the most
important as well as one of the most difficult tasks of the taxonomist (10.4).
Chapter Q The Qualitative and
O Quantitative Analysis
of Variation

Ji-n his work the taxonomist does not encounter species

as such; he works with specimens and samples, and his first task is to
assemble these into groups of similar specimens, phena (Chap. 1). These
in turn he assigns to species. The history of taxonomy, particularly the
evidence supplied by the long lists of synonyms in many groups, proves
that this is not an easy task, owing to the great variability of most species.
A student of inanimate objects establishes classes merely on the basis
of similarity. He would not hesitate to place things as different as the
caterpillar, chrysalis, and imago of a butterfly in three different classes,
while he might place the adults of two similar species of butterflies in
the same class. The biologist, however, knows that he must take into con¬
sideration other factors than mere similarity. The continuity of the genotype
from the fertilized egg through all larval stages to the adult is one such
factor. Sexual reproduction—which results in a genetic cohesion among
all the individuals belonging to the same local population—is another such
factor. All members of a local population are, so to speak, products of
the same gene pool and are thus a single taxonomic entity.
It is the knowledge of these biological phenomena which permits
the taxonomist to assign phena correctly to species. In a species with sexual
dimorphism, for instance, males and females will belong to two different
phena. The fact that one phenon consists entirely of males and the other

144
 The Qualitative and Quantitative Analysis of Variation 145

entirely of females indicates that these phena are not species. Additional
information will now be brought to bear on the situation. For instance,
if the two phena are the only ones in the collection made at a given
locality representing a certain genus and were collected simultaneously in
the same habitat, the probability is high that they represent males and
females of the same species. Breeding tests, the raising of larvae, and the
study of courtship and copulation in nature furnish additional sets of bio¬
logical information permitting the correct assignment of phena. The number
of possible inferences from the available information is usually large. A
knowledge of the nature and amount of sexual dimorphism found in living
(Recent) species often permits the correct assignment of fossil phena to
species, as is the case, for instance, among fossil ostracods.
The greater the amount of information available about the phena
in question, the easier their classification. All the information must be con¬
sidered that is specified in Chap. 6, such as correct locality, habitat (and
other relevant ecological information), and season of capture. The reasons
for precision in this information are twofold:

1. Many aspects of ecology and life history are species-specific (see Chap.
7).
2. The phenotype of animal populations of the same species often varies
according to locality, season, or habitat (see below).

Differences between phena may thus reflect either a species difference

or intraspecific variation. A complete understanding of intraspecific varia¬
tion is therefore necessary before we can make the probabilistic statement
that phenon B belongs to a different species from phenon A. This is the
reason for the immense importance of a thorough understanding of indi¬
vidual and geographic variation.
The taxonomic literature still contains numerous named phena that
have not yet been correctly combined into biological species. This includes
males and females in sexually dimorphic groups of insects (8B.1), workers
and sexual castes in social insects, stages in the life cycle of parasites, and
juvenile stages as well as morphs. It is one of the continuing preoccupations
of the taxonomist to unmask nominal species that are not genuine biological
species.
Basic Questions Concerning Phena. When a taxonomist has before
him two or more phena which he must correctly assign to the species
to which they belong, he must ask himself these questions:

1. Do the phena or samples come from the same locality? If the answer
is yes, only two alternatives are possible. Either the phena are individual variants
of a single species or else they are several good species (Chap. 9A.1). If the phena
are from different localities, there is a third possibility, namely that they represent
different subspecies of a geographically variable species (Chap. 9B).
146 Methods of Zoological Classification

2. Is there any evidence for reproductive isolation between the populations

from which the phena were sampled? This is a much more difficult question to
answer. Collected material may provide only a few clues helpful in this problem.

If the investigator has the answer to these questions, he can read

from the discrimination grid (Table 8-1) what the taxonomic disposition
of the phena ought to be.
Presence or absence of morphological difference as well as geographic
relationship can nearly always be determined in properly labeled material.
It is the lack of information on reproductive isolation which causes most
of the difficulties. By looking at the discrimination grid we can see that
without information on reproductive isolation we shall have difficulty in
choosing between alternatives (1) or (5), (2) or (6), (3) or (7), (4)
or (8). A high percentage of taxonomic errors are the result of a wrong
choice between one of these pairs. What can we do, in the absence of
direct information on reproductive isolation, to avoid error? The experi¬
enced taxonomist knows that a full understanding of both individual and
geographic variation is more helpful in the avoidance of error than is
anything else. Most of the present chapter will therefore be devoted to
a thorough discussion of variability. Even though every kind of animal
has its own pattern of variability, it is apt to fall somewhere into the
framework presented in this chapter.
Phena collected at the same locality (or geological horizon) are either
different biological species or manifestations of individual variation (alterna¬
tives 3 and 7). In order to be able to discriminate between the two alterna¬
tives, in the absence of information on reproductive isolation, it is necessary
to review the total potential of a species for individual variation. Such
variation is far greater than the beginner realizes, and it sometimes deceives
even experienced taxonomists. It is estimated that more than half of all
synonyms owe their origin to an underestimation of individual variation.

Table 8-1. Discrimination Grid

Not reproductively Reproductively

isolated isolated

Morphologically identical:
Sympatric (1) Same population (5) Sibling species
Allopatric (2) Same subspecies (6) Sibling species
Morphologically different:
Sympatric (3) Phena of the same (7) Different species
population
Allopatric (4) Different subspecies (8) Different species
 The Qualitative and Quantitative Analysis of Variation 147

Table 8-2. Major Types of Variation within a Single Population

I. Nongenetic variation
A. Individual variation in time
1. Age variation
2. Seasonal variation of an individual
3. Seasonal variation of generations
B. Social variation (insect castes)
C. Ecological variation
1. Habitat variation (ecophenotypic)
2. Variation induced by temporary climatic conditions
3. Host-determined variation
4. Density-dependent variation
5. Allometric variation
6. Neurogenic color variation
D. Traumatic variation
1. Parasite-induced variation
2. Accidental and teratological variation
II. Genetic variation
A. Sex-associated variation
1. Primary sex differences
2. Secondary sex differences
3. Alternating generations
4. Gynandromorphs and intersexes
B. Non-sex-associated variation
1. Continuous variation
2. Discontinuous variation (genetic polymorphism)

Various forms of individual variation are listed in Table 8-2. The two
most important classes are nongenetic and genetic variation.

8A. NONGENETIC VARIATION

It is of course impossible in a preserved museum specimen to deter¬

mine directly whether a given variant has a genetic basis. Nevertheless,
it is important for the taxonomist to understand that various types of varia¬
tion exist and that in the better-known groups it is possible in most cases
to make a valid inference as to the status of a given variant on the basis
of field observations and available experimental evidence. For a discussion
of the evolutionary aspects of individual variation see Mayr (1963, pp.
138-158). As a broad generalization, one can state that nongenetic variation
adapts the individual, while genetic variation adapts the population and
the species.
Animals, as a whole, are developmentally far better canalized than
plants. In addition, through their power of locomotion and their sensory
148 Methods of Zoological Classification

abilities they have the capacity for habitat selection. As a result, some
well-known exceptions notwithstanding, nongenetic changes of the pheno¬
type are far less of a problem for animal taxonomists than they are for
botanists. However, every zoologist must familiarize himself with the types
of nongenetic variability which he might encounter in his group.

8A.1 INDIVIDUAL VARIATION IN TIME

8A.1.1 Age Variation. Whether they are born more or less developed
or whether they hatch from an egg, animals in general pass through a
series of juvenile or larval stages in which they may be quite different
from adults. The catalogs of any group of animals list numerous synonyms
that resulted from the failure of taxonomists to recognize the relationship
between various age classes of the same species.
In reptiles, birds, and mammals there are no larval stages, but imma¬
ture individuals may be rather different from the adults, particularly in
birds. For example, Linnaeus described the striped immature goshawk (Ac-
cipiter gentilis Linnaeus) as a different species (gentilis) from the adult
(palumbarius) with its crossbarred underparts. Several hundred bird
synonyms are based on juvenal plumages. By finding specimens that molt
from the immature into the adult plumage it is usually easy to clear up
this difficulty.
In many small mammals (e.g., rodents and insectivores), samples col¬
lected in different seasons differ in the proportion of adults and juvenals.
Adults and immatures must be separated before samples from different
localities are compared.
In many fishes the immature forms are so different that they have
been described in different genera or even different families. The immature
stages of the eel (Anguilla) were originally described as Leptocephalus
brevirostris Kaup. The unmasking may be especially difficult in neotenic.
animals, that is, animals that become sexually mature in a larval stage.
The difficulties for the taxonomist are even greater in groups with
larval stages which are so different that they have not even the faintest
resemblance to the adult (e.g., caterpillar and butterfly). The floating or
free-swimming larvae of sessile coelenterates, echinoderms, mollusks, and
crustaceans are often extremely different from the adults. The taxonomic
status of such larval stages can be settled either by assembling a complete
sequence of intermediate stages or by rearing them through a complete
life cycle.
The taxonomic identification of larval stages of parasites is made
particularly difficult in groups where the different stages occur on different
hosts. It is customary in helminthology to assign formal taxonomic names
 The Qualitative and Quantitative Analysis of Variation 149

Fig. 8-1. Difference between adult liver fluke (Fasciola hepatica) and its larval stage (Cercaria) (after
Chandler and Read).

to the larval (Cercaria) stage of flukes (trematodes), in order to facilitate

their ready identification. Such dual nomenclature (Art. 42c) is dropped
as soon as it becomes known to what trematode species a given cercaria
belongs. This can usually be established only through rearing.
Age variation is not restricted to differences between larval stages
and adults but occurs also between “young” and “old” adults. For example,
in various species of deer (Cervus, etc.) it is known that older stags often
have antlers with more points than younger ones. The shape of the antlers
may also change. This age variation must be considered when the antlers
of different species or subspecies are compared. There is probably no more
addition of points (or only an irregular one) after a certain age has been
reached. It would be as futile to try to determine the exact age of a stag
150 Methods of Zoological Classification

by the number of points of its antlers as to determine the age of a rattlesnake

(Crotalus) by the number of rings in the rattle, or the age of a hornbill
(Aceros plicatus Forster) by the number of folds in the casque on the bill.
The taxonomist aims to work with samples that are as homogeneous
as possible. It is much easier to achieve this in animals that have a definite
adult stage (after the larval one) than it is in those that show continuous
growth, such as snakes or fishes, which may reach maturity after having
attained only half their potential size, or less. In such forms it is advisable,
to work with regressions rather than with absolute measurements. Meristic
characters (e.g., number of scales or fin rays) are in many species not
increased after they are formed, in spite of the enormous subsequent growth.
These characters are therefore especially important in herpetology and
ichthyology.
In birds it is generally assumed that final size is reached with the
first completely adult plumage. There is now good evidence that this is not
always so. In the hornbill of the Papuan region, Aceros plicatus, it is well
established that those adults which have only two or three folds on the
bill are younger than those with five or more folds. Birds in adult plumage
with five to eight folds have a bill length of 198 to 227 mm, while birds
in equally adult plumage with two or three folds have a bill of 185-199
mm. In some passerine birds it has been possible through banding to show
that the size of known adult individuals increases slightly over the years.
8A.1.2 Seasonal Variation of the Same Individual. In animals that
live as adults through several breeding seasons, it sometimes happens that
the same individual has a very different appearance in different parts of
the year. Many birds have a bright nuptial dress which they exchange
for a dull (“eclipse”) plumage at the end of the breeding season. Among
North American birds this is true, for example, of many ducks, shore birds,
warblers, tanagers, and others. In many species a change of plumage is
restricted to the males.
In arctic and subarctic birds and mammals, such as ptarmigans
(Lagopus) and weasels (Mustela), there may be a change from a cryptic
white winter dress to a “normally” colored summer dress. In other birds
the colors of the soft parts change with the seasons. In the common egret
(Egretta alba Linnaeus) and in the European starling (Sturnus vulgaris
Linnaeus), the bill may change from yellow to black. Plumage changes
in birds are usually effected by molting, but wear alone may produce striking
changes. In the European starling (Sturnus vulgaris), for example, the
freshly molted bird of October is covered with white spots, and all the
feathers show whitish or buffy margins. During the winter the edges of
the feathers wear off, and in the spring, at the beginning of the breeding
season, the whole bird is a beautiful glossy black without the molt of a
single feather. A similar process of wear brings out the full colors of the
 The Qualitative and Quantitative Analysis of Variation 151

nuptial plumage in the males of many other birds. In arid regions, particu¬
larly in real deserts, the sun bleaches the pigments. A bird before the molt
will look much paler than one in freshly molted plumage.
In all these cases it is the same individual which in different parts
of the year looks very different. Such seasonal variation is particularly com¬
mon among vertebrates, with their elaborate endocrine systems. Many such
seasonal variants were described as distinct species before their true nature
was realized.
8A.1.3 Seasonal Variation of Consecutive Generations. Many spe¬
cies of short-lived invertebrates, particularly insects, produce several genera¬
tions in the course of a single year. In such species it is not uncommon
that the individuals which hatch in the cool spring are quite different
from those produced in the summer, or that the dry-season individuals
are different (e.g. paler) from the wet-season population.
Such seasonal forms can usually be recognized not only by the occur¬
rence of intermediates in the intervening season, but also through identity
of wing venation, genitalia, etc.
Cyclomorphosis. A special kind of seasonal variation is found in certain
freshwater organisms, particularly rotifers and cladocerans. The populations
of a species undergo quite regular morphological changes through the sea¬
sons, in connection with changes in the temperature, turbulence, and other
properties of the water. Many “species” have been named, particularly
in the genus Daphnia, that are nothing but seasonal variants. There are
several recent analyses of this cyclomorphosis by Brooks (cladocerans), by
Buchner (rotifers), and by their associates. In rotifers different morphs
may result from different kinds of food.

8A.2 SOCIAL VARIATION (INSECT CASTES)

In the social insects, such as some bees and wasps, but particularly
among ants and termites, castes have developed. These are definite groups
of individuals within a colony in addition to the reproductive castes (queens,
and males or drones) : workers (sometimes of different types), and soldiers
(also sometimes of different types). In the Hymenoptera, these castes are
most commonly modified females and genetically identical (except for the
workers in some social bees), but in the Isoptera (termites) both sexes may
be involved. The structural types observed may result from different larval
food or may be due to hormonal or other controls. Obviously, taxonomic
names should not be applied to these intracolonial variants; but invalid
species have sometimes been described because it was not realized that there
were different types of soldiers or workers in the same colony.
152 Methods of Zoological Classification

8A.3 ECOLOGICAL VARIATION

8A.3.I Habit Variation (Ecophenotypic). Populations of a single

species that occur in different habitats in the same region are often visibly
different. The taxonomic treatment of such local variants has fluctuated
between two extremes: some authors have described them as different spe¬
cies while others consider them all as nongenetic variants. Actually they
may be (1) microsubspecies (or ecological races) or (2) nongenetic eco-
phenotypes. The latter are particularly common in plastic species, such
as some mollusks.
Dali (1898) gave a very instructive account of all the variations
he observed in a study of the oyster (Crassostrea virginica Gmelin) :

The characteristics due to situs may be partially summarized: When a

specimen grows in still water, it tends to assume a more rounded or broader
form, like a solitary tree compared with its relatives in a crowded grove.
When it grows in a tideway or strong current the valves become narrow
and elongated, usually also quite straight. Specimens which have been re¬
moved from one situs to another will immediately alter their mode of
growth, so that these facts may be taken as established. When specimens
are crowded together on a reef, the elongated form is necessitated by the
struggle for existence, but, instead of the shells being straight they will
be irregular, and more or less compressed laterally. When the reef is dry
at low stages of the tide, the lower shell tends to become deeper, probably
from the need of retaining more water during the dry period. . . . When
an oyster grows in clean water on a pebble or shell, which raises it slightly
above the bottom level, the lower valve is usually deep and more or less
sharply radially ribbed, acquiring thus a strength which is not needed when
the attachment is to a perfectly flat surface which acts as a shield on
that side of the shell. Perhaps for the same reason oysters which lie on
a muddy bottom with only part of the valves above the surface of the
ooze are less commonly ribbed. When the oyster grows to a twig, vertical
mangrove root, or stem of a gorgonian, it manifests a tendency to spread
laterally near the hinge, to turn in such a way as to bring the distal margin
of the valves uppermost, and the attached valve is usually rather deep,
the cavity often extending under and beyond the hinge margin; while the
same species on a flattish surface will spread out in oval form with little
depth and no cavity under the hinge.

In freshwater snails and mussels, such habitat forms are particularly

common. The upper parts of rivers, with cooler temperatures and a more
rapid flow of water, have different forms from the lower reaches, with
warmer and more stagnant waters. In limestone districts the shells are
heavy and of a different shape from those which grow in waters poor
in lime. This dependence of certain taxonomic characters on environmental
 The Qualitative and Quantitative Analysis of Variation 153

factors was, curiously enough, entirely overlooked by some earlier workers,

a fact which resulted in absurd systematics. Schnitter (1922), who largely
cleared up the situation, described these absurdities as follows:

The last step in the splitting of the freshwater mussels of Europe was
done by the malacozoologists Bourguignat and Locard. According to the
shape and the outline of the shell, they split up the few well-known species
into countless new ones. Locard lists from France alone no less than 251
species of Anodonta. On the other hand, two mussels were given the same
name, if they had the same outline of the shell, even though one may
have come from Spain and the other from Brittany. It seems incredible
to us that it never occurred to these authors to collect a large series at
one locality, to examine the specimens, to compare all the individuals and
to record the intermediates between all these forms. It is equally incompre¬
hensible that these people did not see the correlation between environment
and shape of shell, even though they spent their entire lives in collecting
mussels.

All these “species” of Anodonta are now considered to be habitat

forms of two species, and the other names have been sunk into the synonymy
of the two valid species.
Whether a given habitat form is an ecophenotype or a microgeo¬
graphic race is not always evident. It is sometimes necessary to transplant
it or to raise it in the laboratory in order to solve this question. Much
work of this sort still remains to be done.
8A.3.2 Variation Induced by Temporary Climatic Conditions. Some
animals with a highly plastic phenotype may produce year classes that
differ visibly from the norm owing to unusual conditions (drought, cold,
food supply, etc.) in a given year. Fish of a given year class may be stunted
or, on the contrary, have proportions indicative of particularly rapid growth.
Samples of susceptible species must be collected in such a way as to com¬
pensate for distortions caused by this factor (Harrison, 1959; Mayr, 1963,
p. 145).
8A.3.3 Host-determined Variation. Host-determined variations in
parasites of plants and animals provide a potential source of taxonomic
error and permit confusion with microgeographical races or with sympatric
species. This phenomenon is most commonly expressed in size differences
but may involve other morphological or physiological characters.
Gerould (1921) has reported that the braconid wasp, Apanteles
flaviconchae Riley, spins white cocoons when reared from blue-green cater¬
pillars of Colias philodice Godart, but golden cocoons when reared from
yellow-green caterpillars from the same species. In another parasitic wasp,
Trichogramma semblidis Aurivillius, Salt (1941) found that males tend
to be wingless and otherwise modified when they develop in the eggs of
the alderfly, Sialis lutaria (Fabricius) (Megaloptera), but not when reared
from lepidopterous hosts.
154 Methods of Zoological Classification

Another kind of difficulty is cited by H. S. Smith (1942). He states

that the encyrtid wasp, Habrolepsis rouxi Compere, readily parasitizes red
scale on citrus but is unable to do so when the red scale is reared on
Cycas. This apparent immunization by the plant host might well confuse
interpretations which utilize parasites as a taxonomic character.
Host-induced variation is not as common as it was formerly believed
to be. Many so-called host races of the literature have been found in recent
decades to be valid sibling species (Mayr, 1963, chap. 3).
8A.3.4 Density-dependent Variation. The effects of crowding are
sometimes reflected in morphological variation. This is not uncommon
where crowding produces a shortage of food materials. However, density-
dependent variation need not be related to food supply. Uvarov, Kennedy,
and others have shown that gregarious species of locusts exist in various
unstable biological phases. (Kennedy, 1956, 1961; Albrecht, 1962). These
phases differ in anatomy (Fig. 8-2), color, and behavior characteristics
and have often been described as distinct species. When newly hatched
nymphs are reared under crowded conditions, they mostly develop into
the gregarious phase; under less crowded conditions into the transitional
phase; and when isolated and reared separately, into the solitary phase.
Similar phases have been reported by Faure (1943a, 19436) in two species
of armyworms (Lepidoptera), Laphygma exigua (Hiibner) and L. exempta
(Walker).
8A.3.5 Allometric Variation. Allometric growth may result in the
disproportionate size of some structure in relation to that of the rest of
the body. If individuals of a population show allometric growth, animals
of different size will show allometric (heterogonic) variability. This is par¬
ticularly well marked among insects. It involves such features as the heads
of ants (Fig. 8-3), the mandibles of stag beetles (Lucanidae), the frontal

Fig. 8-2. Morphological differences between sedentary and migratory phena of Locusta migratoria
Linnaeus. Pronotum of female in dorsal view, A solitary, B migratory phase (from Uvarov, 1921).
 The Qualitative and Quantitative Analysis of Variation 155

Fig. 8-3. Allometric variability. Neuters of Pheidole inslabilis, showing increase in the relative
size of the head with absolute size of the body (after Wheeler, 1910).

horns and thoraces in scarabs, antennal segments of thrips, etc. Failure

to recognize the nature of such variations has resulted in much synonymy.
The exact causes of much of this variation are unknown. In species
with continuous growth, it is actually a form of age variation (see 8A.1).
Some of it has a genetic basis and is more properly classified under 8B.2.
In holometabolic insects, however, where this phenomenon is particularly
common, it is closely correlated with size, and this in turn is thought to
be the result of variation in food supply, which causes the larva or nymph
to metamorphose at different growth stages. For a recent discussion of
allometry see Gould (1966).
8A.3.6 Neurogenic or Ncurohumoral Variation. Neurogenic or neu-
rohumoral variation is color change in individual animals in response to
the environment. Such changes are accomplished through the concentration
or dispersal of color-bearing bodies known as chromatophores. This type
of variation was first thoroughly studied in the chameleon. It occurs sporadi¬
cally in the lower animals but is best developed among the crustaceans,
cephalopods, and cold-blooded vertebrates (cyclostomes, elasmobranchs,
teleost fishes, amphibians, reptiles). Space will not permit a discussion of
this specialized type of variation. For details the reader is referred to Finger-
man (1963), Gersch (1964), and Waring (1963).

8A.4 TRAUMATIC VARIATION

Traumatic variation occurs with varying frequency in different groups

of animals. The abnormal nature of this type of variation is usually obvious,
but in some cases it is subtle and may be misleading.
156 Methods of Zoological Classification

8A.4.1 Parasite-induced Variation. Aside from such familiar effects

of parasitism as swelling, distortion, and mechanical injury, parasites may
produce conspicuous structural modifications. In the bee genus Andrena,
for instance, parasitism by Stylops frequently results in reduction in the
size of the head, enlargement of the abdomen, and changes in puncturation,
pubescence, and wing venation. It commonly results, also, in intersexes.
Since Andrena is markedly sexually dimorphic, these intersexes have been
a source of taxonomic confusion and synonymy. However, in one case
(Linsley, 1937) a stylopized intersex proved of value in associating the
sexes of a bee which had been described as two different species.
Salt (1927) has made the most comprehensive study of the morpho¬
logical effects of stylopization in Andrena. In females he found reduction
of the pollen-collecting organs, loss of anal fimbriae, changes in relative
length of antennal segments, reduction of facial foveae, reduction of the
sting and accessory organs, paling of ventral abdominal pubescence, acquisi¬
tion of angular cheeks, and yellow on the normally dark clypeus. In males
he reports the development of long hairs resembling the female flocculi,
broadening of the posterior basitarsus, changes in proportions of antennal
segments, loss of cheek angles and some yellow from the clypeus, indications
of facial foveae, and reduction in size of genitalia.
Some strikingly different termite soldiers from the Orient were as¬
signed to a new genus and species, Gnathotermes aurivillii. Later it was
shown that these modified soldiers are nothing but parasitized individuals
from colonies of Macrotermes malaccensis (Haviland).
8A.4.2 Accidental and Teratological Variation. Accidental variation
is usually externally induced, although it may work internally through some
developmental or hormonal system. The external stimuli may be mechanical,
physical, or chemical. Such variation is extremely diverse; in most animals
it may be readily identified because the individuals involved either deviate
so markedly from type as to be recognized as freaks, or because the injuries
or abnormalities involved are asymmetrical. However, in those forms which
undergo metamorphosis, injuries to an earlier stage may produce later ab¬
normalities which are not so easily recognized as such. This is especially
true when the anomalies involve characters which are normally of tax¬
onomic value in the group concerned. For instance, certain types of pupal
injury in beetles may produce symmetrical abnormalities in punctation,
surface sculpturing, or segmentation of appendages; in butterflies, symmetri¬
cal modification of wing patterns. In most cases, however, even with such
subtle difference, the abnormal nature of the variation may be detected
by the specialist without much difficulty.
Teratological variation has been elaborately studied and classified by
Cappe de Baillon (1927) and Balazuc (1948). The student interested in
pursuing this subject further is referred to these works for details and
for further references.
 The Qualitative and Quantitative Analysis of Variation 157

Fig. 8-4. Different degrees of segmentation (A-I) in the pleon of males of the parasitic copepod
Plturocrypta porcellanae. Stage I was originally described as a new species in a different genus
(from Bourdon, 1965).

8A.5 POST-MORTEM CHANGES

The taxonomist must guard against one further type of individual

variation. In many groups of animals it is impossible to prevent post-mortem
changes of preserved specimens. Some extreme cases are known in birds.
The deep orange-yellow plumes of the twelve-wire birds of paradise (Seleu-
cidis ignotus Forster) fade in collections to white. Skins of the Chinese
jay (Kitta chinensis Boddaert), whose plumage is green in life, turn blue
in collections, owing to the loss of the volatile yellow component in the
pigment. Many birds that are clear gray or olive-gray when freshly collected
become more and more rufous through oxidation of the black pigment
158 Methods of Zoological Classification

(“foxing”). Many synonyms have been created in ornithology owing to

the comparison of freshly collected material with old museum specimens.
Other post-mortem changes result from the chemical action of pre¬
servatives or killing agents. A common color change of this nature takes
place when certain yellow insects, especially wasps, are overexposed to
cyanide. The specimens turn bright red, and so far no method has been
found for reversing this reaction without injury to the specimens.
When preserving specimens with evanescent colors (corals, marine
slugs, etc.), it is essential to take full notes and preferably color photographs
or water-color sketches. This will make possible an accurate description
of the living animal.

8B. GENETIC VARIATION

In the cases of variation discussed in the preceding section, the same

individual is actually or potentially subject to a change in appearance.
In addition to this noninherited variation, there is much intrapopulation
variation that is primarily due to differences in genetic constitution. This
genetically induced individual variation can—somewhat arbitrarily-—be di¬
vided into two classes.

8B.1 SEX-ASSOCIATED VARIATION

Among genetically determined variants within a population, there

are many which are sex-associated. They may be sex-limited (express them¬
selves in one sex only) or be otherwise associated with one or the other
sex, or they may involve sex characters or modes of reproduction. Some
of these are as follows:
8B.1.1 Primary Sex Differences. These are differences involving the
primary sex organs utilized in reproduction (gonads, genitalia, etc.). Where
the two sexes are otherwise quite similar, primary sex differences will rarely
be a source of taxonomic confusion.
8B.1.2 Secondary Sex Differences. There is more or less pronounced
sexual dimorphism in most groups of animals. The differences between
male and female are often very striking, as for instance in the birds of
paradise, hummingbirds, and ducks. In many cases the different sexes were
originally described as different species and retained this status until pains¬
taking work by naturalists established their true relationship. A celebrated
 The Qualitative and Quantitative Analysis of Variation 159

case is that of the king parrot Eclectus roratus (Muller) of the Papuan
region, in which the male is green with an orange bill, the female red
and blue with a black bill. The two sexes were considered different species
for nearly one hundred years (1776-1873), until naturalists proved conclu¬
sively that they belonged together.
Striking sexual dimorphism is particularly frequent in the Hymenop-
tera. The males of the African ant Dorylus are so unlike other ants that
they were not recognized as such and were for a long time considered
to belong to a different family. In the mutillid wasps (Mutillidae) the
small wingless female and the large winged male are so different that
some taxonomists use a different nomenclature for the two sexes. Whole
“genera” consist entirely of males, others of females. The best way of deter¬
mining with which female of “genus” B a given male in “genus” A belongs
is to find a pair in copula or to watch a female in the field and catch
the males as they are attracted to her. Once it has been established that
B is the female of A, it is sometimes possible to associate several other
“species pairs” in the same genus by utilizing additional information on
distribution, frequency, color characters, etc.
8B.1.3 Alternating Generations. In many insects there is an alterna¬
tion of generations that is very confusing to the taxonomist. In the genus
Cynips (gall wasps), the agamic generation is so different from the bisexual
one that it has been quite customary to apply different scientific names
to the two (Kinsey, 1930). In the aphids (plant lice) the parthenogenetic
wingless females are usually different from the winged females of the sexual
generations (Fig. 8-5).
8B.I.4 Gynandromorphs and Intersexes. Gynandromorphs are indi¬
viduals that show male characters in one part of the body and female
characters in another part. Thus the two halves of the body may be of
opposite sexes, or the division may be transverse, or the sex characters
may be scattered in a mosaic. In the latter case symmetrical variants may
be produced. Usually gynandromorphs are easily recognized as such and
rarely provide a source of taxonomic confusion. Gynandromorphism is pro¬
duced by an unequal somatic distribution of chromosomes, particularly
the sex chromosomes.
Unlike gynandromorphs, intersexes are likely to exhibit a blending
of male and female characters. Intersexes are generally thought to result
from an upset in the balance between male-tendency and female-tendency
genes. This upset may result from irregularities in fertilization or mitosis,
or from physiological disturbances associated with parasitism. Intersexes
are particularly apt to appear in populations of interspecific or inter-sub¬
specific hybrids. They have been studied in greatest detail in Lymantria
(Goldschmidt, 1933) but are well known in many other animals.
160 Methods of Zoological Classification

Fig. 8-5. Periphyllus californiensis (Shinji). 1, Fundatrix or stemmother; 2, normal apterous

parthcnogcnetic viviparous female; 3, alate of same; 4, smallest spring alate viviparous female
(Essig and Abernathy, 1952).
 The Qualitative and Quantitative Analysis of Variation 161

8B.2 NON-SEX-ASSOCIATED INDIVIDUAL VARIATION

This term is simply one of convenience applied to intrapopulation

variation which is not sex-limited or does not primarily involve sex
characters.
8B.2.1 Continuous Variation. The most common type of individual
variation is that which is due to the slight genetic differences which exist
between individuals. No two individuals (except monozygotic twins) in
a population of sexually reproducing animals are exactly alike, genetically
or morphologically. One of the outstanding contributions of population
genetics has been the establishment of this fact. The differences are in
general slight and are often not discovered unless special techniques are
employed.
The study of this variation is one of the foremost tasks of the tax¬
onomist. It is now evident that no one individual is “typical” of the charac¬
ters of a population. Only the statistics of the whole population can give
a true picture of the population (Chap. 8C). A model study of variability,
based on 2,877 skins of the house sparrow (Passer domesticus) is presented
by Selander and Johnston (1967).
Each character may show a different degree of variability within
a single population. Likewise there are different degrees of variability among
related species. Just why one species should be highly variable and another
one not is not always clear. A taxonomist who has adequate material con¬
cerning one species should not hastily assume that this would permit him
to be certain of the variability of related species.
Some early taxonomists vastly underrated individual variation in many
genera of animals. The species of the snail genus Melania (fresh and brack¬
ish water) have been described largely on the basis of shell characters,
such as the presence or absence of spines and of diagonal and spiral ribs.
However, spined and spineless specimens occur in the species in M. scabra,
M. rudis, and M. costata, sculptured and smooth specimens in M. granifera,
and so forth. In a revision of this genus, no less than 114 “species” were
found to be nothing but individual variants and had to be added to the
synonymy of other species (Riech, 1937).
8B.2.2 Discontinuous Variation (Polymorphism). The differences
between individuals of a population are, in general, slight and intergrading.
In certain species, however, the members of a population can be grouped
into very definite classes, determined by the presence of certain conspicuous
characters. Such discontinuous individual variation is called polymorphism.
Frequently such polymorphism is controlled by a single gene, subject to
simple Mendelian inheritance.
Polymorphism is more pronounced in some groups of animals than
162 Methods of Zoological Classification

Table 8-3. Mimetic Polymorphism in West African Papilio dardanus

Brown (after Goldschmidt, 1945)

Nonmimetic Mimetic
Male Models
females females

Typical dardanus Basic type $ sim¬ hippocoon Fabricius Amauris niavius Lin¬
ilar to O’ naeus
dionysus Doubleday trophonissa Aurivil- Danaus chrysippus
and Hewitson lius Linnaeus
niobe Aurivillius Bematistes tellus
Aurivillius

in others. The spotting in lady beetles (Coccinellidae) is a well-known

example of genetic polymorphism, as is industrial melanism in moths. Poly¬
morphism has great biological importance, since it proves the existence
of selective differences between apparently neutral characters. For a more
detailed discussion of the topic, see Ford (1945, 1966) and Mayr (1963).
Its practical importance to the taxonomist is that it has led to the descrip¬
tion of many so-called “species” that are nothing but polymorphic variants
(morphs). In ornithology alone, about 100 species names were given to
polymorphs. The establishment of their true nature has led to a considerable
simplification of taxonomy.
Perhaps the most spectacular cases of polymorphism are to be seen
in the Lepidoptera and more particularly in certain species of butterflies.
The common alfalfa butterfly, Colias eurytheme (Boisduval), for example,
has two strikingly different female forms, one resembling the orange-colored
male while the other is largely white. The most complicated cases of sex-
limited polymorphism which have been studied genetically are the examples
of mimetic polymorphism in African swallow-tail butterflies of the genus
Papilio. Quite apart from the fact that allopatric populations throughout
Africa show distinct subspecific differences which are correlated with differ¬
ences in the species of butterflies which they mimic, we find that several
distinct female forms exist within a single population. Thus in West Africa
one finds, in the same population of Papilio dardanus Brown, one male
form and five female forms, three of the latter mimicking different models
which belong to the families Danaidae and Nymphalidae (see Table 8-3
and Fig. 8-6). The most remarkable feature of this polymorphism is that
although the various forms are so distinct as to resemble representatives
of three different families of Lepidoptera, breeding experiments have shown
that the differences are caused by a few Mendelian genes. Another cele¬
brated case is that of Pseudacraea eurytus Linnaeus (Carpenter, 1949).
 The Qualitative and Quantitative Analysis of Variation 163

Fig. 8-6. Mimetic polymorphism in the Papilio dardanus complex. A, male of cenea—
also basic type of nonmimetic female, ground color yellow; B, dionysus, nonmimetic
female, group color of forewings white, hind wings yellowish; C, trophonissa, mimetic
female, ground color for forewings white, hind wings brownish; D, hippocoon, mimetic
female, ground color white (redrawn from Eltringham, 1910, by Goldschmidt, 1945).

8G. STATISTICAL ANALYSIS OF INDIVIDUAL

VARIATION

8C.1 TAXONOMY AND ELEMENTARY STATISTICS

It may be useful to a beginner to single out for discussion a few

terms and elementary methods in statistical analysis that are routinely used
in taxonomy. These discussions are not meant as a substitute for the techni¬
cal literature. In order to find out how to calculate standard deviations
or regression coefficients, one should consult Simpson, Roe, and Lewontin
(referred to as SRL in the ensuing discussions) or a more advanced text.
164 Methods of Zoological Classification

The replacement of typological thinking by population thinking has

had a far-reaching impact not only on the concepts of taxonomy but also
on its methods. To consider taxa as populations and aggregates of popula¬
tions led automatically to a statistical approach. The taxonomist deals with
samples from natural populations and can make estimates on the character¬
istics of these populations only with the help of statistics.
A quantitative analysis adds greatly to the precision of a description.
Actual measurements of a series of specimens are far more useful than
the meaningless statement “of medium size.” “Sex comb with seven teeth”
is more precise than merely “sex comb present.” Such precision is all the
more important whenever related species differ not by the presence or
absence of a character, but rather by its size, proportions, or number. When¬
ever we compare two populations or two other taxa, we cannot make a
concrete statement on the degree of difference until we have determined
the amount of variation within the compared entities.
Statistical methods should be used whenever they contribute to the
taxonomic analysis. They are not to be used as window dressing. Taxonomic
papers are occasionally published with highly elaborate statistics that make
no contribution whatsoever to the taxonomic analysis. To undertake the
calculations is pure waste of time in such a case. Statistics cannot improve
heterogeneous original data or unreliable measurements. The simplest
method yielding the desired information is always the best.
A detailed presentation of the principles of statistics and of the ap¬
plication of the various statistical methods is beyond the scope of this man¬
ual of taxonomy. Many excellent texts are available. The best for the pur¬
poses of the animal taxonomist is Quantitative Zoology by Simpson, Roe,
and Lewontin (1960), referred to in this text as SRL. The splendid bio¬
metric analysis of Nigerian ostracods by Reyment (1960, 1963) is a good
introduction to an application of modern methods, particularly of multi¬
variate analysis. Seal (1964) gives an exposition of some of the more com¬
plex multivariate methods. Special applications to paleontology are de¬
scribed in Miller and Kahn (1962). The coming of the computer has
greatly facilitated the use of many of the more elaborate methods. Com¬
puter programs are now available for many methods for which the calcula¬
tions used to be altogether forbidding. See also Sokal (1965).
Statistical methods never produce an automatic answer to questions
of taxonomic evaluation. In this respect the situation is quite different
from that in experiments where the worker is often satisfied with a simple
“yes” or “no” answer to the question whether a change in experimental
conditions has resulted in a “significant” difference from a set of controls.
However, statistical analysis may give us important information on
the nature of our data. Experience has shown that morphological measure¬
ments usually show a normal distribution. A strong deviation from normality
 The Qualitative and Quantitative Analysis of Variation 165

requires that we examine the material for “bias” and the possibility of
“heterogeneity” (see statistics texts). Statistical analysis may also give us
important information as to the weight which we should assign to certain
characters. Highly variable characters, as well as some that are closely
correlated with other characters, are given low weight in classification.
What Statistics Cannot Do. Statistics is no cure-all. It cannot make
taxonomic decisions for us. Statistical methods do not tell us whether two
phena belong to the same population or not, whether two allopatric popula¬
tions belong to the same species or represent two species, or whether two
sympatric phena are individual variants or sympatric species. Reproductive
isolation (species criterion) and degree of morphological difference are
not always closely correlated. Sibling species may be almost identical mor¬
phologically, while intrapopulation variants and subspecies are often strik¬
ingly different. The taxonomist must also keep in mind that what may
be “significant” for the statistician may not be at all significant biologically
or at least taxonomically.
8C.1.I Samples and Sampling. It is rarely, if ever, possible to study
an entire natural population. The taxonomist must be satisfied with a sam¬
ple from which he attempts to reconstruct the properties of the population
from which the sample was drawn. In order to permit reliable conclusions,
the sample should be homogeneous, adequate, and unbiased (Cochran,
1953).
A heterogeneous sample can often be segregated into smaller homo¬
geneous samples by separating the specimens according to age, sex, locality,
or other factors that had introduced heterogeneity. Great care must be
taken when segregating a sample to avoid bias (see below). Homogeneity
is particularly important in comparative studies because samples which
differ in their components owing to heterogeneity cannot be legitimately
compared.
The sample should not be biased; that is, the method of getting the
sample should be such that the variations of the pertinent characters in
the sample occur at the same frequency as in the population. Fossils, for
instance, are sometimes deposited according to size, and a sample drawn
from one of these size classes is not an unbiased representation of the
population from which it is drawn. Collectors sometimes concentrate on
unusual specimens and thus introduce bias. Instead they should adopt the
same safeguards of randomization as experimentalists and pay due respect
to the variation introduced by locality, season, and time of day. One should
never discard part of a collection and keep only those specimens that seem
either typical or particularly interesting for being atypical. In polymorphic
populations special efforts should be made to collect specimens in the true
population frequency. In order to reduce collecting bias, it is often advisable
to employ several different collecting techniques at the same locality.
166 Methods of Zoological Classification

What constitutes an “adequate” sample depends on the nature of

the studied taxon and on the objective of the investigation. In the case
of the living coelacanth, Latimeria, a single specimen was adequate to
prove that this was a new species and genus of a class of vertebrates believed
to have been extinct for more than 70 million years. In some groups of
animals diagnostic species characters arc well defined in only one sex, and
the description of a new species may not be feasible if the collection contains
only representatives of the other. The study of polytypic species requires
a comparison of samples from different populations which may differ only
in quantitative characters. Large samples are a necessity in such studies.
This is also true for the study of variation in polymorphic species. As
a general rule one can say that an adequate sample is a sample which
allows a reasonable estimate of the total variability of a species. Whenever
knowledge of a group has reached the degree of maturity where taxonomic
analysis concentrates on a study of individual and geographic variation
of species, the availability of large samples for study is a necessity. A knowl¬
edge of the variability of species is valuable not only for the taxonomist
but for anyone dealing with the biology of species. The evolutionist in
particular, but also the ecologist and population biologist, are interested
in the nature and the extent of variation within and between populations
of species.
8C.1.2 Measurements and Counts. Only quantitative data can be
subjected to a statistical analysis. In this fact lies the importance of charac¬
ters that can be counted or measured. Meristic (= countable) characters,
such as number of spines, scales, or fin rays, permit greater accuracy than
do measurements. They arc therefore favored wherever possible—by stu¬
dents of echinoderms, fishes, and reptiles, for example. SRL (pp. 20-30)
gives various requirements of good measurements. Most important is that
measurements be standardized (applying to a specified distance) and accu¬
rate. For instance, the length of the bill in birds may be measured in
several ways: (1) from the nostril to the tip, (2) from the beginning
of the feathering to the tip, or (3) from the beginning of the bony forehead
to the tip. Observations have shown that the first can be measured very
accurately but does not give the full length of the bill; the third can
be measured fairly accurately in all birds with a steep forehead; and the
second can rarely be measured with any accuracy. Consequently, in some
genera of birds the third is the preferred measurement, in others the first.
In this case as well as in all similar cases, the record should show which
of several possible measurements was actually taken.
Having to take many measurements of a large scries of specimens
is very time consuming. Automation is beginning to come to the rescue
of the investigator, particularly through a computer-connected recording
of the measurements (Gain and Helmrich, 1967).
 The Qualitative and Quantitative Analysis of Variation 167

It is seldom possible to predict which of a set of possible measurements

will be most important in the comparison of several samples, and so it
is advisable to measure all variates that may possibly be of importance.
Subsequent analysis will show that many of these measurements either fail
to show significant differences or are merely duplications of other data.
In view of the high costs of printing, such superfluous data should not
be published. They may be placed in the archives of a public institution
(museum or library) where they are available to other students.
When measuring an important lot of specimens, or when measuring
specimens before the method of doing so has been completely standardized,
one should measure each variate repeatedly. The duplicate sets of measure¬
ments should be taken on different days and on new record sheets. When
completed, the various sets of measurements should be compared and aver¬
aged. Particularly deviating measurements should be checked for possible
errors in the measuring technique or in recording.
8C.1.3 Measurable Characteristics. Total length is usually a very
important measurement, particularly when if is used as the yardstick for
ratios and proportional measurements. In each case what is meant by total
length should be specified. Is it taken before or after preservation? Does
it include or exclude appendages on the head and the tail? Total length
is most satisfactory in beetles and other rather rigid, hard-shelled animals.
In birds, the wing length (= actually the length of the longest primary)
is a much less variable quantity than is total length measured in the flesh.
The cube root of the weight may, under certain conditions, replace the
total length in calculations of allometric ratios. Body length, i.e. total length
minus tail length, is usually a more accurate measure of size than total
length.
Different measurements are used for nearly every category of animal.
In mammals, for instance, body and tail length are measured, as well as
length of the hind foot and ear and the various dimensions of the skull.
In birds, wing, tail, bill, and tarsus are the most commonly measured vari¬
ates. In most groups of insects not only length but also width and antennal
and tarsal formulas should be given. These data should be recorded as
a routine matter regardless of their immediate diagnostic value. Special
measurements are traditionally given in particular taxonomic groups, such
as the length of the rostrum in Hemiptera, length of the wings in some
Diptera, etc. It is important for comparative purposes to give measurements
that conform with the system which is customary in the group under study
and to present them in a standardized sequence.
8C.1.4 Technical Aspects of Measuring. Zoological measurements
are now universally given in terms of the metric system. However, many
descriptions written in the nineteenth century use inches and lines
(1 line = -jig- in.). (See Table 8-4.)
168 Methods of Zoological Classification

Table 8-4. Conversion of Lines into

Millimeters

1 line = 2. Ilf mm 7 lines = 14,81-f mm

2 lines = 4.23f mm 8 lines = 16.93f mm
3 lines = 6.35 mm 9 lines = 19.05 mm
4 lines = 8.46-f mm 10 lines = 21. 16f mm
5 lines = 10.58f mm 11 lines = 23.28f mm
6 lines = 12.7 mm 12 lines = 25.4 mm

Various measuring tools are used for different groups of animals.

A millimeter rule (often with a “zero stop”) and dividers (calipers) are
used for most larger animals. The eyepiece micrometer is used to measure
microscopic objects. It may be divided into small or large units and may
be arranged as a linear scale or in squares. The individual units must
be translated into the metric system by calibration with a stage micrometer.
Projection devices, such as microscopic projectors, are sometimes use¬
ful. By means of such devices, the specimens can be drawn from a projected
outline, and the various parts can then be measured in the enlargement.
This method is particularly useful for the measurement of relative sizes
and angles. Care should be taken to avoid the danger of distortion in
the two-dimensional projection of the three-dimensional object.
As far as refinement is concerned, it is important to carry measure¬
ments to whatever decimal point may be necessary, but not to waste effort
by an unreasonable desire for accuracy. It would be useless to give the
height of a person as 176.583 cm.
How, then, shall the proper degree of refinement be decided? The
recommended unit of measurement is one-twentieth of the difference be¬
tween the largest and the smallest specimen, if an adequate series is avail¬
able. Thus if the measurements range from 10 to 12 mm, one should mea¬
sure to 0.1 mm; if they range from 40 to 50 mm, to 0.5 mm. If they
range from 70 to 90 mm, no decimal places need be recorded. If fractions
are rounded off, they should consistently be rounded to the nearest full
number, halves to the nearest even number. When fractions are measured,
a bias in favor of integral numbers should be avoided.
8C.1.5 Recording of Measurements. Whenever large numbers of
measurements are taken, it is advisable to enter them on special data sheets.
If adequate samples are available, each sample should be recorded on a
separate sheet. Each specimen should be entered separately, its field or
museum number (if any), age, and sex recorded, and then the various
measurements recorded in separate columns. If there is room, the calculated
ratios between measurements can be entered on the same sheets.
Color differences are important characters in the taxonomy of virtually
all animals. Various instrumentations are used to determine the color values
 The Qualitative and Quantitative Analysis of Variation 169

objectively and to translate them into precise measurements of reflected

light of selected wavelengths (Chap. 10). This permits a statistical analysis.

8C.2 STATISTICS OF A POPULATION SAMPLE

The beginner often asks, “What statistics should I use?” When one
deals with a single character (univariate analysis), the two essential statistics
are the arithmetic mean, M, and the total variance, V (or the square
root of the variance, designated as the standard deviation, SD). With these
two values, one can calculate most other statistics.
If the values of a sufficiently large sample are plotted, it is usually
found that the resulting frequency curve corresponds to the so-called normal
curve (see statistics texts). The reason why most biological characters seem
to show the pattern of variation of the normal curve is probably that
they depend on a great number of genetic factors each of which makes
either a positive or a negative contribution to the phenotype of the charac¬
ter. For example, literally hundreds of genes tend to increase body size
and a similar number to decrease it. The probability is very low that an
individual will have all plus genes or all minus genes; most individuals
will have a balance between the opposing tendencies. This is why the major¬
ity of individuals of a population are relatively close to the mean value.
8C.2.1 Standard Deviation (SD). The standard deviation measures
the variability of a sample. The broader the scattering of values around
the mean, the “flatter” the curve, the greater the standard deviation.
Knowledge of the SD of a population permits predictions as to the observed
range of variation, because in a normal curve

M + 1 SD includes 68.27% of the population

M + 2 SD includes 95.45% of the population
M + 3 SD includes 99.73% of the population

With desk calculators universally available, the standard deviation

is now calculated by a machine method, as described in SRL, p. 87.
8C.2.2 Coefficient of Variability. The numerical value of the SD
is meaningful only in relation to the mean value of the same sample. An
SD of 2 indicates extremely low variability if the mean is 120, but very
high variability if the mean is 8. In order to permit a comparison of degrees
of variability in different species, it is advisable to calculate the coefficient
of variability (CV). This expresses the standard deviation as percentage
of the mean:

SD X 100
CV
M
170 Methods of Zoological Classification

The numerical value of the CV depends on the measured character

and on the particular taxonomic group. There are different CVs for meristic
quantities, linear measurements, and ratios. The number of eyes (a meristic
quantity) in the human species has a CV that is virtually 0; height in
man (even in a homogeneous sample) has a CV exceeding 4. The CV
is often a sensitive indicator of the homogeneity of samples. If, for instance,
the CV of a certain statistic fluctuates around 4.5 in a series of samples,
but is 9.2 in one sample, such a sample should be reinvestigated. It may
include an additional sibling species, wrongly sexed specimens, or some
other alien component. Zones of secondary intergradation between subspe¬
cies are often characterized by a greatly increased CV. The calculation
of CV is particularly useful when comparable samples of the same species
from different localities are investigated or when the variability of different
variates of the same sample is compared. Even though widely used by
taxonomists, the coefficient of variability has the great disadvantage that
there are no statistical tests for comparing different CVs. Lewontin (1966),
following earlier authors, has therefore proposed that the variance (or the
standard deviation) of the logarithms of measurements be used instead.
This measure of intrinsic variability is invariant under a multiplicative
change. This means also that it does not matter what units of measurements
(metric or nonmetric) are used. A simplified approach is possible for CVs
of less than 30 (which includes most that are used in taxonomy). Here
the squared CV equals the variance of the natural logarithms (to the
base of e) and can be used in statistical tests.
8C.2.3 Linear Measurements. Absolute size of the whole body or
of selected measurements of parts is remarkably invariant in adult mammals
and even more so in adult birds.
A series of 49 adult males and 29 adult females of the kingfisher
Halcyon chloris pealei Finsch and Hartlaub from Tutuila Island, Samoa,
had the measurements and coefficients of variability shown in Table 8-5.
In carefully measured homogeneous samples of adult birds, the CV of
wing length is usually between 1 and 2.5, rarely above 3. In mammals
the CV for linear dimensions is usually between 4 and 10, occasionally
between 3 and 4.
In insects which reach the imago stage through molt or metamorpho¬
sis, a small CV of linear measurement might be expected, since there is
no further growth after the sclerotic exoskeleton has hardened. However,
the final size of the imago depends to some extent on the feeding conditions
of the larvae or nymphs, and as a result there is usually considerable vari¬
ability in the size of the imago. Linear measurements are extremely variable
in all animals that continue to grow throughout life, such as fishes and
snakes, not to mention such forms as corals and bryozoans. Employment
of mean values and coefficients of variability are rather meaningless in
 The Qualitative and Quantitative Analysis of Variation 171

Table 8-5. Measurements of a Sample of Halcyon chloris pealei from

Tutuila, Samoa

N Range Mean SD CV

Adult males:
Wing 49 94.0-101.0 97.48 1.71 1.75
Tail 49 63.5- 69.5 66.44 1.33 2.00
Bill 49 31.0- 39.0 34.46 1.56 4.54
Adult females:
Wing 29 95.5-102.5 98.86 0.88 0.90
Tail 29 64.0- 72.0 67.62 1.56 2.29
Bill 28 33.5- 37.5 35.20 0.95 2.69

such cases. Instead, bi- and multivariate analysis, particularly multiple-

regression analysis, yield statistics that are useful in taxonomic comparisons.
8C.2.4 Meristic Quantities. When countable characters, such as the
number of segments, scales, spines, or chaetae vary, we speak of meristic
variation. Some meristic characters are exceedingly constant, such as the
number of eyes in mammals; others may have a greater or lesser variability
which is usually characteristic for a given species. Examples are the number
of scales in snakes or fin rays in fishes. Meristic characters are not always
strictly discontinuous, particularly at the beginning or end of a series, so
that the total number may have to be decided somewhat arbitrarily.
The CV of most meristic characters is smaller than that of linear
measurements, and it is not permissible to compare the CV of the two
kinds of variates. One should only compare relatively equivalent sets of
data, such as the coefficients of linear measurements with those of other
linear measurements, ratios with ratios, etc.
8C.2.5 Ratios. The statistics so far discussed deal with the variation
of absolute size of single characters. Shape and proportions, however, are
often more important in taxonomy than absolute size. Ratios between two
dimensions or two parts of the body are therefore often more informative
than linear measurements. The ratio is most meaningful if it is given in
a formula which expresses the size of the smaller value as a percentage
of the larger:

s X 100
R (ratio) =
/
where s equals the smaller of the two values and l equals the larger. Com¬
monly used ratios are the length of the head against length of the body
(without head), length of the tail against length of the body, width of
172 Methods of Zoological Classification

the skull against length, etc. Such ratios are quickly calculated with the
help of a slide rule. If R is near 100, it may happen that s is larger
than l in some samples. It is obvious that the positions of s and l cannot
be reversed in such cases even when R becomes larger than 100.
Ratios are best demonstrated visually in the form of scatter diagrams
in which one value is plotted on the abscissa, the other on the ordinate
(Fig. 8-7).
It is important that the proper standard of comparison is chosen
when one wants to determine the relative size of an organ or appendage.
For instance, relative head width in insects is calculated against head length
(without rostrum). Relative tail length in birds is usually calculated against
wing length (as standard of general size). However, the wing is not an
accurate yardstick for size in migratory and high-altitude birds, nor in
some birds in which the wing is used in courtship. The cube root (^/ )
of body weight might be a better measure in such species. If an appendage
is calculated against the whole, as tail against body, the appendage should
not be included in the whole; the trunk without the tail should be used
as standard of the “whole.”
A ratio has a typological connotation, as does the mean. It may there¬
fore be a misleading way of expressing dimensional relationships when our

Fig. 8-7. Separation of two subspecies of Parus carolinensis on basis of wing and tail length;
triangles = agilis, circles = atricapilloides, and AB = line of best separation (from Lunk, 1952).
 The Qualitative and Quantitative Analysis of Variation 173

RELATIVE LENGTH OF TAIL

Fig. 8-8. Increase in the depth of the tail fork (in millimeters)
correlated with increase in relative length of tail (in percentage
of wing length) in Dicrurus leucophaeus. Each point indicates
the mean for the available sample of adult males of one of 14
subspecies (from Mayr and Vaurie, 1948).

sample is highly heterogeneous with respect to age and size or when the
various compared body parts display allometric growth. In such cases it
is better to undertake a regression analysis (SRL, p. 213) and in the case
of allometric growth to plot the variation on logarithmic paper. Good
straight-line relations are sometimes obtained by plotting absolute size
against the relative size of a body part (Fig. 8-8).
In the case of ratios and regressions we are comparing two variates.
Whenever we want to deal simultaneously with more than two variables,
we must employ some method of multivariate analysis.
174 Methods of Zoological Classification

Vagrans

Biscutatus

Elegans (Lassen County)

Elegans (Oregon, California

West of Sierran Divide)
Elegans (San Bernadino Mts.)

Hydrophila (Humboldf and

Mendocino Counties)
Hydrophila (Umpqua, Rogue
and Klamath River Basins)

Couchii

Hammondii

Atratus

Ordinoides

Fig. 8-9. Histograms showing head and body length in centimeters of adult males of Thamnophu
ordinoides. Each square represents a specimen (Filch, 1940).

8C.2.6 The t Test. Statistically this is the most satisfactory method

of determining whether or not the difference between two samples is “sig¬
nificant” (see SRL, p. 176). In cases where a multivariate analysis is em¬
ployed, the corresponding test is called the T2 test.
8C.2.7 Comparison of Frequencies. The taxonomist is often con¬
fronted with the problem of having to determine whether two morphs
or other variants occur at the same frequency in two or more populations.
If he had infinitely large samples, he would simply express frequencies
in percentage. With samples of limited size, a test is employed that tells
 The Qualitative and Quantitative Analysis of Variation 175

us what the probability is that the populations from which the samples
are taken are really different or, respectively, what the probability is that
the observed differences are merely due to accidents of sampling. This
test is called the (chi-square) test (SRL, pp. 306-338).
The degree of significance of a given ^2 value is given in P tables,
which can be found in all standard statistical texts. ^2 tests are highly
sensitive to sample size.

8C.3 GRAPHIC PRESENTATION OF QUANTITATIVE DATA

It is often desirable to present numerical data visually. Such a presen¬

tation by graphical methods not only permits a rapid visual survey of all
the data, but actually often brings out fine points that are not apparent
in the raw data. SRL, chap. 14, gives an excellent survey of such graphic

Localities from North to South

Fig. 8-10. Population-range diagram. Variation in the number of vertebrae of the anchovy,
Anchoviella mitchilli. The letters A to O refer to 15 population samples, arranged from north
(A) to south (O). In each sample the vertical line indicates the total variation of the sample;
the broad portion of the line, 1 SD on each side of the mean; the hollow rectangle, twice the
standard error on each side of the mean; and the crossbar, the mean (Hubbs and Perlmutter,
1942).
176 Methods of Zoological Classification

Fig. 8-11. Triangular graph of the length (L), height (H), and distance to maximum down-bulge
(D) of four species of Anthracomys (Burma, 1948).

methods. In the present text only those few simple methods will be men¬
tioned that are most frequently used in taxonomic publications.
8C.3.1 Histograms. Unreduced samples are best shown as histo¬
grams. A histogram consists of a set of rectangles in which the midpoints
of class intervals are plotted on the abscissa and the frequencies (usually
number of specimens) on the ordinate. This presentation has several ad¬
vantages, the principal one being that it presents the original data in mini¬
mum space. Whatever form of statistical analysis a subsequent author may
want to apply, he will find the actual number of specimens given for each
size class. A quick comparison of different populations is made possible
by arranging a series of histograms above one another (Fig. 8-9).
8C.3.2 Population Statistics Diagrams. Even more data can be com¬
pressed into minimum space by giving sample range, mean, one or more
SDs, and two standard errors. This is the method of Dice and Leraas
(1936) (see Fig. 8-10). Several modifications of this method have been
proposed. For instance, one can give the size of the sample (N) with
 The Qualitative and Quantitative Analysis of Variation 177

each bar and replace standard error by the 95% confidence limits of the
mean. Nonoverlap of 1.5 SD (of each compared sample) indicates a degree
of difference usually considered sufficient for subspecific separation, as dis¬
cussed later.
8C.3.3 Scatter Diagrams. The difference between two or more popu¬
lations in respect to two characters is best illustrated by a scatter diagram.
Each individual is indicated by a spot or other symbol which is placed
where the value for one character (read off the ordinate) intersects the
value for the other character (read off the abscissa) ; each population
is indicated by a different symbol (circles, squares, triangles, solid or empty,
etc.-—see Fig. 8-7). Scatter diagrams have many advantages. They help
the student to visualize allometric relationships and facilitate the plotting
of regression lines. They also sometimes disclose errors of measurement
or sexing that might otherwise go undiscovered.
If three characters are involved, triangular charts can be employed.

0-4 5-9 10-14 15-19 20-24 25-29 30-34 35-39 40-44

NUMBER OF HAIRS PER WING

Fig. 8-12. Pictorialized scatter diagram (pictogram) of 25 individuals from a population of stemless
white violets. Two of the characters are indicated along the margins, five by position and
length of rays; filled circles indicate individuals with a heavy blotch on the spur petal (from
Hatheway, 1962, after Anderson, 1954).
178 Methods of Zoological Classification

A = PETAL VENATION INDEX

Fig. 8-13. Weighted scatter diagram, ordering the same 25 individuals shown in Fig. 8-12. The
horizontal scale is a weighted index composed of four aspects of petal venation. The vertical scale
combines five measures of hairiness (from Hatheway, 1962).

In this case the actual values are not plotted, but rather their percentage
contribution to the sum of the characters. For example, if character
a = 80 mm, b = 32 mm, and c = 48 mm,

a b c = 160 mm = 100%

Then a = 50%, b = 20%, and c = 30% of the whole. These percentages

are plotted on the graph, which thus shows proportions rather than absolute
sizes. In each individual case the triangular graph is scaled in such a way
as to produce a maximum spread of the points. As an illustration a triangu¬
lar chart from a paper by Burma (1948) is reproduced in Fig. 8-11.
More than three characters can be presented in a pictorial scattergram
(Anderson, 1949, chap. 6) (Figs. 8-12, 8-13). It is advantageous to select
those two characters for plotting on ordinate and abscissa which result
in the clearest separation of the clusters.
Fig. 8-14. Map of the distribution of the tiger beetle Cicindela punctulata, illustrating the pie-graph method. The size of the sectors of each circle in¬
dicates the relative frequency in the collected samples of three variants: punctulata, chihuahuae, and intermediates. Many populations are pure punctulata,
others are pure chihuahuae (unpublished, courtesy of Dr. M. A. Cozier).
180 Methods of Zoological Classification

8C.3.4 Mapping of Quantitative Data. It is often desirable to illus¬

trate the geographical relationships of populations which differ in quantita¬
tive characters. In the case of continuous characters (size, etc.), the simplest
method is to record the means of the various populations on a base map,
and if there is regularity to draw in the isophenes ( = lines connecting points
of equal expression of a character). For instance, if the means of a series
of populations in a species vary from 142 to 187, it is helpful to draw
in the isophenes of 140, 150, 160, 170, 180, and 190 (see Chap. 3, fig.
3-4). Subspecies borders frequently fall where the gradients are particularly
steep. A method for determining such contour lines is described by Lidicker
1962.
If qualitative or semiqualitative characters are to be plotted, it is
sometimes helpful to choose a different symbol for each class of characters.
The relative size of the symbol can be used to indicate sample size.
To present frequencies of polymorph characters on a map, the “pie
graph” is the most convenient method. The percentage occurrence within
the population is indicated by the size of the sectors (Fig. 8-14).
Chapter Taxonomic Decisions on
the Species Level

j.n his sorting of phena and populations into subspecies

and species the taxonomist is forever forced to make decisions. In this
chapter we shall discuss the various alternatives and suggest appropriate
methods (some statistical) that may help in reaching the right decision. A
study of the discrimination grid (Table 8-1) will facilitate asking the right
questions.
Doubts arise in the analysis of sympatric samples and in the compari¬
son of allopatric (or allochronic) samples. The two parts of this chapter,
9A and 9B, are devoted to these topics.

9A. ANALYSIS OF SYMPATRIC SAMPLES

The taxonomist is frequently in doubt whether certain phena sampled

in the same general area belong to a single species or to several. On the
whole there are three very different kinds of situations that are responsible
for these difficulties: (1) extreme difference of phena belonging to a single
species, (2) extreme similarity of good biological species (sibling species),
and (3) wide variability and phenotypic overlap of two species. The first
hint that—in the case of extreme difference—only one species is involved,

181
182 Methods of Zoological Classification

or—in the case of extreme similarity—more than one species is involved,

is usually provided by behavioral, ecological, or distributional data. The
subsequent analysis serves to confirm (or refute) the earlier supposition.

9A.1 PHENA (INDIVIDUAL VARIANTS) OR DIFFERENT

SPECIES?

There is hardly a species that does not contain several if not dozens
of phena. To add to the complexity of the situation several other species
with a similar assortment of phena may be sympatric. Often a phenon
of one species resembles a corresponding phenon of another species more
than other phena of the same species. For instance, females in certain
species of many genera of birds and insects are more similar to the females
of closely related species than to the males of their own species. There
is nothing in the phenotype of caterpillars that would permit correct asso¬
ciation with imagos. Only breeding can do this, or the careful evaluation
of other biological information.
The correct assignment of many phena is possible on the basis of
a correct interpretation of morphological information. If a large sample
of a population is available, intermediate forms between the more extreme
variants are usually found. Also, certain characters in every group are less
subject to individual variation than others. The genitalic armature in most
insects, the palpus in spiders, the radula in snails, and the structure of
the hinge in bivalves are such characters. If several sympatric phena agree
in their genitalic armatures (or one of the other mentioned characters),
it is very probable that they are conspecific. However, even here one has
to apply a balanced judgment. Although in most genera of Lepidoptera
there are characteristic differences between the genitalia of related species,
there are cases known in which forms have identical genitalia, even though
they are different species by every other criterion. Parasitic animals present
some rather special problems (Manwell, 1957).
The establishment of correlations is often very helpful. If two forms
which differ in character a can be shown to differ also in the less conspicu¬
ous and functionally unrelated characters b, c, and d, it becomes very
probable that they are different species. Some years ago Mayr (1940) found
that among birds identified as the southeast Asiatic minivet (Pericrocotus
brevirostris Vigors), some had the innermost secondaries all black while
others had a narrow red margin on these feathers. A detailed study revealed
that those birds with red on the innermost secondaries had seven additional
minor characters: a more yellowish red of the underparts, a different dis¬
tribution of black and red on the second innermost tail feather, a narrow
whitish margin along the outer web of the first primary, and four other
 Taxonomic Decisions on the Species Level 183

minor characters. Slight though they were, these characters were well cor¬
related with each other and with geographical and vertical distribution.
The conclusion that two full species were involved has since been confirmed
by several authors.
As a general rule, one finds that the decisions of a superior taxonomist,
when based on a careful evaluation of the morphological evidence, are
confirmed wrhen a species recognized by him is subjected to genetic tests
or to an evaluation of nonmorphological characters. A purely phenetic
approach, on the other hand, is unable to discriminate between phena
and species.

9A.2 SIBLING SPECIES

Biological species are reproductively isolated gene pools (Chap. 2).

When two populations (gene pools) become geographically isolated, they
diverge in their isolation genetically and may eventually acquire isolating
mechanisms. As a by-product of the genetic divergence during this process,
species normally also acquire morphological differences suitable for diagnos¬
tic purposes. A few species fail to acquire conspicuous morphological differ¬
ences during the process of speciation. Such very similar, cryptic species
are called sibling species. All the available evidence indicates that minute¬
ness of morphological difference is the only aspect in which they differ
from ordinary species. They are merely those species that are near the
invisible end of the spectrum of morphological species differences. They
grade imperceptibly into species that are morphologically more and more
distinct from one another. Once discovered, and thoroughly studied, sibling
species are usually found to have previously overlooked morphological
differences.
Mayr (1963, pp. 33-58) has shown how widespread in the animal
kingdom sibling species are. Most of them were discovered not during rou¬
tine taxonomic analysis but during the study of species that are medically
(e.g. Anopheles), genetically (e.g. Drosophila, Paramecium), cytologically,
agriculturally, or otherwise of special importance. It is therefore impossible
to indicate what percentage of species are sibling species. In the case of
North American crickets, about 50 percent of the species were discovered
through differences in their sounds (Walker, 1964) and in certain genera
of protozoans (e.g. Paramecium) the percentage of cryptic species seems
to be even higher.
The discovery of sibling species is possible because they may differ
in various attributes even when they are extremely similar in the morpho¬
logical characteristics normally employed in taxonomic analysis. Mayr
(1963, p. 50) has listed a number of characteristics that facilitate the
184 Methods of Zoological Classification

recognition of sibling species. Precise measurements sometimes display bi-

modal characteristics, and the two modes can be correlated with additional
characters. Very often there are differences in the number or structure
of the chromosomes, a fact which has led to the recognition of sibling
species in Drosophila, Sciara, Chironomus, Prosimulium, and other dip-
terans, as well as in orthopterans, beetles, and other insects. Various aspects
of behavior—such as differences in visual and vocal displays, nest construc¬
tion, breeding season, migratory behavior, prey selection, and host prefer¬
ence—have perhaps led to the discovery of more sibling species than any
other characteristic. Sibling species may differ in their pathogenicity (e.g.
Anopheles), or in their susceptibility to parasites and suitability to serve
as hosts. Various biochemical methods, particularly those testing protein
specificity, are suitable for checking on the probability of a real difference
between “stocks” discovered by one of the other methods.
Kohn and Orians (1962) point out that the ecological situation makes
the occurrence of sibling species more probable in some groups than in
others. Sibling species will be common whenever species are able to occupy
different niches in the same community without appreciable differentiation
in those morphological characters that are used by taxonomists in their
classification. This means, in particular, protozoans, small crustaceans, and
insects. Every case of sympatry among sibling species is a case where no
sympatric character divergence, in the sense of Darwin, has occurred in
taxonomically useful morphological characters.
Sibling species are obviously inconvenient to the museum taxonomist.
Specimens of such species cannot in some cases be recognized in preserved
material. However, since species are not the creation of museum taxonomists
but phenomena of nature, it is impossible to ignore their existence. The
museum worker will be unable in many cases to do better than label mu¬
seum specimens from a group of sibling species by the group name, e.g.
Anopheles maculipennis group. Once a sibling species is discovered, morpho¬
logical differences are usually found subsequently which will permit the
identification of preserved material. It is generally possible in a group of
sibling species to identify old type-specimens as to the particular species
to which they belong. Sibling species are fortunately rare among vertebrates
and other groups of special interest to the paleontologist. Whenever the
recognition of a sibling species depends entirely on the behavior of living
animals, it is of course impossible to recognize them in fossil specimens.
The paleontologist is forced to proceed under the assumption that the
genera he studies do not include sibling species.
Sibling species among protozoans are not formally described as species
and not designated with binominals. The approximately 16 species of the
Paramecium aurelia complex, for instance, are designated as varieties.
One particular class of sibling species, polyploids, raises special diffi-
 Taxonomic Decisions on the Species Level 185

culties. Polyploids are unable to engage in normal gene exchange with

individuals of different ploidy, and because of this reproductive isolation
they have the biological characteristics of good species. They may, however,
be morphologically indistinguishable, particularly if they are autopolyploids.
Fortunately polyploidy is rare in the animal kingdom and essentially re¬
stricted to genera with parthenogenesis. Occasionally there may be a secon¬
dary return to sexual reproduction—in the oligochaetes, for instance. It
is up to the specialist to decide whether or not to give taxonomic recognition
to what is, biologically, surely a species. In instances of morphological iden¬
tity such recognition would be unwise.
Until recently, polyploidy was believed to be restricted to invertebrates.
A number of parthenogenetic (or gynogenetic) populations have been
found, however, among fishes (several genera), salamanders (Ambystoma
jeffersonianum group), and lizards (Lacerta, Cnemidopliorus), and at least
some of these populations are triploid. On the basis of their morphological
and ecological differences (and absence of gene exchange), several of these
populations have been described as separate species.

9A.3 VARIATIONAL OVERLAP

Closely related species are sometimes so variable and their variation

so overlapping that no single character seems to have absolute diagnostic
value. A combination of characters usually permits the correct assignment
of all seemingly intermediate specimens. As Anderson (1954) perceptively
pointed out, mechanical reliance on a biometric analysis or on straight
diagnostic characters is often less effective than a purely intuitive approach
based on the totality of characters as revealed by inspection.
A combination of two characters is often sufficient for diagnosis. In
such cases it is simplest to plot the two characters on a “scatter diagram.”
This method usually yields several clusters of points when heterogeneous
material is involved (see Chap. 8C).
A number of multivariate methods are available when the bivariate
scattergram does not yield satisfactory results. Anderson’s pictogram
(8C.3.3) is one type of multivariate scattergram (Fig. 8-12).
In most cases there is no doubt as to the validity of species recognized
by these methods, but there may be uncertainty about certain specimens
which appear to be intermediate. Calculation of a character index is the
simplest quick method for placing such specimens (see Table 9-1). Since
this index was first proposed by Anderson (1936) and Meise (1936) to
evaluate the hybrid nature of intermediate specimens, it was originally
proposed as “hybrid index.” I prefer the broader term character index,
since the method is equally suitable for a quantitative treatment of non-
186 Methods of Zoological Classification

Table 9-1. Weighted Decimal Character Index

(after Meise, 1936)

Character Passer domesticus Passer hispaniolensis

Crown gray, 0 rust brown, 30

Ear coverts gray, 0 white, 10
Sides of nape gray, 0 black, 15
Flanks plain, 0 striped, 10
Other 0 35
Total 0 100

hybridizing species with overlapping variability. The method consists in

providing a series of states for each character in which the two species
differ. The typical condition in species A is always designated as 0, the
typical condition in species B most frequently as 2 and that of intermediate
specimens as 1. For 12 characters, essentially typical specimens of species
A may vary between 0 and 3, typical specimens of species B between 20
and 24, and hybrids between 8 and 16. Two refinements are possible. The
first is to allow for more intermediate states in the case of important charac¬
ters so that specimens may score anywhere between 0 and, let us say, 6
for such a character. The second refinement is to arrange the scale of
values in such a way that the maximal score adds up to 100. In this
way it is possible to express the similarity of intermediate specimens as
percentages. In order to illustrate the variation, it is advantageous to graph
it in the form of histograms. Hatheway (1962) shows how to construct
a weighted hybrid index (Fig. 8-13).
A far more precise method is the multivariate method of discriminant
functions (Fisher, 1936, 1938). It is based on the fact that in the discrimina¬
tion between two taxa, the greater the distance between the means and
the smaller the combined standard deviations of this character, the greater
will be the contribution made by a character. The method then consists
in calculating a factor (61, 62, . . . bx) which, when multiplied by the
numerical value of the character, will result in a maximal value. The calcu¬
lation of the 6 values on a desk calculator is laborious, but there are now
good computer programs which calculate the discriminant function from
the raw data. “Stepwise” programs are now available which permit the
elimination of characters that do not significantly contribute to the
discrimination.
The method of discriminant functions has been used increasingly in
taxonomy in recent years. It is particularly useful in the identification of
doubtful specimens and occasionally in the establishment of a well-defined
 Taxonomic Decisions on the Species Level 187

class of intermediate hybrids. Recent applications were made in studies

of Drosophila (Carson and Stalker), fish (Stone), birds (Storer), and mam¬
mals (Foster, 1965). Biihler (1964) shows, step by step, how a discrimina¬
tion analysis is carried out and how, in his example, it permits the un¬
equivocal assignment to the correct species of every skull belonging to two
exceedingly similar species of shrews. Kim, Brown, and Cook (1966) point
out how to employ discriminant functions to find the diagnostically most
useful characters.
The analysis described in the preceding paragraph makes the assump¬
tion that one already knows the essential characteristics of the respective
species and is mainly concerned in the assignment of dubious specimens.
There is always a residue of cases where the variation is too great and
discordant to lead to an unequivocal species separation. A good example
is provided by spiders of the species group Steatoda punctulata in which
different specialists at one time or another considered that one, two, four,
six, or eight species were involved (Levi, 1959). In such a case, larger
series from single localities are needed as well as the utilization of additional
characters. Each of the two to four species of the Steatoda punctulata
complex varies individually and geographically, and a sophisticated bio¬
metric analysis would be futile until better material permits the study of
the trends of this variation.
Several (on the whole rather complicated) methods of multivariate
analysis have been tested in recent taxonomic publications, particularly
in their application to fossil material. The impression one gets from a
study of these papers is that no method has been found that makes enough
of a contribution to the analysis to justify the very considerable labor in¬
volved in applying it routinely. The T2 test is perhaps most useful.

9B. COMPARISON OF ALLOPATRIC AND ALLOCHRONIC

SAMPLES

The taxonomist has a well-defined interest when he compares samples

drawn from different natural populations. He is looking for evidence that
will facilitate the decision whether or not to include the populations in
the same taxon and, if not, whether to rank them as different subspecies
or species. The question is never whether or not the compared populations
are completely identical. Population geneticists have demonstrated conclu¬
sively that no two natural populations in sexually reproducing animals
are ever exactly alike. To find a statistically significant difference between
several populations is, therefore, only of minor interest to the taxonomist;
he takes it for granted. Even the lowest recognizable taxon, the subspecies,
188 Methods of Zoological Classification

is normally composed of numerous local populations some of which differ

“significantly” in gene frequencies and in the means of a number of variates.
If no statistically significant differences between the samples from
two populations are found, the sampled populations must be referred to
the same taxon. If a difference is established, additional considerations
are required before it can be decided whether or not the populations should
be considered different taxa or not, and, if different taxa, whether they
should be ranked subspecies or species.

9B.1 DIFFERENT SUBSPECIES OR NOT?

In groups of animals in which polytypic species are generally recog¬

nized, it is frequently questioned whether or not the difference between
two populations is sufficiently great to justify their recognition as two differ¬
ent subspecies. The most important prerequisite for making such a decision
is a clear understanding of the nature of the subspecies category (see Chap.
3, also Mayr, 1963, pp. 347-351). If the samples are clearly different in
one or several respects, they qualify as different subspecies (but see below).
A problem arises when the ranges of variation overlap. How much overlap
between two subspecies is permissible? The simplest way of determining
overlap is to plot the linear overlap of the observed samples (Fig. 9-1),
but this method is misleading in two ways: it gives only the overlap of
the samples (which is always smaller than that of the sampled populations),
and it exaggerates the importance of the end points of the range, that
is, the tails of the overlapping curves. Linear overlap is therefore a very
unsatisfactory way of describing the degree of difference between two
populations.
Is there a more satisfactory way? As far as we know, there is no
method that is not vulnerable to objections. Mayr (in Oliver, 1943) pro¬
posed a “coefficient of difference,” which relates the difference between
the means to the standard deviations of the samples.

100 105 B NO 115

Fig. 9-1. Linear overlap of observed samples. A = 99-106, B = 104-114 mm.
 Taxonomic Decisions on the Species Level 189

9B.1.1 Coefficient of Difference (CD). T his statistic is based on

the observation that the less overlap there is between two population curves,
the larger is the difference between the means M when divided by the
standard deviations (SD). The formula for the coefficient of difference
for populations a and b is (b being the population with the larger mean) :

M b — Ma
CD = ----
SD a + SD b

The CD is easy to calculate when one knows the means and standard
deviations of the two populations. However, it allows only a rough approxi¬
mation because it makes various assumptions that are not strictly correct,
e.g. that the distributions are normal and that the sample statistics equal
the population statistics. The latter is clearly not true, particularly for stan¬
dard deviations of comparatively small samples. A number of authors have
therefore suggested various modifications. Ride (1964) suggests calculating
two CD’s, one for the upper and one for the lower confidence limit. Another
suggested refinement is to use the square root of the pooled variances.
Gery (1962) believes that it would be simpler to base degree of difference
between population samples on a t test and supplies tables for this purpose.
Reyment (1960, p. 28) proposes a multivariate extension of the coefficient
of difference.
The reason for the uncertainties is that statisticians have until recently
been mainly interested in determining presence or absence of a difference.
They have paid little attention to methods establishing a degree of differ¬
ence. In taxonomic work, particularly in the recognition of subspecies, so
many additional considerations enter the picture that extreme accuracy
is not important, while a measure that would give the order of magnitude
of the overlap of two population curves is. The CD (or one of the cited
alternate methods) is a simple answer for this need. However, computer
methods have now become available that permit the routine calculation
of measures of distance, like Mahalanobis’ D2, which are far more accurate.
See Seal (1964) for an introduction to this method.
9B.1.2 Degree of Difference and Subspecies Recognition. Degree
of difference is only one of a number of considerations in the recognition
of subspecies (Chap. 3). A yardstick such as the CD will help to achieve
more uniform standards, but other information, such as degree of isolation,
presence or absence of clinal variation, presence or absence of a checker¬
board type of distribution, or discordant variation of different characters,
must be equally taken into consideration.
Widely different standards of subspecies recognition have been
adopted by different authors. Some extreme “splitters” called every popula¬
tion a different subspecies that could be shown by statistical tests to be
190 Methods of Zoological Classification

different. Some “lumpers” insist that every individual of a subspecies must

be diagnostically different. Very few taxonomists hold with either extreme.
A so-called 75-percent rule is widely adopted. According to this, a popula¬
tion is recognized as a valid subspecies if 75 percent of the individuals
differ from “all” (= 97 percent) of the individuals of a previously recog¬
nized subspecies. At the point of intersection between the two curves where
this is true, about 90 percent of population A will be different from about
90 percent of the individuals of population B (to supply a symmetrical
solution). This corresponds to a CD of about 1.28 (Table 9-2). From
this table one can readily see how much overlap there is between curves
at given coefficients of difference.
In the heyday of subspecies splitting in ornithology, some authors
recognized subspecies on the basis of a CD of only 0.675 or even lower.
There is now a tendency to demand a CD of well over 1.28, perhaps
at least 1.5, in order to justify subspecies recognition. In cases involving
irregular distribution patterns, position on a long cline, and discordant
variation of different characters, an even higher CD is sometimes insuffi-

Table 9-2. Percentage of Nonoverlap of Partially Overlapping

Curves Associated with Given Values of the
Coefficient of Difference (CD)

Joint
Values CD nonoverlap,
percent

Below the level of conventional subspecific difference 0.675 75

0.84 80
0.915 82
0.995 84
1.04 85
1.08 86
1.13 87
1.175 88
1.23 89

Conventional level of subspecific difference 1.28 90

Above the level of conventional subspecific difference 1.34 91

1.405 92
1.48 93
1.555 94
1.645 95
1.75 96
 Taxonomic Decisions on the Species Level 191

Fig. 9-2. Discordant geographic variation in the salamander Plethodon jordani. Darkened areas
represent regions where more than 95% have: (A) red cheeks, (B) red legs, (C) dorsal red spots in
newly hatched young, (D) lateral white spots, (E) a dark belly, (F) small dorsal brassy flecks. An
area with small dorsal white spots is indicated in (F) by stippling (from Highton, 1962).
192 Methods of Zoological Classification

Fig. 9-3. Geographic variation in the frequency of 16 color morphs in 7 populations of the wolfspider
Geolycosa xera. Ventral color pattern on legs 1 and 2 of females is shown above. Populations 1-4
belong to a different subspecies from populations 5-7 (from McCrone, 1963). Subspecies often differ
from each other in the relative frequency of phena.

dent. Highton (1962) demonstrates that geographic variation in the

salamander Plethodon jordani is too discordant to justify the recognition
of formal subspecies even though the variation of each individual character
shows a definite geographic trend (see Fig. 9-2). Inger (1961) gives
a well-balanced discussion of various criteria to be used in the recognition
of subspecies. Degree of difference, as expressed in the coefficient of differ-
 Taxonomic Decisions on the Species Level 193

ence, is only one of them. The standards for subspecies recognition are
now much more rigorous than they were a generation ago (see also
Chap. 3).
9B.1.3 Subspecies Borders. Most subspecies are geographical isolates
or former isolates, and their borders are easily established. The delimitation
of subspecies that are connected by primary intergradation is difficult, and
their recognition usually unwise. Such subspecies are often the adaptive
response to regional climatic conditions (particularly of temperature and
humidity) and are no more sharply delimited than the causative climatic
factors. Where substrate races are involved (black lavas, white sands or
limestones, red soils, etc.), subspecies borders are sometimes remarkably
sharp, particularly when reinforced by habitat selection. Lidicker (1962)
plotted on a map the total character change in a kangaroo rat
(Dipodomys merriami) per unit of distance and found that this resulted
in well-defined contour lines. Bands of rapid character changes, as defined
by an index of differentiation, coincided remarkably closely with the previ¬
ously recorded subspecies boundaries. In other species, however, no well-
defined subspecies borders seem to exist, as shown by Hagmeier (1958)
for the marten (Martes americanus) and by Jolicoeur (1959) for the wolf
(Canis lupus), two species in which the geographical variation of every
character seems to be independent of all others.
9B.1.4 Polytopic Subspecies. When subspecies of a species differ
only in a single diagnostic character relating to color, size, or pattern,
it may happen that several unrelated and more or less widely separated
populations independently acquire an identical phenotype. The evolutionist
knows that such populations are not identical genetically, but since the
subspecies is not an evolutionary concept, taxonomists sometimes combine
such visually identical populations into a single subspecific taxon. Such
a geographically heterogeneous subspecies is called a polytopic subspecies.
The only alternative to its recognition is not to recognize any subspecies
in such a species. In the absence of diagnostic differences there is no legiti¬
mate excuse for dividing a single polytopic subspecies into several subspecies
merely on the basis of locality. It must be remembered that any subspecies
is a heterogeneous composite, even when it consists of contiguous
populations.

9B.2 SUBSPECIES OR ALLOPATRIC SPECIES?

Whenever the taxonomist encounters two taxonomically distinct al-

lopatric populations, he must decide whether to consider them species or
subspecies. Various types of evidence are used in making this decision.
Degree of difference per se was sufficient reason for species recognition
194 Methods of Zoological Classification

by adherents of a typological species concept. Adherents of the biological

species concept look for evidence of actual or potential interbreeding and
use degree of morphological difference only to draw inferences on the
probability of potential interbreeding.
The word allopatric is essentially an antonym of sympatric and means,
therefore, distribution without geographical overlap [see Smith (1965) for
the proposal of a more elaborate terminology]. On the basis of geographical
pattern and observed interbreeding, five kinds of allopatry can be
distinguished :

Kinds of Allopatry

I Allopatric populations A and B are in contact. II

Allopatric populations A and B are separated by an unoccupied
area. V
II Populations A and B intergrade or interbreed freely. Ill
Populations A and B do not interbreed, or do so only very
occasionally. IV
III A and B intergrade clinally in a (usually fairly wide) zone of
contact (a)
A and B interbreed completely in a (usually rather narrow) zone
of contact or by hybridization. (h)
IV A and B meet in zone of contact where occasional hybrids occur, (c)
A and B meet in zone of contact but do not interbreed at all. (d)
V A and B do not interbreed because they are separated by a dis¬
tributional gap or natural barrier which prevents contact. (e)

Populations that qualify under (a) and (b) are nearly always to be treated
as subspecies; under (c) and (d) as species; and under (e) as species
or subspecies. The following comments, designated to correspond to the
key above, may be helpful:
Allopatric populations that intergrade clinally with each other belong
to the same species. It depends on the degree of difference whether or
not they are to be considered subspecifically different.

1. Primary Intergradation. There is no clear-cut distinction between inter¬

gradation and allopatric hybridization. In general, we speak of primary intergrada¬
tion when a series of intermediate populations is intercalated between two sub¬
species, each population with approximately the same amount of variability as
any other population of either subspecies. We speak of allopatric hybridization
when the two subspecies meet in a well-defined zone and form there a hybrid
population with greatly increased variability, often containing the entire spectrum
of character combinations from subspecies a to subspecies b. There must be evi¬
dence for random interbreeding in this zone. Allopatric hybridization is sometimes
also referred to as secondary intergradation, because it is a secondary event,
following a breakdown of previously existing extrinsic isolation of the populations.
 Taxonomic Decisions on the Species Level 195

Among North American birds the flickers (Colaptes), juncos (Junco), and towhees
(Pipilo) furnish good examples of hybridization between widely divergent subspe¬
cies. For further details and additional examples, see Mayr (1942, pp. 263-270,
and 1963, p. 118), and Sibley (1961).
2. Secondary Intergradation. Secondary intergradation, that is, the occur¬
rence of zones of contact of previously isolated populations or subspecies, occurs
very commonly in geographically variable species. No taxonomic difficulty arises
when the zone of hybridization is narrow. However, if it is wide and if a well-
defined, stabilized hybrid population with intermediate characters develops, it
is sometimes convenient and justified to recognize the “hybrid” population tax-
onomically. It may be treated as a subspecies if it satisfies the requirements
of the 75-percent rule. The taxonomic recognition of a hybrid population is not
justified if it is highly variable and includes in this variation a range of phenotypes
extending from one parental extreme to the other. If two taxa that were previously
recognized as two allopatric species completely intergrade in a zone of secondary
contact, it proves that they are not reproductively isolated and that they should
be considered subspecies of a single polytypic species.
3. Occasional Hybridization. Allopatric forms that hybridize only occasion¬
ally in the zone of contact are full species. There are a few cases where it
is difficult to decide whether the hybridization is occasional or complete. Much
recent evidence indicates that hybridization has to be fairly complete in order
to restore secondary intergradation. For a treatment of occasional hybridization,
see 2.5 and 13.21.
More difficult to evaluate are cases where two species remain as distinct
species over most of their range but form complete hybrid populations in a
few areas. This happens particularly in regions where human interference in
recent years has badly disturbed the natural ecological balance. It is recommended
that such forms be treated as full species in spite of the occasional free hybridiza¬
tion under the stated conditions (see Mayr, 1963, pp. 119—124).
4. Parapatric Species. Allopatric populations that fail to interbreed, although
in contact, are full species. Failure to interbreed indicates reproductive isolation
and attainment of species rank. The absence of geographical overlap may be
caused by one or the other of two opposite reasons. The zone of contact may
connect two very different ecological areas (e.g. savanna and forest). If one of the
two neighboring species is specialized for one of these habitats and the other species
for the other, the two species cannot invade each other’s range because their
ecological requirements are too different.
The other possible reason for nonoverlap of full species is that their ecologi¬
cal requirements are so similar in every respect that they compete with each
other. On one side of the zone of contact one species is slightly superior, on
the other side the other. Such “competitive exclusion” (for detailed discussion
see Mayr, 1963, pp. 81-82) will result in strict allopatry of full species. It is
important to understand this because allopatry used to be accepted by some
authors as an automatic criterion of conspecificity. Vaurie (1955) showed that
in 225 species of 7 families of Palearctic songbirds, 22 good species had been
treated earlier as subspecies by one or another author owing to their allopatry.
See alsd Kohn and Orians (1962).
196 Methods of Zoological Classification

A careful study of the zone of contact usually reveals areas where increased
habitat diversity actually permits occasional svmpatry of the two species. The
best evidence for species status is provided by the sharpness of phenotypic differ¬
ence in the zone of contact. If two continental species display no evidence at
all of any intergradation in their zone of contact (or close approach), they evi¬
dently do not exchange genes with each other and must be treated as full species,
even though they are allopatric.
5. Isolated Allopatric Populations. Geographically isolated allopatric popula¬
tions may be either species or subspecies. The most important of the species
criteria, the presence or absence of reproductive isolation, cannot be used (except
experimentally, and even then only with reservations) to determine the status
of populations that are separated from their nearest relatives by a distributional
gap. This is the reason why the classification of allopatric populations is so often
subject to a considerable amount of disagreement among taxonomists. Many solu¬
tions for this dilemma have been proposed, but all of them are beset with
difficulties.
Some taxonomists insist that all morphologically distinct, isolated populations
be treated as full species “until it is proved that they are subspecies.” This
solution is of course impractical, because it is impossible in most of these cases
ever to obtain clear-cut proof one way or the other. Furthermore, this solution
overlooks the fact that it is just as serious an error to call a population a species
when it is really only a subspecies, as vice versa.
The second solution is to treat as full species all populations that are
not connected by intergradation. This procedure is founded on the correct observa¬
tion that populations which are connected by intergradation are conspecific; the
reverse conclusion is then drawn, namely, that populations which are not connected
by intergradation are not conspecific. This however is in conflict with the rules
of logic when applied to isolated allopatric populations. Geographical isolation
is not an intrinsic isolating mechanism (Mayr, 1963, p. 91), and there is no
guarantee that the morphological hiatus caused by the temporary stop in the
gene flow is proof of the evolution of isolating mechanisms. The opposite ex¬
treme—considering all related allopatric forms to be conspecific—is equally wrong,
as shown above.

A complete experimental analysis, including studies on mating prefer¬

ence and a cytological examination of hybrids, is usually impossible, and
even where it is possible it may not be conclusive. Ecological preferences
are part of the isolating mechanisms between species, and these cannot
be properly evaluated in the laboratory. For example, the sympatric sibling
species Drosophila pseudoobscura Frolova and D. persimilis Dobzhansky
and Epling always hybridize in laboratory populations, but only a few scat¬
tered Fj hybrids have ever been found in nature.
When direct proof is unavailable, it becomes necessary to decide the
status of isolated populations by inference. Several kinds of probabilistic
evidence are available on w'hich to base such inferences. All rely on the
observation that reproductive isolation is correlated with a certain amount
 Taxonomic Decisions on the Species Level 197

of morphological difference, which is fairly constant within a given tax¬

onomic group. The taxonomist can use this evidence to work out a yardstick
which can be applied to isolated populations. There are three sets of mor¬
phological differences that can be utilized to calibrate such a scale.

1. Degree of Difference between Sympatric Species. Within a given genus

or within a group of closely related genera, there is usually a fairly well-defined
amount of morphological difference between valid sympatric species. This differ¬
ence may be great, as in the case of birds of paradise, or it may be very slight,
as in the case of sibling species. This amount of difference between good species
can be used to determine the status of isolated populations in these same genera.
2. Degree of Difference between Intergrading Subspecies within Widespread
Species. The amount of morphological difference between the most divergent sub¬
species in species of the same genus indicates how much morphological difference
may evolve without acquisition of reproductive isolation.
3. Degree of Difference between Hybridizing Populations in Related Species.
Subspecies or groups of subspecies within a species sometimes become temporarily
separated from one another through the development of a geographical barrier
but merge again after the breakdown of the barrier. Free interbreeding, which
often occurs even after morphological difference of considerable magnitude has
developed, proves conspecificity. Good examples of such free interbreeding of
morphologically strongly differentiated populations are to be found in North
American birds, among some of the juncos (Junco) and flickers (Colaptes) (Short,
1965). This is to be taken into consideration in the ranking of isolated populations
in these genera.

Even after all these criteria have been applied, some doubtful cases
remain. It is preferable for various reasons to treat allopatric populations
of doubtful rank as subspecies. The use of trinominals conveys two impor¬
tant pieces of information: (1) closest relationship and (2) allopatry. Such
information is very valuable, particularly in large genera. Geographical
replacement suggests furthermore that either reproductive isolation or eco¬
logical compatibility has not yet been evolved. To treat such allopatric
forms as separate species has few practical advantages. If further analysis
shows that such a form had been erroneously reduced to subspecific rank,
it can again be restored to full species status.
Chapter The Procedure of
Ji 1/ Classifying

j.n this chapter an attempt will be made to explain

the procedure of sorting species into higher taxa. The theory of biological
classification, on which this procedure is based, has been treated in Chap. 4.
Most of the problems of classification of higher taxa—one might designate
this as macrotaxonomy—are still controversial. Compared to the extraordinary
activity on the species level, there has been until quite recently very little con¬
ceptual ferment on the level of the higher categories. We are still waiting for
a new systematics of macrotaxonomy, even though recent work represents
significant forays into this terra incognita. What is particularly badly needed
at this time is a greater interest among taxonomists in the taxonomic method
as such, in the properties of the evidence on which taxonomic conclusions
are based, and in the process of inference by which conclusions are derived
from the evidence. The new interest in methodology, aroused by Hennig,
Cain, Michener, Simpson, and the numerical pheneticists, is likely to pro¬
duce eventually as much of an advance on the level of macrotaxonomy
as the new systematics did on the species level.
In the past, animal taxonomists have been singularly silent about
their methods and guidelines. Many of their conclusions, no matter how
sound they turned out to be, were reached by “inspection,” by an overall
simultaneous evaluation of all readily accessible lines of evidence. Only
the final result was presented to the reader, rather than the steps that

198
 The Procedure of Classifying 199

led to it. The very fact that this holistic approach resulted so often in
a durable product, confirmed by subsequent researches, demonstrates that
it must have been based on a sound and valid method of evaluation. It
would be most helpful to later workers if specialists would always record
how they obtained and interpreted the evidence. One suspects that they
often reached results intuitively, like the mathematician Gauss who is sup¬
posed to have said “I have the result, but I don’t know yet how I can
arrive at it.”
The time has not yet come to present a well-balanced methodology
of macrotaxonomy. The treatment in this chapter is an attempt to present
a synthesis of conflicting claims and alternative procedures, and, perhaps
most important, a critique of unsound approaches and assumptions. The
literature cited in the various sections of this chapter will permit a deeper
penetration into the field. A careful study of various recent publications
will help the reader to understand the nature of current disagreements
and uncertainties; some of these are Michener (1957), Inger (1958), Simp¬
son (1961), and Mayr (19656).
The activities of the taxonomist may be best characterized by recogniz¬
ing four steps:

A. Preparatory activities
1. The sorting of individuals into phena and these into populations
(Chap. 8)
B. Genuine classification
2. The assigning of populations to species (Chaps. 3, 9)
3. The grouping of species into higher taxa
a. Determination of relationship
b. Formal delimitation of taxa
4. The ranking of taxa in a hierarchy of categories

In the present chapter we are concerned with operations 3 and 4.

Simultaneously with operations 2, 3, and 4 names are assigned to
the recognized taxa (Chaps. 12, 13).

10.1 THE GROUPING OF SPECIES INTO HIGHER TAXA

The above phrase suggests a single operation, but there are actually
at least three involved:

1. The determination of the nearest relatives of each species

2. The searching for gaps which permit the ordering of species into clusters
and groups of clusters, and the decision which of these clusters to recog¬
nize formally as genera and which others merely informally as species
groups
200 Methods of Zoological Classification

3. The arrangement of genera into groups of higher and higher taxa, and
the ranking of these in the proper categories of the taxonomic hierarchy

These three operations merge into each other and cross-affect each
other, as we shall presently see. This complexity of classifying is by no
means revealed by the definition given in 4.1 “Zoological classification
is the ordering of animals into groups on the basis of their similarity and
relationship.” Before the various steps in the classifying procedure can be
discussed, the meaning of the term relationship must be clarified.
Relationship. Unfortunately this term has been used in the taxonomic
literature with three very different meanings. To the pheneticist, relationship
(and its synonym affinity) simply means unweighted similarity; to the
cladist, it means genealogical relationship (4.3.5), i.e. propinquity of de¬
scent; finally, to the evolutionist it means inferred genetic similarity, as
determined both by distance from branching points and subsequent rate
of divergence (4.8). He finds it by evaluating “weighted phenetic simi¬
larity.” The use of the words affinity or relationship, when nothing but
similarity is meant, leads to confusion and should be avoided (Mayr,
19656).
10.1.1 Finding the Nearest Relatives of a Species. What is the near¬
est relative of a given taxon is in many cases so obvious that it requires
no special investigation. Equally often the choice is not a foregone conclu¬
sion, and the question arises, Is taxon B better grouped with taxon A
or with taxon C? In order to find the answer, the following procedure
is usually employed. All characters likely to shed light on the relationship
of B, or, in other words, all characters knowrn or suspected to vary in
the taxon group A, B, and C, are tabulated (see Table 10-1).
An analysis of Table 10-1 shows that B is closer to A in one set
of characters (a, b, c, d) while it is closer to C in another set of characters
(e', /', g', h'). There is a third class of characters (i", k", l", m"), in which B
differs from both A and C.
Two sets of evidence are thus available: One is the relative weight of
characters a, b, c, d, as compared with that of e', /', g', h'. The other is the
information contained in the characters peculiar to B (i", k", l", m").
Are these characters more easily derived from the corresponding characters
of A or of C, or are they perhaps in part ancestral to the equivalent
characters of A or of C? All these questions are merely somewhat more
sophisticated restatements of the traditional question: Is B more similar
to 4 or to C? Determination of similarity then is the key operation in
classification. No matter how widely the theories of classification diverge
(4.3), the actual classifying operations resemble one another. Careful
comparative evaluation of similarities and differences is the first step in
all procedures of classification, regardless of whether the observed similarities
 The Procedure of Classifying 201

Table 10-1. Occurrence of Characters in Three

Related Taxa

Characters Taxon A Taxon B Taxon C

Shared by B and A a a a!
b b b'
c c c'
d d d'

Shared by B and C e ef e'

f f f
8 s' g'
h h! h!

Peculiar to B i i" i'

k k" k!
l l" - V
m m" m!

are directly converted into a phenetic classification or whether they are

carefully evaluated before being used as evidence for the delimitation of
groups considered to have descended from common ancestors. Since in
general the more closely two groups of organisms are related, the more
similar they are to one another, pheneticists and evolutionists share an
interest in “similarity.” They differ, however, in their determination and
evaluation of similarity.
10.1.2 Similarity. The history of taxonomy decisively refutes the
assumption that similarity is self-evident and not in need of careful evalua¬
tion. It requires a great deal of knowledge and experience to be able to
look through superficial similarities and to discount superficial dissimilarities.
The taxonomist forever makes decisions (at a more sophisticated level)
that are best illustrated by an example. Show to a biologically uneducated
person pictures of a shark, a porpoise, and a cow, and ask: “Which two
of these three are most similar?” and often you will get the answer: “Of
course, the two fish!” In this case, the zoologist has no trouble. Yet much
of his work in taxonomy consists in the elimination of spurious similarities
and the approriate weighting of seeming differences. Pheneticists who
are merely interested in identification schemes might be entirely satisfied
with a classification that arranges the porpoise with the fishes and mutatis
mutandis all other phenetic deviations from evolutionary groupings merely
on the basis of superficial similarity.
The evolutionary taxonomist, however, treats similarity merely as evi-
202 Methods of Zoological Classification

dence on which to base inferences on phyletic relationship. He attempts

to penetrate the superficial aspects of the phenotype in order to determine
underlying genetic similarities. He believes that taxa which are based on
inferred genetic similarity have a far greater predictive value than taxa
based on superficial unevaluated similarity.
Similarity established by a procedure in which the probable informa¬
tion content of each character is carefully weighted almost invariably gives
a classification in which the taxa truly are monophyletic descendants of
common ancestors. Bader (1958) points out correctly how little effect the
discovery of an abundant fossil record has had on the classification of
mammals. The conventional method of taxonomy, constructing classifica¬
tions on the basis of carefully weighted similarity, is not often apt to lead
the taxonomist astray.
10.1.3 Causes of Similarity. Two taxa may be similar to each other
for various reasons. The different kinds of similarity have unequal weight
in the construction of natural taxa, although our understanding of these
differences is still incomplete. The combined similarity between two taxa
may be composed of four kinds of similarities (see also Cain and Harrison,
1960).

1. Similarities resulting from joint possession of characteristics shared with

a common ancestor
a. Ancestral characters (symplesiomorphs of Hennig) shared with a
remote ancestor
b. Derived characters (synapomorphs of Hennig) shared with a more
recent ancestor
2. Similarities resulting from joint possession of independently acquired
phenotypic characteristics produced by a shared genotype inherited from
a common ancestor (similarity through parallel evolution)
3. Similarities resulting from joint possession of independently acquired
phenotypic characteristics that are not produced by a genotype inherited
from a common ancestor (similarity through convergence)

As far as giving information on common descent is concerned (and

hence on inferred genetic similarity), these characters must be ranked in
the sequence: 1 b, 2, and 1 a, while 3 is to be eliminated altogether. But
even within the three first groups the similarities are of unequal value
(see below under weighting).
10.1.4 Methods for the Determination of Similarity. The many poor
classifications that have been proposed in the taxonomic history of almost
any group of animals prove that the establishing of natural groups is no
easy matter. Operationally it requires the simultaneous consideration of
numerous characters in numerous taxa to make a classification. One of
the methods employed in this procedure, the ordering of characters into
important and unimportant ones, often led to error. Another (simul-
 The Procedure of Classifying 203

taneously employed) method, to improve the classification by a process

of trial and error leading to an ever closer approximation, though eventually
successful, appeared wasteful in time and effort. These weaknesses of the
traditional approach led to alternate proposals.

10.2 GROUPING BY UNWEIGHTED PHENETIC SIMILARITY

In order to avoid the shortcomings of traditional procedures based

on single characters some taxonomists have been searching for a measure
of “overall similarity.” Serologists thought that protein interaction gave
such an overall measure, while DNA matching (Hoyer et al., 1964) is
the most recent candidate (7.4.9). So far none of these methods has fulfilled
the hopes of its proponents. The pheneticists have chosen a different ap¬
proach. They propose to determine overall similarity quantitatively, by com¬
pounding similarity values derived from a large number of individual char¬
acter comparisons.
10.2.1 Quantification of Similarity. The recent efforts to quantify
similarity are not the first, earlier attempts having been made nearly 100
years ago. More recently Sturtevant (1942) tabulated 33 characters of
56 species of Drosophila and found that the concordance between a given
pair of species in the chosen characters ran from 0 to 25. When a second
tabulation was made of 11 additional characters, based on features not
used in the first tabulation, it was found that species that were similar
on the basis of the first tabulation were also similar with respect to the
second set of characters. The development of electronic computers has
made the utilization of more sophisticated calculations possible. The new
development started with the almost simultaneous publications of Sneath
(1957), Michener and Sokal (1957), and Cain and Harrison (1958), each
author proposing slightly different methods. A detailed presentation of these
and other methods can be found in chaps. 6 and 7 of Sokal and Sneath
(1963), Principles of Numerical Taxonomy. Many additional methods are
now described each year. A special newsletter [T axometrics) is now pub¬
lished by L. R. Hill which contains a current bibliography as well as short
communications concerning new programs and modifications of previously
published ones. Computer programs are now available for most of the
suggested approaches. It cannot be our task to review in detail a field
covered in an entire book and annually in more than 50 journal articles.
The field is changing so rapidly that statements made here may well
be out of date within a year or two. Anyone who wants to use the computer
in classification must try to keep abreast of the rapidly changing literature.
The short presentation given here can serve only as a most elementary
introduction to a complex field.
204 Methods of Zoological Classification

Table 10-2. Data Matrix (after Sokal and Sneath, 1963)

Taxa
Characters
A B C D E F

.1 1 8 1 7 2 5
2 1 6 1 6 1 3
3 6 1 5 1 4 2
4 1 0 1 0 1 4
5 NC 6 3 6 NC 1
6 NC 2 NC 3 1 1
7 8 2 7 2 5 5
8 1 6 1 6 3 4
9 1 8 1 8 2 4
10 6 1 6 1 5 2
11 3 3 3 3 3 3

10.2.2 Data Listing. The first step is the listing of the raw data
in the form of a data matrix (Table 10-2). The vertical columns list the
taxa, while the horizontal rows tabulate the characters. There are two
ways of dealing with characters in such a tabulation. If only the presence
or absence of a character is to be indicated, plus and minus signs are
all that is needed. On the other hand if different character states are
to be recorded, numerals have to be used. The special definition of “charac¬
ters” in computer taxonomy has to be kept in mind (7.1).
Such a matrix can be examined from two points of view. An analysis
of the association of pairs of character (rows) is called the R technique;
it studies the correlation among characters. Conversely, an analysis of the
association of pairs of taxa is called the Q technique. This is the technique
used by the pheneticists when making the simplifying assumption that all
characters have equal information content. Every taxonomist realizes that
this is unrealistic. When Rogers (1963) tested the contribution of various
attributes to the grouping of specimens of Manihot,

... it invariably happened that each group of specimens was set apart
by a certain number of attributes and that the remaining attributes did
not define the groups. Some of the attributes provided definitions of small
groups, some of larger groups, and some were not significant at all at
any level. What had happened, apparently, was that certain of the charac-
terisdcs and attributes were at work in definition and characterization of
the groups; others were not. Those that did indeed separate and define
the taxa had more weight than those that did not.

Yet a recent study by Rohlf (1967) involving an R-type analysis did not
 The Procedure of Classifying 205

apparently lead to an improvement of a classification previously produced

by a Q-type analysis.
There are numerous ways of computing resemblance between taxa,
based on the Q-type approach. Three types of coefficients of similarity,
those of association, correlation, and distance have been proposed. Each
has advantages and disadvantages. In many ways the most satisfactory
method for expressing the difference between two taxa is to calculate total
“distance” in a multidimensional space (Seal, 1964). With electronic com¬
puters available, it is possible to calculate taxonomic distance even when
numerous taxa and characters are involved. This method likewise has deep-
seated flaws that have not yet been overcome. Both Boyce (1964) and
Minkoff (1965) have shown that different mathematical treatments of the
same raw data lead to drastically different classifications.
10.2.3 The Number of Characters. In the old identification schemes
(mislabeled classifications) taxa were characterized monothetically, that
is by single characters. In contrast, empiricists in the eighteenth century
already proposed using the greatest feasible number of characters. This
advice is on the whole sound; however, there tends to be a difference
between characterizing species and higher taxa. Species taxa can often
be recognized on the basis of a single character. For instance, both the
Mikado Pheasant (Syrmaticus mikado) of Formosa and the Ribbon-tailed
Bird of Paradise (Astrapia mayeri) of New Guinea were named on the
basis of a single tail-feather. The shape of the baculum of mammals is
sometimes species-specific (Fig. 7-3), as are genitalic structures in arthro¬
pods, color patterns and songs in birds, and vocalizations in frogs, orthop-
terans, and cicadas. Sometimes a single character may be diagnostic of
a higher taxon. However, higher taxa are groups of species, and they almost
invariably require for characterization a number of traits, because each
character or character complex may have its own independent evolution
in each phyletic line and subline. In such a case a classification based
on few characters, or a single one, will be quite misleading. For instance,
on the basis of the morphology of the placenta and associated fetal mem¬
branes, one author proposed that lemurs be classified with the ungulates
rather than with the higher primates, and the Megachiroptera with the
rodents rather than u'ith the regular bats (Microchiroptera). An ornitholo¬
gist classified owls on the basis of the size and asymmetry of the external
ears. A reexamination showed that this is a food-associated specialization
closely correlated with climatic zones: tropical owls which find their prey
visually have small cars without dermal flaps, while temperate-zone owls
feeding during long winter nights on rodents which are acoustically located
have large ears with wide dermal flaps. Altogether too many early classifica¬
tions were based on a single conspicuous character complex: parasitic bees
versus nonparasitic bees; songbirds with cone-shaped bills (finches) versus
206 Methods of Zoological Classification

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Solitary bees Parasitic bees Social bees

Fig. 10-1. A so-called “practical” classification of bees (after Friese) in which the parasitic species
are treated as a separate group, independent of their close relatives among the solitary and social bees.

songbirds without cone-shaped bills, and so on. Every specialist can cite
similar examples from the taxa with which he is most familiar.
The unreliability of single characters led to the belief that, the more
characters a classification is based on, the more reliable it will be, which
is not necessarily true. Even a large number of unweighted characters is
not always sufficient for unequivocal classification. Rohlfs (1963) two classi¬
fications of the mosquito genus Aedes, one based on 77 adult and the
other one on 71 larval characters, are discordant to such a degree that
a very much larger number of characters would be needed even to hope
for concordance. Also, there is always the problem of defining what is
a character. A large-sized species may differ from a related small-sized
species by hundreds of mensural characters, but all of them may be nothing
more than aspects of a single trait, namely size.
Methods based on the processing of unselected and unweighted char¬
acters require large numbers of characters, preferably more than 100. These
can be readily found in arthropods, particularly insects; it is not surprising
that bees, mosquitoes, and mites were preferentially used to test these meth¬
ods. It has not yet been determined what methods are most suitable for
use in morphologically uniform groups (such as birds or lower fungi) where
there is an embarrassing deficiency of useful characters. Results of princi¬
pal-component analyses suggest that a relatively small number of characters
might be sufficient if one only knew how to select such characters.
The only limit to the number of characters provided by a taxon
is that set by the patience of the investigator. It would seem prudent not
 lEricrgpini) (Melectini)

. Anthophorini
(Epeoloidini Hemisiini
(ProtepeoHm Emphorini
(Neolarrini Eucerini

APIDAE
(TownsendielUni
APIDAE

(Biostini
(Neopasi tini
(Ammobatoidini
(Ammobatim
(CaenoprosopidTni
(Nomadini

Fideliinae
MELITTIDAE

MEGACHILIDAE
Dasypodinae

Mel ittinae
Macropidinae

Ctenoplectrinae
HALICTIDAE

Dufoureinae

Nomiinae

Halictinae Melitturgini
ANDRENIDAE
Panurgini
Chilicolinae

Hylaeinae Andreninae

Euryglossinae
Oxaeinae
COLLETIDAE

Diphaglossinae

Stenotritinae
Caupolicanini
Colletini
Paracol let-ini

Fig. 10-2. A phylogenetic classification of bees (after Michener, 1944) in which the parasitic bees are
placed according to their inferred relationships among the solitary and social bees. Wholly parasitic
groups are circled, partially parasitic groups underlined.

207
208 Methods of Zoological Classification

to use more characters than are sufficient for the establishment of a sound
classification. Most characters have a low information content or are com¬
pletely redundant. What is important is not the number of characters but
their taxonomic weight. Indeed Kendrick and Weresup (1966) and Throck¬
morton (1969) have demonstrated that many, if not most, characters are
merely “noise” (as far as their informational value is concerned) and that
their removal from the list of considered characters may increase efficiency
of grouping.
When a new set of characters is utilized, it will have one of three
possible effects on the existing classification. The new characters either
completely confirm the traditional classification (this being what usually
happens), or lead to a clear improvement of the classification, or introduce
ambiguity and contradiction which can be resolved only by the application
of additional new characters. At the present time it cannot be predicted
which of the three will be realized in a given instance.
10.2.4 Weaknesses of the Purely Phenetic Method. Many recent
writers have pointed out deficiencies in the methods and theory of numerical
phenetics (see recent volumes of Systematic Zoology).
Among the theoretical weaknesses are its inability to distinguish be¬
tween phena and taxa. Under its own terms this is legitimate, since the
end product of the phenetic operation is an identification scheme rather
than a biological theory, and this is what a good classification should be.
Another theoretical weakness is the assumption that a random selection
of components of the phenotype would lead to correct estimation of the
properties of the genotype, provided enough characters are included in
the analysis. The visible phenotype is as small a fraction of the total poten¬
tial phenotype as the visible part of an iceberg is of the whole. It is not
true, for this reason, that the amount of unweighted phenetic similarity
necessarily runs parallel with the amount of genetic similarity. Sibling spe¬
cies prove conclusively that drastic genetic differences may not be reflected
in the visible phenotype. The opposite extreme is demonstrated by groups
like the birds of paradise, where intensive sexual selection (Mayr, 1942;
Sibley, 1957, 1959) has produced enormous phenetic differences between
closely related species that are still fully or largely fertile with each other.
Only appropriate weighting can convert the phenetic distances between
the genera of birds of paradise into a biologically meaningful classification.
This proves the falseness of the assumption that each character is so poly¬
genic that any random sample of characters will accurately reflect the prop¬
erties of the genotype.
It is important here to mention computers and electronic data pro¬
cessing (EDP). Some users of EDP have suggested that thinking and theory
become unnecessary if we merely entrust our fate to the computer. Nothing
could be farther from the truth. The operations of the computer are in¬
finitely faster than our own, and it can handle far more data at one time
 The Procedure of Classifying 209

than we can; but objectivity is not affected by the computer, and the
underlying principles are not modified in the least. The computer is defense¬
less against logical errors of the programmer. It is a pity that so much
of the early work in computer taxonomy was based on unsound taxonomic
theory (see also Ghiselin, 19666).
Equal weighting and random selection of characters are implicitly
based on the assumption that during evolution the genotype as a whole
changes harmoniously and that all components of it change at approxi¬
mately equal rates. Mosaic evolution and many other evolutionary phenom¬
ena, however, show that this assumption is unrealistic (10.4.5).
The phenetic method also encounters various practical difficulties.
One is its inability to produce repeatable results whenever there is a slight
change in the selection of characters or the methods of determining simi¬
larity (Boyce, 1964; Eades, 1965; Minkoff, 1965; Kendrick and Weresup,
1967; Michener and Sokal, 1966; Sokal and Michener, 1967).
The method is, in a way, inefficient because the programming of
large numbers of taxa with about 100 (or more) characters is very time-con¬
suming. A specialist may not have such time available, if he is the only
living specialist for a group of 1,000, 5,000, or 20,000 species. In order
to be able to undertake identification work, and to describe and name spe¬
cies rapidly, he is forced to adopt short-cut methods of classifying. He sorts
numerous specimens and species into “natural groups” by scanning their
total gestalt, based on an evaluation of very many characters, most of which
he does not analyze or record in detail (Anderson 1954). A posteriori
he determines what the most constant and most easily recognizable charac¬
ters of these natural groups are.
By far the greatest practical difficulty in applying a purely phenetic
procedure is the scarcity of taxonomically useful characters in most groups
of organisms. Insects and other arthropods with a highly sculptured exo¬
skeleton are the only conspicuous exceptions to this statement, but even
in these there is often a “desperate need for new characters,” as recently
stated by a specialist in digger wasps. By the time all characters of low
weight have been eliminated (see 10.4), there is often hardly anything
left. In the case of most of the thirty-odd orders of birds, it is still unknown
which other order is the nearest relative. The same is true for the families
of songbirds (Oscines). There are several drastically different classifications
of the sponges depending on the relative weight given to the very few
available characters. Convergent adaptations lead to far greater similarities
in such groups than does the joint possession of ancestral characters. The
finches, tree creepers, titmice, flycatchers, and possibly the ratites are illustra¬
tions among birds of groups based on convergent characters. Many similar
examples will occur to the specialist in rodents, snakes, urodeles, teleost
fishes, bivalve mollusks, and lower invertebrates.
There are two ways to overcome the dilemma of a scarcity of useful
210 Methods of Zoological Classification

characters. One is to look for new characters (7.4), the other is to attempt
to extract more information from existing characters. Bock (1963) has
shown how a deeper analysis of known characters sometimes leads to a
better understanding of the phylogeny of morphologically uniform groups.
A purely quantitative procedure, by which the few useful characters are
diluted by a large number of useless ones, was rightly ridiculed by Adanson
(Stafleu, 1965).
10.2.5 Utility of the Phenetic Approach. Nonweighting is least ob¬
jectionable when applied to groups with immature classifications (particu¬
larly single-character classifications) and those with numerous nonredun-
dant characters. In such taxa it has produced groupings that are clearly
superior to the traditional ones. When there are several competing classifica¬
tions, a phenetic analysis may be illuminating. For instance, the genera
of Megascolecoid earthworms were assigned to very different families and
subfamilies in three almost simultaneously published revisions. In one of
these the higher taxa were based on the position and number of the cal-
ciferous glands; in the second, they were based on the number and position
of the male terminalia, and in the third on the structure of the prostatic

IVORY-BILLED WOODPECKERS

woodpeckers. The genera in the three vertical columns have similar color patterns indicative of true
relationship. The similarities in foot, bill, and tail of the ivory-bills are considered to be due to
convergence (after Bock, 1963).
 The Procedure of Classifying 211

glands and the excretory system. Forty-three characters of 30 species (in

29 genera) were analyzed by Sims (1966) with the help of association
coefficients, and the grouping resulting from this analysis agreed reasonably
well with one of the previously proposed classifications, i.e. that based on
the structure of the prostatic glands. In the mites (Acarina), also a group
with an immature classification, computer techniques have likewise been
very helpful (Sheals, 1965).
The phenetic method is a particularly uneconomical procedure in
groups with either a mature classification or a great deficiency of taxonomi-
cally useful characters. Weighting becomes the more important the higher
the categorical level of the taxon we are attempting to place. When we
sort species within a genus, weighting is relatively unimportant, and much
of it is arbitrary. But the higher in categorical rank two compared taxa
are, the fewer will be the number of characters that indicate relationship.
And the fewer the number of usable characters, the more carefully they
must be weighted.
It is too early to pass final judgment on the phenetic method. Much
of its theory (4.3.3), based on nominalism, is naive and unsuitable as an
appropriate basis for work in evolutionary taxonomy (Mayr, 19656). Yet
computer methods as such have a tremendous potential. The greatest con¬
tribution the pheneticists have made is to harness the computer to the
classifying procedure. The original goal, to automate taxonomic procedure
and to remove from it all subjective evaluations, has not been achieved
in the first ten years. “In the present state of its development, it [phenetics]
is not able to provide stable classification” (Sokal and Michener, 1967).
Other characters, other species in the same genera, other measures of simi¬
larity, and other clustering methods often result in different classifications.
The ultimate answer will almost surely be a combination of classical and
new computer methods. Nevertheless we owe the numerical taxonomists
a debt of gratitude for developing new methods and for trying to find
out which would give the most informative results. Methods of weighting
will undoubtedly play an increasingly important role in computer taxonomy.

10.3 GROUPING BY CLADISTIC APPROACHES

Instead of making “overall similarity” the basis of grouping, one can

also establish groups based on the inferred branching pattern of phylogeny.
All methods which give branching patterns primacy in classification may
be designated as cladistic approaches.
These methods share a common attempt (1) to determine the number
of phyletic splits by which different taxa are separated from each other,
and (2), perhaps more important, to reconstruct the sequence by which
212 Methods of Zoological Classification

ABC D
+ + + +

Fig. 10-4. Hennig’s method of determining

groups derived from the most recent common
ancestor. Such groups are distinguished by the
joint possession of at least one derived char¬
acter (shaded rectangles) such as character 1
for taxon A, 2 for taxon B C -j- D, 3 for
B, 4 for C D. This permits reconstruction
of the sequence of branching (= recency of
common ancestry). Ancestral characters are
designated by open circles or squares (after
Hennig, 1966).

these branchings occurred. This strategy has long been employed by students
of chromosomal evolution in order to determine the sequence in which
chromosomal inversions and other chromosomal mutations must have oc¬
curred. Hennig (1950, 1966) pioneered in systematically applying this ap¬
proach to taxonomy (see Fig. 10-4 for an explanation of this method).
A good example of an application of this method is Wagner’s (1962)
classification of the Delphacidae of Europe. Wilson (1965) discusses a simi¬
lar approach.
A somewhat different cladistic method was developed by Camin and
Sokal (1965) for deducing phylogeny. Human geneticists (Cavalli-Sforza
and Edwards, 1964) and molecular evolutionists (Fitch and Margoliash,
1967) have, largely independently, developed similar methods. All these
approaches have a common objective, namely, to design a phyletic branch-
 The Procedure of Classifying 213

ing pattern that would result from the minimal number of evolutionary
steps by which the various taxa became separated from each other.
After a phyletic line has split into two separate lines, the subsequent
retention of ancestral characters and the acquisition of new, derived charac¬
ters occur independently in the two lines. The basic rationale of all cladistic
methods is that the more recent the common ancestry of two species (or
other taxa), the more characters in common they should have. The same
thought, expressed in terms of taxonomic characters, is that the occurrence
of a relatively new character will be limited to the descendants of the
particular species in which the new character originated. A careful study
of the distribution of characters in the taxonomic hierarchy should therefore
give information on the grouping and ranking of taxa. The determination
of the phyletic age of characters is the key operation in this method.
10.3.1 Ancestral Characters. In view of the stipulation that taxa
should consist of descendants from a common ancestor, a separation of
characters into those similar to or identical with the characters of the ances¬
tor and those that have more or less diverged from the ancestral condition
becomes important. The terms “primitive” and “advanced” for these two
classes of characters were inherited from progessionist preevolutionary the¬
ories (scala naturae, etc.). The term primitive implies simplicity, and it
is therefore often misleading because comparative anatomy and the study
of fossil series have shown that a progressive simplification is one of the
most frequent trends in evolutionary lines. The terms “advanced” and “spe¬
cialized” for the derived condition of a character can be misleading for
the same reason. Hennig (1950) uses the terms plesiomorph for primitive
and apomorph for derived characters or character states. There is a need
for simple, self-explanatory terms that are not committed to any evolution¬
ary theory. The terms ancestral and derived may serve as such simple,
descriptive terms. A character is ancestral if it has not changed materially
from the homologous character in the ancestor, while a character is derived
if it has changed materially.
A given character may be either ancestral or derived depending on
the stages in the phylogeny that are compared. Possession of wings is an
ancestral character for flightless birds, but a derived character for birds
as a w'hole when compared with reptiles. In some cases it is easy to deter¬
mine the ancestral condition of a given character in a phyletic line; in
other cases this is very difficult. Hennig (1950), Maslin (1952), Remane
(1952), Simpson (1961), and Wagner (1962) give criteria facilitating the
correct choice. The reconstruction of the entire phylogeny of the frogs
(Anura) depends on decisions concerning the relative primitiveness of vari¬
ous characters (Inger, 1967).
In some higher taxa, e.g. the primates, many primitive taxa still sur¬
vive, and it is almost possible to reconstruct the probable phylogeny by
214 Methods of Zoological Classification

a concatenation of living forms from the lemurs to the anthropoid apes.

In other groups, like the ungulates, as Simpson points out, there is no
survival of the ancestral types that lived during the early Tertiary period.
Among the species of a genus it is often possible to single out one that
has an assortment of characters which one would expect in the ancestral
species of the genus. In dendrograms such species are often placed at the
point of origin of the genus, but this is somewhat misleading. Only rarely
has such a species literally given rise to the others by budding off peripheral
isolates. More commonly, the recent “primitive” species is that one of the
many descendants of the ancestral species which happens to have retained
the highest proportion of ancestral characters., Such conservative species
are of extreme practical importance in helping to determine the relationship
of taxa. Conclusions regarding relationships depend largely on the reliability
of the determination of primitivity of characters. When a taxon is conserva¬
tive in one character, it very often is also conservative in other characters,
but certainly not always and rarely in all characters. Platypus is the most
primitive of living mammals in several anatomical features, particularly
in its reptilian shoulder girdle, and also its egg-laying habit. However,
it is highly specialized in its dentition, bill, poison spur, and various aquatic
adaptations.
Ancestral characters or character states have various properties that
help in their identification (Hennig, 1950, 1966). They tend to be irregu¬
larly scattered among a broad group of related taxa, while a given derived
character usually characterizes only one particular group of descendants
(Fig. 10-4).
The more widely a character is distributed in the system, the more
distant is the relationship which it indicates (Hennig, 1950, p. 172), and
the more remote the common ancestor. The distribution of the vertebrate
column in all vertebrates and the occurrence of feathers in all birds serve
to illustrate this principle. Yet possessors of a given character are not neces¬
sarily more closely related to each other than they are to other taxa that
lack the character. The reason is parallel evolution in the acquisition of
new or in the loss of old characters. For instance, pentadactyl vertebrates
(amphibians, reptiles, primates, and other mammals) are not more closely
related to each other than they are to perissodactyls, artiodactyls, cetaceans,
and other mammals that are not pentadactyls.
The earlier a phyletic line has acquired a character, or, as we usually
say, the older a character, the more completely it seems to be integrated
into the entire genotype and the more closely it is correlated with other
characters. The assumption that the older a character is, the earlier it
is laid down in ontogeny, is probably true but not proved. If true it would
nevertheless not confirm recapitulation, because there is no evidence that
an organism during ontogeny recapitulates the adult stages of its phylogeny
 The Procedure of Classifying 215

(as postulated by some phylogenists). Indeed all the evidence refutes such
an assumption.
Ancestral characters or character states are the condition from which
various specialized conditions are most easily derived. For instance, cyto¬
geneticists have shown for Drosophila (among other cases) that chromo¬
somal variation among near and distant relatives can be arranged in a
definite sequence of chromosomal mutations (7.4.7). Either one or the
other terminal condition must have been the ancestral one. Collateral evi¬
dence (if not internal evidence) usually permits a definite choice between
the two alternatives. Occasionally an intermediate condition is ancestral,
giving rise to two diverging trends. Various empirical rules help in determin¬
ing the ancestral condition. If one taxon is clearly derived from another
taxon, as birds and mammals are derived from the reptiles, the ancestral
condition may still be largely present in the ancestral taxon. Indeed, most
mammalian and avian characters can be clearly derived from homologous
features in the reptiles.

D. mojuoides

Fig. 10-5. Chromosomal phylogeny in the fasciola subgroup of the Drosophila repleta species group.
Ancestral arrangement in the bottom rectangle. Arrows indicate the sequence of chromosomal
rearrangements. Heterozygous inversions which occur with the standard are indicated by the
inversion shown over a plus sign (from Wasserman, 1962).
216 Methods of Zoological Classification

Various phylogenetic rules such as Cope’s rule, Dollo’s rule, and others
discussed by Rensch (1959, 1960) and Simpson (1959a), are sometimes
helpful. Some of these empirical indications are definitely wrong (4.8),
while others have varying numbers of exceptions. Contrary to the statement
of Dollo’s rule, for instance, lost ancestral structures may occasionally re¬
appear in descendants if the potential for them was retained in the genotype.
Conspicuous body setae and antennal papillae reoccur in the larvae of
a number of specialized groups of Hymenoptera after being lost earlier
in the evolution of this order of insects (Evans, 1965; see also Maslin,
1952). A careful evaluation of every character as to whether or not it
represents an ancestral condition is a help not only in the delimitation
of taxa, but even more so in ranking them as needed for the construction
of a taxonomic hierarchy (10.5), and in arranging them in a linear
sequence.
10.3.2 Auxiliary Evidence. When all the other evidence on relation¬
ship is ambiguous, one can sometimes employ two other sources of evidence.
Parasites evolve with their hosts but are sometimes more conservative.
An existing relationship may be more evident in the parasites than in their
hosts (7.4.12). The difficulty is that a parasite sometimes shifts to a new
host (more or less unrelated to the old host), which deprives the parasitolog¬
ical evidence of some of its value. In ambiguous cases the concordance
among different kinds of parasites must be analyzed. Most animal groups
have several kinds among the following parasitic taxa: protozoans (particu¬
larly in the blood), cestodes, trematodes, nematodes, acanthocephalians,
copepods, mites, lice, fleas, and mallophaga. Some of these parasitic taxa
shift rather easily to new hosts, while others are remarkably host-specific.
The relationship between man and the African anthropoids (Pan), for
instance, is indicated by four different groups of parasites. See Baer (1957)
for a discussion of various aspects of this method.
Geographic distribution is another important clue to relationship. Hen-
nig (1950) presents numerous cases where the study of distribution patterns
has resulted in improvements in classification. This evidence has been pre¬
sented in detail in Chap. 7 (7.4.13).
During the continuous faunal turnover on all continents older elements
are often forced to retreat to the southern continents (Australia and South
America) or to such islands as Madagascar and New Zealand. These relicts
of formerly more widespread taxa often possess exactly the ancestral charac¬
ters from which the character states of more modern types can be derived.
The study of relict types now found only at these peripheral locations
is thus often of particular importance for the reconstruction of the inferred
ancestral characters.
10.3.3 Utility of the Cladistic Method. The careful analysis of every
taxonomic character which the cladistic method demands is of considerable
 The Procedure of Classifying 217

heuristic value. Even taxonomists who reject the ranking criteria of cladism
(see 10.5.2) often find it useful to try to infer the phyletic age of each
character, since this makes such a large contribution to its taxonomic weight.
In the grouping step of the classifying procedure cladistic analysis is indeed
most useful, but when it comes to drawing inferences from cladistically
produced evidence, one must make two reservations.
First, an operational one—none of the methods listed above has been
sufficiently tested to permit judgment as to its efficiency and reliability.
Second (and more important)—none of these methods allows sufficiently
for mosaic evolution, for parallel evolution, and, particularly, for a combina¬
tion of both. The cladistic approach has a strong typological element, de¬
manding “yes” or “no” decisions and assuming all changes of characters
to be unique events. Other difficulties are caused by evolutionary reversals
and by the occasional impossibility of determining the ancestral condition.
The probabilistic considerations that will have to be built into the cladistic
approach to make it more reliable will inevitably bring it nearer to the
phenetic approach, and it is conceivable that the best elements of both
approaches can eventually be combined in a “synthetic” method. The classi¬
cal method of taxonomy, of course, has essentially employed such a “syn¬
thetic” procedure.

10.4 INFERRING RELATIONSHIP BY WEIGHTING SIMILARITY

Experienced taxonomists have always insisted that characters differ

in the contribution they make to the soundness of a classification. Darwin
(1859, pp. 414-—417), for instance, gives some good empirical rules concern¬
ing the usefulness of certain characters. With the failure of the phenetic
and cladistic approaches to produce automatically and objectively sound
classifications, there has been a renewed interest in the traditional methods
of taxonomy. The weighting of similarities as evidence for relationship
is the key operation in this approach. When well done, it leads to classifica¬
tions of lasting value; when poorly done, it leads to much changing and
controversy. Owing to their crucial importance the weighting criteria must
be discussed in detail.
10.4.1 A Priori and A Posteriori Weighting. The Aristotelians and
their successors often assigned a priori weights to certain characters. Cain
(1959a) has pointed out the fallacy of this approach. Neither function
nor conspicuousness nor any other known aspects of a character gives it
a priori a greater weight than other characters. Indeed the very same
218 Methods of Zoological Classification

structural difference may have high weight in one taxon and low weight
in a related taxon. Nor should the taxonomist confuse weight of a character
with its usefulness in a diagnostic key. Any rigid following of the dictates
of a few arbitrarily chosen a priori characters leads inevitably to a classifica¬
tion which is not “natural,” that is, which has low predictive value.
Adanson and the empirical taxonomists of the ensuing period rejected
a priori weighting and replaced it by an empirical process, perhaps best
called a posteriori weighting. Adanson was satisfied with merely eliminating
useless and redundant characters; later authors, when the natural groups
of animals were ever better understood, attempted increasingly to assess
the relative merits of each character. The value of this approach, in spite
of individual errors, is best substantiated by the fact that all existing good
classifications are the result of such a posteriori weighting.
The acceptance of evolutionary theory after 1859 did not change
this method as such. It did, however, provide scientific justification. It
now became evident why some characters are better indicators of natural
groups than others. Different characters contain very different amounts
of information concerning the ancestry of their bearers. Weighting, then,
can be defined as a method for determining the phyletic information content
of a character. If character a indicates assignment of a species to genus
A and character b to genus B, we must determine which of the two charac¬
ters (a and b) has the higher information content. It is neither necessary
nor even possible to give a precise numerical value to the relative weight
of each character. Qualitative statements are usually more important than
quantitative ones. In order to assign a species to the correct phylum, to
know that it has a chorda is more important than a thousand measurements.
The scientific basis of a posteriori weighting is not entirely clear,
but difference in weight somehow results from the complexity of the rela¬
tionship between genotype and phenotype. Characters which appear to
be the product of a major and deeply integrated portion of the genotype
have a high information content concerning other characters (which are
also products of this genotype) and are thus taxonomically important. Other
kinds of characters, such as monogenic and oligogenic characters, as well
as superficial similarities, convergences, and narrow adaptations, have low
information contents concerning the remainder of the genotype and are
thus of low value in the construction of a classification. What the descen¬
dants of a common ancestor share is not an aggregate of independent
characters but a whole well-adapted harmonious genotype. Such a genotype
has a considerable evolutionary inertia, and it appears that adaptively
needed modifications can be superimposed on it without destroying it. In¬
deed one might speculate that the characters in a phyletic line which are
most intimately tied up with the basic well-integrated genotype are the
most conservative.
 The Procedure of Classifying 219

Simpson (1962&) has described good taxonomic characters as “readily

observable characters that are believed to be fairly constant within taxa
but different between taxa at any pertinent level.” This is a good empirical
description, but it requires painstaking study of large series of specimens
to demonstrate that characters are “fairly constant” and a delimitation
of the taxa (based on a study of characters!) before it can be shown
that the characters are different between them! Nor does this description
cope with the difficulties produced by the various types of similarity (10.1.2)
and with certain other difficulties to be mentioned later (10.4.5). Indeed,
it is precisely this problem of how to determine what is a good character,
a character with high weight, that has been a main source of concern
among taxonomists.
Methods of a posteriori weighting have been discussed by Hennig
(1950, 1966), Remane (1952), Maslin (1952), Cain and Harrison (1960),
Simpson (1961, pp. 82-106), and Throckmorton (1969). No overall treat¬
ment of this subject has yet been published, and it is still cloaked in uncer¬
tainty. There has been an increasing interest in methodology and a hope
that computer programs can be developed that will facilitate a posteriori
weighting. Actually, there is already a certain amount of implicit weighting
in most existing methods of numerical taxonomy.
The classical approach has been to work backward from classifications
that produce natural groupings and study the characters which delimit
such natural groups. Tooth structure in mammals, wing venation and struc¬
ture of the genitalia among insects, and the structure of the bony palate
among birds are taxonomic characters that are fairly constant within groups
that appear to be natural and are therefore given high weight. Each of
these favored taxonomic characters fails occasionally and in certain groups
frequently. The number of cervical vertebrae (seven) is a class character
in mammals, while it is not even a generic character in birds; for instance,
it fluctuates from 23 to 25 in the genus Cygnus (swans). The only reason
why high weight is given to certain characters is that generations of taxono¬
mists have found these characters reliable in permitting predictions as to
association with other characters and as to the assignment of previously
unknown species.
It has been said with good reason that the trial-and-error method
of improving classification is ponderous and uneconomical. To undertake
a successful a posteriori weighting of characters requires a thorough knowl¬
edge of the history of previous classifications of a given group and an
ability to make value judgments. Yet no clearly better method has so far
been found. Beginnings are being made in the development of computer
methods that endeavor to select characters of special weight in classification
(e.g. Throckmorton, 1969). Until such methods have been further worked
out and more thoroughly tested, a survey may be presented of the traditional
220 Methods of Zoological Classification

considerations in weighting. This will continue to remain important in

all groups with a paucity of taxonomic characters.
10.4.2 Characters with High Weight. A few generalizations can
be made about characters with high taxonomic weight. These include the
following:
Complexity. Complex structures have greater weight than do simple
structures, even when there are more of the latter. This is one of the
reasons why genital armatures in arthropods are of such high taxonomic
importance. They are usually highly complex and differ even in closely
related species, sometimes quite drastically. The probability that species
which are only distantly related would become similar in such structures by
convergence is extremely low. Complex ornamentations, complex cusp pat¬
terns of teeth, complex color patterns, etc., all fall under this category.
Joint Possession of Derived Characters. Hennig (1950 and later) has
articulated a principle followed by taxonomists for generations but appar¬
ently never before spelled out succinctly. It states: “Taxa ought to be defined
on the basis of shared derived characters (synapomorphy) and not of shared
ancestral ones.” The strength of this principle is self-evident. If two forms
share a primitive (ancestral) character, this may be due to the fact that
they have not yet lost the ancestral trait rather than that they are closely
related. The acquisition by two taxa of the same evolutionary novelty is
almost invariably the result of close relationship, rarely of convergence.
Constancy. A character that is constant “throughout large groups of
species” (Darwin, 1859, p. 415) has higher weight than a variable character.
Farris (1966) suggests that low phenotypic variability be given very high
weight. Since monogenic characters sometimes have low variability, this
principle is probably valuable only with relatively complex (polygenic)
characters.
Consistency. A character that is consistently present in one group
and equally consistently absent in related groups has obviously higher weight
than a character occurring sporadically in several groups and differing
merely in the frequency of its occurrence.
The Darwin Principle. Taxonomists have long stressed the relative
importance of characters that do not serve a specific ad hoc adaptation
but are merely, so to speak, an indication of an underlying basic genetic
similarity. As Darwin (1859, p. 414) put it, “The less any part of the
organization is concerned with special habits, the more important it becomes
for classification.” This is why color pattern in birds, a particular configura¬
tion of cusps on molars in mammals, a particular pattern of reduction
or fusion of wing veins and of elaboration of sclerotic structures in the
genitalia in insects are of such high value in classification. It is possible
that the special configuration of some of these structures has an adaptive
significance that has not yet been discovered. In most instances it is more
 The Procedure of Classifying 221

probable that the genotype responsible for the structural configuration was
brought together by natural selection and has adaptive significance as a
whole. The principle stating that a character which is the product of the
general genotype is more likely to have high taxonomic weight than a
character that represents an ad hoc specialization has wide application.
Characters Not Affected by Ecological Shifts. Most higher taxa include
subtaxa (species in genera, genera in families, etc.) that have made an
ecological shift. Any character not affected by such a shift has a higher
weight than characters affected by it. In the mergansers (Mergus), to be
discussed in the following section, the color pattern of the downy young,
the courtship, and other characters not affected by the shift into the fish-eat¬
ing food-niche have higher weight than characters affected by the shift.
Correlated Characters. Taxonomists are in general agreement that
they rely more on correlated character complexes than on any other clue
to relationship. Yet in the weighting of characters a strict distinction must
be made between two kinds of correlations, one of very high and the other
of very low weight. Of low weight are characters that are functionally
correlated and do not deserve to be treated as separate characters because,
being members of a single functional complex, they are redundant (see
10.4.3). Of high weight are characters that are not functionally correlated
but are found in members of the same taxon because these characters
are phenotypic manifestations of the ancestral well-integrated gene complex,
let us say like the diagnostic characters of the Deuterostomia or the Chor¬
data. One might call this kind of concordance phyletic correlation. When
comparing the species of two well-defined higher taxa, let us say taxon
A and B, one always finds that the species within either taxon share certain
functionally uncorrelated characters not displayed by the species of the
other taxon and vice versa. The species of taxon A may have characters
a, b, c, and d, the species of taxon B characters e, f, and g. We infer
that the joint possession of characters by species of a higher taxon is the
result of common descent and assign such concordant, that is, phyletically
correlated, characters (like a, b, c, d and e, f, g) high taxonomic weight.
The character that is concordant with the greatest number of other charac¬
ters has the highest weight. Each species of such a higher taxon, of course,
has many additional characters that do not belong to such correlated charac¬
ter complexes.
Whether an association of characters represents functional or phyletic
correlation is not necessarily evident at first sight. It sometimes requires
a careful functional analysis. In a few cases, indeed, there is no sharp
line between them. Many phyletically correlated character complexes may
have originally started as a functional complex in which the genetic integra¬
tion was retained even after the functional correlation had broken down
owing to a change of function of individual components.
222 Methods of Zoological Classification

10.4.3 Characters with Low Weight. Every specialist knows charac¬

ters which in his group are “unreliable,” that is, poor indicators of relation¬
ship. They have low weight in the grouping of taxa. The characters here
enumerated are usually considered as belonging to this category.
All conditions that represent the converse of characters with high
weight (see 10.4.2) have low weight, other things being equal.
High Variability. All highly or erratically variable characters are in
this category. The pattern of branching in arteries of vertebrates may be
different not only in different individuals of the same populations but even
in the left and the right side of the body. Differences in arterial pattern
are not nearly as helpful for classification as some authors thought they
were. Wing venation provides important characters for the classification
of insects, but Sotavalta (1964) showed that there was far more variation
in this character in the tiger moths (Arctiinae) than known and that
the traditional generic arrangement of the family, based on wing venation,
is thoroughly in need of revision.
Characters known to vary greatly within groups of clearly related
species and to show similarly great variation in other only distantly related
groups. The presence or absence of bands in snails or large body size in
certain groups of mammals are such characters.
The term variable character may mean several things. It may mean
that it is either present or absent in members of the same population or
species. It may mean that it shows many different degrees of expression
in the same species or population. But it also may mean that, although
constant within a given species, it is either present or absent in a natural
group forming a higher taxon. Variability in the characters of higher taxa
may mean something very different from variability in populations.
Totally invariable characters have low weight; for instance, “posses¬
sion of two eyes” in taxa belonging to the vertebrates.
Characters that are too difficult (or too time-consuming) to be deter¬
mined, such as number of hairs in mammals or certain physiological con¬
stants, have low weight.
Monogenic or Oligogenic Characters. A simple monogenic character,
like those involved in balanced polymorphism, usually varies independently
of other characters. Such a character has low phyletic information content.
The distribution of monogenic characters (like albinism) in the zoological
system is often rather haphazard.
Regressive (Loss) Characters. Any kind of regressive character is usu¬
ally of low taxonomic weight, because such losses may happen independently
in more or less distantly related phyletic lines. Taxa based on the loss
of eyes, loss of wings or wing veins in insects, loss of toes or other appendages
in mammals and birds, loss of teeth in mammals, or of segments in seg¬
mented animals, are often unnatural. Trends toward simplification are often
 The Procedure of Classifying 223

erratically realized in different phyletic lines. Losses may be the result of

special environmental conditions (eyes in caves, wings on islands in stormy
oceans), or they may represent tendencies in a higher taxon that are realized
many times independently in subordinate lower taxa. The Sanderling
(Calidris alba), for instance, has lost the rudimentary hind toe which is
still present in related sandpipers and was for this reason placed in a special
genus (Crocethia). Yet this bird is more closely related to certain species
of the genus Calidris (minutilla and pusillus) than these species are to
other species of Calidris like C. acuminata or melanotos. Michener (1949)
has shown how the independent loss of characters leads to spurious simi¬
larities among saturnid moths. Neoteny (sexual reproduction in a preadult
state) has also sometimes led to spurious similarities and proposals for
unnatural taxa. Since the loss of a structure or feature may occur repeatedly
in independent lines, taxa based on the absence of characters are often
polyphyletic and unnatural.
Narrow Specializations. Characters reflecting a single selection pres¬
sure, such as loss of an organ, or desert eoloration in desert birds and
mammals, have lower taxonomic weight than their conspicuousness would
suggest. As Darwin expressed it (1859, p. 414): “Nothing can be more
false” than to assume “that those parts of the structure which determined
the habits of life, and the general place of each being in the economy
of nature, would be of very high importance in classification.” (See also
10.4.2 under Darwin Principle.)
An example may illustrate this. A group of fish-eating ducks, the
mergansers (Mergus, etc.), acquired—in connection with their fish-eating
habits—a number of ad hoc adaptations, such as a streamlined body and
head and a long thin bill with numerous homy teeth. Some ornithologists
placed them in a separate family. Later on it was shown that the mergansers
agree with the Golden-eye group (Bucephala) in the essentials of courtship
display, color of downy young, internal anatomy, and in protein characters.
The two groups also frequently hybridize with each other. The taxonomic
value of the fish-eating adaptations was consequently drastically downgraded
and mergansers and golden-eyes combined in a single tribe. Characters
associated with shifts in the food niche are particularly susceptible to a
rapid attainment of conspicuous differences or, conversely, of convergent
similarities.
Structural adaptations subsequent to the invasion of a new food niche
may happen very rapidly and generally have low taxonomic weight. In
birds, for instance, particularly in the songbirds, the bill is highly plastic,
and flycatcher bills, finch bills, warbler bills, thrush bills, and shrike bills
have developed many times independently. Different subspecies of the same
species may have drastically different bill types, indicating how quickly
the shift from one type to the other may occur. Teeth and jaw structures
224 Methods of Zoological Classification

Megocereso
Neocornegia
Anitoto
Oryocompo
Psilopygoides

Sphingicompo
Bouvierino
Syssphim

Arsenuro
Rhescynfis

Fig. 10-6. Phylogenetic tree of the Saturniidae of the New World, in which all lines where eye
size is greatly reduced are marked with a black spot. Normal eye upper left (from Michener, 1949).

of cichlid fishes in African lakes likewise illustrate the principle of the

rapidity of nutritional adaptations. This is equally true for any other charac¬
ter difference that is the result of a single selection pressure, such as loco-
motory specialization or secondary sexual characters resulting from sexual
selection (e.g. birds of paradise).
 The Procedure of Classifying 225

Redundant Characters. As pointed out in 10.2.3, it is not necessarily

true that the more characters a classification is based on, the better it
is. There is a law of diminishing returns, because different components
of the phenotype ( = different characters) may be pleiotropic or functionally
correlated aspects of the same information in the genotype. A character
is redundant if it is a necessary correlate of other characters. Large size
alone may make species A appear more similar to species B than to a
third small species C even though elimination of the size factor and its
universal correlated effects may reveal that A is actually far more closely
related to C than to B. Proportions are often far more sensitive indicators
of relationship, although the effects of allometry must be evaluated carefully
(Gould, 1966). A tendency toward an increase in meristic elements may
actually be a single factor, even though it may affect all parts of the body
and all appendages that show meristic factors. To score each of them sepa¬
rately would grossly distort actual similarity.
The same peril of redundancy is ever-present with functional char¬
acter complexes. Verheyen (1956), on the basis of an analysis of over
50 characters, placed the diving petrels (a family of Tubinares) in the
same order as the auks (actually related to the gulls) because nearly all
the characters used by him were taken from locomotory adaptations for
wing-diving. The degree of inferred relationship among the anthropoid
apes was distorted until quite recently because the classification was based
almost entirely on arboreal locomotory and vegetarian masticatory adapta¬
tions. Bill, tongue, soft palate, bony palate, jaw muscles, and other structures
in the skull of birds are all adapted to the particular food niche which
a given species utilizes. Individual aspects of these structures cannot neces¬
sarily be scored as so many independent characters.
One can count and evaluate in classification only those characters
that are reasonably “independent” of each other. Just exactly what “inde¬
pendent” is, and how this can be determined, is still controversial. To
reduce the weight of characters according to their degree of correlation
is part of the answer, but not the entire answer, because characters may
be correlated for two entirely different reasons, function or phylogeny.
It is not yet quite clear when to consider phyletically correlated char¬
acters as redundant. A mild amount of redundancy may be useful in con¬
firming a proposed classification. Beyond this point it will be misleading
or, at best, will cause a waste of time. Various mathematical tools, such
as correlation analysis and principal-component analysis, are sometimes very
helpful. In his use of taxonomic characters the taxonomist must always
watch for redundancy and try, instead, to sample as many independent
manifestations of the genotype as possible.
10.4.4 Summary. The experience which taxonomists have gained
from weighting can be summarized in the statement that a classification
226 Methods of Zoological Classification

based on phyletic weighting has numerous advantages. It is the only known

system that has a sound theoretical basis; it has greater predictive value
than other kinds of classification; it stimulates a character-by-character
comparison of organisms believed to be phylogenetically related; and it
encourages the study of additional characters and character systems in order
to improve the soundness of the classification, hence its information content
and predictive value. Finally, it leads to the discovery of interesting evolu¬
tionary problems. Thus classifications based on phyletic weighting not only
have scientific advantages but are actually best able to answer the demands
of the practice by having a greater total information content than artificial
systems.
10.4.5 Difficulties in Weighting and Phyletic Grouping. A number
of difficulties in arriving at an appropriate weight for a given character
were mentioned in the two preceding sections. Some additional ones will
now be discussed.
Reversal. It is not known to what extent reversibility may cause con¬
fusion. Only few exceptions are known to the “irreversibility” rule that
a structure or organ once lost is not reacquired in the same way. The
few exceptions do not seriously weaken the validity of this rule. Far more
frequent however are cases where, owing to a loss of specialization, an
advanced form acquires secondarily a spurious similarity to a primitive
ancestor. Generally such despecialization affects only a single character
or character complex and does not lead to confusion with primary
primitiveness.
Convergence. The secondary acquisition of a character by two taxa,
not derived from a common ancestor showing that character, is called
convergence. Does it ever happen that different taxa, through adaptation
to the same mode of life, become so similar that their different origin
is not recognizable? Convergence is believed by many taxonomists to be
the greatest obstacle against achieving a truly phyletic classification. Actually
the difficulties caused by convergence have been exaggerated. Even though
convergence involving particular characters, entire organs, or general shape
and proportions occurs quite frequently, convergence between the entire
phenotypes of unrelated organisms is rare, if not absent. I am not aware
of a single case among higher organisms. No flying mammal, reptile, or
arthropod could ever be mistaken for a bird. Nor can a whale, a seal,
or a manatee be mistaken for a fish. Convergence is a problem only among
fairly close relatives in rather uniform groups, particularly where classifica-
tory schemes are based on few characters.
Convergence involves almost invariably an adaptation for a similar
niche utilization. That the lagomorphs (hares and relatives) were classified
with the rodents was because of their similar gnawing incisors. When several
kinds of apparently unrelated seed-eating songbirds were combined in the
 The Procedure of Classifying 227

a b a b

Recent Separation

a b

Convergent Evolution

Fig. 10-7. Three alternate reasons for similarity of taxa a and b (from Mayr, 1964).

family Fringillidae, it was because of their cone-shaped bill. Other familiar

examples are the distantly related but similar-appearing families of water
beetles which share a streamlined form; the strikingly similar structure
of the forelegs in mantids (Mantodea) and mantispids (Neuroptera) ; and
the superficially similar ectoparasites of vertebrates, which belong to at
least six different orders of insects. Extreme convergence in response to
adaptive needs occurs also in many physiological properties, as, for instance,
228 Methods of Zoological Classification

those of the blood pigments of marine invertebrates and of neurotoxins

in certain fishes and salamanders. Most cases of convergence are restricted
to a single character or functional character complex. The utilization of
new characters invariably leads to the unmasking of unnatural assemblages.
Mosaic Evolution. The unequal rate of evolution of different features
among descendants from a common ancestor, often combined with parallel
evolution, is perhaps the most frequent cause of difficulties in classification.
Let us assume that there are seven evolutionary trends in a major phyletic
assemblage. In each subline, however, the rate at which these trends are
realized will be different (Table 4-1). When this happens, one may fail to
find correlated groups of characters—an almost indispensable basis for the
clustering of taxa. Also, each taxon will show a different combination of
primitive and advanced characters. This raises the question of which special¬
izing trend should be emphasized in the grouping of taxa and also makes it
quite impossible to arrive at a noncontroversial sequence of taxa. Among
specialists there will always be disagreement as to which of these trends of
specialization has led farthest away from the ancestral condition. There is
no simple solution to this dilemma.
Incongruence among the Information Contents of Different Phena.
Separate classifications based on different phena in a species, let us say
one classification based on males and another on females, or one on larvae
and another on imagoes, often lead to different results. Since the different
phena are either genetically identical (juveniles and adults) or differ only
by sex or morph genes, such a discrepancy indicates shortcomings in the
analysis of similarity. A conflict between larval and adult characters usually
arises from the fact that either stage has acquired certain secondary special¬
izations or regressions and only a deeper analysis will give the right answer
(7.4.5). If it cannot be found, new characters (biochemical, behavioral,
chromosomal) will have to be employed to break the deadlock.
10.4.6 Grouping Species by Weighted Similarity. The long preced¬
ing discussion on how to determine the kind of similarity which indicates
common descent places us now in the position to go back to our original
question: What species shall we unite into a given group or cluster? The
performance of this operation is a consequence of the fact that each higher
taxon (from the genus up) consists of a cluster of species separated by
minor or major gaps from other clusters. (A monotypic taxon is a “cluster”
of a single species.) A species belongs to a cluster because it has had largely
the same evolutionary history as the other species of that cluster and is
therefore—in an evolutionary sense—more similar to them than to other
species. The formal taxonomic recognition of such clusters raises a number
of practical and operational problems that will be discussed in the next
section.
 The Procedure of Classifying 229

10.5 CLASSIFICATION AND INFORMATION RETRIEVAL

Determination of similarity between species is the first step in the

classifying procedure. It would also be the next-to-last step if this would
invariably produce groups of species that are well defined and as isolated
as hummingbirds, penguins, or bats. The question whether a given mammal
is a bat or not never arises. Even flying squirrels or flying lemurs could
not possibly be mistaken for bats. Unfortunately evolution is not so orderly
as to produce only well-defined taxa, neatly arranged like the elements
in the periodical table of chemistry. In almost any assemblage of species
one can distinguish subgroups and supergroups of species, dense and loose
clusters, and a liberal proportion of intermediate or very isolated species—in
other words, one finds gaps of every width. To translate this variation
optimally into a hierarchy of higher taxa is part of the art of taxonomy.
It is during this part of the classifying procedure that the taxonomist
must be most keenly aware of the fact that a classification is an information
storage and retrieval system. The complex relationship of species with each
other and the varying rates of branching and of evolutionary divergence
must be translated into a system of taxa, conveniently ranked in the appro¬
priate categories, in such a way that it forms as efficient as possible a
system of information storage. One must remember that it is possible to
translate the same information about relationship into very different classi¬
fications. It is the purpose of the ensuing discussions to make the taxonomist
aware of various practical and theoretical considerations that will permit
him to produce a classification meeting two objectives: grouping nearest
relatives near each other and facilitating information retrieval. It must
be the principal concern of the classifier to delimit and rank the taxa
in such a way as to meet both objectives.
10.5.1 The Formal Recognition of Taxa. A taxon is defined (1.2)
as a group of species receiving a definite rank in the hierarchy of categories.
The inevitable consequence of this definition is that the two operations,
the formal recognition of taxa and their categorical ranking, are inseparable
in the classifying procedure and must be treated together.
This particular step in classification, formal recognition and ranking
of taxa, is the most controversial one in taxonomy and still subject to
widely diverging opinions. Even though there may be a fair amount of
agreement on similarity among species, the translation of this evidence
into formal classifications shows wide variation. This is apparent from three
lines of evidence:

1. Historical Changes. The genus, the family, and the other higher categories
have changed their value during the history of taxonomy. The various species
230 Methods of Zoological Classification

recognized by Linnaeus are in most cases still listed as species today, while the
status of his genera has completely changed in most cases. Most Linnaean genera
of animals have since been raised to the rank of families or even higher categories.
2. Splitters and Lumpers. There is usually a wide difference of opinion
among contemporary authors on the average size they favor for taxa (see 10.5.3)
and the rank they assign to them. Contemporary treatments of the same higher
taxon may show drastically different categorical assignments. For instance, in
Parker and Haswell’s textbook of zoology (1940) the insects are classified as
a class of the phylum Arthropoda, and the Orthoptera as an order with four
suborders. In Handlirsch’s treatment of the insects in Kiikenthal’s Handbuch der
Zoologie (1926-1936), the insects are listed as a subphylum, and the Orthoptera
are arranged in two superorders and four orders.
3. Different Standards in Different Groups. Extremely different standards
of categorical assignment are traditionally adopted in different branches of the
animal kingdom. The recent birds, for instance, are classified by various authors
into from 20 to 50 orders. There is perhaps less difference between some of
these orders than there is between some of the families of insects or of mollusks.
Likewise the families in the order Passeres (songbirds) are much less distinct
than are the families in most other groups of animals. We find similar inequalities
throughout the animal system. Obviously the higher categories, particularly order
and family, do not have the same meaning for specialists in different areas of
zoology.

The reasons for the complete absence of a uniform standard of ranking

criteria is that the higher categories, in contrast to the species category,
lack a nonarbitrary definition. They have only a formal definition which
states that “a higher category designates the rank of a taxon which consists
of a group of taxa of the next lower categorical rank and which is separated
by a discontinuity from other taxa of the same categorical rank.” There
is no reliable criterion to tell us whether a cluster of species should be
ranked in the hierarchy of categories as a genus, a tribe, a subfamily,
a family, etc., and as a result hardly any two authors agree in the delimi¬
tation and the ranking of taxa.
10.5.2 Ranking Procedures. To get away from the traditional sub¬
jectivity and arbitrariness, two methods have been proposed by which to
establish categorical rank automatically: cladistic and phenetic ranking.
Cladistic Ranking. The cladists (4.3.5) base categorical rank on the
geological time at which the ancestor of a given taxon branched off from
the parental stem (or from a sister branch). This method fails because
it ignores differences in the rates of evolution and confuses genealogical
with genetic relationship. As Darwin has rightly said, the degrees of modi¬
fication which the different groups have undergone are very different. Even
though the family Limulidae (horse-shoe crabs) is older than the mam¬
malian order Carnivora, it does not mean that the categorical rank of
these two taxa has to be reversed. Rapidly evolving taxa often branch
 The Procedure of Classifying 231

off from slowly evolving taxa, but it would be absurd to demand that
the subsequent fate of the daughter taxon should determine the categorical
rank of the parent taxon.
The bird-crocodilian-reptile example (Fig. 4-4) best illustrates this
problem. In almost any assemblage of related taxa, there is one or another
that has started off on a new evolutionary path. Although the branching
point is no more remote in time than that of related taxa, the taxon has
diverged more strongly. This is the “ex-group problem” of Michener
(1957). Whenever the divergent taxon has truly entered a new major adap¬
tive zone, as birds compared to reptiles, or the hominids compared to the
pongids, placing the divergent taxon in a higher category is justified. Such
a separation of a single deviant evolutionary pioneer from another taxon
with which it shares the nearest branching point is not in conflict with
the principle of monophyly (see 4.3.5; also Mayr, 1965a). In most cases,
however, the categorical separation of an aberrant species (as a separate
genus) or an aberrant genus (as a separate family) leads to a fractioning
of the system and lowers its information-retrieval capacity.
Cogent arguments against cladistic ranking are also provided by the
chronology of parasites and their hosts (Osche, 1960). For many taxa of
vertebrate parasites, for instance, it may be very probable that they origi¬
nated together with their hosts, that is, at the same geological time. Should
a genus of cestodes be raised to the rank of a family or order because
it parasitizes a family or order of vertebrates? Should the superfamily As-
caroidea of the nematodes be given the same categorical rank as the class
Cestoda because both invaded the vertebrates at the same time? For other
arguments against the cladistic approach see 4.3.5 and 10.3.
Phenetic Ranking. The phenetic approach is to calculate a measure
of overall similarity between taxa, to set arbitrary numerical levels of differ¬
ence to designate categorical levels, and thus to obtain “automatically”
the correct rank of each taxon. Unfortunately, this unbiological approach
has numerous weaknesses.
Firstly, it is fundamentally unsound to quantify similarity in a com¬
parison of entities as highly heterogeneous as the character complexes of
different taxa (Ghiselin, 19665). The conspicuous phenetic differences
caused by some taxonomically unimportant ad hoc specializations prove
this point convincingly. Striking differences in food-getting specializations
of closely related species and conspicuous secondary sexual characters, as
in male birds of paradise, lead to impressive phenetic differences that have
very little taxonomic weight (10.4).
The second reason, in part a consequence of the first, is that genetic
differences between species and higher taxa are very unevenly reflected
in the phenotype (sibling species!). Also the amount of phenotypic differ¬
ence between higher taxa of presumably similar genetic difference varies
232 Methods of Zoological Classification

from order to order and from class to class. Osche (1960) points out,
for instance, that the subdivisions of the tapeworms (cestodes) differ far
more from each other morphologically than do the taxonomic subdivisions
of the nematodes. Adoption of a uniform phenetic yardstick for the recogni¬
tion of categories would lead to a badly distorted and unbalanced system.
Finally, the particular numerical level of similarity (“phenon level”
of the pheneticists) depends strongly on the particular numerical method
applied in a given case and is often not repeatable when a different com¬
puter program is used. The reasons why unweighted phenetic similarity
is unable to reflect the underlying genetic program correctly were dis¬
cussed earlier (10.2).
Nevertheless there are situations where the phenetic approach is help¬
ful, if not indispensable. The successful application of the weighted ap¬
proach (see below), in which clustering and ranking are carried out by
inspection and overall evaluation, requires experience and a deep under¬
standing of the taxon. The beginner, or someone dealing with a little-studied
group, will have considerable difficulties in determining the proper weight¬
ing criteria. He should by all means employ, as a first step, one of the
computer clustering methods. However, he must remember that an auto¬
mated method is a short cut and that the subsequent decisions (size and
ranking of taxa to be formally recognized) must be based on a careful
weighting of the manifold considerations discussed below (10.5.3).
It has not yet been determined which computer clustering method
gives the best results. Sokal and Sneath (1963) favor the nearest-neighbor
method. Wirth et al. (1966) are experimenting with “graph clustering”
(also based on the nearest-neighbor approach), while Watson, Williams,
and Lance (1967) prefer “centroid sorting.” Further testing will soon reveal
which method is most advantageous under what circumstances. The relation
between grouping, in the classical sense, and clustering, in the sense of
numerical taxonomy, is not yet clear.
Weighted Ranking. Owing to the deficiencies of the automated ap¬
proaches, most taxonomists continue to favor the classical approach of a
careful weighting of numerous separate factors and considerations. The
experienced taxonomist knows that every classification is a compromise
between different requirements, some of which are occasionally in conflict
with each other.
The most frequent conflict is between the practical objectives of any
information retrieval system and our scientific understanding of relation¬
ships. For instance, specialists are often able to determine the probable
relationship of species down to the level of very small groupings. Yet to
establish a separate genus for each of these groups, in extreme cases for
each well-defined species, would completely destroy the value of the classi¬
fication as an information retrieval system.
 The Procedure of Classifying 233

10.5.3 Criteria for Delimitation and Ranking. Scrutinizing the clas¬

sifications of successful taxonomists, it seems that they base the recognition
of taxa on the relative merits of five considerations:

1. Distinctness (size of gap)

2. Evolutionary role (uniqueness of adaptive zone)
3. Degree of difference
4. Size of taxon
5. Equivalence of ranking in related taxa

All five factors must be weighted before each decision (see also Michener,
1957). When a taxonomist, during the classifying procedure, encounters
a group of species that would seem to merit recognition as a new higher
taxon, he will ask questions concerning the stated five points, such as:
Is the new taxon sufficiently different from the taxon with which it is
most nearly related? Is the new taxon of a size that is convenient for
information retrieval? Only after all these questions have been answered
in the affirmative should the taxon be formally recognized.
Distinctness (Size of Gap). The greater the gap between two clusters
of species, the greater the justification for recognizing both clusters as sepa¬
rate taxa. Size of the gap is measured not merely in terms of phenetic
distance but, more important, in terms of the biological significance of
the difference (see following section on the evolutionary role).
Concerning the evaluation of the gap, there is a profound difference
between the recognition of taxa on the level of the species and on that
of the higher taxa. The mere existence of a very special kind of gap (“repro¬
ductive isolation”) is the necessary and sufficient criterion of taxonomic
recognition on the species level. No other condition has to be met, as proved
by the recognition of sibling species. Above the species level the presence
of a gap is only one of several requirements, since not every species is
placed in a higher taxon of its own.
The gaps between taxa are the result of evolution. There is no factual
or conceptual conflict between evolutionary continuity and the existence
of these gaps. Numerous evolutionary processes generate discontinuities in
spite of the complete continuity of populations in time (as far as they
are in an ancestor-descendant relationship). Speciation, extinction, adaptive
radiation, unequal rates of evolution, and other evolutionary phenomena
are responsible for the existence as well as for the unequal size of the
gaps separating higher taxa. It must be emphasized that not only the gaps
but also the clusters of species separated by the gaps are realities of nature.
When there is an appreciable gap between two taxa, the terms “degree
of difference” and “size of gap” mean essentially the same thing. However,
when clusters of species are large and heterogeneous and not separated
by a clear-cut gap, their mean values may be quite different and yet they
234 Methods of Zoological Classification

may not be very distinct. It is not justifiable to select two extreme species
in a widely scattered array of species and make them the types of genera,
if there is not some sort of discontinuity between them. Indeed, the discon¬
tinuity should be reasonably wide and well-defined, in order to preclude
excessive splitting. Michener (1957; 1963, p. 153) discusses the problem
of gap evaluation.
Evolutionary Role (Nature of Adaptive Zone). There has been an
increasing tendency in systematic zoology to give weight in ranking to the
evolutionary potential of a taxon. Almost any prosperous higher taxon is
descended from a founder species (or species group) that succeeded in
shifting to a promising new adaptive zone (birds, beetles, whales, etc.).
Consequently any taxon that has succeeded in entering a new adaptive
niche or zone usually receives higher rank than a taxon that lacks such
ecological significance. It is one of the aims of the new systematics to
investigate the ecological significance of taxa and to use this information
in the weighting of the evidence (5.3.2).
Well-delimited higher taxa almost invariably have a definite ecological
meaning. Cats, dogs, horses, woodpeckers, etc., fill well-defined ecological
niches or adaptive zones in nature. A consideration of its adaptive and
evolutionary role is thus an important element in the categorical ranking
of a higher taxon (7.4.11; see also Gisin, 1964).
The relationship between the hominids and the anthropoids, and its
translation into ranked taxa, is a further illustration of the importance
of evolutionary considerations. The study of chromosomes, of numerous
biochemical characteristics, and of their parasites has revealed so much
similarity between Man and the African apes (Pan) that some authors
have suggested placing them in a single family. Yet Man has entered so
unique and strikingly distinct an adaptive zone that Huxley even suggested
recognizing for him a separate kingdom (Psychozoa). This would seem
to go too far, but the evolutionary distinctness of Man surely justifies recog¬
nition of a separate family.
Evolutionary considerations are also very important for the ranking
of taxa along a single phyletic line (without intervening branching). Evi¬
dence is mounting that the polytypic species Australopithecus africanus
(sensu lato) is the immediate ancestor of Homo. Yet, the man ape africanus
with a mean brain size of 450 cc filled such an entirely different niche
from sapiens with a brain of ± 1,500 cc that generic separation is abun¬
dantly justified.
It is the difference in the utilization of the environment, the difference
in the occupation of an adaptive zone, that is responsible for the width
and the sharpness of the gap between taxa.
Degree of Difference. When speaking of the degree of difference be¬
tween two clusters of species, the taxonomist usually thinks of the “distance”
between the means of the two groups of species. This value can be employed
 The Procedure of Classifying 235

only in conjunction with two other data, the scatter of the cluster and
the amount of discontinuity (gap) between the cluster and others. The
denser and more uniform a cluster of species, the more justified one usually
feels in recognizing it formally. The greater the difference between
the most distant species of a cluster, the greater is the justification for a
splitting of the taxon, other things being equal. This is an important con¬
sideration in the evaluation of the relative weight of cluster and of gap
characteristics (Fig. 10-8). In the upper two clusters, the mean difference
(A-B) between the clusters is greater than in the lower clusters
but the size of the gap (a-b vs. a'-b') is smaller.
Optimal Size of Taxon. The number of species included in a taxon
constitutes its size. A genus with many species is called large, one with
few species small. Since a classification functions like a filing system, its
subdivisions (the taxa) should fulfill the demands of greatest possible effi¬
ciency, that is, they should ideally be of approximately equal size in order
to facilitate information retrieval. The amount of material included under
one heading should be neither too voluminous nor too scanty. As a taxono¬
mist once remarked, all one can say about the genus is that it should
be neither too large nor too small. There are a number of biological reasons
that make it impossible to achieve this ideal.
The fact that the two processes of phylogeny, branching and diverg¬
ence, are independent of each other is responsible for the observed inequality

m x n X
X X X
X
X
X
X x b
•e
x
X
X X
X
X X
X

(1)

X
X •A1 X
X
X X X

(2)
Fig. 10-8. Mean distance of clusters and size of gap are independent. Above the mean distance
between the clusters A and B and their diameters is great, but the gap is narrow. Below
the clusters arc nearer and more compact, but the gap {a'-b') is greater.
236 Methods of Zoological Classification

in the size of taxa. Continuous speciation (branching) without visible diver¬

gence results in the production of large taxa. Divergence without branching
leads to the occurrence of monotypic taxa, in an extreme case to a series
of superimposed monotypic higher taxa. When the line which includes
the ancestor of the Aardvark (Orycteropus afer) split off from its parental
line (Condylarthra?), it diverged steadily. Whatever branching (speciation)
occurred in former times has since been obliterated by extinction. This
genus became sufficiently different to be ranked as a separate family
(Orycteropodidae) and eventually deserved to be ranked as a separate
order (Tubulidentata). The higher taxa in this line (genus, family, and
order) satisfy the definition of “classes” in the language of logic, each
with a single member, and in spite of their different rank, each with the
same member, Orycteropus afer.
The taxonomist would say that the distinctness of the species
Aardvark is of such an order of magnitude that recognition of the separate
order Tubulidentata is justified (and with it the subordinate categories
family and genus), while the logician would say that the Aardvark has
a series of properties which would qualify successively as defining the names
Orycteropus, Orycteropodidae, and Tubulidentata (Buck and Hull, 1966).
The independence of branching and divergence results in a frequency
distribution of genera and species usually referred to as “hollow curve”
(Fig. 10-9; see Mayr, 1942, p. 288). When the number of species per
genus is plotted for a family or an order, it is found that many genera
have only one or twro species, while a few genera contain a large or very
large number of species. Excessively large genera as well as an excessive
number of monotypic genera reduce the usefulness of a classification for
information retrieval. This is the reason for the recommendation that the
size of the gap justifying the separation of a higher taxon should be inversely
correlated to the size of the taxa (5.4, 5.5). The recommendation to strive
for taxa of equal size unquestionably leads to value judgments, therefore
it is opposed by those who want to automate classifying.
Most higher taxa of organisms contain some 20 to 30 percent mono¬
typic genera. The Okapi, the Ostrich, and many other well-known animals
are monotypic and in many cases the only representatives of an entire
family or order. Yet the taxonomist must make every effort to keep the
number of monotypic taxa as low as possible. Slightly aberrant species
should whenever possible be included in the same genus as the most nearly
related species. The same advice applies to higher taxa. General similarity
should be given far greater weight than conspicuous difference in a single
character. A highly unbalanced classification of the African plantain eaters
(Musophagidae) resulted from the recognition of three monotypic genera
based almost entirely on slight differences in the shape of the nostril
(Moreau, 1959; Fig. 10-10). Mayr (1942, pp. 280-289) has pointed out
 The Procedure of Classifying 237

1,500
|
' No. of genera No. of species
1,400
1,446 1
1,300 539 2
511 3-4
1,200 350 5-8
288 9-16
1,100
179 17-32
1,000 99 33-64
64 65-128
900 17 129-256
14 257-512
800
3 >513
700

600

500

400
\
-
300
-\
200
V
100
.
100 200 300 400 500

No. of species
Fig. 10-9. Example of a hollow curve: The number of species in 3,510 genera of weevils (Cur'
culionidae) (after Kissinger, 1963).

how unhelpful for information retrieval are classifications that contain too
many monotypic taxa.
The converse is equally true. There are some well-defined natural
genera with over 1,000 or even over 2,000 species. Among well-known
large taxa are the North American freshwater fish genus Notropis with
about 120 good species, certain genera of weevils (Curculionidae) with
more than 1,000 species each, in addition to the entire family of weevils
with perhaps more than 100,000 species, and the genus Drosophila with
more than 1,000 species (Fig. 1-1).
A consideration of size becomes important whenever there is a question
whether or not to split a taxon. Splitting a genus of more than 1,000
species requires much less of an excuse than splitting a small genus. The
larger a cluster of species, the smaller the permissible gap between it and
other clusters. For instance, in a taxon with 1,000 species a narrow gap
separating 400 species from 600 other species may be a better justification
for a split than a much larger gap in a species cluster separating a single
species from three others.
Instead of breaking up such a well-known entity as the genus Droso-
238 Methods of Zoological Classification

tauraco TAURACO t
M. MACRQRRYNCHUS C COR YTM AIX

f
o

T. ERYTHROLOPHUS T. BANNE R MANI

T. C.POPPHYREOLOPHUS T. J. JOHNSTON I

Fig. 10-10. Types of beak and nostril in tauracos (Musophagidae). Nostrils may be concealed or open,
round, oval, or slitlike (from Moreau, 1959).

phila, it is more useful to introduce subdivisions. The subgenus is available

under the genus, but the liberal use of the informal category species group
is an even more useful device, as a relief for one’s memory. Drosophila
specialists have adopted a few subgenera, but on the whole they get along
extremely well by speaking of the repleta group, the willistoni group, the
melanogaster group, etc. Nothing would be gained by chipping off a few
aberrant species from such a large group and making them the types of
monotypic taxa. The species group is the most neglected category in many
areas of systematic zoology.
Splitters and Lumpers. No nonarbitrary method is known by which
the proper size of taxa can be determined unequivocally, except in the
case of very isolated species clusters. This is the reason for the eternal
argument between splitters and lumpers, two types of taxonomists found
 The Procedure of Classifying 239

among the specialists of almost any group of organisms. It is the ideal

of the splitter to express every shade of difference and degree of relationship
through the formal recognition of separate taxa and their elaborate categori¬
cal ranking. It is the ideal of the lumper to express in the classification
the fact that the higher taxa and categories express relationship and that
too fine a division of taxa leads to an intolerable burden on the memory.
The author of this volume believes in broad taxa and sides, therefore,
with the lumpers.
Among the reasons for splitting are the following. The splitter is
aiming at the ideal of recognizing only homogeneous groups. However,
evolution being disorderly, it is virtually impossible to find a polytypic taxon
in which some species do not differ more from each other than do others.
Removal of these species into separate taxa leads to an ever finer pulveriza¬
tion of the system until finally most taxa become monotypic. Splitters almost
without exception classify characters rather than groups of organisms.
Whenever a splitter finds a species with an aberrant character, he establishes
a separate genus for such a species. He fails to remember that since the
time of Linnaeus it has been realized that it is the taxon that gives the
character and not the reverse (5.4.2). Roewer’s splitting of the Opiliones
is a notorious case. He placed 1,700 species in 500 genera, of which more
than half (300) contain only one or two species. Subsequent work has
shown that most of these “genera” are meaningless biologically, being some¬
times based even on different individuals of the same population. In the
Hominidae no less than 30 generic names have been proposed for what
at most are three taxa: Homo, Australopithecus, and Paranthropus.
Splitting is particularly deleterious on the generic level. The generic
name is part of the scientific name of an organism and can therefore be
employed more advantageously to indicate affinity than can the name of
any of the other higher categories. McKevan (1961) and Sailer (1961)
have described particularly well the disadvantages of excessive splitting.
The specialist must have a regard for the general zoologist; he must always
remember that he is not writing merely for three or four cospecialists.
The difference in the attitude of lumpers and splitters may be illus¬
trated with a concrete example. Figure 10-11 represents a dendrogram
(10.7) of the 36 living species of river ducks of the genus Anas (sensu
lato). There is comparatively little difference of opinion among ornitholo¬
gists as to the grouping of these species. It is agreed that there are several
more closely bunched clusters of species, each of which forms a natural
group, such as species 16-21 (mallard group) and species 30-36 (blue¬
winged teal-shoveler group). It is also agreed that certain species, such
as 1, 2, 7, 22, and 23, are rather isolated, and that some of the species
and species groups are somewhat intermediate between others. Furthermore,
ornithologists agree that the whole group is fairly well isolated from the
240 Methods of Zoological Classification

landica, 13 = gibberifrons, 14 = caslanea, 15 = bernieri, 16 = melleri, 17 = rubripes, 18 = platy-

rhynchos, 19 = undulata, 20 = poecilorhyncha, 21 = luzonica, 22 = specularis, 23 = specularoides,
24 = acuta, 25 = georgica, 26 = bahamensis, 27 = erythrorhyncha, 28 = versicolor, 29 = punctata,
30 = querquedula, 31 = discors, 32 = cyanoptera, 33 = platalea, 34 = smithi, 35 = rhynchotis,
36 = clypeata.

other genera of the tribe Anatini. The congeneric status of this group
of 36 species is supported by their biochemical and anatomical similarity,
by a remarkable interspecific hybrid fertility and by the joint possession
of many behavioral characteristics (Johnsgard, 1965, p. 131).
The “splitters” interpret this evidence as indicating that this group
of species should be divided into at least 12 genera, many of them mono-
typic. They insist that this is the only way of indicating the existence of
the various groups of species. The lumpers, on the contrary, feel that rela¬
tionship within the duck family, and a balanced presentation of the relative
distinctness of other genera of ducks, is best accomplished by including
all 36 species in a single genus.
 The Procedure of Classifying 241

Lumping, however, can also be carried too far. When lumping be¬
comes fashionable in a previously oversplit group, a few authors invariably
go too far in their zeal and combine elements of such heterogeneity that
their genera become almost as useless as the virtually monotypic genera
of the splitters. Excessive lumping tends to produce a good deal of subjective
(secondary) homonymy which is difficult to handle nomenclatorially. Fi¬
nally, it results in very long and rather heterogeneous lists of synonyms.
These drawbacks of lumping would be very minor if it were not for the
preceding creation of useless names by the splitters.
Every branch of systematic zoology seems to go through a cycle. As
more and more species are discovered, the taxa are split more and more
finely, and a steady process of escalation of the categories to higher rank
seems to occur. Finally, a saturation point is reached in the splitting, and
a vigorous period of lumping follows. Let us take the case of birds. Linnaeus
arranged the 554 species of birds known to him in 63 genera (8.8 species
per genus). Some ornithologists in the 1920s allowed no less than 7,000
to 8,000 genera for the 8,600 species of birds now known. In the 1930s
these were reduced to about 2,300 genera, and now perhaps 1,700 genera
are recognized (5 species per genus).
Equivalence of Ranking in Related Taxa. A very important considera¬
tion in ranking is balance. Categorical rank in related taxa should be equiva¬
lent. For instance, the genus should designate a similar category in the
Carnivora and Rodentia—in fact, as far as this is possible, even in birds
and reptiles. The family (if possible) should have a similar significance
in flies and beetles, and so forth. To achieve such equivalence Crowson
(1958) has proposed that norms be adopted for each taxonomic group.
One might, for instance, adopt the Rodentia as yardstick for a mammalian
order. No other group of species of mammals should be recognized as
an order unless it has a combination of qualities which make it equivalent
to the rodents. The time has not yet come to implement this suggestion,
but it seems to come far closer to a realistic solution of the problem of
ranking than the adoption of an arbitrary numerical value of phenetic
similarity.
In the past, splitters have too often concentrated their attention on
a given taxon, raised it in rank, and then subdivided it finely. As a result
such a taxon is completely out of step with all equivalent related taxa
(not similarly treated by the splitter). Arkell and Moy-Thomas (1940)
cite some typical examples of this ill-advised procedure:

Examples of flagrant disregard of this rule [a uniform scale of values]

are Buckman’s innumerable genera made by splitting up contemporary spe¬
cies of the single good Liassic ammonite genus Dactylioceras, and by Heinz’s
pulverization of the Cretaceous lamellibranch genus Inoceramus. Out of
what was originally a single genus Inoceramus Heinz created a whole sys¬
tematic hierarchy, comprising 2 families, 24 subfamilies, 63 genera, and
242 Methods of Zoological Classification

27 subgenera; and even so he did not take the Jurassic forms into account.
[Even if this minute subdivision were taxonomically justified] it should be
carried out downwards in the scale, starting with the genus Inoceramus
and proceeding through subgenera to groups and sections. All the advantages
of minute subdivisions can thus be achieved without disturbance to the
whole classification of Mollusca.

In order to determine how many categorical levels are needed to

express degrees of relationship, a careful analysis of morphological similarity
and also of behavioral and ecological information is needed. To reach
the proper compromise between too little or too much is not always easy.
For instance, there are only two formal categories (suborder and super¬
family) between order and family. Yet recently acquired biological informa¬
tion indicates that there are three levels of difference between the order
Orthoptera and the included families: the Saltatoria and Cursoria within
the Orthoptera, the Ensifera and Caelifera within the Saltatoria, and the
Tettigonioidea and the Grylloidea within the Ensifera. The Tettigonioidea,
in turn, are divided into several families, including the Tettigoniidae and
Gryllacrididae. Utilization of the category “infraorder” is indicated in such
a case.
The same levels of (genetic) relationship are frequently correlated
in different higher taxa with different degrees of morphological change.
Thus in the Nymphalidae, a family of butterflies, the subfamilies are sepa¬
rated by smaller gaps and show less morphological diversity than do the
subfamilies of another family, the Papilionidae. In order to achieve equal
standards of morphological difference the splitter would be inclined to
raise the subfamilies of Papilionidae to family rank and thereby generate
a categorical escalation throughout the system of the Lepidoptera. If fam¬
ilies and genera of cestodes were based on the same amount of differences
as in the nematodes, it would lead to a great deal of splitting of the former.
In short, the recognition of taxa and their ranking must be undertaken
in such a manner as not to upset the equivalence of categories in related
taxa.
10.5.4 Special Problems. A number of special problems require sepa¬
rate discussion.
Intermediate Species. If a species is intermediate between two genera
(or other higher taxa), it should be referred to that genus which includes
its nearest relatives. In practice this rarely causes difficulties. What does
occur, however, is that a species l of genus A may be more different from
the most distant species m of its own genus [Fig. 10-8, (1) ] than from
the nearest species n in genus B even though separated from it by a decided
gap. Nevertheless, since the most distant species of genus A are connected
by a whole array of intermediate species, while genera A and B are sepa¬
rated by a well-defined gap, there is no contradiction.
 The Procedure of Classifying 243

Very large species groups may be recognized as separate taxa even

if there are a few intermediate species. For instance, the characteristic
families of Chrysomelidae (leaf beetles) and Cerambycidae (long-horn
beetles) are connected by a few intermediate genera. Yet the two groups
are so large and the connecting element so small in comparison with the
divergent element that it would completely defeat the objectives of sound
classification to combine the two families into one (Michener, 1957). In
such a case one may have to assign the intermediate species somewhat
arbitrarily to one or the other of the related higher taxa.
Parallel Evolution. A particular difficulty is caused by parallel evolu¬
tion. “Parallelism is the development of similar characters separately in
two or more lineages of common ancestry and on the basis of, or channeled
by, characteristics of that ancestry” (Simpson, 1961). The readiness with
which taxa in different (but closely related) phyletic lines respond to the
same selection pressure with similar phenotypic changes permits important
inferences on the degree of similarity of their genotypes.
Parallel evolution is extremely widespread, since most of the genotype
remains untouched during speciation and phyletic branching. When orangs
(Pongo) and gibbons (Hylobates) acquired certain structural similarities
in connection with a brachiating arboreal mode of life, this was largely
an activation of dormant tendencies already present in the genotype in¬
herited from the common ancestor. Nasute soldiers evolved twice indepen¬
dently in the termite subfamily Nasutitermitinae; strongly asymmetrical
soldier mandibles evolved in three genera of the subfamily Termitinae;
phragmotic heads (a defensive adaptation for plugging holes in the nest)
evolved in three different genera of the family Kalotermitidae. A secondary
jaw articulation originated at least 14 times in the class Aves (Bock, 1963).
The innate potential is often realized in a rather haphazard manner.
Hennig (1950, p. 176) cites stalked eyes as an example; these occur widely
among the acalyptrate dipterans in scattered species and genera, but no¬
where else in the large order Diptera. The potential for acquiring such
a character indicates relationship, but this does not justify combining into
a taxon those species in which the potential manifests itself in the visible
phenotype.
The phenomenon of parallel evolutionary changes in related groups
is particularly often encountered by paleontologists. The emergence of mam¬
mals from various lines of therapsid reptiles, the repeated emergence of
teleosts from holostean fishes, the independent evolution of similar func¬
tional types from separate lines of rodents, and many cases described by
invertebrate paleontologists (e.g. Arkell and Moy-Thomas, 1940), are fa¬
miliar illustrations.
The possibility exists in such cases that the morphological changes
in related lineages parallel each other to such an extent that they will
244 Methods of Zoological Classification

go through the same evolutionary stages or grades, as Huxley (1958) has

called them. When such grades of different lineages resemble each other
far more closely than do stages in a single phyletic line, the taxonomist
faces a dilemma. Should he adopt a “horizontal” classification by combining
similar grades in a single taxon? There is no easy answer. The redefinition
of monophyly (4.3.5) and the realization that we aim to classify on the
basis of inferred genetic similarity make us increasingly inclined to favor a
certain amount of horizontal classification in such cases (quite independently
of additional practical advantages). The delimitation of the mammals
against the therapsid reptiles is an illustration. The experienced taxonomist
must use his judgment as to what particular balance between horizontal
and vertical classification he wants to adopt (see MacIntyre, 1966).
In view of the important function of a classification as an information
retrieval system, we recommend that genetic relationship resulting from
parallel evolution be given the appropriate weight in the delimitation of
higher taxa. Horizontal classifications are permissible only when based on
all considerations, not just on one single character complex. A particularly
strenuous effort must be made to exclude all similarities caused by conver¬
gence.
Evolutionary Continuity. If the fossil record were complete, a delimita¬
tion of taxa against each other would be a near impossibility (2.5.1). Fortu¬
nately for the taxonomist (as regrettable as this is for the evolutionist),
the fossil record is so full of gaps as to permit a clear-cut delimitation
of taxa in most cases. However, the record is beginning to be sufficiently
complete to raise some real difficulties. The delimitation of the fossil
Creodontia from the derived fissipede carnivores is such a case (MacIntyre,
1966). The separation of the mammals from the mammal-like reptiles is
another case. Eohippus is about as good an ancestor for tapirs and rhi¬
noceroses as for the Equidae. Again, the taxonomist must base his ultimate
decision on a careful evaluation of cladistic and phenetic considerations
as well as on the question of what classification would yield the most infor¬
mation.
Summary’. The discussion of the many criteria affecting decisions on
the delimitation and ranking of taxa should have made clear why there
is so much disagreement among specialists. It depends on tradition, on
the maturity of the existing classification in a given area, on the richness
of species in a group, on the degree to which the phenotype reflects (or
exaggerates) changes of the genotype, and on the number and scope of
the specialists, among other factors. Presumably there will always be special¬
ists who tend toward splitting and others who tend toward lumping. Some
want to express degree of difference in their classification, others want
to emphasize relationship. Also, there will be those who stress recency of
common ancestry (cladistic aspects) and those who simply rely on degree
of “similarity.” Nevertheless, there has been an unmistakable trend in recent
 The Procedure of Classifying 245

decades toward more uniform standards. These have already developed

within restricted taxa (birds, Coleoptera, isopods), and the inequality of
standards is, on the whole, not a severe impediment to communication.

10.6 IMPROVEMENT OF ESTABLISHED CLASSIFICATIONS

Discussions of taxonomic procedure tend to create the impression

that the act of classifying is always a de novo operation, as if the taxonomist
always had to classify huge piles of unassorted species. Actually, the greater
part of our zoological system is mature, sound, and stable. The student
of mammals does not have to discover the differences between marsupials
and placentals or between primates and rodents. The entomologist has
no trouble in delimiting Odonata, Lepidoptera, or Coleoptera; there has
been no difficulty about this since long before Linnaeus. What then are
the real problems of classification in modern taxonomy? The answer de¬
pends on the particular group being studied.
Most acts of classification in recent organisms are minor rearrange¬
ments of existing classifications. For instance A (a -f- b -f- c) and B{d -f- e)
are reclassified into A'(a-\-b) and B'(c d -|- e) or into A"(a-\-e)
and B"{b -f- c -f- d). The recent classification of the echinoderms (Table
10-3) may be given as a case in which the primary classifying criterion
was replaced. Such rearrangements of taxa occur at all levels of the hier¬
archy of categories. An entire breaking-up of existing taxa into much smaller
pieces and their reassortment into a totally new set of higher taxa happen
rarely and only in poorly known groups. The reclassification of the rhab-
docoels (Turbellaria), or some proposals for a reclassification of the sponges
(Porifera) discussed by Levi (1957) and Hartman (1958), are examples
of rather drastic reclassifications.
In the older classifications the turbellarians were classified entirely
on the basis of the presence and configuration of the intestinal tract. How¬
ever, the single feature of body size has a dominant influence on the mani¬
festation of the intestines. They are often absent in very small species but
highly elaborate in the larger species (to keep the ratio of intestinal surface
to body volume reasonably constant). The Rhabdocoela of the older litera¬
ture, turbellarians with tube-shaped intestines, were a rather heterogeneous
assemblage. The replacement of this single-character classification by an¬
other one, yolk content of the eggs and associated mode of cleavage, has
not been a full success. This specialization is indicative of a grade and
may well have been acquired repeatedly. Which subclass one adopts for
the orders of turbellarians depends entirely on the primary character chosen.
The reconstruction of the ancestral characters and of the common ancestor
as a whole is still controversial (Ax, 1961).
246 Methods of Zoological Classification

Table 10-3. Effect of Alternate Classifying Criteria on the

Classification of the Echinodermata
(from Fell 1965)

Conventional classification based Classification based on additional char¬

on growth form acters, unrelated to growth habits

Subphylum Pelmatozoa: Subphylum Homalozoa:

Class Carpoidea Class Carpoidea
Subphylum Crinozoa:
Class Cystoidea Class Cystoidea
Class Eocrinoidea Class Eocrinoidea
Class Paracrinoidea Class Paracrinoidea
Class Blastoidea Class Blastoidea
Class Edrioblastoidea Class Edrioblastoidea
Class Crinoidea Class Crinoidea
Subphylum Echinozoa:
* Class Helicoplacoidea *
Class Edrioasteroidea Class Edrioasteroidea
Subphylum Eleutherozoa:
Class Ophiocistoidea Class Ophiocistioidea
Class Echinoidea Class Echinoidea
Class Holothuroidea Class Holothuroidea
Subphylum Asterozoa:
Class Asteroidea Class Stelleroidea (including
Class Ophiuroidea the subclasses Somasteroidea,
Asteroidea and Ophiuroidea)

* Hclicoplacoidea unknown before 1963.

Table 10-4. Inconclusiveness of Classifying Characters in Two

Genera of Calcareous Sponges (from Vacelet, 1961*)

Nucleus of Trirad iate

Taxon Larva
choanocyte spicules

Subclass Calcaronea Apical Amphiblastula Sagittal

Genus Murrayona Basal ? Sagittal
Genus Minchinella ? Filling cell Parenchymula Sagittal
Subclass Calcinea Basal Parenchymula Equiangular

* Syst. Zool ., 10:45-47.

The genera Murrayona and Minchinella of the Pharetronida seem to have a mixture of the characters
of the two subclasses of the Calcarea (Calcaronea and Calcinea).
 The Procedure of Classifying 247

10.6.1 Classificatory Activities. The problems of classification can

perhaps be tabulated as follows (without claiming to exhaust all possibilities) :

1. The assignment of a newly discovered species to the right genus by

asking these questions:
a. Is it possible to include it in an established genus?
b. Is it necessary to erect for it a new genus and possibly for this genus
a still higher new taxon?
2. The transfer of an incorrectly placed taxon to its proper position. This
includes the shift of a species to a different genus, the shift of a genus
to a different subfamily or family, and so forth.
3. The splitting of a taxon into several of the same rank (genus, family,
etc.), either by cleaving a heterogeneous assemblage of species into several
smaller, more homogeneous ones, or by removing an alien element from
an otherwise homogeneous taxon.

When breaking up too large a taxon, certain rules must be obeyed in

the ranking and naming of the resulting new taxa.

a. The rank of the original taxon is to- be maintained if at all possible.

Finer discrimination is possible by the elaboration of subtaxa. For in¬
stance, it is usually less desirable to raise a heterogeneous family to the
rank of superfamily and then recognize as families all the previously
recognized subfamilies, than to develop a finer subdivision into tribes
and genus-groups.
b. No taxon should fall out of step with its sister-groups. The classifica¬
tion of fossil man by certain anthropologists is a warning illustration
of an unbalanced classification (see 10.5.3).
c. A minimal number of names is desirable. By adopting informal group¬
ings such as “species-groups” (instead of new genus or new subgenus)
and “genus-group” (instead of new family, new subfamily, or new
tribe), the same information can be conveyed without a burden on
the memory and without disturbing the balance of the hierarchy of
categories (10.5.3).
d. Subdivide an inconveniently large taxon only if it can be “cleaved,”
that is, if it can be divided into taxa of approximately equal size.
Splitting off a number of monotypic genera from a genus with 500
species would only impede information retrieval. This advice does not
concern the removal of clearly alien elements from currently recog¬
nized taxa.
4. The raising in rank of an existing taxon, a genus to a subfamily, a
subfamily to a family, etc.
5. The fusion of several taxa of the same rank and the synonymizing of
those with junior names.
6. The reduction in rank of a taxon as for instance that of a genus to
a subgenus, a family to a subfamily, etc.
There is very little doubt that certain groups of animals are badly
oversplit and that natural taxa in these groups are ranked in higher
248 Methods of Zoological Classification

categories than necessary. This is true of both birds and fishes. Even the
specialists concerned admit that there is little justification for having 412
families of fishes and 171 families of birds. But which of these should
be reduced to the rank of subfamilies? There is no easy answer.
For instance, there is a well-defined group of songbirds in the Old
World tropics, the drongos or crow shrikes. In spite of their wide dis¬
tribution and numerical abundance, they consist of only 2 genera and
about 20 species. Up to now, not a single good morphological character
diagnostic for this group has been found, and yet in general habitus and
in behavior they stand reasonably well apart from all other songbirds.
Ornithologists would be perfectly willing to consider the drongos a sub¬
family or perhaps a tribe of some other songbird family, but as yet no
character is known that would help in finding that family. Among the
families that have been suggested are the Campephagidae (cuckoo-
shrikes), the shrikes (Laniidae), the Muscicapidae (flycatchers), the
Paradisaeidae (birds of paradise), and the Stumidae (starlings). In
desperation ornithologists finally raised the drongos to the rank of a
family, the Dicruridae, while perfectly willing and ready to reduce this
rank as soon as additional information becomes available. The same is
true for at least twenty other families of birds.
7. The creation of a new higher taxon not by raising the rank of a taxon
(e.g. a family to superfamily rank) but by making an entirely new group¬
ing of taxa of the next lower rank. The proposal of a new superfamily
for a number of existing families or a new order for a series of families
illustrates this procedure.
8. The search for the nearest relative of an isolated taxon and, if it is
successful, the study of the question whether a new taxon of higher
rank should be created for the newly established group of relatives.
For instance, behavioral and anatomical researches indicate that the
Tubinares (shearwaters, etc.) are the nearest relatives of the penguins
(Impennes). Should one establish a superorder for these two orders?

10.6.2 Stability. During such minor improvement activities a deter¬

mined effort must be made to disturb the stability of the currently prevailing
classification as little as possible and to maintain, if not improve, its informa¬
tion retrieval qualities. The usefulness of a classification as a communication
system stands in direct relation to its stability, which is one of the basic
prerequisites of any such system. The names for the higher taxa serve
as convenient labels for the purpose of information retrieval. Terms like
Coleoptera or Papilionidae must mean the same to all zoologists in order
to have a maximum of usefulness. This is even more true for the genus,
the name of which is included in the scientific name (13.1).
When a well-established taxon is found to be somewhat heterogeneous,
it is often inadvisable to split it into several taxa of the same rank if
the components are each other’s nearest relatives.
 The Procedure of Classifying 249

For example, if cytological or biochemical investigation reveals that a certain

genus consists of two groups of species, each with a different series of
chromosome numbers or biochemical reactions, it is not necessary, in order
to record and study this phenomenon, for the genus to be split into two
separate genera, even if certain characters are correlated with the two groups.
Instead, one can deal quite satisfactorily with the situation by leaving the
limits and nomenclature of the genus intact, and by making a special classi¬
fication of its species into the two groups, giving them ad hoc designations
such as group A and group B [or species group A and species group 5],
thus enabling one to refer to them and study them just as satisfactorily
as if one had split the genus into two—with the great advantage of leaving
the general classification intact, stable, and perfectly serviceable for its pur¬
pose. (Gilmour, 1961, p. 37).

The currently adopted zoological system contains numerous taxa sus¬

pected of being polyphyletic owing to convergence. Among the birds there
are many such groups, particularly among the songbirds (titmice, warblers,
babblers, flycatchers, shrikes, finches, etc.). As long as it remains unknown
what the nearest relatives of the components of such unnatural groups
are, it is far better to retain these groups provisionally, for ease of reference.
Such provisional classifications must be abandoned, however, as soon as
the true relationship of the components is established.

10.7 PRESENTATION OF A CLASSIFICATION

When publishing the classification which has resulted from one’s tax¬
onomic studies, one must present it either as a printed list or as a diagram.
Both methods of presentation raise problems.
10.7.1 The Printed Sequence. The technology of printing enforces
a linear, one-dimensional sequence upon any printed classification. One
species will have to come first, another species last, while all others will
have to be listed sequentially between the first and the last. How can
we determine the simplest, most convenient sequence of species?
When the classification of a group is still entirely obscure and catalogs
consist merely of lists of nominal species, an alphabetical sequence is often
most convenient for information retrieval. However, an alphabetical listing
lacks the heuristic value of a classification arranged according to inferred
genetic relationship. By not placing closest relatives near each other, such
a listing makes it difficult to undertake evolutionary studies. Finally, there
is no stability because it necessitates a change in the sequence every time
the name (synonymy!) or the rank (e.g. shift from species to subspecies)
of a taxon changes (Mayr, 1965c). Species in all better-known groups
should be listed according to their relationship with each other. However,
this raises various difficulties.
250 Methods of Zoological Classification

The multidimensional phylogenetic tree with the dimensions of time,

space (longitude and latitude), and adaptational divergence must be con¬
verted into a single dimension. In order to do this the taxonomist inevitably
must compromise between various considerations. Most important among
these are the following three:

1. Continuity. Each species is to be listed as near as possible to its closest

relatives.
2. Progression. Each series of species or higher taxa should begin with the
one closest to the ancestral condition (“the most primitive one”) to be followed
by derived taxa which deviate increasingly from the ancestral state.
3. Stability. One should not make “experimental” changes from previously
adopted sequences, unless the latter are proved unequivocally wrong. A classifica¬
tion is a reference system, and it greatly helps, particularly in a comparison of
faunal lists, if different authors use the same sequence.

The three principles are often in conflict with each other (particularly
1 and 2). It is sometimes possible to establish a well-defined morphological
sequence without being able to state which end of the sequence is the
more primitive. In other cases, there is a dual progression from a group
of primitive species toward tw7o specialized extremes (Fig. 10-12). Instead
of dividing the closely related primitive species into two groups, one leading
to one extreme and the other to the other extreme, as might be demanded
by the progression principle, it usually results in greater continuity if one
starts at one specialized end and establishes a single sequence by first de¬
scending to the most primitive species and then ascending again to the

Fig. 10-12. Arranging 23 species on a dendrogram in a linear sequence. Other alternatives are: 1-13,
17, 16, 15, 14, 18-23, or 1-4, 10-13, 5-9, 17-14, 18-23.
 The Procedure of Classifying 251

other extreme. This avoids a more or less arbitrary split through the middle
of the group of primitive species.
Because of mosaic evolution, most groups show several trends of spe¬
cialization at one and the same time. In such cases, the decision regarding
which specialization is considered most advanced may be entirely arbitrary.
Among birds, for instance, we find four particularly conspicuous specializa¬
tions of the wing:

1. Increased flight efficiency: (a) primaries reach functional peak (swifts,

hummingbirds), (b) secondaries reach functional peak (albatrosses)
2. Loss of flight: (a) wing changed to swimming paddle (penguins), (b)
locomotory function of wing lost in flightless birds (e.g. Ratites).

It is entirely a matter of convention which of these four groups of taxa

is considered “high” or “low” in the system. Most of the so-called primitive
animals, for instance the monotremes among the mammals, are highly spe¬
cialized in certain aspects. Consideration of the totality of characters usually
permits a decision as to the most logical or convenient sequence, but in
highly uniform groups such as birds no clearly superior sequence is obvious.
The taxon that is highest in wing development may be lowest in the develop¬
ment of the central nervous system, and vice versa.
10.7.2 Graphic Representations. Species living at the present time
are the current end points of innumerable phyletic lines. A linear listing
of species, genera, and families cannot even begin to convey an impression
of the various lines of descent in a complex phylogeny. Nor can it show
to what extent the grouping of species is the result of divergent, convergent,
and parallel evolution. It is this deficiency of the printed sequence which
has induced taxonomists to attempt to express their conclusions on relation¬
ship in the form of diagrams.
Up to the eighteenth century naturalists and philosophers arranged
the diversity of nature in the form of a single scala naturae ascending
from inanimate matter in an undivided continuous line up to man. The
continued emphasis on “lower” and “higher” in our classifications is a rem¬
nant of this kind of thinking. As more and more types of organisms became
known, the linearity became increasingly suspect. Pallas, Buffon, and
Lamarck (Voss, 1952) suggested ramifications or anastomoses of the scala
naturae, to satisfy the principle of plenitude. Cuvier’s recognition of four
major branches of the animal kingdom, each totally independent of the
others, completed its demise. Darwin (1859, opposite p. 117) proposed
a purely theoretical diagram of genealogy, but it was Haeckel (1866) who
made the first successful attempt at presenting the relationships of all ani¬
mals phylogenetically (Fig. 4-3). Since that time it has been traditional
among taxonomists to present their concepts of relationships in the form
of phylogenetic trees or similar diagrams (Jepsen, 1944). In spite of their
252 Methods of Zoological Classification

Ser ieos|om>tid< *5 RhyKophilidje

?■= Iff?; -
3rs. Hydropsy- £ I ^ ° -| -I
* ~ chid* S' -g E a-
S -S -Z——. I « -i ° -S
¥ -sor o £

Present time

Remote past

(Morphological similarity)

Fig. 10-13. A three-dimensional dendrogram representing the evolution of caddisworm case con¬
struction (Milne and Milne, 1939).

obvious shortcomings such diagrams are useful summaries of taxonomic

knowledge and permit an easily accessible presentation of the author’s ideas
on the evolutionary history of a group. Such diagrams have great heuristic
value because they tempt later students to test them on the basis of new
characters and subsequently discovered taxa.
Haeckel’s original phylogenetic trees were indeed drawn as trees.
Many subsequent constructions have attempted to convey the impression
of a three-dimensional structure (see Fig. 10-13). In most cases so-called
 The Procedure of Classifying 253

phylogenetic trees are far more diagrammatic and often concentrate on

two dimensions representing time and difference, with special attention
to the branching points. A diagram of the phylogeny of the Equidae (Fig.
10-14) is a typical example. In this case the diagram is to a large extent
based on the rich fossil record of this family of ungulates. Even in this

Fig. 10-14. Phylogeny of the Equidae (after Stirton).

254 Methods of Zoological Classification

arctica

Fig. 10-15. Star-type phenogram of six species of ground beetles of the

genus Miscodera. Figures indicate number of characters (in a total of
20) shared by each pair of species (from Lindroth, 1961).

case the interpretation is largely a matter of inference. It is true that the

fossil record gives substance to phylogenetic trees, but the gaps in the record
are still sufficiently large even in the best-known groups to require much
conjecture.
Diagrams of Relationship. Numerous methods of representing graphi¬
cally the similarity or relationship of taxa have been proposed by taxonomists
over the years. They differ by the kind and amount of information which
they convey and whether or not they include inferred phylogeny. There
are various criteria on the basis of which these diagrams can be classified.
The theory of classification underlying the various schemes is perhaps best
brought out in the following arrangement:
Phenograms. Any diagram which expresses “affinity” as indicated by
degree of similarity but does not aim to convey any other information
is a phenogram. Serologists have long used such phenograms to indicate
relationships as indicated by cross reactivity. Plant geneticists have used
them to indicate degrees of interfertility between species, taxonomists to
express morphological similarities (Lindroth, 1961, Fig. 10-15). This method
has two virtues—it gives the degree of similarity between each species and
all the other species considered, and it does not make any inferences as
to probable lines of descent.
In contrast to these topologically uncommitted phenograms is the
phenogram adopted by the pheneticists. It has the form of a phyletic den-
 The Procedure of Classifying 255

drogram, with a regular branching of lines just as in a phylogenetic tree

(Fig. 10-16). As shown by Boyce (1964) and MinkofF (1965), the branch¬
ing points are very sensitive to changes in the methods of computation,
so that each method of computation may result in a different phenogram.
Rapidly diverging lines have to be inserted below the branching points
of slowly evolving lines even though the phyletic branching took place
later in time. For all these reasons the adoption of the dendrogram pattern
for phenograms would seem inefficient and misleading (Mayr, 19656).
Cladogram. In a cladogram (Fig. 10-17) the ordinate gives the esti¬
mated time, while the abscissa gives degree of difference. Degree of relation¬
ship is determined by the position of the branching points. B, for instance,
is said to be more closely related to C and D than to A, because B, C,
and D share branching point To while A branched off earlier (at 7\).
The different amounts of evolution between 7\ and To as against T2
and T3 are ignored (see cladism, Chap. 4). Since the fossil record is rarely
sufficiently complete to indicate the actual position of the branching points
in space and time, they are reconstructed through a careful evaluation
of the similarities and differences of taxa (Hennig, 1960).
Phylogram. In an orthodox phyletic dendrogram (Fig. 10-18) three
kinds of information are conveyed, degree of difference in the abscissa,
geological time in the ordinate, and degree of divergence by the angle

100 A B C

90

80

70

60

50

40

30

Fig. 10-16. A phenogram. The figures on the ordinate indicate degrees of dif¬
ference (or similarity) between the taxa listed (at arbitrary distances) on the
abscissa. The horizontal connecting lines of the taxa do not indicate phylogenetic
branching but merely levels of phenetic difference.
256 Methods of Zoological Classification

A B CO

Fie. 10-17. A dadogram. The branching points at Tv 7.. and Tt

determine the classification of the taxa A, B, C, and D.

of divergence. This diagram presents a maximum of information, but two

of the three sets of information (branching time and rate of divergence)
are inferred. A1 though this diagram shares with the phenogram the emphasis
on degree of observed similarity, it can cope better with problems of con¬
vergence and parallelism and particularly with problems produced by differ¬
ent rates of evolution.
The phylogeny represented in a phylogram is only inferred and is
unlikely to be an exact representation of the real phylogeny. Likewise,
mosaic evolution, that is, different rates of evolution for different compo-

* b c D E F

<->
decree of difference

Fig. 1U-18. The phyletic dendrogram (phylogram) of the evolutionary

taxonomist. The slower a line evolves, the more it approaches the vertical;
the more rapidly it evolves (as Tj to F), the more it approaches the
horizontal.
 The Procedure of Classifying 257

nents of the phenotype, makes it virtually impossible to represent phylogeny

adequately in any diagram.
Curiously, such diagrams are particularly helpful where the classifica¬
tion must deviate from the ideal of a purely cladistic classification. It aids
understanding in cases where parallel evolution requires horizontal classifica¬
tion (Chap. 4) or, as in the birds-crocodiles case, where a particularly rapid
evolution of a single line has resulted in the evolution of a higher taxon
with all other branches remaining at a lower categorical level. Finally,
such multidimensional diagrams are a useful supplement to the linear se¬
quence of the printed classification.
Chapter Taxonomic Publication

N. ro taxonomic study is complete until its results have

been published. The advances in any area of science are made known
through publication, an inseparable part of research. This is particularly true
for a vast field like systematic zoology where the number of workers is so
small. Applied biology is to a large extent dependent on the advances made
in the correct classification of organisms.
The Strategy of Publishing. A research worker in any branch of science
must ask himself at regular intervals whether or not he is pursuing the
most productive line of research (1.4.4). Should he shift his emphasis, should
he switch to a different group of organisms? What sort of research is most
needed in the particular group of organisms for which he is a specialist?
Should he concentrate on alpha, beta, or gamma taxonomy? The answer
in most cases is that a judicious balance of various approaches is the most
rewarding strategy.
A major taxonomic revision is very time-consuming and may require
years of work prior to publication. The question inevitably arises whether
there should be preliminary papers and whether the specialist should pre¬
pare at the same time a series of shorter papers, perhaps dealing with
different taxa, in order to maintain a publication schedule. Obviously there
is no excuse for duplication in publication for the sake of increasing the
number of titles in one’s bibliography. On the other hand the field work

258
 Taxonomic Publication 259

and research required for the preparation of a comprehensive revision may

result in biological, evolutionary, and biogeographical findings which are
best published separately.
Some taxonomists find it difficult to complete their manuscripts. They
should realize that there is a law of diminishing returns and that no research
is ever “complete.” There will always be areas that have not yet been
collected and collections in remote museums that have not yet been studied.
The same is true in experimental research, where results are published even
though more and perhaps slightly different experiments might well have
been conducted. The number of specialists is very limited, and they have
a real obligation to publish their results as soon as a reasonably reliable
summary of their researches is possible.
Documentation and Information Retrieval. The names of 8,000 con¬
temporary animal taxonomists are included in a recent listing (Blackwelder
and Blackwelder, 1961), but many are inactive. Every year many new
authors start publishing. To keep track of their output has become a for¬
midable task. Sooner or later some system for automatic information re¬
trieval will be introduced. The animal taxonomist is fortunate in having
a relatively complete listing of all new names and taxonomic revisions in
the Zoological Record.
Every effort should be made by the taxonomist to facilitate the task
of information retrieval and to avoid anything that might increase its diffi¬
culty. The latter includes name changing in violation of the principles
laid down in the Preamble of the Code; and unnecessary splitting or lump¬
ing on the generic level; any change in a widely adopted taxonomic se¬
quence without compelling reason; and, finally, the proposal of new species
not connected with complete revisions.

11 A. KINDS OF PUBLICATIONS

Taxonomic publications range all the way from the short description
of a new taxon, covering only part of a page, to lengthy monographs and/or
handbooks that may consist of several volumes. This includes works pri¬
marily useful for the purpose of identification as well as revisionary works
and new classifications. In some works the nomenclatural aspects are
stressed, in others biology, distribution, or illustrative material. There is
a time in the history of the study of each group of organisms in which
one or the other kind of publication is most useful. A specialist must fully
understand the function of each kind of publication and must select one
of them as the form of his own next publication in the field. The following
comments on these different types of taxonomic publication may be helpful
to the beginner.
260 Methods of Zoological Classification

Many different titles are used for more comprehensive taxonomic

publications. Unfortunately, the meaning of such titles is not always the
same for different groups of organisms. A checklist in ornithology is a
very careful, critical revision, usually with extensive synonymy and a detailed
elaboration of the geographic distribution of each species and subspecies.
A checklist in entomology is generally only a list of names, often in alpha¬
betical sequence. Words like handbook and review likewise have different
meanings in different fields. The subsequent discussion must be regarded
in the light of this heterogeneity of usage. Many published works combine
features of several of the kinds of publications described below.
11A.1 Description of New Taxa. The isolated description of new
subspecies, species, and genera, divorced from revisional or monographic
work, is the least desirable form of taxonomic contribution, except in well-
known groups. The preparation of such a description does not permit as
careful a comparison of all related species as does a revision; in many
cases it results merely in the addition of still another name to the already
long list of nominal species. The specialist who eventually prepares the
revision must search out the type, which is often quite inaccessible, and
lose much valuable time in getting it properly identified. In many (if not
in most) cases the description of isolated new species in poorly known
groups of animals is a handicap rather than a help to subsequent workers.
Such isolated descriptions are justified only when names are needed in
connection with biological or economic work or when a group has been re¬
cently revised and the new species can be readily fitted into the classification.
Theoretically, in order to provide an adequate description of even
a single species an author must undertake much of the work basic to the
preparation of a revision, and he could carry his work to that point with
a little extra effort. All too frequently, however, the isolated description
results from a rather superficial acquaintance with previous work, and a
far greater percentage of synonyms is created through isolated descriptions
than in more substantial revisions.
11A.2 Synopses and Reviews. Synopses and reviews are brief sum¬
maries of current knowledge of a group, and the inclusion of new material
or new interpretations is not necessarily implied. They serve the utilitarian
purpose of bringing scattered information on a group together in one place,
perhaps as a basis for some future revisional or monographic study. Exam¬
ples of a synopsis and a review are as follows :
La Rivers, Ira. 1947. A synopsis of the genus Endrodes (Coleoptera: Tene-
brionidae). Ann. Ent. Soc. Amer., 40:318-328.
Ross, H. H. 1946. A review of the Nearctic Lepidostomatidae (Trichoptera).
Ann. Ent. Soc. Amer., 39:265-291, 37 figs.

I1A.3 Revisions. Revisions are presentations of new material or new

interpretations that have been integrated with previous knowledge through
summary and reevaluation. They vary greatly in completeness of treatment.
 Taxonomic Publication 261

Some revisions are monographic in approach yet fall short of being mono¬
graphs because of inadequate material. Others are limited to a new arrange¬
ment of a group. Most of the important current taxonomic contributions
in groups where new species are still constantly being discovered fall in
this category. Such revisions may deal with a whole family (or part of one),
with a genus, or with a species group. Generic revisions, illustrated by
the following example, are the most common type of such work.
Sommermann, K. M. 1946. A revision of the genus Lachesilla north of Mexico
(Corrodentia: Caeciliidae). Ann. Ent. Soc. Amer., 39:627-657, 4 plates.

11A.4 Monographs. Monographs are complete systematic publica¬

tions. They involve full systematic treatment of all species, subspecies, and
other taxonomic units and a thorough knowledge on the author’s part
of the comparative anatomy of the group, the biology of the species and
subspecies included, the immature stages in groups exhibiting metamorpho¬
sis, and detailed distributional data. For the student of evolution such mono¬
graphic treatises are the most rewarding type of taxonomic publication.
They permit a detailed treatment of geographic variation, of relationships,
and of distributional history. Generalizations on the structure of species,
modes of speciation, nature of taxonomic categories, and the like are based
on such monographs. They have the disadvantage that they require more
complete material than other kinds of taxonomic papers. However, with
the growth of collections in the museums of the world, it is becoming
increasingly feasible to prepare monographs. Some papers which fully qual¬
ify as monographs are published as revisions or under some other title.
Unfortunately, in the present state of our knowledge of many groups, espe¬
cially among the invertebrates, few taxonomic papers can justify the title
monograph. Monographs are more frequently possible among vertebrates.
Three fairly typical examples of monographs are:
Hubbell, T. H. 1936. A monographic revision of the genus Ceuthophilus (Orthop-
tera, Gryllacrididae, Rhaphidophorinae). Fla. Univ. Pub., Biol. Sci. Ser., vol. II,
no. 1, 551 pp., 38 plates.
Linsley, E. G. 1961ff. The Cerambycidae of North America, Univ. Calif. Publ. Ent.,

vol. 18 (97 pp., 35 pi.), vol. 19 (102 pp., 1 pi.), to be continued.

Usinger, R. L. 1966. Monograph of Cimicidae (Hemiptera-Heteroptera). Thomas

Say Foundation Publications, vol. 7, xi + 585 pp.

11A.5 Atlases. In recent times the need has been felt for complete
illustrations of the species of various taxonomic groups. This is a reflection
of the inadequacy of the printed word as a means of conveying a mental
picture of the general facies of an animal. The idea of an atlas grew
also out of the need for taxonomic data which are strictly comparable
from one species to another. Since the purpose of an atlas is purely tax¬
onomic, semidiagrammatic drawings are commonly used, though full half¬
tones or colored plates have been employed when dealing with such groups
as butterflies and birds.
262 Methods of Zoological Classification

Examples of this type of treatment are as follows:

Ferris, G. F. 1937-1950. Atlas of the scale insects of North America. Stanford Uni¬
versity Press, Stanford University, Calif., 5 vols.
Ross, E. S., and H. R. Roberts. 1943. Mosquito atlas. American Entomological
Society, Philadelphia; part 1, 44 pp., part 2, 44 pp.

11 A.6 Faunal Works. In faunal works the scope is dictated by the

limits of a geographic region rather than by the limits of a taxon. There
are different kinds of faunal works of varying usefulness. On the whole,
the broader the area covered the more useful is the faunal work.
Examples are:

Fauna of British India. Taylor and Francis, London. Many volumes covering most
groups of animals, published from 1888 on.
Biologia Centrali-Americana, 1879-1915, parts 1-215. Dulau and Co., London.
Faune de France. 1921-1967 et seq., vols. 1-68. Office Centrale de Faunistique
Paris.
Fauna SSSR. Zoologicheskii Institut Akademii Nauk SSSR (some 90 vols. published).

An example of a local list is:

Brown, H. E. 1939. An annotated list of the species of Jassinae known to occur

in Indiana (Homoptera, Cicadellidae). Amer. Midland Nat., 21:663-673.

Reports of expeditions and voyages also belong to this category. These

were useful publications in the early history of taxonomy, but such reports
on individual collections are no longer looked at with favor. They never
permit the careful revisionary work necessary for the exact placement of
specimens, and they often lead to the proposal of new names later found
to be synonyms. Faunistic reports on the whole are among the most inefficient
kinds of taxonomic papers. The time required to run down all the locality
records in a local fauna can be spent far more productively in preparing
a revision or a manual.
A separate class of publications consists of identification works devoted
to a local fauna (see below). Local faunas, in the case of well-known
organisms, give the local naturalist an opportunity to provide ecological
and behavioral information, often based on many years of field work. Such
publications are possible only when the knowledge of the fauna has passed
the level of alpha taxonomy.
11A.7 Field Guides and Manuals. Certain works, although taxo¬
nomic, are designed primarily or exclusively for field identification. In such
cases new species are expressly excluded, and emphasis is placed on clear-cut
key characters or recognition characters. Examples of this are cited in Chap.
6.
11A.8 Flandbooks. The term handbook is sometimes used for field
identification guides but nowadays more commonly for comprehensive works
on a group of organisms with particular emphasis on their biology and
 Taxonomic Publication 263

distribution. An example is:

Palmer, R. S. (ed.). 1962. Handbook of North American birds, vol. 1. Yale Uni¬
versity Press, New Haven, 567 pp.

More broadly the term handbook (traite) is used for surveys of large areas of
zoology, like Kukenthal’s Handbuch der Zoologie or Grasse’s Traite de Zoologie.
The word treatise is also sometimes used for such handbooks as Moore, R. C.
(ed.). 1953. Treatise on invertebrate paleontology. Geological Society of America
and University of Kansas Press.

11A.9 Catalogs and Checklists. A catalog is essentially an index

to published taxa arranged in such a manner as to provide a complete
series of references for both zoological and nomenclatural purposes. Species
and genera are often listed in an alphabetical sequence, because most cata¬
logs are nothing but uncritical (prerevisionary) listings of nominal species.
The value of a catalog depends on its degree of completeness, and its
preparation is therefore a highly technical task requiring great patience
and an intimate knowledge of bibliographic resources and methods.
A checklist is an abbreviated synopsis. It provides a skeleton classifica¬
tion of a group and serves as a convenient source of reference for the
naming of specimens and the arrangement of collections. Checklists vary
greatly in their elaborateness. A list of names deserves the name checklist
(in contradistinction to catalog) only if a careful distinction is made therein
between valid names and synonyms. The name of each species is usually
accompanied by a more or less detailed indication of the range of each
species. Checklists are most useful in the better-known groups of animals.
One in which species are critically evaluated and carefully distinguished
from synonyms qualifies as “primary zoological literature” (13.24), whereas
an uncritical listing of names or nominal species does not.

Stone, A. et al. 1965. A catalog of the Diptera of America north of Mexico.

Agricultural Research Service, U.S. Dept, of Agriculture, Washington, D.C.,
1696 pp.
McDunnough, J. 1938-1939. Checklist of the Lepidoptera of Canada and the

United States of America. Part 1, Macrolepidoptera; part 2, Microlepidoptera.

South. Calif. Acad. Sci. Mem., 1, 1-275; 2(1), 1-171.
Mayr, E. 1941. List of New Guinea birds. American Museum of Natural History,
New York, 260 pp.

11 A. 10 Revisions of Higher Taxa. As a consequence of the increas¬

ing interest in macrotaxonomy, numerous works have appeared in recent
years which deal exclusively with genera and still higher taxa. Frequently
they contain an evaluation of the characters which have been used by
the author to infer relationship and phylogeny and to determine ranking.
Such taxonomic works are:

Corliss, T. O. 1961. The ciliated protozoa: Characterization, classification, and

guide to the literature. Pergamon Press, New York.
264 Methods of Zoological Classification

Edmunds, G. F. 1962. The principles applied in determining the hierarchic level

of the higher categories of Ephemeroptera. Syst. Zool., 11:22-31.
Ehrlich, P. R. 1958. The comparative morphology, phylogeny, and higher classifica¬

tion of the butterflies (Lepidoptera: Papilionoidea). Univ. Kansas Sci. Bull.,

39:305-378.
Fell, H. B. 1965.The early evolution of the Echinozoa. Breviora, 219:1-17.
Greenwood, P. H., D. E. Rosen, S. H. Weitzman, and G. S. Myers. 1966.

Phyletic studies of Teleostean fishes with a provisional classification of living forms.

Bull. Amer. Mus. Nat. Hist., 131:341—455.
Honigberg, B. M., et al. 1964. A revised classification of the phylum Protozoa.

J. Protozool., 11:7—20.
Marcus, E. 1958. On the evolution of animal phyla. Quart. Rev. Biol., 33:24-58.
Marx, H., and G. B. Rabb. 1965. Relationships and zoogeography of the viperine

snakes (Family Viperidae). Fieldiana-Zool., 44:161—206.

Mayr, E., and D. Amadon. 1951. A classification of recent birds. Amer. Mus.

Novitates, 1496:1—42.
Michener, C. D. 1952. The Saturniidae (Lepidoptera) of the Western Hemisphere.

Bull. Amer. Mus. Nat. Hist., 98:339-501.

Newell, N. D. 1965. Classification of the Bivalvia. Amer. Mus. Novitates, 2206:

1-25.
Rosen, D. E., and R. M. Bailey. 1963. The Poeciliid fishes (Cyprinodontiformes),
their structure, zoogeography, and systematics. Bull. Amer. Mus. Nat. Hist.,
126:1-176.
Simpson, G. G. 1945. The principles of classification and the classification of
mammals. Bull. Amer. Mus. Nat. Hist., 85:i-xvi, 1-350.
Stunkard, H. W. 1963. Systematics, taxonomy and nomenclature of the Trematoda.

Quart. Rev. Biol., 38:221-233.

11A.11 Evolutionary and Biological Publications. It is inefficient

to include too much heterogeneous information in a single publication.
This is particularly true for taxonomic monographs. Such works are usually
read only by a few specialists; any information of broader biological interest
published there will not come to the attention of the general biologist
and is therefore bound to be ignored. Almost every author of a taxonomic
revision makes interesting ecological, evolutionary, or zoogeographical dis¬
coveries which would be of great interest to other biologists. Instead of
burying these findings in the introduction of a monograph, he should publish
them in a general journal like Ecology, Evolution, or Systematic Zoology,
where, in turn, these articles will draw attention to the monograph. The
fact that taxonomists have increasingly adopted this policy partly explains
the noticeable improvement in the general image of taxonomy in recent
years. No other group of biologists can make such important contributions
to all problems relating to the diversity of the organic world. The taxonomist
has an obligation to make his knowledge broadly available, not to bury
it in a monograph or revision.
In addition to these by-products of monographs and revisions, more
and more taxonomists publish short contributions to evolutionary or be¬
havioral biology or to ecology, these being the result of ad hoc investigations.
 Taxonomic Publication 265

For instance, every volume of Systematic Zoology contains papers dealing

with cytological or chemical attributes of species (and higher taxa), dis¬
tribution patterns, behavior, taxometrics, variability, endemism, extinction,
serology, geographic variation, speciation, rates of evolution, climatic rules,
allometry, etc. Yet this is only one of a large number of journals publishing
the results of evolutionary research conducted by taxonomists. In addition
to journal articles there are such books as:

Huxley, Julian (ed.). 1940. The new systematics. Clarendon Press, Oxford.
Mayr, Ernst.1942. Systematics and the origin of species. Columbia University
Press, New York.
Rensch, B. 1960. Evolution above the species level. Columbia University Press,
New York.

11 A. 12 Theory and Methods of Systematics. Publications devoted

to this area comprise a growing part of the taxonomic literature. They
are cited in all chapters of this book and are listed in the terminal
bibliography.

11B. MAJOR FEATURES OF TAXONOMIC PUBLICATIONS

Most taxonomic publications contain a definite set of major compo¬

nents. These form the basic substance of these publications and therefore
deserve thorough discussion. Many excellent (and a few outworn) traditions
have developed in the past 200 years of taxonomic research. Every taxono¬
mist must know what they are and why he should follow certain standard¬
ized procedures if he is to facilitate quick retrieval of information.

11B.1 DESCRIPTIONS

The chief objective of a description is to aid subsequent recognition

of the taxon involved. It was realized at an early date that different kinds
of descriptions approach this goal in a different manner. Linnaeus distin¬
guished clearly between the general descriptio (character naturalis) on
one hand and the polynominal differentia specifca (character essentialis)
on the other (Svenson, 1945). The latter contained “the essential characters
by which the species is distinguished from its congeners.” It corresponds
to what is nowadays called a diagnosis.
The functions of the two kinds of descriptions, the general description
and the diagnosis, are by no means identical. The diagnosis serves to dis¬
tinguish the species (or whatever taxon is involved) from other known
similar or closely related ones. The general description has a broader func¬
tion. It should present a general picture of the described taxon. It should
266 Methods of Zoological Classification

give information not only on characters that are diagnostic with relation
to previously described species, but also characters that may distinguish
the species from yet unknown species. It should also provide information
that may be of interest to others besides taxonomists.
Linnaeus and many taxonomists since his time have stressed the ex¬
treme practical importance of a short, unambiguous diagnosis. It can only
rarely be combined successfully with the general description. The latter,
in turn, no matter how exhaustive it is, cannot always provide a substitute
for a type specimen (see 13.48) or, in many cases, for illustrations.
There is still considerable confusion in the literature concerning the
meaning and usage of the terms description and diagnosis. In describing
animals the taxonomist should achieve two objectives, that of diagnosis
and that of delimitation. Diagnosis1 is the art and practice of distinguishing
between things; delimitation is the art and practice of setting limits to
things. “Both enter into taxonomy . . . they are essentially different and
their complementary roles should be clearly understood” (Simpson, 1945).
Although the formal diagnosis in taxonomic work sometimes assists in the
delimitation of a taxon, this function is mainly performed by the general
description. The two terms, diagnosis and description, may then be used
as follows:
Description. A more or less complete statement of the characters of
a taxon without special emphasis on those characters that distinguish it
from coordinate units.
Diagnosis. A brief listing of the most important characters or character
combinations that are peculiar to the given taxon and by which it can
be differentiated from other similar or closely related ones.
The direct comparison of a species (or other taxon) with other spe¬
cifically mentioned species (or other taxa) is usually called a differential
diagnosis. Such a comparison with other species is of great practical help
to students who have no material of the newly described form. It also
forces the author of a new form to review all the evidence for and against
the establishment of the new taxon (Rensch, 1934) and thus ensures that
the diagnostic characters of the new form are mentioned. If the nearest
relatives are rare or poorly known, it is also helpful to make a comparison
with a well-known, if not so closely related, species (13.19).
1 IB. 1.1 Original Description. The description published at the time
of proposal of a name for a new species, genus, or other taxon is called
the original description. It has two primary functions. The first, as stated
above, is to facilitate subsequent recognition and identification; the second
is to make the new name available by fulfilling the requirements of Arts.
11-16 of the Code (13.16).

1 Ultimately from the Greek SiayiyyUffKeiVt to distinguish between two (things).

 Taxonomic Publication 267

The importance of preparing a proper description cannot be over¬

emphasized. The describer is forced to rely on words to convey his meaning.
Yet words, no matter how carefully chosen, are rarely adequate to give
an accurate and complete mental picture of the appearance of an organism.
Nevertheless the description should enable a subsequent worker to identify
specimens without reference to the type. In most cases this goal can be
achieved by the careful worker, particularly when the description is properly
coordinated with illustrative material.
A good description requires on the part of its author (1) a thorough
knowledge of the group of organisms concerned, (2) a knowledge of struc¬
ture and terminology, (3) an ability to evaluate differences and similarities,
(4) an ability to select and emphasize the important, (5) a full understand¬
ing of the precise meaning of the words and the correct usage of the
grammar of the language employed, and (6) a concern for the future
worker. Ferris (1928) stated, “If [the describer’s] work of recording the
data has been properly done those data are available for re-examination
and re-evaluation. His conclusions can be checked, they can be extended
or modified or rejected as appears desirable, all without the necessity of
recourse to his types.”
In the less well-known groups much of the taxonomist’s time is spent
in comparing and contrasting one description with another. This task is
difficult under any circumstances, but it is easier when the descriptions
approximate one another in style, arrangement, and form. This does not
mean that a completely standardized description is always possible, because
the factors which influence the order of presentation, form, and style vary
from group to group. Within a particular group, however, much can be
done to standardize descriptions and thus increase their effectiveness and
utility. (See also 1 IB.1.3.)
1 IB. 1.2 Style. The style generally used in descriptions as well as
in diagnoses is telegraphic and concise. It is usually characterized by the
elimination of articles and verbs and by the selection of adjectives and
nouns of explicit meaning. It further involves proper use of capitals and
punctuation and adherence to a logical sequence of presentation. Thus
the telephonic-style statement, “The head is one-third longer than it is
wide, the antennae are shorter than the body, and the outer segments
are serrate” becomes simply

Head one-third longer than wide, antennae shorter than body, outer
segments serrate.

The descriptive style of the second statement has lost none of the
precision or clarity of the first yet is only one-half as long and may be
both read and understood more quickly.
268 Methods of Zoological Classification

1 IB. 1.3 Sequence of Characters. The recommended sequence of

characters is different in a diagnosis and a description. In a diagnosis it
is customary to present characters in the order of their diagnostic impor¬
tance (or what the author regards as the order of importance). This will
facilitate rapid recognition. In the full description the material should be
arranged in a standardized natural order—for instance, describing the body
parts from anterior to the posterior, first on the dorsal and then on the
ventral surface. The details may be varied to fit the groups yet still maintain
a natural and readily comparable order. For instance, the sequence of
presentation for a dorsoventrally flattened animal group would be different
from that for either a laterally compressed or a radially symmetrical group
because of the different methods of orientation during study. The standard¬
ized sequence of characters helps ensure that nothing important is being
overlooked and that the description is comparative. It is very frustrating
to try to use a taxonomic paper in which half a dozen species are described
independently of one another, details being given, for example, of the an¬
tennae of one species, the pronotum of a second, and the elytra of a third.
Such a procedure makes comparison quite impossible. Authoritative mono¬
graphs usually adopt a standardized sequence of characters, and subsequent
describers should follow it as far as possible.
The utility of a description may be increased by the use of devices
enabling the reader to locate quickly the particular characters he is looking
for. One such device is the use of paragraphs to break up the description
according to main body divisions (e.g., in insects: head, thorax, abdomen,
wings, genitalia, etc.). Where paragraphing is undesirable, the same effect
may be gained by italicizing these same key words. If the author has fol¬
lowed a natural sequence of presentation, either method will permit the
reader to orient himself quickly at some particular point in the description
without having to read the w’hole description.
Dictated Descriptions. When preparing revisions, most modern taxono¬
mists dictate the descriptions into an automatic recording device, in order
to speed up the tedious task of describing. A typist can subsequently tran¬
scribe the dictation. When both hands are needed for viewing specimens
through the microscope, dictating equipment with a foot pedal and a desk
microphone is used. The describer has before him a checklist of characters
in standardized sequence which he follows when dictating the descriptions
of each taxon. The time saved by this method is considerable.
1 IB. 1.4 What to Include in a Description. An exhaustive description
of an organism would fill many volumes, as evidenced by works on the
morphology (physical anthropology) and anatomy of the human species.
It is therefore obvious that even the so-called “detailed description” of
a taxonomic species is highly selective and in the nature of an expanded
diagnosis. Furthermore, in these days of steadily increasing printing costs
 Taxonomic Publication 269

it becomes increasingly important not to include triva. How much subject

matter should be included in a description depends on the group concerned
and the state of knowledge of that group. Excessively long descriptions ob¬
scure the essential points; excessively short ones omit pertinent data. While
the diagnosis serves to distinguish a species from other known species,
the description should be detailed enough to anticipate possible differences
from as yet undescribed species. The description should therefore be more
detailed in poorly known groups, because it is impossible to predict which
characters will distinguish a new species from those that are still undis¬
covered. It should pay particular attention to character complexes that
are variable in the given group. On the other hand, subspecies in a well-
known species of birds may differ from one another in so few characters
that an extensive description would be a repetition of the species description.
In such a case the description may not differ at all from a diagnosis—for
instance, “Like subspecies alba but larger, upper parts blackish gray, not
ash gray” (followed by a tabulation of the measurements).
As far as practicable, descriptions should include all characters, both
positive and negative, which are known to be useful or potentially useful
in distinguishing other taxa at the same categorical level. However, charac¬
ters of higher categories should be omitted except where they are anomalous
or where the rank of the taxon is in doubt. For example, the description
of a subspecies of Song Sparrow should not include reference to characters
that are typical for all Song Sparrows (or worse, for all sparrows!). Viola¬
tion of this rule is not only uneconomical but distracts attention from
the essential features of the taxon concerned.
Beyond the above generalizations, there is little to guide the describer
other than his own good judgment. More than almost any other aspect
of taxonomy the description provides a permanent record of the author’s
ability to observe accurately, record precisely, select and interpret intelli¬
gently, and express clearly and concisely the facts which are before him
(see also Cain, 195%).
The description should include a statement of the differences between
the sexes and, if only one sex is available, a frank statement of that fact
(e.g., “female unknown”). Likewise the characters of immaturity should
be discussed, as well as larval stages. Available behavioral, ecological and
other biological data should be presented. In the case of sibling species,
such information is often more important than morphological characters.
Whether or not the description should be based exclusively on the
type is a much-disputed point. Those who favor this method argue that
all too often it has eventually turned out that the original material—and
consequently also the description—was a composite of several species,
which makes it very difficult to disentangle the characters of the various
species. They argue that it is much safer to restrict the description to
270 Methods of Zoological Classification

the type and have it followed by a discussion of the variability of the

rest of the material.
Others believe that such treatment favors the erroneous typological
view that the type has a special significance as far as the characters of
the species are concerned. They prefer the description to be a composite
drawn from a consideration of the entire material (Simpson’s hypodigm)
and propose to mention at the end by what characters (if any) the type
specimen differs from the rest of the material.
Actually both methods agree that (1) the entire variability of the
species material should be described, and (2) it is advisable to mention
the special features of the holotype. Different authors may use different
methods to achieve these objectives.
1 IB. 1.5 Description of Coloration. Differences in coloration are
among the most important diagnostic characters in many groups of animals.
A detailed description of the general pattern of coloration and of the precise
tone of the various colors is therefore essential in many taxonomic groups.
Subspecific differences in birds, mammals, and butterflies are often largely
a matter of coloration. Many attempts have therefore been made to stan¬
dardize color descriptions, since words like rufous or tawny do not neces¬
sarily suggest the same shade of color to every taxonomist. It is for this
reason that color keys are widely used in taxonomy. Those of Ridgway
(1912), Maerz and Paul (1950), and Villalobos-Dominguez and Villalobos
(1947) are specially recommended. When fine shades of color are involved,
a direct comparison with topotypical material is advisable. Measuring de¬
vices (spectrophotometers) give objectivity and standardization in such
comparisons and permit quantification, as needed for statistical evaluation
(Lubnow and Niethammer, 1964; Selander et al., 1965).
11B.1.6 Numerical Data. The recording of a set of precise measure¬
ments is an integral part of a well-rounded description. If the new form
differs from its relatives in its proportions, such proportions should also
be recorded (see Chaps. 7 and 9). Exact data should be given of numeri¬
cally variable features of structure or pattern, such as numbers of spots,
spines, scales, tail feathers, and so forth. The reasons for including such
data are stated in Chap. 9.
1 IB. 1.7 Descriptive Treatment. A full descriptive treatment of a
species may take the following form:

iScientific Name and Its Author

Bibliographic reference to place, date and author of original description
Type (including type locality and repository)
Synonymy (if any) (see below)
Diagnosis and differential diagnosis (brief statement of essential differences
from nearest relatives, see above)
Description
 Taxonomic Publication 271

Measurements and other numerical data

Range (geographical)
Habitat (ecological notes) and horizon (in fossils)
Discussion
List of material examined

Since the type is the name-bearer, it follows immediately after the

name (and that of each synonym) and not after the description.
In case the species is new, the form of the description is as follows:

X-us albus, new species

Type (including statement of type locality and repository)

Diagnosis, etc., as in a redescription

1 IB. 1.8 Illustrations. Illustrations are in most instances vastly su¬

perior to verbal descriptions. Anything that can be made clearly and suffi¬
ciently visible in a picture should be illustrated. The value of illustrations
is recognized in the International Rules, since- a scientific name given to
a published illustration was held to be valid (prior to January 1, 1931)
even when not accompanied by a single word of description [Art. 16a(i),
(vii) ]. Such a naming of illustrations was quite customary in the days
of Linnaeus. In our day, however, sound taxonomists always present a
diagnosis and full description together with the illustrations. See 11B.4
for a discussion of illustrations.
1 IB. 1.9 Redescriptions. The redescription of hitherto poorly de¬
scribed forms is an extremely important element of revisional and other
taxonomic w'ork. In the present state of our knowledge of many animal
groups, it is of greater importance than the description of new forms. Ferris
(1928), in commenting on this phase of systematic entomology, has stated
that:

... a distressingly large percentage of the named species, in almost every

group of the insects, cannot be recognized positively or even at all, on
the basis of the existing literature. It is more important, for the advancement
of our study, to redescribe such forms than it is to describe new species.
The redescription of such forms should be regarded by the student as
an essential part of his work upon any group which he may elect to study.
The fact that a species has been named should make no essential difference
in the wray in which it is treated. . . . The proper aim is not to name
species but to know them. The writer who contributes to the genuine knowl¬
edge of species is accomplishing far more than one who merely names
them. The fact that the author’s name accompanies the names of the new
species which he describes should not be allowed to influence his activities.

With this view the author heartily concurs.

The specimen or specimens on wffiich a redescription or illustration
272 Methods of Zoological Classification

are based should be clearly indicated because, in the event that the species
has been wrongly identified, a new species may be proposed for X-us albus
Jones, not Smith. In such a case the type specimen of the new species
is the specimen (or is selected from the specimens) on which the redescrip¬
tion or illustration was based.
If a good description, correctly and adequately stated, is readily avail¬
able in the literature, it is wasteful to republish copies of it again and
again. A bibliographic reference is sufficient.
11B. 1.10 Description of a Higher Taxon. Much of what has been
said about the description of species taxa is equally true for higher taxa.
However, the description of a new higher taxon traditionally stresses diag¬
nostic features. Citation of the type species (in case of a genus) or of the
type genus (in case of a family) reduces the amount of descriptive material
that needs to be included.
In the case of higher taxa of vertebrates it is good practice to utilize
characters of the skeleton that would be diagnostic in fossils. In mollusks
and those other groups of invertebrates that have a fossil record, the modi¬
fications of the hard parts, used for diagnosis, ar usually the same in Recent
forms and in fossils.
The form of presentation is as follows:

X-us, new genus

Type species
Diagnosis, description, list of included lower taxa, discussion

1 IB. 1.11 Summary'. Recommendations on the preparation of de¬

scriptions may be summarized as follows:

1. The taxonomic characters should be treated in a standardized sequence.

2. The most easily visible characters should be featured.
3. A direct diagnostic comparsion with the nearest relative or relatives should
supplement the description.
4. Since words alone can seldom give an adequate picture of the diagnostic
characters of a taxon, appropriate illustrations should be provided when¬
ever possible.
5. The description should provide quantitative data, supplemented with
information on geographical range, ecology, habits, etc.
6. Species in poorly known genera should be fully described.
7. The formal description should be followed by an informal discussion
of the variable characters.
8. The description should be accompanied by full information on the type
specimen and other material before the author.
9. Characters that are common to all members of the next higher category
should be omitted from the description.
 Taxonomic Publication 273

11B.2 SYNONYMY

Different names given to the same taxon are synonyms (13.23). In

the early taxonomic history of any group of organisms the correct establish¬
ment of synonymies is perhaps the most important task. All other tasks
(such as the elaboration of a classification and the preparation of keys)
depend on the correctness and completeness of the synonymies. A complete
synonymy of every species and genus is therefore a necessity when a higher
taxon is monographed or revised for the first time or because the previous
treatment has become obsolete.
Unfortunately, the exhaustive preparation not only of a complete
synonymy but also of a listing of all references to previous publications
with all possible binominal combinations (in case of generic transfers) has
become the misplaced ideal of scholarship for some taxonomists. The ex¬
pense of printing endlessly repeated names and references (often a separate
line for each reference!) is altogether out of proportion with the benefits
conveyed. Friedmann (1955), illustrated in Part II of Ridgway’s Birds of
North America, what this system leads to in a taxonomic group with a
rich literature. Other examples are provided by monographs on mammals
and fishes.
It has now become customary in the better-known groups of animals
to list in synonymies only names that were not at all or not correctly listed
in the previous standard treatments. For example, in Peters’ Checklist of
the Birds of the World (1931 et seq.) synonyms are not listed that can
be found in previous standard works, such as Catalogue of Birds of the
British Museum (1873-1892), the Handlist of Birds (1896-1910), for
American species Hellmayr’s Catalogue (1918—1944), and for Palearctic
species Vaurie (1959-1965). More recent checklists do not repeat synonyms
correctly cited in Peters. Only genuine synonyms are listed, not mere new
combinations. There is perhaps no other section of a taxonomic paper
in which more economies can be achieved than in the so-called synonymies.
There are many groups of animals, particularly insects, in which only
alphabetical lists of nominal species exist at the present time. Obviously
the first revision in such a group must give a complete synonymy. However,
it is not necessary even in poorly known groups to repeat again the synonyms
correctly placed in the last previous standard treatment, even if it is other¬
wise out of date. Since manuscript names (nomina nuda) have no existence
in nomenclature, their listing in synonymy is only confusing and should
be avoided (13.16).
New synonymy is most usefully cited with the following sequence
of data: (1) scientific name (in its original form), (2) author, (3) date
274 Methods of Zoological Classification

of publication, (4) reference, (5) type-locality, (6) present location of

type (optional). For example,

Oncideres rhodostictus Bates

Oncideres rhodosticta Bates, 1885, Biol. Cent.-Amer., Coleopt., 5:367. [Lerdo, Mex.;
British Mus. (Nat. Hist.)]
Oncideres trinodatus Casey, 1913, Mem. Coleopt., 4:352. [El Paso, Tex.; U.S.
Natl. Mus.] New Synonymy.2

The above form is recommended for a revision of a well-cataloged

group. In poorly known groups, as described above, a full synonymy (i.e.,
a list of scientific names, incorrect and correct) may be required. This
should include all references which have nomenclatural or zoological signifi¬
cance, arranged chronologically under the actual name (correct or incor¬
rect) by which the author actually referred to them.
Many authors use the convenient device of a comma inserted between
the specific name and the author [.X-us albus, Smith (not Brown)] to
distinguish between a misidentification, which has no nomenclatural status,
and a homonym [X-us albus Jones (not Brown)], which has. A full bib¬
liographical synonymy, as used in a first revision of a poorly known genus,
would appear as follows:

Oncideres rhodostictus Bates

Oncideres rhodosticta Bates, 1885, Biol. Cent.-Amer., Coleopt., 5:367 [type: Lerdo,
Mex.; British Mus. (Nat. Hist.)]; Linsley, 1940, /. Econ. Entomol., 33:562
synon., distr.); Linsley, 1942, Proc. Calif. Acad. Sci., (4) 24:76 (distr.); Dillon
and Dillon, 1945, Sci. Pub. Reading Mus., no. 5:xv (key); Dillon and Dillon, 1946,
loc. cit., 6:313, 382 (revis.).
Oncideres putator, Horn (not Thomson, 1868), 1885, Trans. Amer. Ent. Soc.,
12:195 (key, distr.); Schaeffer, 1906, Can. Ent., 38:19 (key).
Oncideres cingulatus, Hamilton (in part) (not Say, 1826), 1896, Trans. Amer. Ent.
Soc., 23:141 (distr.).
Oncideres trinodatus Casey, 1913, Mem. Coleopt., 4:352 [type: El Pasco, Tex.; U.S.
Natl. Mus.].
Oncideres sp., Craighead, 1923, Can. Dept. Agr. Bull. 17 (n.s.), p. 132 (larva, hosts).
Oncideres pustulatus, Essig (not Le Conte, 1854), 1926, Insects of Western North
America, p. 460, fig. 368 (habits, distr.).

Note that type-locality and location of type are recorded for all genuine
synonyms.
The above synonymy might appear in an abbreviated checklist as
follows:
Oncideres Serville, 1835

1. rhodostictus Bates, 1885 So. Calif, to Tex.

trinodatus Casey, 1913 No. Mex.
L. Calif.

2 This synonymy was published as new in /. Econ. Entomol33:562, 1940. Its use as an example
here and elsewhere in the present discussion is not to be interpreted as a nomenclatural change.
The words NEW SYNONYMY are usually printed in caps or small caps to draw attention.
 Taxonomic Publication 275

When a checklist contains a terminal bibliography, its usefulness may

be increased by giving page references which may then be located by author,
date, and page in the bibliography—thus, rhodostictus Bates, 1885:367.
If it is desirable to indicate the various combinations under which
each name has appeared, this may be accomplished by taking the oldest
specific name and following it through its various combinations, then the
next oldest, etc., as follows:

Megacyllene antennata (White)

Clytus antennatus White, 1855, Cat. Coleopt. Brit. Mus., 8:252 [type: “W coast
of America”; British Mus. (Nat. Hist.)].
Cyllene antennatus, Horn, 1880, Trans. Amer. Ent. Soc., 8:135 (descr., syn., distr.) ;
Craighead, 1923, Can. Dept. Agr., Bull. 27, p. 33 (larva, biol.); Hopping, 1937,
Ann. Ent. Soc Amer., 30:411, pi. 1 (revis.).
Megacyllene antennata, Casey, 1912, Mem. Coleopt., 3:348, 351 (descr.).
Arhopalus eurystethus LeConte, 1858, Proc. Acad. Nat. Sci. Phila., 1858:82 [type:
Sonora, Mex.; Mus. Comp. Zool., Harvard]; LeConte, 1859, in Thomson, Arcana
Naturae, p. 127, pi. 13, fig. 9.

In the above example, the comma between the specific combination

and the author’s name has again been used, this time to distinguish between
a new combination (Cyllene antennatus, Horn, 1880) and an original com¬
bination (Clytus antennatus White, 1855).
Generic synonymy is handled in much the same way as a specific
synonymy, except that in the case of new synonymy or full bibliographic
treatment, the generic type (and its designator, if any) is cited in place
of the type locality and type location. The synonymy of the genus Dicrurus,
as cited in Vaurie’s (1949) revision of the Dicruridae, may be listed as
an example.

Dicrurus Vieillot, April 14, 1816, Analyse d’une nouvelle ornithologie elementaire,
p. 41. Type, by subsequent designation, Corvus balicassius Linnaeus (G. R. Gray,
1841, A list of the genera of birds, ed. 2, p. 47).
Edolius Cuvier, Dec. 7, 1816, Le regne animal, vol. 1, p. 350. Type, by subsequent
designation, Lanius forficatus Linnaeus (G. R. Gray, 1855, Catalogue of the
genera and subgenera of birds, p. 58).
Drongo Tickell, 1833, Jour. Asiatic Soc. Bengal, vol. 2, p. 573. Type, by monotypy,
Drongo caerulescens Tickell = Lanius caerulescens Linnaeus.
Chibia Hodgson, 1836, India Rev., vol. 1, p. 324. Type, by subsequent designation,
Edolius barbatus J. E. Gray = Corvus hottentottus Linnaeus (G. R. Gray, 1841, A
list of the genera of birds, ed. 2, p. 47).
Bhringa Hodgson, 1836, India Rev., vol. 1, p. 325. Type, by original designation
and monotypy, Bhringa tectirostris Hodgson.
Bhuchanga Hodgson, 1836, India Rev., vol. 1, p. 326. Type, by subsequent designa¬
tion, Bhuchanga albirictus Hodgson (Sharpe, 1877, Catalogue of birds in the
British Museum, vol. 3, p. 245).
Chaptia Hodgson, 1836, India Rev. vol. 1, p. 326. Type, by monotypy, Chaptia
muscipetoides Hodgson = Dicrurus aeneus Vieillot.
Dissemurus Gloger, 1841, Gemeinniitziges Hand- und Hilfsbuch der Naturgeschichte,
p. 347. Type, by monotypy, Cuculus paradiseus Linnaeus.
276 Methods of Zoological Classification

Musicus Reichenhach, 1850, Avium systema naturalc, pi. 88, fig. 9. Figure of generic
details, no species included, cf. Bonaparte, 1854, Compt. Rend. Acad. Sci. Paris,
vol. 38, p. 450. Type, ljy tautonymy, Dicrurus rnusicus Vieillot = Corvus adsimilis
Bechstein.
Dicranostreptus Rcichenbach, 1850, Avium systema naturale, pi. 88, fig. 12. Figure
of generic details, no species included. Type, by subsequent designation, Edolius
megarhynchus Quoy and Gaimard (G. R. Gray, 1855, Catalogtie of the genera
and subgenera of birds, p. 58).

11B.3 KEYS

The purpose of a key is to facilitate identification of a specimen

(4.3.1 and 7.3). This goal is achieved by presenting appropriate diagnostic
characters in a series of alternative choices. The worker finds the correct
name of his specimen by making the appropriate choice in a series of
consecutive steps.
The procedure involved is somewhat analogous to that of the physician
who, by means of a series of questions and examinations, arrives, by a
process of elimination and confirmation, at the diagnosis of the ills of a
patient, or to the elimination method in culture identification of bacteria.
The use of keys in identification is old indeed. Much of Aristotle’s
classification of animals was presented in the form of simple dichotomous
alternatives (“bloodless versus with blood,” etc.). Voss (1952) gives an
interesting history of the development of keys in systematic biology. Metcalf
(1954) provides some hints on constructing keys. In the Aristotelian proce¬
dure keys were an instrument of classifying logic. Typologically an object
was either A or not-^4. To use a key typologically for classifying purposes
is misleading in view of the polythetic nature of many taxa. On the other
hand, if a key is used as a purely pragmatic device, the taxonomist can
cope with the existence of a polythetic taxon quite easily, by keying it
out repeatedly. This would be fatal for a classification but does not consti¬
tute weakness in an identification procedure.
Keys are also a tool for taxonomic analysis since in their preparation
one must select, evaluate, and arrange taxonomic characters. In this sense
keys are an integral part of taxonomic procedure, as well as a means of
presenting findings.
The construction of keys is a laborious and time-consuming task, in¬
volving the selection and sifting of the most useful and most clearly diagnos¬
tic characters. Ideal key characters apply equally to all individuals of the
population (regardless of age and sex) ; are absolute (two scutellar bristles
versus one scutellar bristle) ; are external, so that they can be observed
directly and without special equipment; and are relatively constant (with¬
out excessive individual variation). Unsuitable key characters include those
that require a knowledge of all ages and stages of a species (c.g., “sexual
dimorphism present” versus “sexual dimorphism absent”; “male larger than
 . j-ivmnm v Poiikacss

trzzAje~ "versTs ''rzAte aa*Tw thAr. V~ Hal aji; .-vcc r-zz ettaas
Ta_. =>“>-: PAtta! zr-Az-'r : • v -* • tv._ a>c • - a. -x v.
. ...

«*» 1 v T •: •' .- -- . • _ -' tz

Iz za^c rases the ziti perarct the ficoof zc se-secrt! tfiyaatB
*ac tre x^ncus nr—azt. azc sxnxiz zhrcsszcis zc the tr I: > free zz±:
tze wnser s oJk: «PCC XZ • > z*
tne rxsc SMK7KMT zhArazaers At the tr-vels. If he ts xzrz bt«>«a
Z • ' • c - • : zazat a~ . "o • - - • .-r;- • •.• .- -. -.- ~ x

or orosr of sehseczjernt trrarzjez: '<*1 tAkr zc re z>ce- Hz w

*hez rzzvzf a tec ia a pore in: crz>_p z' rz*r- zrvhss.—tci
speoes ;: is use- tz ArrAzpe the ce* in sac* a zaaansrr tzar zOzse* rr-Atrc
speoes tec ozt near zoe Arcchrr 7ht? tArTzAzes the sdhsecKJfcac trserzoz
of specses. as well as lie decssoz whether zc tszc a speeszs is rxr»
It is fzrztzAze vbs: the xnacesi&I pesntts the occsawczjzc zc a ke- • Tci
alaws tbe ptnMM «f rMnb% «Am terfeng atiA Ac cm
ftmzrszct. thai cc ezszzziz: :orr.zhzA:jzc. \r> zzcacracrjzc. > a zczor\£-_zr
::: --.-r_.cz zzrrrzz: . Ate rc a r. az r T.\:s 5 z.zr z.: zzr z\a~
possfw ttses zc the zzcrpttter tz. ttoarcsoMz Rztrt aL Ii*hf
A ?xc kr>. s senzz'- fbzhzczcDza&. ro: zeeznz rrvee zzaz r*c Alrr—A-
rves az arc pxrzs Akouhns shouc be ppecse. Iftaz the scAtrcfertts
sz. .■: --t z; - “ zr~ - re tz -— : fez: hzAtszc : r. ' t. - •.-. • : •
•>»tthzftt zererezee tz zcher srteejes Iz *r« raw. .oez.zzcAfzc. AraM hr
prss. rue «Aoal zefezrzjze ?z the copastr sex zc rz .rzzzAr_re setfes. Thrst
shzctl'f he tzeAtez z ftzezez: kev^ -hez: asasKcpbszt s rv' Zirez Iz zisr
a tec dunoer acadr sepAZAtes the szc-z :> z z -
eaczeoc fzc cos zc >e sens thit rr zirrzieiai* zc vizihe. i< f_rr
Ketrnnur tz aadohe these va-aH sprzss .z hzch st-X.-.-ssrcs. A pet
ioeoes
■ zaztt tzsr
•• Ans irorr
* « rezeAtetl'r
* - aa a ce^
--» The tarcoedstze
• %hozi
—the qcictess azc zdzss: zzAzt'h;z->z<zs oer.zhzAZ<ziz shx. c hr Af.~ccA.t
Oz.-azv Dr« sziez^es shzc_'e zee he hestczAtec as sazh tz a tec. Ahzv
:t s naalh cttscrzctzAz* rz czznt Aztbccttses rtvez iprz.fr zAtzes tz ie>< zz ess
these Ate art rethorz elsewhere tz the att.v>.
The sc* <e of kev? ts *tt»tcntiaf. Tie that « da.T.xkci azc the
phrases Are «saalh se-pAZAtezt b saKokei zverr. thxzzT the pezraev c.'vz-
z-astztc zhAzazzers d cazi cwtpl« zzac he ctAttvsh: Arc oe-zzfcwr. sttA'
asttA donnen Ate oestrAtle tz the oet: tZAt the trztitx zhArazter
ztAv noc he zjeArh ctsoerzjei cc the scevzzsez zzr« hr izz.tei zc rtocztec
iz az ■BMisfKMn zzAzzer Ctee zc the zrosr sArscaztJzcc zaechxh fzc
jsszzt tt cAtA fzc the ezoszrjztx'z zc a kr-i ts sb>z'«z aa az exart.hse

k » A hi ■iTC-'CN i »
V-
278 Methods of Zoological Classification

Table 11-1. Arrangement of Key Characters*

Name of Antennal Tarsal Leg

Wings Antennae Eyes
species color segments color

smithi clear filiform black entire linear black

completa opaque serrate black entire linear black
emarginata opaque serrate black emarginate linear black
rufipes opaque filiform black entire linear red
nigripes opaque filiform black entire linear black
flavicornis clear filiform yellow entire bilobed black
ruficornis clear filiform red entire linear black
calif ornica clear filiform black entire bilobed black

* Characters used in examples are italicized.

of the method and the subsequent analysis given in Table 11-1. This exam¬
ple is oversimplified in order to demonstrate the method more clearly.
Several types of key are used in taxonomic papers, all being dichoto¬
mous and based on a series of choices. By far the most commonly used
is the dichotomous-bracket key. The other is the indented key. The latter
has the advantage that the relationship of the various divisions is apparent
to the eye. It has the disadvantages, especially in a long key, that the
alternatives may be widely separated and that it is wasteful of space. For
these reasons it is generally used only for short keys, keys to higher taxa,
or comparative keys (keys which not only serve the purposes of identification
but also treat the same comparative characters at each level for each
group). An indented key based on the hypothetical data given in Table
11-1 might be as follows:

A. Wings opaque
B. Antennae serrate
C. Eyes entire. complcta
CC. Eyes emarginate. emarginata
BB. Antennae filiform
C. Legs red. rufipes
CC. Legs black. nigripes
AA. Wings clear
B. Tarsal segments linear
C. Antennae black. smithi
CC. Antennae red. ruficornis
BB. Tarsal segments bilobed
C. Antennae black. californica
CC. Antennae yellow. flavicornis
 Taxonomic Publication 279

The second type of key, which is used almost exclusively by most

taxonomists, is the bracket key. This key has the advantage that the
couplets are composed of alternatives which are side by side for ready
comparison, and that it is more economical of space because it is not in¬
dented. When properly constructed it may be run forward or backward
with equal facility by following the numbers, which indicate the path that
the various choices follow. This is the type which best fulfills the diagnostic
purpose of a key. Its main disadvantage is that the relationship of the
divisions is not apparent to the eye. An example based on the same data
previously used is as follows:

1. Wings opaque. 2
Wings clear. 5
2. (1) Antennae serrate. 3
Antennae filiform. 4
3. (2) Eyes entire. completa
Eyes emarginate. emarginata
4. (2) Legs red. rufipes
Legs black. nigripes
5. (1) Tarsal segments linear. 6
Tarsal segments bilobed 7
6. (5) Antennae black. smithi
Antennae red. ruficornis
7. (5) Antennae black. calif arnica
Antennae yellow. flavicornis

Pictorial keys deserve mention among various other kinds of keys

designed for special purposes. The pictorial key is of value for field identi¬
fication by nonscientists. During World War II, for example, malaria crews
based their control operations on the results of field identifications of
anopheline mosquito larvae (Fig. 11-1). The fact that critical characters
were illustrated as well as described made the keys usable by such persons
as medical corpsmen and engineers as well as by entomologists. Other ex¬
amples of pictorial keys are one by Corliss (1959) of the higher taxa of
Ciliates and one of the Rotatorian genus Ptygura by Edmondson (1949).
Pictorial keys have been employed also in field guides to vertebrates and
flowering plants.
Those who employ them must remember at all times the utilitarian
purpose of keys and of the characters used in them. The character on
which a given taxon “keys out” may be of no particular biological or
phylogenetic significance for this taxon. It is simply the character which
gives the best assurance for correct identification. Keys are not phylogenies.
280
PICTORIAL KEY TO ANOPHELINE LARVAE OF UNITED STATES

fresh water brackish water

breeder breeder
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281
282 Methods of Zoological Classification

11 B.4 ILLUSTRATIONS

The old Chinese proverb says: One picture is worth a thousand words.
This is all too true for illustrations in taxonomic papers. It is often quite
impossible to describe adequately in words a complex structure, let us say
the genital armatures of an insect or the palps of a male spider. Descriptions
without illustrations are of rather limited value for many organisms.
The taxonomist must decide in each case what kind of illustration
would be most useful. Except for special purposes an accurate but somewhat
diagrammatical line drawing is usually the most informative illustration
for a taxonomic publication. A scientist is fortunate if he happens to be
endowed with talent as an artist. However, the scientist who lacks it need
not be discouraged, because clear-cut diagrammatic drawings are perfectly
satisfactory and, in some cases, superior to artistic drawings for scientific
purposes. Ferris (1928) called this type of drawing drafting and expressed
the opinion that any conscientious scientist could learn to make satisfactory
drawings of this kind. Several books or manuals have been published on
the subject, among which may be mentioned Kuhl (1949), Staniland
(1953), Papp (1968), and particularly Zweifel (1961).
Pencil sketches should be made with a soft pencil, and bilaterally
symmetrical animals should be “corrected” for symmetry by tracing one
half on the other with thin semitransparent paper.
The original outline may be sketched freehand, but, at least with
microscopically small organisms, it can be done more quickly and accurately
by one or another mechanical means. Perhaps the most popular of these
devices is the camera lucida, which, by means of prisms and a mirror,
projects the microscope image on a piece of paper. With this apparatus
it is possible to look in the microscope and see the specimen superimposed
on the reflection of the paper. By careful adjustment of the light, one can
draw an outline with specimen and pencil point both clearly in view (super¬
imposed on each other). Another method of obtaining the outline is the
direct projection of an image on a screen or paper by a microprojector
attached to the microscope. Still another technique is to photograph the
specimen and print an enlargement on a dull mat paper. The outline
can be inked directly on the photograph, after which the photographic
emulsion can be washed off. Some workers prefer to sketch freehand on
a crosshatched paper, guided by a grid in the ocular of the microscope.
The type of pencils, crayons, pens, and papers suitable for various
drawings are described in the cited literature. Better stationery and artists’
supply stores can give advice on the suitability of various kinds of materials.
Maps, often an important kind of illustration for taxonomic papers,
 Taxonomic Publication 283

are best done by using available outline maps (base maps) on which trans¬
parent paper cutouts are pasted with dots, letters, figures, and crosshatchings
of various kinds and sizes.
11B.4.1 Reproduction. In the preparation of all figures, bear in
mind that their size is usually reduced (sometimes drastically) in the process
of reproduction for publication. An experienced author knows that all sym¬
bols and letters will have to be sufficiently large in the original to remain
easily legible after reduction. Editors and some authors have reducing lenses
which show them how the illustration looks after reduction.
In the layout of the illustration due consideration must be given to
format and size of type bed of the publication (book or journal) for which
the illustration is intended. Proper proportions for the original drawing
may be obtained by expanding on a diagonal line through a rectangle
drawn to page or column size (Fig. 11-2).
Leave room at the bottom of the page for the legend. Figure numbers,
letters, abbreviations, etc. should be put on neatly. In order to be legible,
letters should be 1^, 2 X V« or ^2 *n- high; depending on the amount
of reduction. Freehand lettering is rarely satisfactory. Numbers and letters
may be clipped from old calendars or from standard characters printed
on gummed paper, or they may be made by various mechanical lettering
guides. When determining the amount of reduction the editor is limited
by the size of the page, the need for captions, and other considerations,
and he is often not in a position to follow the instructions of the author.
This is particularly true in the case of large figures. Magnification or reduc¬
tion should therefore not be stated on the figures themselves, but rather
in the captions.
Where many illustrations are used, grouping is often required for
economical reproduction. For zinc etchings, drawings may be assembled
into plates by merely arranging and pasting on a cardboard sheet. Colorless
paste or rubber cement should be used. The paper edges of individual
drawings will not show. For halftones, however, trimmed edges will show,
and when several drawings are to be fitted together for a plate, a mechanical
paper-cutter should be used for trimming. Slight discolorations, especially
when yellowish, also become conspicuous in reproduction. It is usually more
satisfactory to draw numbers and letters directly on the original rather
than paste them on. However, characters printed on transparent gummed
paper are also available for halftone. Photographs should be mounted with
smooth edges touching and symmetrical, so that the engraver can rout
out neat, straight lines.
Curves and graphs are reproduced as zinc etchings and largely used
as text figures. In preparing them for publication, the same instructions
as to size, proportions, and lettering apply as for drawings. However,
284 Methods of Zoological Classification

Fig. 11-2. Method of calculating proportions for enlarge¬

ment or reduction of illustrations. If one side AC of an
illustration is enlarged to AD or reduced to AB, the
corresponding length of the other side (AD' or AB') can
be easily determined as the point of intersection with the
diagonal.

they should be made either on white paper or blue-lined coordinate paper,

never on green-, black-, red-, or yellow-lined paper unless the coordinates
are to be reproduced.
Line drawings are usually reproduced as zinc etchings. The illustration
is photographed on a zinc plate and the background is etched away with
acid. This is the least expensive method of reproducing illustrations and
the most satisfactory for line drawings, graphs, charts, etc.
Where fine detail is needed, also the representation of delicate shades
of light and dark, as in the reproduction of most photographs, so-called
halftones have to be prepared. Here the picture is broken up into thousands
of tiny dots through a process of screening. In photolithography the illustra¬
tion is photographed on gelatin and not screened, thus becoming a fulltone
process. It therefore shows even greater detail than a halftone but lacks
 Taxonomic Publication 285

some of the contrast. Halftones and photogelatin plates are often printed
on glossy paper or at a different printing establishment, and for this reason
it is frequently easiest to assemble them together at the end of an article.
Zinc etchings are usually printed on the same paper as the text and may
therefore more readily be distributed through the article as plates or text
figures. The latter are most economical when printed the same width as
the printed page.
Photography for scientific publication requires special approaches and
setups. Hints on equipment and lighting are given by Blaker (1965).
Colored illustrations are the most complicated and most expensive
to reproduce. A screen similar to halftone screens is used, and several sepa¬
rate colors are used in printing, each one superimposed on the previous
impressions.
Some scientific journals charge the author for copper halftone plates
and for the glossy paper if halftones are to be printed in a journal which
normally employs a rough eggshell-surface paper. This extra cost may in¬
clude not only the paper but the hand labor involved in tipping in or
pasting in the extra pages. When a journal makes page charges on a flat-rate
basis, no special charges are made for illustrations.
Offset printing obviates most of these difficulties because the entire
page, including printed (or typed) matter and line cuts, is photographed
on a plate and then rolled onto a second roller before it is printed on
the final paper. Photographs are made separately because of differences
in contrast and then “stripped in” on the negative of the photolithographed
page. By this method illustrations cost no more than printed matter.
It is a wise precaution to retain good, clear, photographic copies
of all illustrations in the event that they are lost in the mail. A good
photograph of a drawing is only slightly inferior to an original as a basis
for reproduction.

11B.5 REFERENCES AND BIBLIOGRAPHY

References in taxonomic papers are generally treated either in foot¬

notes, or in parentheses in the text, or in a terminal bibliography. Footnotes
are useful when only a few references are involved and when repeated
reference to the same bibliographical item is unnecessary. However, since
they are costly to handle in typesetting and printing and may add materially
to publishing costs, parenthetical references are to be preferred. When refer¬
ences are numerous, they are most frequently handled in a terminal bibliog¬
raphy. In most cases this bibliography should be as short as is consistent
with its purpose, and the items included should be selected. Frequently
the value of the terminal bibliography may be greatly increased by including
286 Methods of Zoological Classification

parenthetical comments on the nature of the subject matter covered. Un¬

verified references may be included when necessary for completeness, but
they should be marked with an asterisk or some other device to indicate
the fact that the author has not seen them.
Bibliographical items should receive full citation, including author,
date, title, publication, volume, pages, etc. The references to the terminal
bibliography may be made by enclosing an author’s name and date (some¬
times also page) in parentheses. Two or more references to publications
by a single author in the same year may be designated by appended letters
(Smith, 1940a; Smith, 19406). The author-date system of bibliographical
reference is far more satisfactory than the straight numbering system which
is sometimes used. The number system tells nothing about the reference;
moreover, the author-date system permits the addition of references during
the preparation of the manuscript without the necessity of renumbering
all references beyond the point of insertion.
The designation “Bibliography” implies completeness of coverage of
the subject. “Literature Cited” indicates restriction of references and is
self-explanatory. For illustrations, see the terminal bibliography of this book.
An example of footnote style1 is given here. In the typescript, foot¬
notes are often entered beneath a marginal line in the text wherever they
occur, rather than at the bottom of the page, because in the final publication
pagination is entirely different from the pagination of the typed pages.
Numerous exceptions will be encountered, especially in various govern¬
ment documents, but a majority of literature citations will fit into one
or other of the simple styles illustrated. It is becoming standard practice
to list year of publication immediately after the author’s name, since this
sequence agrees with that of the author-date system of reference.
If the work cited is by several authors, only the first need be reversed
for alphabetical purposes. Thus,

McAtee, W. L., and J. R. Malloch. 1922. Changes in names of American

Rhynchota, chiefly Emesinae. Proc. Biol. Soc. Wash., 35:95-96.

The original style of capitalization and italics may be followed. How¬

ever, many titles are set entirely in capitals, others entirely in lowercase,
except that the first word and scientific and place names are capitalized.
The full title should be given in all but the briefest of footnotes, because
readers obtain valuable leads in this way. Abbreviations of journals should
follow such standard works as the World List of Scientific Periodicals.
The citation: (Wheeler, 1889a) is specific enough because it refers
to a two-page paper. On the other hand, it may be necessary to refer
to a particular page of a larger work. In this case the page is indicated

1 W. M. Wheeler, 1889. Amer. Nat., 23:644-645 [sample footnote].

 Taxonomic Publication 287

in the citation (Wheeler, 1910, p. 263, or 1910:263), and the complete

work is listed in the bibliography.

11C. FORM AND STYLE OF THE TAXONOMIC ARTICLE

Much scientific literature is of a rather ephemeral nature, except

for a few classical papers. This is not true in taxonomy, where publications
(particularly if they contain new names) are consulted for many genera¬
tions, indeed back to 1758 and earlier. The taxonomist must try to give
his publications a quality which lives up to this timelessness. Innumerable
details require attention, and many of them escape the beginner’s attention.
There are various guides to the preparation of scientific papers (Trelease,
1951; Hurt, 1949), as well as style manuals, such as A Manual of Style
of the University of Chicago Press, the U.S. Government Printing Office
Style Manual, the Style Manual for Biological Journals, published by the
A.I.B.S., and the American Museum Style Manual, the latter with particu¬
lar attention to the practices of the taxonomist. There are also some useful
hints in appendix E of the International Code (Chap. 12). Before sub¬
mitting a manuscript to a journal, study the instructions to authors which
nearly every journal has and the particular style it prefers for citations,
bibliography, etc. This will save a great deal of retyping.
11C.1 Title. The title is the first part of the paper encountered
by the reader, although it is often the last item to be added in the prepara¬
tion of the paper. Its bibliographical prominence and significance warrant
much care in its selection. The title should be long enough to be specific
as to the contents of the paper but brief enough for easy indexing. Short
words are preferable to long ones. The most important nouns should be
near the beginning of each series of words. The title should contain key
words which in indexing will classify the article. This is particularly impor¬
tant today when some abstracting services are based entirely on titles. Punc¬
tuation should be avoided unless essential to meaning. Among the essential
elements of a title are (1) a clear indication of the field involved (taxonomy,
morphology, ecology, etc.), (2) the scientific name of the taxon treated,
(3) indications of the order and family either by means of scientific names
(which may be in parentheses) or, rarely, by a well-known common name,
and (4) the geographical area, fauna, or locality. The following are exam¬
ples of good titles:

A Taxonomic Revision of American leafhoppers (Homoptera, Cicadellidae)

A Checklist of the Birds of Alabama
Geographical Variation of Hippodamia convergens in Southern California
(Coleoptera, Coccinellidae)
288 Methods of Zoological Classification

Two New Species of Wood Rats (Neotoma) from the Rocky Mountain
Region

The following are a few examples of poor titles for taxonomic papers.
On the basis of the above-enumerated principles the objections to these
are obvious.

New Hymenoptera
Notes on Mammals
The Western Biota
A Collecting Trip to Texas
Additions to the Fauna of Nebraska
Studies in the Mollusca
A Newr Acanthiza

Titles need not be as bad as these, however, to cause difficulties for

catalogers, abstractors, reviewers, and other bibliographers. No author has
cause for complaint that his work is being overlooked if it masquerades
under an incomplete, ambiguous, or misleading title.
11C.2 Author’s Name. The author’s name follows the title. Biblio¬
graphical problems are simplified if an author always uses the same form
of his name. The entomologist Laporte sometimes published under the
name Laporte, sometimes as the Comte de Castelnau. The bibliographical
confusion which resulted still persists in modern literature. Women taxono¬
mists who begin publication before marriage frequently avoid confusion
by continuing to publish under their maiden name or by a system of hy¬
phenation, e.g., Dorothy McKey-Fender. It is customary in America to
omit degrees and titles from the author’s name, although these are used
in many European journals. The author’s correct postal address should
follow his name in order to facilitate correspondence. The address can
also be given at the end of the article or in a footnote.
When more than one author is involved, the order of names is deter¬
mined by the nature of the contribution each has made. When the work
has been more or less equally shared, the names are usually arranged in
alphabetical order. When the work has been disproportionately divided
or there is a marked discrepancy in age or experience, the name of the
author to be considered “senior author” appears first. If the same coauthors
publish several papers in a single year, it is advisable to change the sequence
of names in order to facilitate ease of reference.
11C.3 Introduction. Every taxonomic paper should include an intro¬
ductory paragraph stating the scope of the paper and, where pertinent,
the reasons for the study, as well as the nature of the studied material.
Frequently a brief historical review is appropriate. These features serve
to orient the casual reader and the new student of the group, as well
as to refresh the minds of other workers in the field.
 Taxonomic Publication 289

11C.4 Acknowledgments. Acknowledgements may be included in the

introduction when they can be treated as part of the natural sequence
of exposition. Some authors place them in a footnote appended to the
author’s name. This system is regularly used in the Annals of the Entomolog¬
ical Society of America and certain other journals which are primarily
taxonomic in content. Sometimes the acknowledgements are placed at the
end preceding the summary.
Giving proper credit is one of the most important responsibilities of
the scientist. Acknowledgment should be made of all unpublished observa¬
tions, determinations, and data derived from others. If an unpublished
statement is quoted, the author of the statement should be allowed to
prepare it especially for that purpose. Previously published data should
never be utilized in such a way as to appear original.
Acknowledgment should be made of borrowed and donated specimens
studied. The methods of doing so vary with the amount of material received
from any one source and the general plan of presentation of the paper.
A way can be found for such acknowledgement even in complex cases.
Photographs, drawings, and other illustrative material lent or donated
by others should be credited. Credit should also be given to artists and
photographers for their work, whether or not they received payment for
their efforts. Good drawings or good photographs are scientific contributions
on a par with descriptive work and are frequently far more accurate and
useful.
Full credit should be given to the collector, who, after all, is the
real discoverer of the material and not the describer.
Assistance in outlining a research program (including the help of
a major professor or senior colleague) should be acknowledged, as well
as help in the preparation of a manuscript by means of critical reading.
Finally acknowledgment should be made of financial grants or of
institutional aid, such as the use of laboratory facilities, libraries, etc. Fre¬
quently such help is a primary factor in making a particular taxonomic
research project possible. Grant-giving agencies usually stipulate that the
code number of the grant be recorded in all publications resulting from
the grant.
11C.5 Methods Used and Materials Studied. In a revisional or
monographic work it is desirable to include a statement about methods
utilized and collections, specimens, or other materials studied. This enables
the reader to evaluate conclusions and to judge the thoroughness of the
work. Standard methods for measuring, mounting, staining, special prepara¬
tions, etc., may be referred to by name and reference. Only new methods
need to be described in detail.
1IC.6 Body of the Text. The material comprising the body of the
text will, of course, depend on the scope and objectives of the particular
290 Methods of Zoological Classification

paper. It is perhaps sufficient to mention that a complete systematic paper

includes (1) a delimitation of the highest included taxon (family, tribe,
etc.), (2) a key (or keys) to all intermediate taxa treated (genera), (3)
synonymies and descriptions of the taxa of intermediate rank (genera),
(4) statement of the type species, (5) comparison with other genera, (6)
keys to the species of each genus, (7) synonymies and descriptions of each
species, and (8) statements as to type localities and to location of types,
general distribution, hosts and other significant biological data, comparisons
with other species, etc. (for details on preparation of descriptions and keys,
see 11B.3.
IIC. 7 Summary. A summary is usually unnecessary in a strictly
taxonomic paper. If required, it should be brief but not in telegraphic
style. It should be written as a series of short paragraphs and should be
specific, not written in broad general terms.

1 ID. PREPARATION OF THE MANUSCRIPT FOR

PUBLICATION

Aside from matters involved in the actual organization and construc¬

tion of a taxonomic paper, there are some points which should be kept
in mind in order to facilitate editorial handling after the paper has been
submitted for publication. Editors are much more apt to accept readily
(and publish quickly) papers which are in good form and require a mini¬
mum amount of editing. Most publications have special form requirements,
and much editorial time can be saved by careful study of the instructions
given in the journal in which the paper is to be published. The A.I.B.S.
style manual for biological journals is very helpful (see 11C).
IID. I Typing. Manuscripts submitted for publication must be
typed. The original drafts may be on yellow paper, but the final copy
should be on standard (8^ by 11 in. or 8 by 10^ in.) white paper,
entirely double-spaced (some publications require triple spacing), and with
a wide margin for adding proof marks and for editing. If approximately
the same number of lines is typed on each page, the editor can conveniently
estimate the size of the final printed paper. However, some editors require
that pages end with completed paragraphs. Pages should be numbered
consecutively in the upper right-hand corner. Inserted pages are numbered
alphabetically (e.g., 65a). Whole sheets should be used for insertions, re¬
gardless of length of inserted matter. When it becomes necessary to cut
and rearrange, sheets should be assembled by pasting, not by pinning. Some
journals request two copies of each manuscript, the original and one carbon.
All tabular material should be typed on separate sheets, since it is
usually set in a different type from the text.
11D.2 Underlining. Underlining indicates that the material so
marked is to be printed in italics. In a taxonomic manuscript submitted
 Taxonomic Publication 291

for publication, underlining should be limited to scientific names of genera

and species which appear in the text. New names should not be underlined,
because the editor will usually mark these with a wavy line to indicate
boldface. Indications of style or sizes of type for titles, headings, subheadings,
sideheadings, and the like should be left to the editor. In general, marks
which the author makes merely interfere with the editor’s work, though
marginal notes as to the relative rank of headings may be helpful.
11D.3 Legends and Text Citations to Illustrations. Titles and
legends should be self-explanatory. The manuscript of these titles should
be typewritten, double-spaced on separate sheets (several titles on a single
sheet), and assembled in numerical order at the end of the manuscript
following the bibliography. A short identifying title may be placed on each
plate for purposes of identification, but this title will not be printed. Usually
in the process of handling, titles and legends go to the typesetter with
the rest of the manuscript, whereas illustrations are sent to the engraver.
The printer may never see the original drawings.
The place of insertion of the illustrations should be marked in the
manuscript and also in the galley proof. Illustrations are usually numbered
starting with each article, but some journals number plates consecutively
throughout a volume. In any event, a new series of figure numbers or
letters should be used on each plate. Many journals designate figures with
Arabic numbers, plates with Roman numerals. All figures should be referred
to in the text by number.
11D.4 Revision of the Manuscript. A few authors have sufficient
mastery of the English language so that they can write directly in final
form for publication. Other equally competent scientists find it necessary
to revise page after page not once but many times. T. D. A. Cockerell
was an example of the former type of writer, while by his own testimony
Charles Darwin was an inveterate reviser.
Trelease (1951) recommends careful reading of the manuscript ten
times, each time for one of the following: (1) consistency, (2) sentences,
(3) clarity, (4) repetition, (5) connectives, (6) euphony, (7) punctuation,
(8) style, (9) accuracy, (10) length. Authors of taxonomic papers seldom
follow the details of this recommendation, but most papers would benefit
from more revisions than they usually receive. It often helps to put a
manuscript aside for a while before the final revision is made. It is always
advisable to have other persons read a manuscript before it is submitted
for publication. A fully corrected carbon copy of the manuscript should
be kept by the author for use in case the original is lost.
11D.5 Proofreading. Most scientific journals permit the author to
read proof on his papers before publication. A few journals place the entire
burden of the proofreading on the author and hold him responsible for
typographical or other errors which may pass undetected. In any event,
where the author sees the proof, proofreading becomes a very important
292 Methods of Zoological Classification

part of his scientific responsibility. The scientific value of his paper can
be greatly lessened by unfortunate typographical errors. Such errors are
sometimes obvious to the reader, others are insidious and may be wholly
misleading.
In general, proof is submitted to the author to permit the elimination
of printer’s errors. Author’s errors are his own responsibility, and many
publications charge authors for corrections other than the printer’s errors.
Changes in proof are costly and therefore should not be made unless neces¬
sary, or unless the author is willing to assume the cost of the change.
Proofreading cannot always be done satisfactorily by one person. It
is advisable to supplement the personal reading by having someone else
read slowly from the original manuscript, while the proofreader (preferably
the author) carefully reads the proof. Special attention should be given
to punctuation, spelling of scientific names, numbers, and dates of all kinds.
When corrections are necessary, they should be made according to the
standard system of proofreader’s marks, as given in most dictionaries and
style manuals.
Most authors see only galley proofs of their papers. These are long
sheets with the text continuous and not broken into pages. For most journals
a galley is the equivalent of about three printed pages. Some publications
also submit page proofs to the authors. In such cases proofreading cannot
be restricted to individual words which were corrected in the galley proof
but must include the whole line in which the correction was made. Modern
linotype machines cannot change a single letter in a word but must reset
the whole line. If a word was inserted, it may have been necessary to
reset several lines or perhaps the remainder of the paragraph. The author
should carefully check everything which has been reset. It is advisable
also to read the top and bottom line of every page, because in the conversion
of the galley into page proof mishaps sometimes occur in these lines. Cor¬
rected proof should be returned at once to the editor or printer in order
to avoid delay in publication. The printing of an entire issue of a periodical
may be held up by a single tardy author.
11D.6 Reprints. Reprints must be ordered at the time the proof
is returned to the editor. It is advisable to order a larger than usual number
of reprints of papers that deal with general principles, because requests
for these will come not only from cospecialists.
A good reprint collection is particularly valuable in taxonomic re¬
search, where so many publications are only one or two pages long. With
the help of modern copying methods (Xerox, etc.) an author can build
up rapidly and inexpensively a specialized library of the entire literature
of a taxon (except for a few very popular groups).
Letters requesting reprints from other authors should be specific. Most
authors resent a request for “a set of reprints of your papers,” except
 Taxonomic Publication 293

under unusual circumstances. Few authors have an inexhaustible supply

of their papers, and the majority prefer to distribute their limited stock
to those who will obtain the greatest use from them.
If the author wishes to send all his reprints, he is still free to do
so. The receipt of requested reprints should be acknowledged, and in some
cases, especially where the expense of shipment is significant, the recipient
should offer to refund the postage involved.
Part TTT Principles and
m Application of
Zoological
Nomenclature
Chapter The Rules of Zoological
1 Nomenclature

T-A. he zoologist deals with an enormous

of items; each species, genus, and higher taxon is a different item. He
number

would not be able to refer to them if each one did not have a separate
name. The term nomenclature comes from the Latin words nomen (name)
and calare (to call) and means literally to call by name. It is the role of
nomenclature to provide labels for taxa at all levels, in order to facilitate
communication among biologists. The scientific names for species of organ¬
isms and for the higher taxa in which they are placed form a system of
communication, a language; they must fulfill the same basic requirements
as any other language.
It is most unfortunate that some taxonomists take a far greater interest
in the names of animals than in the animals themselves. Nomenclature
is an area in which one can operate even if one has only a minimal knowl¬
edge of biology. The result has been the elaboration of all sorts of complex
rules and regulations, often without a clear understanding of the underlying
principles. Many leading taxonomists, such as Hubbs, K. P. Schmidt, and
Simpson, have admonished taxonomists not to make a game of their occupa¬
tion with names, but to remember at all times that nomenclature is a
means to an end and not an end in itself.
What are the important requisites of any system of communication,
scientific nomenclature included? Surely there are many, but three seem
to be outstanding.

297
298 Principles and Application of Zoological Nomenclature

Uniqueness. A classification is a filing system, an information retrieval

system. The name of an animal, like the index number of a file, gives
immediate access to all the known information about the particular taxon.
Every name has to be unique because it is the key to the entire literature
relating to this species or higher taxon. If several names have been given
to the same taxon, there must be a clear-cut method whereby it can be
determined which of them has validity. Normally (but see 13.1, 13.3) pri¬
ority decides in cases of conflict.
Universality. Scientific communication would be made very difficult
if we had only vernacular names for animals; specialists would have to
learn the names of taxa in innumerable languages in order to communicate
with each other. To avoid this, zoologists have adopted by international
agreement a single language, a single set of names for animals, to be used
on a worldwide basis.
Stability. As recognition symbols the names of objects would lose much
of their usefulness if they were changed frequently and arbitrarily. It would
surely create confusion if we were to call an object a spoon today but
an apple next week. Yet this basic principle of communication has been
constantly violated by zoologists. Altogether too much name-changing has
occurred in zoological taxonomy during the past 200 years.
These three major objectives of the communication system of taxono¬
mists are singled out for attention in the Preamble of the Code: “The
object of the Code is to promote stability and universality in the scientific
names of animals, and to ensure that each name is unique and distinct.
All its provisions are subservient to these ends.”
The Preamble stresses another vitally important principle, that none
of the provisions of the Code shall “restrict the freedom of taxonomic
thought or action.” This principle is of relevance in Arts. 11, 17, 36, 49,
59b, 64, and 67-69 of the Code. It means that no taxonomist shall be
forced to accept a particular classification, or the delimitation of a taxon,
or be guided in his original choice of the type of a taxon by any but
zoological reasons (see 13.2).

THE INTERNATIONAL CODE

The valid rules of zoological nomenclature are contained in an au¬

thoritative document entitled the International Code of Zoological Nomen¬
clature. Its most recent version, adopted by the Fifteenth International
Congress of Zoology (London), came into force on the day of its publica¬
tion, November 6, 1961. A few minor modifications, affecting Arts. 11,
31, 39, and 60, were adopted by the Sixteenth Congress at Washington
in 1963 and are incorporated in the 1964 edition of the Code. This Code
 The Rules of Zoological Nomenclature 299

is a stage in a long history described in part in the introduction to the

Code (pp. ix-xix) and by Linsley and Usinger (1959).
Linnaeus himself was the father of a set of rules of nomenclature
published in the Critica botanica (1737) and the Philosophia botanica
(1751). As the Linnaean authority faded out during the nineteenth century,
new and local nomenclatural traditions developed and so did local sets
of scientific names, owing to a lack of communication among taxonomists.
As Strickland said in 1842,

If an English zoologist, for example, visits the museums and converses

with the professors of France, he finds that their scientific language is
almost as foreign to him as their vernacular. Almost every specimen which
he examines is labeled by a title which is unknown to him, and he feels
that nothing short of continued residence in that country can make him
conversant with her science. If he proceeds thence to Germany or Russia,
he is again at a loss; bewildered everywhere amidst the confusion of nomen¬
clature, he returns in despair to his own country and to the museums
and books to which he is accustomed.

To stem this confusion zoologists and paleontologists in England,

France, Germany, Russia, and America drafted formal rules of nomencla¬
ture. What was particularly needed, however, was an International Code.
This was adopted in principle at the Fifth International Zoological Congress
(Berlin, 1901) but was not formally issued until after the Sixth Congress
(Bern, 1904) as the Regies Internationales de la Nomenclature Zoologique
(Paris, 1905).
Certain of the articles of the original Code were amended by the
later congresses at Boston (1907), Graz (1910), Monaco (1913), Budapest
(1927), and Padua (1930). Far more extensive changes were discussed
at the three postwar congresses of Paris (1948), Copenhagen (1953), and
London (1958). By this time it had become apparent that a drastic reediting
of the rules was necessary. The sequence of the articles in the old Code
was rather arbitrary, and much nomenclatural tradition was not codified
at all. A large body of case law had also accumulated in the meantime,
published in the Opinions of the International Commission on Zoological
Nomenclature. An editorial committee under Norman R. Stoll edited the
version of the new Code as it had been adopted by the previous congresses,
particularly the London Congress.
The new Code, like any human document, is not perfect. We shall
point out in Chap. 13 some aspects that would seem to be still in need
of improvement. Nevertheless, this Code is a document of extraordinary
clarity and comprehensiveness. A great deal that was left to implication
in the old rules is now explicitly stated in the new Code. The rules are
a code of law, which makes it necessary to state matters precisely and
without explanation. In order to improve understanding, Chap. 13 of this
300 Principles and Application of Zoological Nomenclature

book attempts to provide explanatory comments, numbered 1-66, on all

those provisions of the Code, the understanding of which can be facilitated
by explanations. Some very valuable explanations are also found in the
“Bradley draft” (Bulletin of Zoological Nomenclature, vol. 14, pp. 1-286),
which formed the basis for the deliberations of the London Congress. How¬
ever, the Bradley draft has no official status, and some of its provisions
were rejected by the London Congress.
The relations between the Zoological Congresses and the International
Commission on Zoological Nomenclature are frequently misunderstood. The
Congress is a legislative body, and all provisions of the Code as well as
the Constitution of the Commission were adopted by vote of the Congress.
The Commission is a judicial body elected by the Congress as provided
in the Constitution and By-laws of the Commission (see 13.59).

EVOLUTION OF THE THEORY OF NOMENCLATURE

Many provisions of the Code are the results of compromise between

conflicting principles. Such compromises go back to Linnaeus, whose nomen¬
clature embodied on one hand the principles of Aristotelian logic and on
the other hand some very practical considerations. The conflicts between
stability and priority, between taxonomic freedom and nomenclatural rigid¬
ity, between typification through type-fixation and through type-restriction
are some other examples. Equally important are certain changes in the
last 150 years in the basic concepts of taxonomy. These include the concept
of taxa as populations rather than types, the nature of the type as a name
bearer, the categorical status of infrasubspecific names, and finally the appli¬
cation of some basic legal principles to the laws of nomenclature, such
as the impropriety of a retroactive application of laws and the stabilizing
effect of statutes of limitation.
All good law is living law. This is as true for the Code as for all
other codes of law. The International Rules have prevented the develop¬
ment of chaos in the naming of animals and have greatly helped to stan¬
dardize taxonomic procedures. Any change of these rules is upsetting and
should not be undertaken lightly. Yet the rules should not become rigid,
losing contact with the conceptual evolution of taxonomy. It is the rules
that will have to be adjusted to the conceptual development of taxonomy
and not the reverse.
An unresolved difficulty is posed by the drastically different needs
of different taxonomic groups. The needs that exist in popular groups like
birds or mammals, in which a limited number of species are referred to
by their scientific names hundreds or thousands of times annually, are
very different from the needs in an obscure group of arthropods attended
 The Rules of Zoological Nomenclature 301

to by a single specialist, with many species not mentioned in the literature

more frequently than once every 30 or 50 years. Different problems exist
for the parasitologist, who must make tentative assignments of stages in
the life cycle of parasites (e.g. cercariae) even though the connection with
the adult stages can be established only by experiment, and for the paleon¬
tologist, who may be forced to adopt form genera until the true taxonomic
status of the objects placed in these form genera is fully established. Dissen¬
sion and controversy will inevitably result if specialists in one group of
organisms are oblivious of the needs of specialists in other groups.

TEXT OF THE INTERNATIONAL CODE OF ZOOLOGICAL

NOMENCLATURE

The 1964 Code consists of a Preamble, 86 Articles (Article 31 was

repealed in 1963), five Appendices, an official Glossary, and a detailed
Index, all items in parallel English and French versions. The excerpt here
reprinted is limited to the English version of the preamble, articles, and
recommendations, needed as the basis for the commentary in Chap. 13
(comments 1—66). Since the French version, as well as the Appendices,
the official Glossary, and the official Index are indispensable for a full
understanding of the Code, every zoologist, whether a taxonomist or not,
is urged to acquire an official copy of the Code. It can be ordered from
the International Trust for Zoological Nomenclature, 14 Belgrave Square,
London S.W.l, at the cost of $3.00.

INTERNATIONAL CODE OF ZOOLOGICAL NOMEN¬

CLATURE ADOPTED BY THE XV INTERNATIONAL
CONGRESS OF ZOOLOGY, LONDON, JULY 1958.
Published in 1961. Revised edition 1964
Excluding Preface, Introduction, Appendices, Glossary,
and Index.

PREAMBLE

The International Code of Zoological Nomenclature is the system of rules

and recommendations authorized by the International Congresses of Zoology.
The object of the Code is to promote stability and universality in the scien- 1*
tific names of animals, and to ensure that each name is unique and distinct.
All its provisions are subservient to these ends, and none restricts the freedom 2,43
of taxonomic thought or action.
The numbers in the margin refer to the numbered comments in Chap. 13.
302 Principles and Application of Zoological Nomenclature

Priority is the basic principle of zoological nomenclature. Its application,

however, under conditions specified in the Code, may be moderated to pre- 3,4
serve a long-accepted name in its accustomed meaning.
When stability of nomenclature is threatened in an individual case, the
strict application of the Code may under specified conditions be suspended 50,60
by the International Commission on Zoological Nomenclature.

I. ZOOLOGICAL NOMENCLATURE

Article 1. Zoological nomenclature is the system of scientific names applied 5

to taxonomic units of animals (taxa; singular: taxon) known to occur in
nature, whether living or extinct. This Code is concerned with such names
in the family-, genus-, and species-groups [VIII, IX, X; for work of an ani¬
mal, see Art. 16a(viii)]. Names given to hypothetical concepts, to terato- 6,17
logical specimens or to hybrids as such, to infrasubspecific forms as such, or 21,41
names proposed for other than taxonomic use, are excluded.
Article 2. Independence of zoological nomenclature. Zoological no¬
menclature is independent of other systems of nomenclature in that the 5
name of an animal taxon is not to be rejected merely because it is identical
with the name of a taxon that does not belong to the animal kingdom.

(a) Transfer of taxa to the animal kingdom. If a taxon is transferred to the

animal kingdom, its name or names enter into zoological nomenclature
with the original date and authorship.
(b) Removal of taxa from the animal kingdom. If a taxon is removed from
the animal kingdom, its name or names continue to compete in homonymy
with names in the animal kingdom.
Recommendation 2A. Names already in use outside the animal kingdom.
It is preferable not to propose for a genus of animals a name already in use for a
genus outside the animal kingdom.

Article 3. Starting point. The 10th edition of Linnaeus’s Systerna Naturae

marks the beginning of the consistent general application of binominal 8,9
nomenclature in zoology. The date 1 January 1758 is arbitrarily assigned
in this Code as the date of publication of that work and as the starting point
of zoological nomenclature. Any other work published in 1758 is to be
treated as having been published after that edition.

II. NUMBER OF WORDS IN ZOOLOGICAL NAMES 10,14

Article 4. Taxa of rank above the species-group. The name of a taxon 11

of higher rank than the species-group consists of one word (uninominal).
Article 5. Species and subspecies. The name of a species consists of two 11,12,
words (binomen) and that of a subspecies of three words (trinomen); in each 13,14
case the first word is the generic name, the second word is the specific name, 40
and the third word, when applicable, is the subspecific name.
 The Rules of Zoological Nomenclature 303

Article 6. Subgenus. The name of a subgenus, when used in combina- 11

tion with a generic and a specific name, is placed in parentheses between
those names; it is not counted as one of the words in the binominal name of
a species or trinominal name of a subspecies.

Ill: CRITERIA OF PUBLICATION 15

Article 7. Application. The provisions of this Chapter apply to the publi- 22

cation not only of a new name but also to any other information that affects
nomenclature.
Article 8. What constitutes publication. To be regarded as published 15
within the meaning of this Code, a work when first issued must

(1) be reproduced in ink on paper by some method that assures numerous

identical copies;
(2) be issued for the purpose of scientific, public, permanent record;
(3) be obtainable by purchase or free distribution; and
(4) not be reproduced or distributed by a forbidden method (Art. 9).
Recommendation 8A. Mimeographing and similar processes. Zoologists
are strongly urged not to use mimeographing, hectographing, or similar pro¬
cesses for a publication containing a new name or a statement affecting nomen¬
clature.

Article 9. What does not constitute publication. None of the following 15

acts constitutes publication within the meaning of the Code:

(1) distribution of microfilms, or microcards, or matter reproduced by similar

methods;
(2) distribution to colleagues or students of a note, even if printed, in explana¬
tion of an accompanying illustration;
(3) distribution of proof sheets;
(4) mention at a scientific or other meeting;
(5) labelling of a specimen in a collection;
(6) mere deposit of a document in a library; or
(7) after 1950, anonymous publication.

IV. CRITERIA OF AVAILABILITY

Article 10. When a name becomes available. A name becomes avail¬

able, and takes date and authorship, only when it satisfies the provisions of 16
Article 11; in addition, names published after certain dates must satisfy also
the conditions of Articles 12, 13, 14 and 15. 17

(a) Interrupted publication. If publication of the data relating to a new

nominal taxon is interrupted and continued later, the name becomes avail¬
able only when it satisfies all the relevant provisions.
Recommendation 10A. Divided description. Editors should not permit the
description of a new taxon below the family-group to be so divided that portions
appear at different times.
304 Principles and Application of Zoological Nomenclature

(b) Infrasubspecific names. A name first established with infrasubspecific

rank becomes available if the taxon in question is elevated to a rank of the 41
species-group, and takes the date and authorship of its elevation.

Article 11. General requirements. A name, to become available, must

satisfy the following provisions:
(a) Publication and date. It must have been published, in the meaning of
Chapter III, after 1757. 22
(b) Language. The name must be either Latin or latinized, or, if an arbi¬
trary combination of letters, must be so constructed that it can be treated 28
as a Latin word [VII],
(i) The letters “j”, “k”, “w”, and “y”, more common in Neo-Latin,
may be used in zoological names.
(c) Binomial nomenclature. The author must have consistently applied
the principles of binominal nomenclature [II] in the work in which the 11
name is published.
(i) Uninominal genus-group names published before 1931 without
associated nominal species are accepted as consistent with the prin¬
ciples of binominal nomenclature, in the absence of evidence to
the contrary.
(ii) Names published before 1931 in the index to a work, if they satisfy
the relevant provisions of this Article and of Articles 4, 5 and 6, are
available, even if the author did not use binominal nomenclature in
the body of the work, provided that there is a clear bibliographic
reference to a description, indication, or figure of the animal in
question, or, if it is a family-group name, provided that it is founded
on an included nominal genus [Art. 16a(ii)].
(d.) Publication in synonymy. A name first published as a synonym is not
thereby made available unless prior to 1961 it has been treated as an avail- 18
able name with its original date and authorship, and either adopted as the 2
name of a taxon or used as a senior homonym.
(e) Names of the family-group. A family-group name must, when first
published, be based on the name then valid for a contained genus, and 33
must be a noun in the nominative plural.
(i) The name must clearly be used to denote a suprageneric taxon,
and not merely be employed as a plural noun or adjective referring
to the members of a genus.
(ii) A family-group of which the suffix is incorrect is available with its
original date and authorship, but in properly emended form
[Art. 29],
Example. Latreille [1802-1803], proposed a family TIPULARIAE, based on
Tipula Linnaeus, 1758. The name must be corrected to Tipulidae and attributed
to Latreille [1802-1803], not to the author who first corrected the spelling.
(iii) A family-group name published before 1900 in accordance with the
above provisions of this Section, but not itself fully latinized, is
available with its original date and authorship, provided that it has
been latinized by later authors and that it has been generally ac¬
cepted by zoologists interested in the group concerned as dating
from its first publication in vernacular form.
Example. The family name TETRANYCHIDAE is generally attributed to
Donnadieu, 1875. He published the name as “Tetranycides,” but in view of the
general acceptance of his name, it is to be attributed to his work and date, not to
Murray, 1877, who first latinized it.
 The Rules of Zoological Nomenclature 305

(/) Names of the genus-group. A genus-group name must be a noun in the 38

nominative singular or be treated as such.
(g) Names of the species-group.
(i) A species-group name must be a simple word of more than one 40
letter, or a compound word, and must be or be treated as
(1) an adjective in the nominative singular agreeing in gender
with the generic name (e.g. Felts marmorata), or
(2) a noun in the nominative singular standing in apposition to
the generic name (e.g. Fells leo), or
(3) a noun in the genitive case (e.g. rosae, sturionis, thermopylarum,
galliae, sanctipauli, sanctaehelenae, cuvieri, merianae, smithorum), or
(4) an adjective used as a substantive in the genitive case, derived
from the specific name of an organism with which the animal
in question is associated (e.g. Lernaea lusci, a copepod parasite
on Gadus luscus).
(ii) A species-group name must be published in combination with a
genus-group name, but the latter need not be valid or even avail¬
able.
(iii) A species-group name must not consist of words related by a con¬
junction, nor include a sign that cannot be spelled out in Latin.
Example. Expressions like rudis planusque and 1-album are not admissible as
specific names.

Article 12. Names published before 1931. In addition to satisfying the

provisions of Article 11, a name published before 1931 must have been ac- 16
companied by a description, definition, or indication [Art. 16].
Article 13. Names published after 1930.
(a) Names in general. In addition to satisfying the provisions of Article 11,
a name published after 1930 must be either
(i) accompanied by a statement that purports to give characters differ- 19,38
entiating the taxon; or 40
(ii) accompanied by a definite bibliographic reference to such a state- 47
ment; or
(iii) proposed expressly as a replacement for a preexisting available 47
name.
(b) Genus-group names. A genus-group name published after 1930 must, 38
in addition to satisfying the provisions of Section (a), be accompanied by 52
the definite fixation of a type-species [Art. 68].
(i) The provisions of this Section do not apply to names of collective
groups [Art. 66].
Article 14. Names published after 1950. After 1950, a new name pub¬
lished anonymously is not available.
Article 15. Names published after 1960. After 1960, a new name pro¬
posed conditionally, or one proposed explicitly as the name of a “variety” or
“form” [Art. 45<r], is not available. 20
Article 16. Indications.
(a) What constitutes an indication. The word “indication” as used in this
Chapter applies only to the following:
(i) a bibliographic reference to a previously published description,
definition, or figure;
(ii) the inclusion of a name in an index to a work, provided that the
provisions of Article llc(ii) are satisfied;
:i(Mi Principles and Application of Zoological Nomenclature

(iii) the substitution of a new name for a previously established name;

(iv) the formation of a new family-group name from the stein of the
name of a genus, which thus becomes the type-genus;
(v) the citation, in combination with a new genus-group name, of one
or more available specific names;
(vi) a single combined description of a new nominal genus and a new
nominal species, which provides an indication for each name;
(vii) the publication of a new genus- or species-group name in connec¬
tion with an illustration; or
(viii) the description of the work of an animal, even if not accompanied
by a description of the animal itself.
(/;) What docs not constitute an indication. The following arc not “indica¬
tions” in the meaning of this Chapter:
(i) mention of a vernacular name, type-locality, geological horizon,
host, or a label or specimen in a collection; or
(ii) citation of a name in synonymy [see also Art. 11 d\.

Article 17. Conditions that do not prevent availability. A name is or

remains available even though
(1) it becomes a junior synonym: such a name may be re-employed if the
synonymy is judged to be erroneous, or if the senior synonym is found to
be invalid or unavailable; or
(2) in the case of a species-group name, it is found that the original description
relates to more than one taxonomic unit, or to parts of animals belonging
to more than one taxon, or to an animal and animals later found to be
hybrid; or
(3) in the case of a species-group name, the genus-group name with which it
is first combined is invalid or unavailable; or
(4) it is based only on any part of an animal, sex of a species, stage in life-
history, one of several dissimilar generations, or one form of a polymorphic
species; or
(5) it was originally proposed for an organism now but not then considered
an animal; or
(6) it was incorrectly spelled with respect to any of the provisions of Chapter
VII, in which case it is to be corrected and the correct spelling is the avail¬
able one [Art. 32c]; or
(7) before 1951, it was published anonymously; or
(8) before 1961, it was proposed conditionally; or
(9) before 1961, it was proposed as a “variety” or “form.”

Article 18. Unallowable causes for rejection

(a) Inappropriatcncss. A genus- or species-group name, once established,

cannot afterwards be rejected, even by its own author, because of in¬
appropriatcncss.
Examples. Names such as Polyodon, Apus, albus, sinensis, etc., once published, are
not to be rejected because of a claim that they denote a character or distribution
not possessed by the animal in question.
(b) Tautonymy. A name is not to be rejected because of tautonymy, that is,
because the species-group name or names are identical with the generic
name.
Examples. Bison bison; Apus apus apus.
 The Rules of Zoological Nomenclature 307

Article 19. Status of emendations and errors. In the meaning of the

Code, an emendation, whether justified or unjustified, is an available name,
but an incorrect spelling, whether original or subsequent, has no standing in
nomenclature and is not an available name [Art. 32c, 33],
Article 20. Genus-group names ending in -ites, -ytes, or -ithes given
to fossils. If an existing genus-group name has been modified by substitut¬
ing -ites, -ytes, or -ithes for its original termination, the modified name if
applied only to fossils is not available, except for the purposes of the Law of
Homonymy [Art. 56b], unless there is clear evidence of intent to establish a
distinct genus or subgenus.

Example. The generic names Pectinites and Tellinites Schlotheim, 1813, used to
denote fossil shells thought to belong to the Recent genera Pecten Muller, 1767,
and Tellina Linnaeus, 1758, are available only for the purposes of the Law of
Homonymy.

V. DATE OF PUBLICATION

Article 21. Interpretation of date. The date of publication of a work and

of a contained name or statement affecting nomenclature is to be interpreted 22
in accordance with the provisions of this Article.

(a) Date specified. The date of publication specified in a work is assumed

to be correct in the absence of evidence to the contrary.
(b) Date incomplete. If the date of publication is not completely specified,
it is to be interpreted as the earliest day demonstrated by evidence, but
in the absence of such evidence, as
(i) the last day of the stated month, when month and year, but not the
day, are specified; and as
(ii) the last day of the year, when only the year is specified.
(c) Date incorrect. If the date of publication specified in a work is found to
be incorrect, the date is to be interpreted as the earliest demonstrated by
the evidence.
(d) Dates of work issued in parts. If parts of a work were published on
different days, the date of each is reckoned independently.
(e) Range of dates. If the specified date of publication contained within a
work is a range of dates, the work is to be dated from the latest day within
that range; but if evidence proves that one or more parts were issued before
that day, it or they are to be interpreted as dating from the earliest day
demonstrated by the evidence.
(/) Date not stated. In the absence of internal evidence of its date of publi¬
cation, a work is to be dated in whole or in part from the earliest date
demonstrated by external evidence, such as mention in another work.
Recommendation 21A. Responsibility of editors, publishers, and authors.
Editors and publishers should not put any copy or part of a work into circulation
in advance of the specified date of publication. Authors should not distribute
reprints (separata) in advance of such publication.
Recommendation 21B. Dating of publications. Editors and publishers
should take care to state the exact date of issue of each component part of a serial
308 Principles and Application of Zoological Nomenclature

publication or of any work issued in parts. A completed volume containing parts

brought out separately should state the exact day of publication of each part,
and the exact pages, plates, maps, etc., that constitute it.
Recommendation 21C. Responsibility of librarians. If a work deals with
zoology, librarians should not remove covers that bear information relative to
the date of publication and content of the work or its parts, or to the dates of their
receipt in the library.
Recommendation 21D. Information on reprints and preprints. Reprints
(separata) should contain information sufficient for complete citation, including
pagination and date of publication coinciding with the source-publication. Pre¬
prints should be definitely identified as such.

Article 22. Citation of date. The date of publication of a name, if cited,

follows the name of the author with a comma interposed.

Recommendation 22A. Method of citation. In citing the date of publication

of a name, an author
(1) should not enclose the date in either parentheses or square brackets if the
work containing it specifies the date of publication;
(2) should enclose the date, or a part of it, in parentheses if it is determined by
evidence derived from the volume concerned other than in (1); or
(3) should enclose the date, or a part of it, in square brackets if it is deter¬
mined only from external evidence.
Recommendation 22B. Date in a changed combination. If the original date
of publication is cited for a species-group name in a changed combination, it
should be enclosed within the same parentheses as the name of the original author,
separated by a comma [Art. 51 d].
Example. Arion ater (Linnaeus, 1758).

VI. VALIDITY OF NAMES

Article 23. Law of Priority. The valid name of a taxon is the oldest
available name applied to it [taking into consideration the provisions of
Sections (<f)(i) and (e), below], provided that the name is not invalidated
by any provision of this Code or has not been suppressed by the Commission.

(a) Exceptions. A name that is not the oldest available name is nevertheless
the valid name of the taxon in question
(i) if it is conserved under Section (b) of this Article; or
(ii) if the Commission has expressly validated it.
(b) Limitation. A name that has remained unused as a senior synonym in
the primary zoological literature for more than fifty years is to be considered
a forgotten name (nomen oblitum).
(i) After 1960, a zoologist who discovers such a name is to refer it to
the Commission, to be placed on either the appropriate Official
Index of Rejected Names, or, if such action better serves the stabil¬
ity and universality of nomenclature, on the appropriate Official
List.
(ii) A nomen oblitum is not to be used unless the Commission so directs.
(iii) This provision does not preclude application to the Commission for
the preservation of names, important in applied zoology, of which
the period of general usage has been less than fifty years.
 The Rules of Zoological Nomenclature 309

(c) Change of rank. The priority of the name of a taxon in the family-, 35
genus-, or species-group is not affected by elevation or reduction in rank
within the group.
(d) Family-group names. 33
(i) A family-group taxon formed by the union of two or more taxa of 25
that group takes the oldest valid familv-group name among those
of its components, with change of termination if required.
(ii) if a zoologist observes that the strict application of the Law of 73
Priority to two or more synonymous family-group names would
upset general usage, he is to request the Commission to decide 1
which name is to be accepted for the Official List of Family-Group 63
Names in Zoology.
(e) Genus- and species-group names.
(i) A genus-group taxon formed by the union of two or more genus- 25
group taxa takes the oldest valid name among those of its com¬
ponents.
Example. The valid name of a genus formed by the union of genus A-us, 1850,
and subgenus B-us, 1800, is B-us, 1800.
(ii) A species-group taxon formed by the union of two or more species- 40
group taxa takes the oldest valid name among those of its com¬
ponents.
(iii) If the name of a genus or species having subordinate taxa is found
to be invalid or unavailable, it must be replaced by the next oldest 23
valid name among those of the included co-ordinate taxa, in¬
cluding synonyms.
Example. Genus A-us, 1850, contains the subgenera A-us, 1850, C-us, 1900,
and D-us, 1860. If the name A-us is found to be a homonym, it is replaced as the
name of the genus by D-us, 1860, the next oldest valid name.
(/) Spelling. For the application of the Law of Priority to the spelling of
names, see Chapter VII.

Article 24. Interpretation of oldest name 23

(а) Names published simultaneously. If more than one name for a single 26
taxon, or identical names for different taxa, are published simultaneously,
whether in the same or different works, their relative priority is determined
by the action of the first reviser.
(i) The expression “first reviser” is to be rigidly construed. In the case 4
of synonyms, an author must have cited two or more such names,
must have made it clear that he believes them to represent the same
taxonomic unit, and must have chosen one as the name of the taxon.
Recommendation 24A. Action of first reviser. In acting as “first reviser” 4
in the meaning of this Section, a zoologist should select the name that will best 26
ensure stability and universality of nomenclature. If none of the names has an
advantage of this sort, nor has any special appropriateness, he should select the
name that has precedence of position in the work in question.
(б) Names founded on any part or form of an animal or on its work. The
Law of Priority applies
(i) when any part of an animal is named before the whole animal; or
(ii) when two or more generations, forms, stages, or sexes of a species
are named as different taxa; or
(iii) when, before 1931, a name is founded on the work of an animal 27
before one is founded on the animal itself.
310 Principles and Application of Zoological Nomenclature

VII. FORMATION AND EMENDATION OF NAMES

Article 25. Formation of names. Zoological names must be formed in 28

accordance with the provisions of Articles 26 to 30.

Recommendation 25A. Transliteration and latinization. Zoologists form¬

ing new names should, in the interest of proper usage and universality of nomen¬
clature, consult Appendices B to D in order to ensure correct transliteration and
latinization.

Article 26. Compound names.

(a) Acceptable compounds. If a name based on a compound name is pub¬

lished as two separate words in a work in which the author duly applied
the principles of binominal nomenclature, the component words are to be 28
united without a hyphen, and the name is to be treated as though it had
been originally published in that form. [For treatment of hyphenated com¬
pounds, see Article 32c(i).]

Examples. Coluber novae hispaniae Gmelin is corrected to Coluber novaehispaniae,

Calliphora terrae novae Macquart to Calliphora terraenovae, and Cynips quercus phellos
Osten Sacken, based on the binominal name of the host species, to Cynips quer-
cusphellos.
(b) Numerals in compounds. A number or numerical adjective or adverb
forming a part of a compound name is to be written in full as a word and
united with the remainder of the name (e.g. decemlineata, not 10-lineata).
(c) Latin letters in compounds. If the first element of a compound species-
group name is a Latin letter used to denote a character of the taxon, it is
connected to the remainder of the name by a hyphen (e.g. c-album).

Article 27. Diacritic and other marks. No diacritic mark, apostrophe,

or diaeresis is to be used in a zoological name; the hyphen is to be used only 29
as specified in Article 26c.
Article 28. Initial letters. Names of the family- and genus-groups must
be printed with a capital initial letter and names of the species-group with a
lower-case initial letter.
Article 29. Formation of family-group names. A family-group name is 30
formed by the addition to the stem [see Glossary] of the name of the type-
genus, of -IDAE in the case of a family and -INAE in the case of a subfamily
[see Appendix D, Table 2, Part A],

Recommendation 29A. Superfamilies and tribes. It is recommended that

the termination -OIDEA be adopted for the names of superfamilies and -INI
for the names of tribes.
(a) Generic name of classical origin. In zoological nomenclature, if the
name of a type-genus either is or ends in a Greek or Latin word, or ends in 30
a Greek or Latin suffix, the stem is found by deleting the case-ending of 9
the appropriate genitive singular [see Appendix D, VII],
(i) Where the word “Latin” is used in the Code, it includes ancient,
mediaeval, and modern Latin, but the word “Greek” refers only to
ancient Greek [see also Article lli(i)].
 The Rules of Zoological Nomenclature 311

(6) Generic name not of classical origin. If the name of a type-genus is or

ends in a word not Greek or Latin, or is an arbitrary combination of letters,
the stem is determined by the zoologist who first publishes a family-group
name based on that nominal genus [see Appendix D, VII],
(c) Generic name latinized from Greek. If the name of a type-genus is or
ends in a Greek word latinized with a change in termination, the stem is
that appropriate for the latinized form.
Example. For Leptocerus, of which the second part is from keras, the stem for the
formation of a family-group name is Leptocer-, not Leptocerat-.

Article 30. Agreement in gender. A species-group name, if an adjective 31

in the nominative singular, must agree in gender with the generic name with
which it is at any time combined, and its termination must be changed, if
necessary, when the species transferred to another genus. The gender of a
genus-group name is determined by the following provisions.

(a) Genus-group names consisting of or ending in a Greek or Latin word

or ending in a Greek or Latin suffix.
(i) A genus-group that is or ends in a Greek or Latin word takes the
gender given for that word in the standard Greek or Latin diction¬
aries, unless the Commission rules otherwise.
Examples. Greek nouns transliterated without change into Latin as the whole
or as part of a name, e.g. Hoplites, from dxXirris, masculine; Ichthyornis, ending in
-opvis, masculine; Waltonithyris, ending in Oipis, feminine; Names ending in -ops, if
from Greek op (“voice,” or rarely “face”; but see under (2), below), -opsis opis,
-gaster yaorfip, -caris nap is, or -lepis \ems, are feminine. Tuba from Latin tuba is
feminine. Names ending in certain Latin nouns in -us are feminine (e.g. -alvus,
-humus, -vanus, -fraus, -laus, -acus, -colus, -domus, -tellus). Names ending in -ceras
(xepas), -soma (au/xa), -stigma (orLypa), or -stoma (aropa) are neuter.
(1) A name is to be regarded as a Greek or Latin word of the same spelling,
unless its original author states otherwise.
(2) A noun of variable gender, masculine or feminine, is to be treated as mas¬
culine, unless its author states, when he first publishes the name, that it is
feminine, or so treats it in combination with an adjectival specific name.
Examples. Compound Latin nouns ending in -cola, such as Sylvicola, are con¬
sidered as masculine. Names ending in -ops derived from the Greek dip (“face”
or “eye”), of which the usual classical gender is masculine, are to be treated as
masculine unless the author indicated otherwise or unless, failing such indication,
zoologists have generally treated them as feminine. [For names derived from op,
see example under (i) above.]
(3) If a genus-group name is a Greek word latinized with a change of termina- 31
don, it takes the gender appropriate to that termination.
Examples. Names ending in -us, latinized from the Greek endings -os (os)
(masculine or feminine), -a (a) (neuter), or -ov (on) (neuter), are masculine, e.g.
-echinus t\i-vos, echinos), -cephalus (KtpaXrj, kephale), -crinus (k/hvov, krinon), -stomus
(cTopa, stoma), -somus (auipa, soma), -cheilus (and -chilus) (x«Xos, cheilos),
-gnathus (yvados, gnathos), -rhamphus (pappos, rhamphos), -rhynchus (pvyxos,
rhynchos), or -stethus (ot-qQos, stethos). Names ending in -cera (nepas, keras) or
-metopa (ptronrov, metopon) are feminine.
(4) If a genus-group name is a Latin word of which the termination has been
changed, it takes the gender appropriate to the new termination.
Example. Dendrocygna, feminine, although partly formed from cygnus, mascu¬
line.
312 Principles and Application of Zoological Nomenclature

(ii) A genus-group name that ends in a Greek or Latin suffix, or in a

letter or letters considered as such, takes the gender appropriate to
its ending.
Examples. Names ending in -ides, -isles, -ites, -odes, or -aides are masculine.
Names such as Scatella and Oculina are feminine because of the suffix, although
derived respectively from the Greek neuter noun <tku>p, okcitos and the Latin
masculine noun oculus. A name based on a word in a non-classical language or on
an arbitrary combination of letters, with the addition of a Greek or Latin suffix,
takes the gender appropriate to the suffix, e.g. Buchia (from von Buch), Cum-
mingella (from Cumming), Velletia (from Vellet), Dafila, and the anagram Solubea
are all treated as feminine, but the anagram Daption as neuter.
(b) Genus-group names entirely of non-classical origin.—
(i) A genus-group name that reproduces exactly a word in a modern
Indo-European language having genders takes the gender of that
word.
Example. Pfrille, from the German “die Pfrille,” is feminine.
(ii) A genus-group name consisting of a word that is not Greek, Latin,
or modern Indo-European, or that is an arbitrary combination of
letters, takes the gender expressly attributed to it by its author, or
implied by an originally associated species-group name. If no
gender was assigned or implied, the name is to be treated as mascu¬
line, except that if the ending is clearly a natural classical feminine
or neuter one, the gender is that appropriate to the ending [Art.
30a (ii)].
Examples. Dacelo (anagram of Alcedo) is feminine as treated by its author, but
Vanikoro, Gekko, Abudefduf, and the anagram Milax are treated as masculine.
Recommendation 31A.* Species-group names formed from modern per¬
sonal names. A species-group name, if a noun formed from a modern personal
name, should usually end in -i if the personal name is that of a man, -orum if of
men or of man (men) and woman (women) together, -ae if of a woman, and
-arum if of women [see Article llg(i)(3) and Appendix D III].

Article 32. Original spelling.

(a) Correct original spelling. The original spelling of a name is to be re¬
tained as the “correct original spelling,” unless
(i) it contravenes a mandatory provision of Articles 26 to 30; or
(ii) there is in the original publication clear evidence of an inadvertent
error, such as a lapsus calami, or a copyist’s or printer’s error (in¬
correct transliteration, improper latinization, and use of an inap¬
propriate connecting vowel are not to be considered inadvertent
errors); or
(iii) in the case of a family-group name, there has been a necessary
correction of termination (other than necessitated by a change in
the rank of the taxon), or a justified emendation [Art. 33a] in the
stem of the name of the type-genus.
(b) Multiple original spellings. If a name is spelled in more than one way
in the original publication, the spelling adopted by the first reviser is to be
accepted as the correct original spelling, unless the adopted spelling is sub¬
ject to emendation under the provisions of Articles 26 to 30.
(c) Incorrect original spelling. An original spelling that docs not satisfy
the provisions of Articles 26 to 30, or that is an inadvertent error [Art.
* This recommendation replaces Art. 31, deleted by the XVIth Congress of Zoology, 1963.
 The Rules of Zoological Nomenclature 313

32a (ii)], or that is one of the multiple spellings not adopted by a first
reviser [Art. 326], is an “incorrect original spelling” and is to be corrected
wherever it is found; the incorrect spelling has not separate status in
nomenclature, and therefore does not enter into homonymy and cannot
be used as a replacement name.
(i) A name published with a diacritic mark, apostrophe, diaeresis, or 29
hyphen is to be corrected by the deletion of the mark concerned and
any resulting parts are to be united, except for one specified use of
the hyphen [Art. 26c], and except that when, in a German word,
the umlaut sign is deleted from a vowel, the letter “e” is to be in¬
serted after that vowel.
Examples. The name terrae-novae is corrected to terraenovae, d’urvillei to durvillei,
and nunezi to nunezi; but miilleri becomes muelleri and is not a homonym of mulleri
[Art. Sid].

Article 33. Subsequent spelling. 32

(а) Emendations. Any demonstrably intentional change in the original

spelling of a name is an “emendation.”
(i) A “justified emendation” is the correction of an incorrect original
spelling, and the name thus emended takes the date and author¬
ship of the original spelling.
(ii) Any other emendation is an “unjustified emendation”; the name
thus emended has status in nomenclature with its own date and
author, and is a junior objective synonym of the name in its original
form.
(б) Incorrect subsequent spellings. Any change in the spelling of a name, 32
other than an emendation, is an “incorrect subsequent spelling”; it has no 9
status in nomenclature and therefore does not enter into homonymy and
cannot be used as a replacement name.

Article 34. Endings.

(а) In names of the family-group. In names of the family-group, the

ending must be changed when a taxon is raised or lowered in rank.
(б) In names of the species-group. In names of the species-group, the
ending must be changed, if necessary, to conform with the gender of the 40
generic name with which the species-group name is at any time combined
[Art. 30],

VIII. TAXA OF THE FAMILY-GROUP AND THEIR NAMES 33

Article 35. Categories and their names.

(а) Categories included. The family-group includes the categories tribe,

subfamily, family, and superfamily, and any supplementary categories
required.
(б) Basis. Each taxon of the family-group is defined by reference to its type-
genus [see XIII, XIV],
(c) Names. A family-group name is to be formed and treated in accordance
with the relevant provisions of Art. 29.
314 Principles and Application of Zoological Nomenclature

Article 36. Categories co-ordinate. All categories in the family-group

are of co-ordinate status in nomenclature, that is, they are subject to the 35
same rules and recommendations, and a name established for a taxon in any
category in the group, and based on a given type-genus, is thereupon avail- 2
able with its original date and author for a taxon based on the same type-
genus in each of the other categories, with appropriate change of suffix.

Example. The proposal of HESPERIIDAE Latreille, 1809 (as HESPER-

IDES), based on Hesperia Fabricius, 1793, thereupon makes available, from the
year 1809, the superfamily name HESPERIOIDEA and the subfamily name
HESPERINAE, even though the former was first used by Comstock, J. H. and
A. B., 1904, and the latter by Watson, 1893.

Article 37. Subordinate taxa. The subordinate taxon that contains the
type-genus of a subdivided family-group taxon bears the same names as the 25,36
latter, except for suffix, and is termed the “nominate” subordinate taxon
(e.g. nominate subfamily, nominate tribe).

Example. The nominal family TIPULIDAE (type-genus Tipula Linnaeus,

1758) is divided into a number of subfamilies, each named after its own type- 35
genus. The subfamily TIPULINAE, containing Tipula, is the nominate subfam¬
ily of the TIPULIDAE.

Article 38. Homonymy between family-group names. See Article 55. 37

Article 39. Homonymy of the type-genus. The name of a taxon of the
family-group is invalid if the name of its nominal type-genus is a junior
homonym.
Article 40. Synonymy of the type-genus. When, after 1960, a nominal 1
type-genus is rejected as a junior synonym (objective or subjective), a fam¬
ily-group name based on it is not to be changed, but continues to be the 33
valid name of the family-group taxon that contains both the senior and
junior synonyms.

(a) Conservation of certain names. If a family-group name, changed be¬

fore 1961 because of such synonymy, has won general acceptance, it is to 1
be maintained in the interests of stability.
(i) In the event of divergent interpretation of the expression “general
acceptance,” reference is to be made to the Commission.
(b) Date of conserved name. A name adopted by virtue of the provisions
of Section (a) takes the date of the rejected name, of which it is to be con¬
sidered the senior synonym.
Example. If an original type-genus A-us is a junior synonym of B-us:
(1) When before 1961, A-IDAE has not already been replaced by B-IDAE, or
B-IDAE has not yet won general acceptance, then A-IDAE is to be con¬
tinued as the name for the family;
(2) But if A-IDAE has already been replaced by B-IDAE, and the latter has
won general acceptance, then B-IDAE is to be accepted as the correct
name, and is to considered as the senior synonym of A-IDAE, with the
same date as that of A-IDAE.
Recommendation 40A. Citation of author and date. If author and date
are cited [see Appendix E9], a family-group name adopted under the provisions
 The Rules of Zoological Nomenclature 315

of Article 40 should be cited with its own author and date, followed by the date
of the replaced name in parentheses.

Article 41. Misidentified type-genera. If stability and continuity in the

meaning of a nominal family-group taxon are threatened by the discovery 1
that its nominal type-genus is based on a misidentified type-species, or by 49
the discovery of an overlooked type-designation, the case is to be submitted to
the Commission [Arts. 65, 70].

IX. TAXA OF THE GENUS-GROUP AND THEIR NAMES

Article 42. Taxa of the genus-group 38

(a) Categories included. The genus-group, which is next below the family-
group and next above the species-group in the hierarchy of classification,
includes the categories genus and subgenus.
(b) Basis. Each taxon of the genus-group is objectively defined only by
reference to its type-species [XV],
(c) Collective groups. The names of certain biological groupings known as
“collective groups” [see Glossary] are to be treated as generic names in the 39
meaning of the Code, but collective groups require no type-species.
Examples. Agamodistomum, Agamofilaria Cysticercus, Diplostomulum, Glaucothoe,
Sparganum.
(i) Wherever the terms “taxon” or “name” are used in this Code at the
level of genus, the provision in question is to apply also to a collective
group or its name, unless there is a statement to the contrary, or unless
such application would be inappropriate.
(d) Subdivision of genera. A uninominal name proposed for a primary
subdivision of a genus, even if the subdivision is designated by a term such
as “section” or “division,” has the status in nomenclature of a subgeneric
name, provided the name satisfies the relevant provisions of Chapter IV.

Article 43. Categories co-ordinate. The categories in the genus-group

are of co-ordinate status in nomenclature, that is, they are subject to the
same rules and recommendations, and a name established for a taxon in 35
either category in the group, and based on a given type-species, is thereupon
available with its original date and author for a taxon based on the same
type-species in the other category.

Article 44. Subordinate taxa.

(a) Nominate subgenus. The subgenus that contains the type-species of a 25,38
subdivided genus bears the same name as the genus and is termed the
“nominate” subgenus.
(4) Change of nominate subgenus. When the invalid name of a subdivided
genus is replaced by the name of a different subgenus [Art. 23? (iii)], the
latter then becomes the nominate subgenus.
(c) Citation of the subgenus. See Art. 6.
Recommendation 44A. Citation of synonyms. In order to avoid misunder¬
standing, a synonym, or any term other than subgenus, should never be cited
between the generic and specific elements of a binomen.
316 Principles and Application of Zoological Nomenclature

X. TAXA OF THE SPECIES-GROUP AND THEIR NAMES

Article 45. Taxa of the species-group.

(a) Categories included. The species-group, for the purposes of this Code,
includes the categories species and subspecies.
(b) Basis. Each taxon of the species-group is objectively defined only by
reference to its type-specimen.
(c) Infrasubspecific forms. Infrasubspecific forms are excluded from the
species-group and the provisions of this Code do not apply to them [Art. 1;
see also Art. 10 (6)].
(d) Determination of subspecific or infrasubspecific status. The original
status of any name of a taxon of lower rank than species is determined as
(i) subspecific, if the author, when originally establishing the name,
either clearly stated it to apply to a subspecies or, before 1961, did
not clearly state its rank [see also Art. 45(e) below], and as
(ii) subspecific, if the author, when originally establishing the name,
stated the taxon to be characteristic of a particular geographical
area or geological horizon and did not expressly refer it to an infra¬
subspecific category; but as
(iii) infrasubspecific, if the author, when originally establishing the
name, either expressly referred the taxon to an infrasubspecific
rank, or, after 1960, did not clearly state that it was a subspecies.
(e) Interpretation of the terms “variety” and “form”
(i) Before 1961, the use of either of the terms “variety” or “form” is
not to be interpreted as an express statement of either subspecific
or infrasubspecific rank.
(ii) After 1960, a new name published as that of a “variety” or “form”
is to be regarded as of infrasubspecific rank.

Article 46. Categories co-ordinate. The categories in the species-group

are of co-ordinate status in nomenclature, that is, they are subject to the
same rules and recommendations, and a name established for a taxon in
either category in the group, and based on a given type-specimen, is there¬
upon available with its original date and author for a taxon based on the
same type-specimen in the other category.

Article 47. Subordinate taxa.

(a) Nominate subspecies. The subspecies that contains the type-specimen

of a subdivided species bears the same name as the species and is termed
the “nominate” subspecies.
(b) Change of nominate subspecies. If the invalid name of a subdivided
species is replaced by the name of a different subspecies [Art. 23«(iii)],
the latter then becomes the nominate subspecies.

Article 48. Binomina.

(a) Change of genus. After its original establishment, a specific name be¬
comes part of another binomen whenever it is transferred to a different
genus.
(b) Generic name conditional. If a zoologist publishes a binomen, using a
 The Rules of Zoological Nomenclature 317

previously established generic name in combination with an available

specific name, but at the same time conditionally proposes a new generic
name in combination with the specific name, he is considered to have
established two binomina, of which the former has priority.
Example. Lowe in 1843 published a new species, gracilis, in the genus Seriola
Cuvier, 1816, but at the same time conditionally proposed a new genus
Cubiceps for that species. He is considered to have established first the
binomen Seriola gracilis, and second the binomen Cubiceps gracilis.

Article 49. Misidentifications. The specific name used in an erroneous 2

specific identification cannot be retained for the species to which the name 49
was wrongly applied, even if the two species in question are in, or are later
referred to, different genera [see Art. 706],

Example. It is found that Smith, 1850, had recorded as “A-us b-us Dupont,
1800” a species different from that actually so named by Dupont. The specific
name b-us cannot be used to denote the species that was before Smith, even if
placed in a different genus from the true b-us Dupont.

XI. AUTHORSHIP 42

Article 50. Author of a name. The author (authors) of a scientific name

is (are) the person (persons) who first publish(es) it [III] in a way that satis¬
fies the criteria of availability [IV], unless it is clear from the contents of the
publication that only one (or some) of the joint authors, or some other person
(or persons), is alone responsible both for the name and the conditions that
make it available.

(a) Exception for names in minutes. If the name of a taxon is established

by publication in the minutes of a meeting, the person responsible for the
name, not the secretary or other reporter of the meeting, is the author.
Recommendation 50A. Information in minutes. Secretaries and other re¬
porters of meetings should not include in their published reports new names of
taxa or any information affecting nomenclature.
(b) Change in rank. Change in rank of a taxon within the family-, genus-,
or species-group does not affect the authorship of the nominal taxon.
(c) Justified emendation. A justified emendation is attributed to the
original author of the name [Art. 33a].
(d) Unjustified emendation. An unjustified emendation is attributed to
the author who published it [Art. 33a].

Article 51. Citation of name of author. 42

(a) Optional use. The name of the author does not form part of the name
of a taxon and its citation is optional.
(b) Form of citation. The original author’s name, when cited, follows the
scientific name without any intervening marks of punctuation, except as
noted in Section (d) and Recommendation 51 A.
(i) The name of a subsequent user of a scientific name, if cited, is to be
separated from it in some distinctive manner, other than by a
comma.
318 Principles and Application of Zoological Nomenclature

Example. Reference to Cancer pagurus Linnaeus as used by Latreille may be

cited as
Cancer pagurus Linnaeus sensu Latreille,
Cancer pagurus: Latreille,
or in some other distinctive manner, but not as
Cancer pagurus Latreille, nor as
Cancer pagurus, Latreille.
Recommendation 51A. Anonymous authors. If the name of a taxon was
published anonymously, but its author is known, his name, if cited, should be
enclosed in square brackets to show the original anonymity.
(c) Citation of contributors. If a scientific name and its validating condi¬
tions are the responsibility not of the author(s) of the publication contain¬
ing them, but only of one (or some) of the authors, or of some other
zoologist, the name of the author(s), if cited, is to be cited as “B” in “A”
or “B” in “A & B,” or whatever combination is appropriate.
(d) Use of parentheses in new combinations. If a species-group taxon was
described in a given genus and later transferred to another, the name of
the author of the species-group name, if cited, is to be enclosed in paren¬
theses.
Example. Taenia diminuta Rudolphi, when transferred to the genus Hymenolepis,
is cited as Hymenolepis diminuta (Rudolphi).
(i) The use of parentheses here applies only to transfers from one nomi¬
nal genus to another, and is not affected by the presence of a sub¬
generic name, or by any shifts of rank or position within the same
genus.
Recommendation 51B. Citation of author of new combination. If it is
desired to cite the names both of the original author of a species-group name and
of the reviser who transferred it to another genus, the name of the reviser should
follow the parentheses that enclose the name of the original author.
Example. Limnatis mlotica (Savigny) Moquin-Tandon.

XII. HOMONYMY 43

Article 52. Homonymy. In the meaning of the Code, homonymy is the

identity in spelling of available names denoting different species-group taxa
within the same genus, or objectively different taxa within the genus-group
or within the family-group.
Article 53. Law of Homonymy. Any name that is a junior homonym of 16
an available name must be rejected and replaced.
Article 54. Names that do not enter into homonymy. The following
names do not enter into homonymy:

(1) names that are unavailable in the meaning of the Code, except as noted
in Arts. 20 and 56b;
(2) names that have never been used for a taxon in the animal kingdom;
(3) names that are excluded from zoological nomenclature [Art. 1]; and
(4) incorrect spellings, both original and subsequent.

Article 55. Family-group names. Family-group names that are based 33

on different type-genera and that are identical, or differ only in suffix, are 37
homonyms.
 The Rules of Zoological Nomenclature 319

(a) Homonymy from similar generic names. If homonymy between names 43

in the family-group results from similarity but not identity of the names of
their type-genera, the case is to be referred to the Commission.
Example. Two generic names, Merope (Insecta) and Merops (Aves), each
resulted in the family name meropidae. To avoid the homonymy, the Commis¬
sion ruled that Merope should form the family name meropeidae (Opinion 140).
(b) Homonymy from incorrect spelling. A family-group name is not to be
rejected as a junior homonym if the homonymy results from the incorrect
spelling of the earlier name.
Example, psilopinae Cresson, 1925, based on Psilopa Fallen, is not to be re¬
jected as a homonym of psilopinae Schiner, 1868, which was based on Psilopus
Meigen and should have been written psilopodinae.

Article 56. Genus-group names. The Law of Homonymy applies to all

names in the genus-group, including those of collective groups. 43
%

(a) One-letter difference. Even if the difference between two genus-group

names is due to only one letter, these two names are not to be considered
homonyms.
Example. Two genera of Diptera, Microchaetina Van der Wulp, 1891, and
Microchaetona Townsend, 1919, are not homonyms; but see Appendix D3.

(b) Names ending in ites, ytes, or ithes given to fossils. A genus-group name
formed for use in paleontology by substituting -ites, -ytes, or -ithes for the
original termination of a generic name, and applied only to fossils, enters
into homonymy [Art. 20]. 44
(c) Precedence of genus over subgenus. Of two homonymous genus-group
names of identical date, one proposed for a genus takes precedence over 45
one proposed for a subgenus.

Article 57. Species-group names. The Law of Homonymy applies to

species-group names originally published in (primary homonyms), or later 43
brought together in (secondary homonyms) the same genus or collective 46
group, except as noted in Art. 59c.

Example. As separate proposals of new names, the following nominal taxa

called intermedius are primary homonyms of one another:

A-us intermedins Pavlov,

A-us intermedius Dupont,
A-us albus intermedius Black, and 46
A-us concolor intermedius Schmidt.

(a) Subgeneric name. The presence of a subgeneric name does not affect
homonymy between species-group names within the same genus.
Example. A-us (B-us) intermedius Pavlov and A-us (C-us) intermedius Dupont are
primary homonyms, but A-us (B-us) intermedius Pavlov is not a primary homonym
of B-us intermedius Black.
(b) Differences in spelling. Species-group homonymy within a given nomi¬
nal genus is not obviated by any emendation or incorrect spelling of the
generic name.
(i) Differences in termination that are due solely to gender are to be
disregarded in determining whether adjectival species-group names
are homonyms.
(c) Identical names in homonymous genera. Homonymy does not exist 43
320 Principles and Application of Zoological Nomenclature

between two identical species-group names originally or subsequently 44

placed in different genera that bear homonymous names.
Example. Noctua (Insccta) and Noclua (Aves) are homonyms, but Noctua
variegata Jung (Insccta) and Noctua variegata Quoy and Gaimard (Aves) arc not.
(d) One-letter difference. Except as specified in Art. 58, the difference of a
single letter is sufficient to prevent homonymy.
Examples. Raphidia londinensis and Raphidia londonensis (derived from Londinium
and London, words of the same origin and meaning), and Chrysops calidus and
Chrysops calhdus (derived from words of different origin and meaning) are not
homonyms; but see Appendix D5.
(e) Precedence of species over subspecies. Of two homonymous species- 45
group names of identical data, one proposed for a species takes precedence
over one proposed for a subspecies.

Article 58. Variable spelling. Two or more species-group names of the

same origin and meaning and cited in the same nominal genus or collective
group are to be considered homonyms if the only difference in spelling con¬
sists of any of the following (for diacritic and other marks, see Arts. 27 and
32c):

(1) the use of ac, oe, or e (e.g. caeruleus, coerulcus, cerulcus);

(2) the use of ci, i, or y (e.g. chciropus, chiropus, chyropus);
(3) the use of c or k (e.g. microdon, mikrodon);
(4) the aspiration or non-aspiration of a consonant (e.g. oxyrhynchus, oxy-
ryncus);
(5) the presence or absence of c before t (e.g. auctumnalis, autumnalis);
(6) the use of a single or double consonant (e.g. litoralis, littoralis);
(7) the use of /or ph (e.g. sulfureus, sulphureus);
(8) the use of different connecting vowels in compound words (e.g. nigri-
cinctus, nigrocinclus);
(9) the transcription of the semi-vowel i as y, ei, cj, or ij;
(10) the termination -i or -ii in a patronymic genitive (e.g. smithi, smithii);
(11) the suffix -crisis or -iensis in a geographical name (e.g. timorensis, timorien-
sis); and
(12) three pairs of names treated as special cases: saghalincrtsis and sakhalincnsis;
sibcricus and sibiricus; tianschanicus and tianshanicus.

Article 59. Primary and secondary homonyms [Art. 57], 43

(a) Primary homonyms. A species-group name that is a junior primary 16

homonym must be permanently rejected.
(b) Secondary homonyms. A species-group name that is a junior secondary 2
homonym must be rejected by any zoologist who believes that the two 44
species-group taxa in question arc congeneric.
(c) Revival of secondary homonyms. A name rejected after 1960 as a 9
secondary homonym is to be restored as the valid name whenever a zoolo- 43
gist believes that the two species-group taxa in question are not congeneric,
unless it is invalid for other reasons.
(i) In such a case the name proposed in replacement of the secondary
homonym becomes a junior objective synonym of the latter.
Example. A-us niger Smith, 1960, is believed to be congeneric with B-us niger
Dupont, 1950, and when transferred to B-us becomes a junior secondary homo¬
nym, and is renamed B-us atcr Jones, 1970. If subsequently A-us niger Smith, 1960
 The Rules of Zoological Nomenclature 321

is no longer believed to be congeneric with B-us niger Dupont, 1950, the former
specific name is again to be used. B-us ater Jones, 1970, then becomes a junior
objective synonym of A-us niger Smith, 1960.

Article 60. Replacement of rejected homonyms. A rejected homonym

must be replaced by an existing available name or, for lack of such a name, 47
by a new name.

(a) Junior homonyms with synonyms. If the rejected homonym has one
or more available synonym(s), the oldest of these must be adopted, with
its own authorship and date.
(i) A subjective synonym retains eligibility as a replacement name only
so long as it is regarded as a synonym of the rejected name.
(b) Junior homonyms without synonyms. If the rejected homonym has no
known available synonym, it must be replaced by a new name which will
then compete in priority with any synonym recognized later.

XIII. THE TYPE-CONCEPT 48

Article 61. Relationship of the type to the taxon. The “type” affords
the standard of reference that determines the application of a scientific name.
Nucleus of a taxon and foundation of its name, the type is objective and does
not change, whereas the limits of the taxon are subjective and liable to
change. The type of a nominal species is a specimen, that of a nominal genus
is a nominal species, and that of a nominal family is a nominal genus. Each
taxon has, actually or potentially, its type. The type of any taxon, once fixed
in conformity with the provisions of the Code, is not subject to change except
by exercise of the plenary powers of the Commission [Art. 79], or, excep¬
tionally in species-group taxa, under the provisions of Art. 75.

(a) Types of nominate subordinate taxa. The type of a taxon is also the
type of its nominate subordinate taxon, if there is one, and vice-versa.
Therefore, the designation of one implies the designation of the other.
(i) If different types are designated simultaneously for a nominal taxon
and for its nominate subordinate taxon, the designation for the
former takes precedence.
(b) Types and synonymy. If two taxa are based on the same type, their
names are objective synonyms. If two taxa with different types are sub¬
jectively united into a single taxonomic unit, their names are subjective
synonyms.

XIV. TYPES IN THE FAMILY-GROUP 33

Article 62. Application. The provisions of this Chapter apply equally

to all categories in the family-group.
Article 63. Types of family-group taxa. The type of each taxon of the
family-group is that nominal genus upon [the name of] which the family-
group name is based [Arts. 35b, 39, 40],
322 Principles and Application of Zoological Nomenclature

Article 64. Choice of type-genus. A zoologist establishing a new family-

group taxon is free to choose as type-genus any included nominal genus, not
necessarily that bearing the oldest name.

Recommendation 64A. Type-genus should be well-known. So far as possi¬

ble, a zoologist who establishes a family-group taxon should base it on a genus
that is both well known and representative of the family.

Article 65. Identification of the type-genus.

(a) Correct identification assumed. It is to be assumed that an author

publishing a new family-group name has correctly identified the nominal
type-genus of the taxon in question.
(b) Misidentification or altered concept. If the nominal type-genus of a
family-group taxon is found to be based on a misidentified type-species
[Art. 70], or if a subsequent fixation of the type-species of a nominal type-
genus has confused the accepted meaning of a family-group name, the
case is to be referred to the Commission.

XV. TYPES IN THE GENUS-GROUP

Article 66. Application. The provisions of this Chapter apply equally to

the categories genus and subgenus but not to collective groups, which
require no type-species [Art. 42c].

Article 67. General provisions.

(а) Types of genus-group taxa. The type of each nominal genus is a nomi¬
nal species known as the “type-species” [Art. 426].
Recommendation 67A. Terminology. Only the term “type-species” or a
strictly equivalent term in another language should be used in referring to the
type of a genus. The term “genotype” should never be used for this purpose.
(б) Kinds of type-designation. The type-species of a nominal genus is
termed “type by original designation” if it is definitely designated in the
original publication [Art. 68a], “type by indication” if determined by the
application of provisions (6) to (d) in Art. 68, and “type by subsequent
designation” if designated after the establishment of the nominal genus
[Art. 69].
(c) Designation. The term “designation” in relation to the fixation of a
type-species must be rigidly construed; a designation made in an am¬
biguous or qualified manner is invalid.
(i) Mention of a species as an example of a genus does not constitute a
type-designation.
(ii) Reference to a particular structure as “type” or “typical” of a genus
does not constitute a type-designation.
Examples. A statement such as any of the following is not to be regarded as a
type-designation in the meaning of this Section: “A-us b-us may possibly be re¬
garded as the type of A-us"; “A-us b-us is a typical example of the genus A-us";
“the venation of the anterior wings of A-us b-us is typical of the genus A-us."
(d) Types of nominate subgenera. See Art. 61a.
(e) Objective synonymy of the type-species. If a nominal species, type of
 The Rules of Zoological Nomenclature 323

a genus, is found to be a junior objective synonym, the senior synonym is

to be cited as the name of the type-species of the genus in question.
Example. Astacus marinus Fabricius, 1775, one of the species originally included
in the genus Homarus Weber, 1795, was designated as the type-species of Homarus.
However, Astacus marinus is a junior objective synonym of Cancer gammarus Lin¬
naeus, 1758, which is therefore to be cited as the type-species of Homarus.
(/) Actions of original author. Only the statements or other actions of the 56
the original author when establishing a new nominal genus are relevant in
deciding
(i) whether the type-species has been designated or indicated in con¬
formity with provisions (a) to (d) of Art. 68, and
(ii) which species are the originally included species in the meaning of
'Art. 69a.
(g) Incorrect reference to establishment of genus. If, in designating the
type-species for a nominal genus, an author refers the generic name to an
author or date other than those denoting the first establishment of the
genus or the first express reference of nominal species to it, he is neverthe¬
less to be considered, if the species was eligible, to have designated the
type-species correctly.
Example. A-us Dupont, 1790, established without a designated or indicated
type-species, is best known from the work of a later author, Smith, 1810. If sub¬
sequently b-us is designated as the type-species of “A-us Smith, 1810,” that desig¬
nation is to be accepted as valid for A-us Dupont, 1790, if b-us was eligible for
designation as type-species of the latter.
(A) Exclusions. A nominal species that was not included, or that was cited
as a species inquirenda or species incertae sedis when a new nominal genus
was established, cannot be validly designated or indicated as the type-
species of that genus.
(i) Replacement names. If a zoologist proposes a new generic name ex¬
pressly as a replacement for a prior name, both nominal genera must have 50
the same type-species, and, subject to (i) below, type-fixation for either
applies also to the other, despite any statement to the contrary.
Example. B-us Schmidt, 1890, is proposed expressly as a replacement name
for a junior homonym, A-us Medina, 1880, non Dupont, 1860. If x-ar is the desig¬
nated type-species of A-us, it is ipso facto the type-species of B-us.
(i) The type-species must be a species eligible for fixation as the type
of the earlier nominal genus.
(ii) An emendation of a generic name, whether justified or unjustified,
is an objective synonym of the original name and therefore has the
same type-species.
(j) Misidentified type-species. If a designated or indicated type-species
is later found to have been misidentified, the provisions of Art. 70 apply. 49
(A) Union of genera. The union of two or more nominal genera to form a
single taxonomic genus does not change the type-species of any nominal 25
genus involved [XIII], and the type-species of the combined genus is that
of the senior component nominal genus.

Article 68. Type-species fixed in the original publication. The pro¬

visions of this Article apply in the following order of precedence.

(a) Original designation. If one nominal species is definitely designated

as the type-species of a new nominal genus when the latter is established,
324 Principles and Application of Zoological Nomenclature

that species is the type-species, regardless of any other consideration (type

by original designation).
(i) The formula “gen.n., sp.n.,” or its exact equivalent, applied before
1931 to only one of the new nominal species included in a newly
established nominal genus, is to be interpreted as original designa¬
tion if no other type-species was designated.
(b) Use of typicus or typus. If, when a new nominal genus is established,
one of the included new species is named typicus or typus, that species is
the type-species.
(c) Monotypy. A genus originally established with a single nominal species
takes that species as its type, regardless of whether the author considered
the genus to contain other species that he did not name, and regardless of
cited synonyms, subspecies, unavailable names, and species that are doubt¬
fully included or identified (type by monotypy).
(d) Tautonymy. If a newly established nominal genus contains among its
originally included nominal species one possessing the generic name as its
specific or subspecific name, either as the valid name or as a cited syno¬
nym, that nominal species is ipso facto the type-species (type by absolute
tautonymy).
(i) If, in the synonymy of only one of the species originally included in
a nominal genus established before 1931, there is cited a pre-1758
name of one word identical with the new generic name, that nominal
species is construed to be the type-species (type by Linnean tau¬
tonymy).

Article 69. Type-species not fixed in the original publication. The

provisions of this Article apply in the following order of precedence only to
nominal genera that were established before 1931 without an originally
designated or indicated type-species.

(a) Subsequent designation. If an author established a nominal genus but

did not designate or indicate its type-species, any zoologist may subse- 4
quently designate as the type-species one of the originally included nomi- 56
nal species, or, if there were no original nominal species, one of those first
subsequently referred to the genus (type by subsequent designation).
(i) In the meaning of this provision, the “originally included species”
comprise only those actually cited by name in the newly estab¬
lished nominal genus, either as valid names (including subspecies,
varieties, and forms), as synonyms, or as stated misidentifications
of previously established species [Art. 70(6)].
(ii) If no nominal species were included at the time the genus was
established, the nominal species-group taxa that were first subse- 56
quently and expressly referred to it are to be treated as the only
originally included species.
(1) Mere reference to a publication containing the names of
species does not by itself constitute the inclusion of species
in a nominal genus.
(2) If only one nominal species was first subsequently referred
to a genus, it is ipso facto the type-species, by subsequent
monotypy.
(3) If two or more nominal species were simultaneously referred
to a nominal genus, all are equally eligible for subsequent
type-designation.
 The Rules of Zoological Nomenclature 325

(iii) In the absence of a prior valid type-designation for a nominal

genus, an author is considered to have designated one of the origi¬
nally included nominal species as type-species, if he states that it is
the type (or type-species), for whatever reason, right or wrong, and
if it is clear that he himself accepts it as the type-species.
(iv) If an author designates (or accepts another’s designation) as type-
species a nominal species that was not originally included, and if,
but only if, at the same time he synonymizes that species with one
of the originally included species, his act constitutes designation of
the latter as type-species of the genus.
(v) A nominal species is not rendered ineligible for designation as a
f type-species by reason of being the type-species of another genus.
(vi) A subsequent designation first made in a literature-recording pub¬
lication is acceptable, if valid in all other respects.
Recommendation 69A. Preference for figured species. In designating a
type-species for a genus, a zoologist should give preference to a species that is
adequately figured.
Recommendation 69B. Other considerations in designating type-species.
In the subsequent designation of a type-species a zoologist should guide himself
by the following precepts, listed in order of precedence:
(1) In the case of Linnean genera he should designate the most common
species or one of medical importance (Linnean aphorism 246, Critica
Botanica, 1737).
(2) If the name or a synonym of one of the originally included nominal species
is virtually the same as the generic name, or is of the same origin or mean¬
ing, that species should be designated as the type-species (type by virtual
tautonymy), unless such designation is strongly contra-indicated by
other factors.
Examples. Bos taurus, Equus caballus, Ovies aries, Scomber scombrus, Sphaerostoma
globiporum; contra-indicated in Dipetalonema dipetalum because only one sex was
described, based on a single specimen not studied in detail.
(3) If some of the originally included nominal species have been removed to
other genera, preference should be given to a remaining species, if any such are
suitable (choice by elimination).
(4) A species based on a sexually mature specimen is generally preferable to
one based on a larval or otherwise immature specimen.
(5) Preference should be given to a species named communis, vulgaris, medi-
cinalis, or officinalis.
(6) Preference should be given to the best described, best figured, best known,
or most easily obtainable species, or to one of which a type-specimen is accessible.
(7) If more than one group of species is recognized in a genus, preference
should be given to a species that belongs to as large a group of species as possible
(de Candolle’s rule).
(8) In genera of parasites, preference should be given to a species that para¬
sitizes man, or an animal of economic importance, or a common and widespread
host-species.
(9) All other things being equal, preference should be given to a species well
known to the author of the nominal genus, prior to publishing the generic name.
(10) If an author habitually placed a leading or typical species first as “chef
de file,” and described others by comparison with it, that fact should be con¬
sidered in the designation of a type-species.
(11) If an author is known to have denoted type-species by their position
326 Principles and Application of Zoological Nomenclature

(“first species rule”), the first nominal species cited by him should be designated
as the type-species.
(12) All other things being equal, preference should be given to the species
cited first in the work, page, or line (position precedence).
Recommendation 69C. Citation of type-species. When designating a type-
species for a nominal genus established before 1931, a zoologist should cite the
name of that species first in its original binomen and then in its current binomen,
if this is different. He should give a bibliographic reference to the work where the
species was established.

Article 70. Identification of the type-species. It is to be assumed that

an author correctly identifies the nominal species that he either (1) refers to
a new genus when he establishes it, or (2) designates as the type-species of a
new or of an established genus.

(a) Misidentified type-species. If a zoologist considers that such a species

was misidentified, he is to refer the case to the Commission to designate as
the type-species (by use of its plenary powers if necessary [Art. 79]) which¬
ever species will in its judgment best serve stability and uniformity of
nomenclature, either
(i) the nominal species actually involved, which was wrongly named
in the type-designation; or
(ii) if the identity of that species is doubtful, a species chosen in con¬
formity with the usage of the generic name prevailing at the time
the misidentification is discovered; or
(iii) the species named by the designator, regardless of the misidentifi¬
cation.
(b) Deliberate use of misidentification. If the type designated for a new
nominal genus is a previously established species, but the designator states
that he employs its specific name in accordance with the wrong usage of a
previous author, the type-species is to be interpreted as the one actually
before the designator, not the one that correctly bears the name.
(i) In such a case, the author of the new nominal genus is considered
to have established also a new nominal species, with the same specific
name as the misidentified species, in the new nominal genus.
Example. If Jones, 1900, designated as type-species of C-us, gen. n., a species
that he cites in some such manner as A-us b-us Dupont sensu Schmidt, 1870, the
type-species of C-us is that which was before Jones, not that named by Dupont,
and its name is to be cited as C-us b-us Jones, 1900.

XVI. TYPES IN THE SPECIES-GROUP

Article 71. Application. The provisions of this Chapter apply equally to

all categories in the species-group.
Article 72. General provisions.

(a) Types of species-group taxa. The type of each taxon of the species-
group is a single specimen, either the only original specimen or one desig¬
nated from the type-series (holotype, lectotype), or a neotype [Art. 45A].
(A) Type-series. The type-series of a species consists of all the specimens on
which its author bases the species, except any that he refers to as variants,
 The Rules of Zoological Nomenclature

or doubtfully associates with the nominal species, or expressly excludes

from it.
(c) Specimens that are already types. The fact that a specimen is already
the type of one nominal species does not prevent its designation as the
type of another.
(</) Types of replacement nominal species. If an author proposes a new
specific name expressly as a replacement for a prior name, but at the same
time applies it to particular specimens, the type of the replacement nomi¬
nal species must be that of the prior nominal species, despite any contrary
designation of type-specimen or different taxonomic usage of the replace¬
ment name.
(e) Types of nominate subspecies. See Art. 61(a).
(/) Value of types. Holotypes, syntypes, lectotypes, and neotypes are to be
regarded as the property of science by all zoologists and by persons re¬
sponsible for their safe-keeping.
Recommendation 72A. Institutional custody. A zoologist who designates a
holotype or lectotype should deposit it in a museum or other institution where it
will be safely preserved and will be accessible for purposes of research. Deposit
of neotypes in a museum or other institution is mandatory [Art. 75c(6)].
Recommendation 72B. Labelling. A zoologist designating a holotype, lecto¬
type, or neotype should unmistakably label the specimen in a way that will
clearly indicate its status.
Recommendation 72C. Information on labels. When designating a holo¬
type, lectotype, or neotype, a zoologist should publish all information that ap¬
pears on the labels accompanying the specimen, so as to ensure the future recog¬
nition of the specimen.
Recommendation 72D. Institutional responsibility. Every institution in
which types are deposited should
(1) ensure that all are clearly marked so that they will be unmistakably
recognized;
(2) take all necessary steps for their safe preservation;
(3) make them accessible for study;
(4) publish lists of type-material in its possession or custody; and
(5) so far as possible communicate information concerning types when re¬
quested by zoologists.
Recommendation 72E. Type-localities. An author who either designates or
restricts a type-locality should base his action on one or more of the following
criteria:
(1) the original description of the taxon;
(2) data accompanying the original material;
(3) collector’s notes, itineraries, or personal communications; and
(4) as a last resort, localities within the known range of the species or from
which specimens identified with the species have been taken.
If a type-locality was erroneously designated or restricted, it should be corrected.

Article 73. Holotypes and syntypes.

(a) Single specimen. If a nominal species is based on a single specimen,

that specimen is the “holotype.”
(6) Specified type. If an author states in the description of a new nominal
species that one specimen and only one is “the type” or uses some equiva¬
lent expression, that specimen is the holotype.
(c) Syntypes. If a new nominal species has no holotype under the provision
328 Principles and Application of Zoological Nomenclature

of (a) and (b), all the specimens of the type-series are “syntypes,” of equal
value in nomenclature.
(i) Syntypes may include specimens labelled “cotype” (in the meaning
of syntype), “type,” or by some other term, or with no identifying
label, or specimens not seen by the author but which were the
bases of previously published descriptions or figures upon which he
founded his taxon in whole or in part.
Recommendation 73A. Original designation. A zoologist when describing
a new species should clearly designate a single specimen as its holotype.
Recommendation 73B. Procedure. If a zoologist, in basing a new nominal
species on specimens before him, subjectively associates with it specimens that he
believes to have been misidentified by another author, he should designate his
holotype from the former.
Recommendation 73C. Data on the holotype. A zoologist in establishing a
new species should publish at least the following data concerning its holotype, in 52
so far as they are relevant and known to him:
(1) the size;
(2) the full locality, date, and other data on the labels accompanying the
holotype;
(3) the sex, if the sexes are separate;
(4) the developmental stage, and the caste, if the species includes more than
one caste;
(5) the name of the host species;
(6) the name of the collector;
(7) the collection in which it is situated and any collection- or register number
assigned to it;
(8) in the case of a living terrestrial species the elevation in metres above sea-
level at which it was taken;
(9) in the case of a living marine species the depth in metres below sea-level
at which it was taken;
(10) in the case of a fossil species, its geological age and stratigraphical posi¬
tion, stated if possible, in metres above or below a well-established plane.
Recommendation 73D. Paratypes. After the holotype has been labelled,
each remaining specimen (if any) of the type-series should be conspicuously
labelled “paratype,” in order clearly to identify the components of the original
type-series.
Recommendation 73E. Avoidance of “cotype.” To avoid misunderstand¬
ings, a zoologist should not use the term “cotype.”

Article 74. Lectotypes.

(a) Designation of a specimen. If a nominal species has no holotype, any 4

zoologist may designate one of the syntypes as the “lectotype.” 54
(i) The first published designation of a lectotype fixes the status of the 56
specimen, but if it is proved that the designated specimen is not a 57
syntype, the designation is invalid.
(b) Designation by means of a figure. Designation of a figure as lectotype
is to be treated as designation of the specimen represented by the figure; if
that specimen is one of the syntypes, the designation as lectotype is valid
from the nomenclatural standpoint.
(c) Designation to be individual. Lectotypes must not be designated col¬
lectively by a general statement; each designation must be made specifi¬
cally for an individual nominal species, and must have as its object the
definition of that species.
 The Rules of Zoological Nomenclature 329

Example. A published statement that, in the type-series of all species described

by a particular author, the specimen bearing the author’s determination label,
or the only surviving syntype, is to be treated as the lectotype, is not a valid
designation of lectotypes.
Recommendation 74A. Agreement with previous restriction. In desig¬
nating a lectotype, a zoologist should in general act consistently with, and in any
event should give great weight to, previous valid restrictions of the taxonomic
species, in order to preserve stability of nomenclature.
Recommendation 74B. Figured specimen. A zoologist should choose as
lectotype a syntype of which a figure has been published, if such exists.
Recommendation 74C. Data on the lectotype. A zoologist who designates a
lectotype should publish the data listed in Recommendation 73C, besides de¬
scribing any individual characteristics by which it can be recognized.
Recommendation 74D. Syntypes in several collections. When possible, a
lectotype should be chosen from syntypes in the collection of a public institution,
preferably of the institution containing the largest number of syntypes of the
species, or containing the collection upon which the author of the nominal species
worked, or containing the majority of his types.
Recommendation 74E. Paralectotypes. A zoologist who designates a lecto¬
type should clearly label any remaining syntypes with the designation “para-
lectotype.”

Article 75. Neotypes. Subject to the following limitations and condi¬ 55

tions, a zoologist may designate another specimen to serve as the “neotype”
of a species if, through loss or destruction, no holotype, lectotype, or syntype
exists.

(a) Cases admitted. A neotype is to be designated only in connection with

revisory work, and then only in exceptional circumstances when a neotype
is necessary in the interests of stability of nomenclature.
(i) The words “exceptional circumstances” refer to those cases in which
a neotype is essential for solving a complex zoological problem, such
as the confused or doubtful identities of closely similar species for one
or more of which no holotype, lectotype, or syntype exists.
(b) Cases excluded. A neotype is not to be designated for its own sake, or
as a matter of curatorial routine, or for a species of which the name is not in
general use either as a valid name or as a synonym.
(c) Qualifying conditions. A neotype is validly designated only when it is
published with the following particulars:
(1) a statement of the characters that the author regards as differen¬
tiating the taxon for which the neotype is designated, or a biblio¬
graphic reference to such a statement;
(2) data and description sufficient to ensure recognition of the specimen
designated:
(3) the author’s reasons for believing all of the original type-material to
be lost or destroyed, and the steps that have been taken to trace it;
(4) evidence that the neotype is consistent with what is known of the
original type-material, from its description and from other sources:
however, if a nominal species is based on a sex or an immature stage
that lacks good diagnostic characters, the neotype may differ in
that respect from the original material;
(5) evidence that the neotype came as nearly as practicable from the
330 Principles and Application of Zoological Nomenclature

original type-locality, and where relevant, from the same geological

horizon or host-species as the original type-material;
((>) a statement that the neotype is, or immediately upon publication
has become, the property of a recognized scientific or educational
institution, cited by name, that maintains a research collection, with
proper facilitites for preserving types, and that makes them acces¬
sible for study.
(il) Priority. The lust neotype-designation published for a given nominal
specie* in accordance with the provisions of this Article is valid, and any
subsequent designation has no validity unless the first ncotype is lost or
destroyed.
Recommendation 7HA. Consultation with specialists, before designating a
neotype, a zoologist should satisfy himself that his proposed designation does not
arouse objections from other specialists in the group in question.
(r) Status of previously designated neotypes. A neotype-designation pub¬
lished before 1961 takes effect from the time when it fulfills all the pro¬
visions of this Article.
Recommendation 7.r>R. Validation. A zoologist who published an invalid
neotype-designation before 1961 should be given an opportunity to validate it
before another zoologist designates a neotype for the same nominal taxon.
Recommendation 75C. Preference for earlier neotypes. If an invalid nco¬
typedesignation was published before I 961, the specimen then designated should
b<- given preference when a neotype for the same nominal taxon is validly
designated.
(/) Status of rediscovered type-material. If, after the designation of a
neotype, original type-material is found to exist, the case is to be referred to
flic (lommission.

XVII. THE INTERNATIONAL COMMISSION ON

ZOOLOGICAL NOMENCLATURE

Article 7(>. Status. The International Commission on Zoological Nomen¬ 59

clature is a permanent body which derives all its power from the Inter¬
national Congresses of Zoology.
Article 77. Duties. The Commission is charged with the following
tluties:

(1) to consider for a period of at least one year in advance of a Congress (or
for such less time as the Commission may agree) any proposal for a change
in the (lode;
(2) to submit to the Congress recommendations for the clarification or modi¬
fication of the Code;
(3) to render between successive Congresses Declarations (i.e. provisional
amendments to the Code) embodying such recommendations;
(4) to render ()pinions and Directions on questions of zoological nomenclature
that do not involve changes in the Code;
(5) to compile the Official Lists of accepted, and the Official Indexes of re¬
jected, names and works in zoology; 63
(ft) to submit reports to the Congresses on its work; and
(7) to discharge such other duties as the Congresses may determine.
 The Rules of Zoological Nomenclature

Article 78. Exercise of powers. The Commission has the power, when an
application is referred to it by any zoologist, to interpret the provisions of
the Code and to apply such interpretation to any question of zoological
nomenclature.

(a) Declarations. If a case before the Commission involves a situation that

is not properly or completely covered by the Code, the Commission is to
issue a Declaration (a provisional amendment to the Code) and to propose
to the next succeeding Congress adoption of this amendment in the manner
prescribed in Article 87.
(i) A Declaration remains in force until the next succeeding Congress
ratifies it, either in its original form or in a modified form (where¬
upon the Code is accordingly amended), or rejects it; the Declara¬
tion is thereupon deemed repealed for all except historical purposes.
(b) Opinions. If the case in question involves the application of the Code
to a particular situation relating to an individual name, act, or publica¬
tion, the Commission is to render a decision, termed an Opinion, and
either
(i) to state how the Code is to be applied or interpreted; or
(ii) acting in the interest of stability and universality, to exempt, under
its plenary powers [Art. 79], the particular case from the application
of the Code, and to state the course to be followed.
(c) Effective date of Opinions. Opinions have force immediately upon
publication of the ruling of the Commission, and are to be reported to the
next succeeding Congress.
(</) Directions. Decisions completing earlier rulings, and formal instruments
required under automatic provisions of the Code, are called Directions.
They have the same status as Opinions.
(e) Construction. All decisions are to be rigidly construed and no con¬
clusions other than those expressly specified are to be drawn from them.
(/) Official Lists and Indexes. Names and works that are accepted or re¬
jected in Opinions are to be entered on the relevant Official Lists or In¬
dexes, whereupon the Opinions concerned are deemed repealed for all
except historical purposes.
(g) Review by Congress of Commission’s decisions. A motion to modify
or reject any decision of the Commission is not to be considered by a Con¬
gress until notice of at least one year (or such less period as the Commission
may agree) has been given to the Commission.

Article 79. Plenary powers. The Commission is empowered to suspend,

on due notice as prescribed by its Constitution, the application of any
provisions of the Code except those in the present and the next succeeding
Chapter, if such application to a particular case would in its judgment
disturb stability or universality or cause confusion. For the purpose of pre¬
venting such disturbance and of promoting a stable and universally accepted
nomenclature, it may, under these plenary powers, annul or validate any
name, type-designation, or other published nomenclatural act, or any publi¬
cation, and validate or establish replacements.

(a) Guiding principles. In exercising its plenary powers, the Commission

is to be guided as follows:
(i) a name suppressed so as to validate the use of the same name pub-
332 Principles and Application of Zoological Nomenclature

lished at a later date in another sense, is to be suppressed for the

purposes both of the Law of Priority and of the Law of Homonymy;
(ii) a name suppressed so as to validate a later name given to the same
taxon is to be suppressed for the purposes of the Law of Priority but
not for those of the Law of Homonymy;
(iii) if the Commission refuses to use its plenary powers in a given case,
the Opinion rendered is to specify the name(s) to be used in the
case in question, and the action (if any) to be taken.

Article 80. Status of case under consideration. When a case is under 61

consideration by the Commission, existing usage is to be maintained until
the decision of the Commission is published.
Article 81. Exemption. The Commission is under no obligation to search
out violations of the Code, or to supplement or verify information contained
in applications submitted to it, or to initiate any action within its field of
competence, although it may, at its discretion, do any of these things.
Article 82. Constitution and Bylaws. Regulations dealing with the 59
membership of the Commission, its governing board, elections, voting pro¬
cedures, meetings, and related matters are incorporated in the Constitution
and Bylaws of the Commission.

(a) Amendments to the Constitution. Changes in the Constitution can be

made only by the Congresses, on recommendation by the Commission, in
the same manner as amendments to the Code [Art. 87],
(b) Amendments to the Bylaws. Changes in the Bylaws can be made by the
Commission under procedures set forth in its Constitution.

XVIII. REGULATIONS GOVERNING THIS CODE

Article 83. Title. The title of these rules and recommendations is: “IN¬
TERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE
ADOPTED BY THE XV INTERNATIONAL CONGRESS OF ZOOL¬
OGY, LONDON, JULY 1958.”
Article 84. Effective date. This Code comes into force on the day of its
publication, and all previous editions of the International Rules of Zoologi¬
cal Nomenclature are thereby superseded.

(a) Previous decisions affecting the Code. All amendments affecting the
Code, adopted by the Congresses prior to the XV Congress, are no longer
valid unless reaffirmed herein, and then only as here expressed.

Article 85. Language of the official texts. The official French and
English texts of the Code are equivalent in force, meaning, and authority.
If it appears that there is a difference in meaning between the two texts, the
problem is to be referred to the Commission for decision, and its interpreta¬
tion is final.
Article 86. Application. The provisions of the Code apply to all zoologi¬
cal names and works, published after 1757, that affect zoological nomen¬
clature.
 The Rules of Zoological Nomenclature 333

(a) Previous decisions of the Commission. No decision taken by the Com¬

mission in relation to a particular name or work, prior to the effective date
of this Code, is to be set aside without the consent of the Commission.

Article 87. Amendment to the Code. Amendment to this Code can be

made only by an International Congress of Zoology, acting on a recom¬ 62
mendation from the Commission presented through and approved by the
Section on Nomenclature of that Congress [Arts. 77 and 78a].
Chapter 1 'y Interpretation of the Rules
of Nomenclature

any code of law, the International Code

presents the rules starkly and without explanation. Even the experienced
specialist is sometimes in doubt how to interpret the rules, and the beginner
is often totally bewildered. The difficulty of understanding individual
provisions is aggravated by the unavailability of a well-articulated presenta¬
tion of the basic principles, which the individual rules attempt to imple¬
ment. To fill this void is one of the aims of this chapter. It is hoped that
the 66 numbered comments presented here will help the taxonomist when
applying the rules to his particular problems. Over and above this purely
practical objective, I hope that these comments will help the taxonomist
to acquire a better understanding of the entire field of nomenclature. Stress
throughout will be on principles rather than on practices. These comments
are numbered consecutively, and the numbers refer to the equivalent
numbers in the margin of the Code in Chap. 12. MLU refers to Mayr,
Linsley, and Usinger, 1953, Methods and Principles of Systematic Zoology;
BZN to the Bulletin of Zoological Nomenclature; SZ to the journal Syste¬
matic Zoology.
Where the application of the rules to a specific situation is involved,
consultation of the Index of the Code is often very helpful.

334
 Interpretation of the Rules of Nomenclature 335

Table of Topics Discussed in Chapter 13

1. Stability 35. Coordinate status of categories

2. Freedom of taxonomic thought 36. Subordinate taxa
3. Priority 37. Homonymy between family
4. First-reviser principle group names
5. Range of authority 38. Genus group names
6. Application of names 39. Collective groups
7. Recommendations 40. Names of taxa in the species
8. Important dates group
9. Retroactivity 41. Infrasubspecific names
10. Vernacular nomenclature 42. Authorship
11. Scientific nomenclature 43. Homonymy
12. Advantages and disadvantages 44. Actuality principle
of binominal nomenclature 45. Replacement name (first re¬
13. Name changing viser)
14. Alternate systems of nomen¬ 46. Homonymy among subspecies
clature 47. Replacement of rejected
15. Criteria of publication homonyms
16. Availability 48. The type method
17. Kinds of names 49. Correcting the misidentification
18. Publication in synonymy of types
19. Differential diagnosis 50. Type of replacement name
20. Indication 51. Kinds of type-specimens
21. Names given to hybrids 52. Type designation
22. Date of publication 53. Type-material
23. Validity 54. Lectotypes
24. Statute of limitation 55. Neotypes
25. Combined and divided taxa 56. Types and first revisers
26. Page and line precedence 57. Type-localities
27. Work of an animal 58. Nominal taxon
28. Formation of names 59. International Commission
29. Diacritic marks 60. Applications to the Commission
30. The stem of a family name 61. Cases under consideration
31. Gender of generic names 62. Amendments to the Code
32. Original spelling and emenda¬ 63. Official Lists and Indexes
tion 64. Appendices
33. Family names 65. General recommendations
34. Names of higher taxa 66. Glossary and Index

1. STABILITY

The statement in the Preamble of the Code that “the object of the
Code is to promote stability” is perhaps the most important provision in
the entire rules of nomenclature. The International Congress of Monaco
in 1913 first accepted the principle, since frequently reaffirmed, that “when
336 Principles and Application of Zoological Nomenclature

stability of nomenclature is threatened in an individual case, the strict

application of the Code may under specified conditions be suspended by
the International Commission on Zoological Nomenclature” (Preamble).
In such cases any zoologist has the right to appeal to the Commission
and ask for the protection of the threatened name.
More specific stability-promoting provisions in the Code can be found
in Arts. 23b (statute of limitation), 23<^(ii), 40, 40a (family names),
41 (misidentified genera), 70a (misidentified type-species), and 79 (plenary
powers, for all actions on behalf of stability, including the designation
of neotypes). The wise application of these provisions has greatly con¬
tributed to the stabilization of zoological nomenclature and to the refuta¬
tion of the claim that taxonomists indulge in reckless and irresponsible
name-changing.
Even if all nomenclatural name changes were eliminated (see 13.3),
and even if the binominal system were replaced (see 13.14), certain name
changes are inevitable. These include cases where:

1. Several species (e.g. sibling species) had been referred to by a single

name prior to a more penetrating analysis
2. Several intraspecific phena were named in the belief that they represented
different species
3. Several authors (especially in different countries) had named the same
taxon, unbeknown to each other

Since the names of animals are part of a system of communication, total

stability is the obvious ideal. Inevitable shortcomings in taxonomic analysis
will make it impossible to achieve this goal at the present time, but every
effort should be made to minimize interference with the stated ideal.
I shall give only a single example (supplied by Corliss, BZN,
24:155-185) for the need for stability in the biological literature. In a
period of 27 years about 1,500 papers were published on the physiology,
etc., of the protozoon pyriforme under the generic name Tetrahymena
Furgason 1940. To replace this generic name for the sake of priority would
obviously be a calamity as far as information retrieval is concerned. It
was therefore quite legitimate for Corliss to ask for the suppression of
ten senior synonyms of Tetrahymena that were totally or virtually unused
in the vast biological literature of this genus.

2. FREEDOM OF TAXONOMIC THOUGHT

It is axiomatic that the freedom to make taxonomic decisions should

not be restricted by rules of nomenclature. This is now expressly guaranteed
in the Preamble, which states that none of the provisions of the Code
 Interpretation of the Rules of Nomenclature 337

“restricts the freedom of taxonomic thought or action.” This basic principle

is applied to specific situations throughout the Code. In Art. lid it ac¬
knowledges that the subjective action of an author in placing a name
in synonymy should not determine the nomenclature to be used by others;
in Art. 59b it permits a zoologist to continue using a name considered
by some other author to be a junior homonym; in Art. 64 it authorizes
the zoologist to choose (as the type of a new family) that genus which
he considers the most typical or characteristic genus of that family but
“not necessarily that bearing the oldest name.” The same is true for type
designation on the generic level (Arts. 67-69).
Conversely, taxonomic mistakes should not affect nomenclature except
where this is expressly authorized by the Code. For instance, the erroneous
assignment of a species to a genus does not constitute the proposal of
a new specific name in that genus (Art. 49) ; the naming of individual
variants that do not form a taxon has no validity in nomenclature (Art.
45c) ; the erroneous placing of a name in synonymy does not invalidate
it [Art. 17 (i) ].
It is possible that the principle of freedom of taxonomic thought
should be extended to some additional articles. For instance, the provisions
of Art. 64 are not applied to the choice of a type-genus of a new superfamily
which has been created by a taxonomist by uniting a number of previously
described families into an entirely new taxon (see 13.35).

3. PRINCIPLE OF PRIORITY

The most controversial problem in nomenclature arises whenever two

names have been given to the same taxon. Fairness and common sense
would seem to dictate a simple solution: let the prior name prevail. As
soon as the oldest name for each taxon has been found, it is often asserted,
complete stability of nomenclature will have been established automatically.
This seemingly persuasive argument has, however, created great difficulties
in many areas of systematic zoology and has been vigorously opposed by
distinguished zoologists. In order to show why there is a controversy, the
arguments pro and con will be presented in detail.
Linnaeus (1753) endorsed priority in principle. In Aphorism 243
he said, “If a generic name is suitable, it is not allowable to change it,
even for another which is more fitting.” In practice, however, he was an
inveterate name-changer, perhaps because he considered many names not
“suitable.” For this he was upbraided by his friends. For instance, Peter
Collinson wrote him in 1754, “But my dear friend we that admire you
are much concerned that you should perplex the delightful science of botany
with changing names that have been well-received.” Even in his own works
338 Principles and Application of Zoological Nomenclature

he shifted the names of taxa from one edition to the next, and indeed
the rule of the first reviser (13.4) seems to have been more important
to Linnaeus and his followers than the rule of priority. Fortunately, the
last (12th) Linnaean edition (1766-67) of the Systema Naturae was ac¬
cepted by his successors as the standard of reference for the next 50 or
more years and provided for great stability during the ensuing period.
Some of his successors, such as Fabricius among the entomologists, exerted
a similar authority, ensuring great stability. As this authority weakened
and as more and more zoologists entered the field of taxonomy, it occurred
increasingly often that one and the same animal was given several names.
During the French Revolution and the Napoleonic wars communication
among scientists was difficult, and taxonomists in one country were often
unaware of the new' species and genera described by taxonomists in other
countries. Equally or even more annoying was the rise of a generation
of pedants who changed names the formation of which they considered
not classically correct or not precisely descriptive. When a bird called capen-
sis (in the belief that it came from the Cape of Good Hope), was found
to have come from Java, it was renamed javensis. If a blue-green bird
was called uiridis but a subsequent author considered it more blue than
green, he would rename it caeruleus. The result was nomenclatural confu¬
sion, if not anarchy.
Eventually the situation became so critical that the British Association
for the Advancement of Science appointed a committee to draw up a
general set of rules for zoological nomenclature. The resulting code (Strick¬
land, 1842), often referred to as the “Strickland Code,” formed the basis
of all future codes and was the beginning of stabilization. Priority was
considered to be the best method of achieving stability, and the changing
or replacing of an earlier name merely because it was incorrectly formed
or misleading, or for other personal, aesthetic, or even scientific reasons,
was outlawed. It is evident from much of the contemporary literature on
the subject that “priority,” as conceived by these authors, was a priority
of usage rather than a priority of publication. Yet priority of usage is
subjective, and in the subsequent codification of the rules it was replaced
by priority of publication in the endeavor to achieve objectivity. Unfortu¬
nately, while gaining objectivity, the nomenclaturists abandoned one of
the most important objects of nomenclature, stability. As a result of the
upheaval caused by a strict application of the principle of priority of publi¬
cation, zoologists soon began to rebel against “priority.” As early as 1849
Darwin wrote to Strickland with regard to cirriped nomenclature, “I believe
if I were to follow the strict rule of priority more harm would be done
than good. . . .” And this conviction was shared by an increasing number
of zoologists. MLU (pp. 215-216) presents data on the percentage of name
changes resulting from priority in various taxonomic groups.
 Interpretation of the Rules of Nomenclature 339

In spite of serious objections, when the first International Code was

adopted, it incorporated “strict priority” as a basic law. This started an
extraordinary period of searching through the earliest post-Linnaean zoologi¬
cal publications and resulted in the discovery of countless totally forgotten
names. Worse than that, it placed an undeserved emphasis on names that
had been deliberately ignored by contemporaries because the descriptions
were too poor for identification, contradictory, or misleading. It led to
a search for still existing specimens (often erroneously called “types”) of
these early authors and deflected taxonomists from biological research into
bibliographic archeology. Four types of discoveries were often made as
the result of these searches: earlier synonyms, earlier homonyms, earlier
fixations of generic types, and misidentified type specimens. None of these
discoveries advanced our knowledge of animals or their classification. As
Michener (1963, SZ, 12:163) has rightly said: “In other sciences the work
of incompetents is merely ignored, in taxonomy, because of priority, it
is preserved.”
A rebellion against the unrestricted application of priority soon arose.
An opinion poll among Scandinavian zoologists in 1911 showed that only
2 were in favor of a strict interpretation of the rule of priority while 120
were against it. Zoologists in other countries likewise voted in favor of
limitations to priority, and the plenary powers decision (Art. 79) was
adopted at Monaco in 1913.
Many zoologists felt that this still did not go far enough in protecting
well-established, familiar names. It was pointed out that priority is a means
to an end, the end being stability. Where the end (stability) is in conflict
with the means (priority), it is the end which should have primacy. The
feeling of exasperation with the strict application of priority is well expressed
by a statement of the Chicago group of taxonomists (Schmidt, 1950) :

The whole business of elaborate argument over rule and “validity” of

names has been a disgrace to zoology and has contributed more than any
other single factor to the low repute of systematics among zoologists as
a whole. Any system seeking continuity rather than change would have
been infinitely preferable to the elaborate search for priority that was estab¬
lished as an international game. . . . The fundamental requirement is a
reformation in attitude. Strict following of the law of priority, regardless
of the consequences, must be set aside as the guiding principle in nomen-
clatural procedure, both in the thinking of the systematist and in the working
of the Commission. In its stead, there must be substituted a proper regard
for the convenience of zoologists generally and a determination, by stabilizing
names and current use, to avoid change and chaos.

The succeeding international congresses (Copenhagen and London)

paid attention to these sentiments and adopted the important statement
in the Preamble that even though “priority is the basic principle of zoologi-
340 Principles and Application of Zoological Nomenclature

cal nomenclature, its application however, under conditions specified in

the Code, may be moderated to preserve a long accepted name in its accus¬
tomed meaning.” Article 23 deals specifically with priority and its
exceptions.
“When stability of nomenclature is threatened in an individual case,
the strict application of the Code may under specified conditions be sus¬
pended by the International Commission on Zoological Nomenclature.”
Of particular importance are the limitations to priority specified in Arts.
23b, 23c?(ii), 41, and 79.

4. FIRST-REVISER PRINCIPLE

A basic Linnaean principle which was widely followed by his disciples

was that the action of the first reviser should be adopted in all equivocal
nomenclatural situations. This principle, although sanctioned by usage, was
almost entirely ignored in the drafting of the 1901 Code. It has been incor¬
porated, however, in numerous provisions of the 1961 Code.
Linnaeus himself was an inveterate first reviser. The reviser principle
to him was quite clearly more important than the principle of priority.
In later publications he again and again changed what he had done in
his own earlier publications, and his students and followers invariably ac¬
cepted his last action rather than that which had priority. This is why
the generations following Linnaeus accepted the 12th edition (1766) of
the Systema Naturae as the standard of reference rather than the 10th
edition (1758). This is why, for at least the first 100 years after Linnaeus,
the type-species of a genus was determined by the method of elimination
rather than that of fixation. It was the almost complete disregard of the
first-reviser principle in the original Code of Zoological Nomenclature
(1901) that was responsible for much of the name changing in the first
half of the century.
There are numerous ways by which a first reviser can help to stabilize
nomenclature. In the case of simultaneously published names (Art. 24a
and Recommendation 24a) he can select the name which is better known
rather than that which has line or page precedence (which is not priority!).
If a new name is spelled in more than one way in the original publication,
Art. 2>2b permits the first reviser to accept that which is most commonly
used. If no type-species is fixed in the publication in which a new genus
was described prior to 1931, the first reviser can select the species whose
designation is in the best interest of stability (Art. 69). In the case of
a composite name, based on a series of syntypes consisting of several species,
the first reviser can help stability by making either of two choices (Art.
74a). He can designate as lectotype a specimen of the species to which
 Interpretation of the Rules of Nomenclature 341

the name has always been applied in the past, or if it is a name that
is best suppressed, for the sake of stability, he can designate a lectotype
belonging to a species for which a senior synonym is available. (See 13.56
for all first-reviser actions concerning types.) Finally, the first-reviser prin¬
ciple is important in the designation and restriction of type-localities (see
13.57).

5. RANGE OF AUTHORITY OF THE CODE

The Code applies to both living and extinct animals (Art. 1). If
a still-living species was first named on the basis of fossil material, that
name is also valid for the living species. If a generic name has been used
for a fossil animal, it cannot be used again for a different genus of living
animals and vice versa. Zoological nomenclators therefore contain the names
given to both living and extinct animals so that the possible occurrence
of homonymy can be determined.
There are, however, separate codes for botanical and bacteriological
nomenclature, and Art. 2 of the Zoological Code spells out the relation
between the codes and the status of names transferred between the king¬
doms. Familiarity with the botanical rules is important for authors working
with protists.

6. APPLICATION OF NAMES

Names are given only to taxa, and all taxa are populations or sets
of populations. Consequently only populations are named. Names are given
to individuals only as representatives of populations. Names given to indi¬
viduals as such, or to phena within populations, have no official status.
“Name given to . . . infrasubspecific forms as such . . . are excluded”
from zoological nomenclature (Art. 1; see also 45c). Any “variety” name
published after 1960 is an unavailable name [Art. 45e(ii)]. Likewise un¬
available are all names given to hypothetical concepts, to teratological speci¬
mens as such, or names proposed for other than taxonomic use (Art. 1).
The absence of a convenient non-Linnaean nomenclature for single
specimens is a considerable handicap in paleontology, particularly in the
study of fossil man. To have a reference name many anthropologists created
a generic and specific name for every new specimen they found. Simpson
(1963) has exposed the fallacy of this custom. When dealing with indi¬
viduals one must adopt some sort of vernacular nomenclature. Let us use
“Olduwai LI” or “Trinil D7” rather than a set of scientific names which
imply a nonexistent zoological status. The same is true for infrasubspecific
names (13.41).
342 Principles and Application of Zoological Nomenclature

7. RECOMMENDATIONS

Recommendations indicate the best procedure in cases not covered

by the strict application of the rules. They are designated by the number
of the article with which they are associated, followed by an appropriate
letter. Although compliance with the recommendations is not mandatory,
it is nevertheless highly advisable as representing good practice.

8. IMPORTANT DATES

The most important dates in zoological nomenclature are summarized

on page xviii of the 1961 Code (xx in the 1964 edition). The starting
date for zoological nomenclature is January 1, 1758, and the tenth edition
of the Systema Naturae has been conveniently decreed to have been pub¬
lished on that date (Art. 3). The only exception is that spider nomenclature
is considered to have started in 1757 [Clerck, C. 1757. Aranei suecici
(Svenska Spindler). Stockholmiae]. See BZN, 4:319.

9. RETROACTIVITY

It is one of the basic concepts of good law that no new law should
be applied retroactively. An article stating this principle explicitly was
adopted by the London congress but was eliminated by the editorial com¬
mittee with approval of the majority of the Commission. This has led
to some difficulties, particularly with respect to the emendation of family
names (Art. 29), to the universal adoption of unjustified emendations (Art.
33), to the types of replacement names (Art. 72d), and the revival of
secondary homonyms (Art. 59c). See BZN, 18:323 and elsewhere.

10. VERNACULAR NOMENCLATURE

Vernacular names for animals and plants exist in all languages. A

primitive tribe of Papuans in the mountains of northwestern New Guinea
has 137 different names for the 138 species of local birds. Hunting peoples
usually have a better knowledge of nature and consequently a richer taxon¬
omic nomenclature than agricultural or particularly urban peoples. The more
conspicuous species of mammals, birds, fishes, and insects have names in
all the languages in Europe. In English many of our so-called “common
names” are of Anglo-Saxon origin. For better-known animals they are usu-
 Interpretation of the Rules of Nomenclature 343

ally uninominal. For example, among British birds, raven, rook, jay, (mag)-
pie, (jack) daw, robin, redwing, thrush, linnet, nightingale, partridge, and
many others; and among the butterflies, monarch, greyling, ringlet, peacock,
comma, etc. In other cases group names exist, like bear, frog, woodpecker,
and species names are formed by modifying these group names with a
descriptive noun or adjective, thus polar bear, brown bear, etc. Still other
names are polynominal, such as small pearl-bordered fritillary, dark green
fritillary, etc. These more complex polynomials, consisting of a generic
name and a qualifying epithet, occur in many native languages all over
the world.

11. SCIENTIFIC NOMENCLATURE

Latin was the international language of European scholars in the

Middle Ages, and up to the eighteenth century the majority of scientific
treatises were written in that language. The Swede Linnaeus, to make
his work more widely available, wrote mostly in Latin. In the Systema
naturae he introduced a system of scientific names for animals and plants,
based on Aristotelian logic, which is still the official system of zoological
nomenclature, 200 years later. The Linnaean method of naming (see Chap.
4) has been excellently explained by Svenson (1945) and by Cain (1958).
The generic name was the important element in the Linnaean nomencla¬
ture, always consisting of a single word. The differentia which followed
the generic name specified the characters of the given species within the
genus. It could be up to 12 words long and often had to be changed
when new species or characters were discovered. To cope with this complex¬
ity and instability Linnaeus invented the trivial epithet for each species—a
single specific term, now called the specific name and, as it turned out,
of immense practical value. Subsequent workers abandoned the differentia
together with the Aristotelian logic but retained as the scientific designation
of a species a combination of a generic and a specific name, a binomen
(see Arts. 5, 6, 11c). Linnaeus consistently applied this system to animals
for the first time in 1758 in the tenth edition of his Systema naturae.
There are slight differences of meaning between the terms binominal,
binomial, and binary. Binary refers to designations consisting of two kinds
of names. The Linnaean combination of generic name and differentia was
polynominal, yet binary. The nomenclature of many early post-Linnaean
authors was binary, having a uninominal generic name, as for instance
that of Brisson. The 1961 Code uses only the term binominal nomenclature
even though admitting that the name of a subspecies is a trinomen. Binomi¬
nal is an amendation of the formerly used term binomial.
344 Principles and Application of Zoological Nomenclature

12. ADVANTAGES AND DISADVANTAGES OF BINOMINAL

NOMENCLATURE

The Linnaean binominal nomenclature is a compromise between Aris¬

totelian logic and a simple information retrieval system. The genera adopted
by Linnaeus were extremely wide in scope and truly an aid to the memory.
Not only a zoologist but even an educated lay person could memorize
the 312 generic names which Linnaeus used for all animals. The scientific
name thus consisted of two components, a group-designating generic name
as an aid to the memory, and a specific name emphasizing distinctness
and uniqueness.
Since Linnaeus’ time this system has lost much of its usefulness. There
are far more kinds of organisms than Linnaeus ever imagined. The total
number of genera of animals now recognized may well exceed 50,000.
A generic name no longer tells much to a zoologist except in a few popular
groups of animals; it has become a tool for the specialist. The degrading
of the usefulness of the generic name has been made worse by the extreme
generic splitting so prevalent in many areas of systematic zoology (see 10.5).

13. NAME CHANGING

The worst aspect of the binominal system is its extreme instability.

Every change in the delimitation of a genus or in the assignment of a
species to a genus will lead to a change in one or several scientific names.
Genera are split, genera are lumped, and species are frequently shifted
from one genus to another. For instance, most newts were at one time
included in the genus Triturus. Subsequently it was discovered that this
so-called genus was actually a highly artificial aggregate of very different
kinds of newts. What was once Triturus viridescens became Notophthalmus
viridescens, and Triturus rivulosus became Taricha rivulosa. Zoologists who
had long used these species for embryological studies were understandably
upset when their experimental material suddenly had a different name.
In a group of bees of the tribe Paracolletini, a single revision resulted
in a change of 288 out of a total of 332 binomina and, even worse, in
the need for renaming 16 species owing to secondary homonymy (Michener,
1964). In these examples we are not dealing with the results of excessive
splitting or any sort of arbitrariness, but with a serious weakness of the
entire system of binominal nomenclature. Increasingly often the question
is therefore asked whether the entire system of binominal nomenclature
has not outlived its usefulness.
 Interpretation of the Rules of Nomenclature 345

14. ALTERNATE SYSTEMS OF NOMENCLATURE

Two sets of suggestions have been made to meet the difficulties men¬
tioned in 13.13. Herrera (1899) and other authors in the nineteenth and
early twentieth century suggested various prefixes and suffixes to the generic
name in order to indicate to what order and class of animals the genus
belongs. For instance, it was suggested modifying the generic name with
initial letters indicating class and order, and terminations indicating sub¬
kingdom (us equals Vertebrata, a equals Invertebrata, um equals Protozoa).
Thus Papilio would become Ylpapilia (Y equals Insecta, l equals Lepidop-
tera). A number of other schemes to facilitate the ready placing of a
generic name in the animal system were published.
More radical proposals suggest replacement of binominalism by
uninominalism. Michener (1964), for instance, proposes freezing the origi¬
nal scientific name of a new species for all time by connecting generic
and specific names with a hyphen. The disadvantage of this proposal is
a perpetuation of all early errors of judgment. Thus Leioproctus jenseni
of the bee family Colletidae would continue to be listed under its original
name, Nomia-jenseni, even though Nomia is a genus of a different family
(Halictidae). A rather high percentage of the names proposed in a poorly
studied group would probably be misleading when the group became better
understood. The obvious advantage of this system is that any name can
be retained for all time regardless of any taxonomic shifts. Michener very
perceptively also points out that many improvements in classification are
quietly resisted by taxonomists when they realize the nomenclatural havoc
the change would produce. Among North American ornithologists, for in¬
stance, it has long been realized that the genera Dendroica (1842) and
Parula (1838) are not separable. Yet Dendroica with over 25 species is
so well known that no one dares to list these species under the older name,
Parula, particularly since this, in turn, may be a synonym of the still older
name Vermivora.
A scientific name serves two functions in the current zoological nomen¬
clature—that of an identification tag (like a social security number) and,
being a binominal, that of a classificatory device. There was no conflict
between the two functions, as far as Linnaeus was concerned. According
to his essentialist concept of the genus, once a species was correctly identified
as to genus, there was little danger of it ever changing its name. Also
the number of taxonomists was so small that Linnaeus and his associates
did not foresee a major problem with synonymy. Both assumptions proved
to be wrong, and the result has been a disastrous instability in the Linnaean
system of nomenclature.
More and more voices are now heard recommending either that the
346 Principles and Application of Zoological Nomenclature

two nomenclatural functions be separated or that they be expressed nu¬

merically, or both. Instead of a binomen there might be a running identifica¬
tion number (for example, 968-1, 968-2) for all species described in 1968,
the numbering to be done in a central international office. To facilitate
information retrieval, it would also be desirable to have a numerical scheme
for the hierarchy of higher taxa in which a given species is classified. Various
authors have made such proposals, for instance Hull (1966) and Bullis
and Roe (1967). With a steadily increasing need for easy information
retrieval and ease of programming for computers, there is little doubt that
taxonomists will have to adopt some uninominal system sooner or later.
This will presumably exist, at least at first, side by side with the Linnaean
nomenclature, but it may eventually replace it. Obviously we need a system
of identifying symbols for species that can be used by nonspecialists and
is suited for mechanical information-retrieval devices. Yet all numerical
systems so far proposed have failed to cope fully with the difficulties pro¬
duced by the discovery of new taxa and the inevitably continuing shifts
in classification.

15. CRITERIA OF PUBLICATION

No nomenclatural action is valid until it has been published. What

constitutes publication is spelled out in Art. 8, and what does not in Art.
9. In the eighteenth and early nineteenth century many authors thought
they could make a name available by labeling specimens in their collection
with that name or by demonstrating such specimens at scientific meetings.
The Code is quite specific in requiring a printing method of publication
by which numerous identical copies can be reproduced. Although not ex¬
plicitly rejected by the Code, replication methods such as mimeographing
are considered by most zoologists as unsuitable for nomenclatural
publications.
Editors of scientific journals must be familiar with the provisions
of the Code if they permit the publication of new names of taxa. Corliss
(19626) stated perceptively:

It seems to me that exercise of just a little more editorial care . . . would

have reduced impressively the number of improper names now in need of
mandatory emendation, the hundreds of inappropriate and undesirable names
in the literature which cannot be changed, the many incorrectly cited dates
and authorships, erroneous combinations, double publications of a new
name as new, and myriads of inaccurate and incomplete bibliographic cita¬
tions of taxonomic and nomenclatural importance.

No author should ever submit the description of a new' taxon simultaneously

to several journals. This inevitably causes difficulties in dating and
synonymy.
 Interpretation of the Rules of Nomenclature 347

16. CONCEPT OF AVAILABILITY

In order to be available, a name must have been published in a

way to satisfy the requirements detailed in Arts. 8-20. The term available,
as used in the Code, means “legitimate” or, as the botanists put it, “effec¬
tively published.” A name may be “available” in this technical sense even
though, as a junior primary homonym, it is automatically invalid at the
moment of publication (Arts. 53, 59a). It is most important to understand
clearly the difference between “available” and “valid.” To have this distinc¬
tion made clear is one of the major advances of the 1961 Code. To employ
the name of a taxon one must make sure that it is available (properly
published) as well as valid (not preoccupied by a senior synonym or
homonym).
A name published without satisfying the conditions of availability
is generally called a nomen nudum, particularly if it fails to satisfy the
conditions of Arts. 12-16. A nomen nudum has no standing in zoological
nomenclature and is best never recorded, not even in synonymy. There
is always the danger that such a listing provides an indication in the sense
of Art. 16 and thus inadvertently makes the manuscript name available.

17. KINDS OF NAMES

In addition to available names and nomina nuda the Code recognizes

directly or by implication several other kinds of names. Pre-Linnaean names,
that is, names published prior to January 1, 1758, have no status. A nomen
dubium is an available name which cannot be assigned to a definite taxon
owing to shortcomings in the original diagnosis or the type material. Since
most nomina dubia are a potential threat to the validity of universally
accepted names, zoologists frequently apply to the Commission to place
such names on the official index of rejected names. For nomina oblita
see 13.24 (statute of limitation). Names to be excluded from zoological
nomenclature are specified in Art. 1. Finally, rejected names are included
in the official indexes of rejected names published by the Commission (see
13.63).

18. PUBLICATION IN SYNONYMY

Article 11 d states that a name first published as a synonym is not

thereby made available. This is a wise and important provision. Many
compilers of name lists had the habit of introducing manuscript names
into synonymy. Their taxonomic judgment was frequently faulty, so that
348 Principles and Application of Zoological Nomenclature

they listed the manuscript name in the synonymy of a species to which

it did not belong. Most of these names were ultimately made available
by proper publication and were then universally adopted as names of the
taxa to which they really belong. To consider such names as preoccupied
by the erroneous synonymation would be a violation of the guarantee of
taxonomic freedom (13.2). Any available name can be placed into and
withdrawn from synonymy without permanently affecting the availability
or validity of other names (Art. 17(1)). Placing a name into synonymy
is a taxonomic action; it is a nomenclatural one only as long as the tax¬
onomic decision is considered correct. However, if a name, first published
as a synonym, has been (universally) treated prior to 1961 as an available
name, such a name shall be considered as available, with its original date
and authorship, and be either adopted as the name of a taxon or be used
as a senior homonym (Art. 1 Id, 1964 ed.).

19. DIFFERENTIAL DIAGNOSIS

The 1901 Code specified that in order to qualify for availability a

name had to be accompanied by a description, definition, or indication.
So many flimsy and meaningless so-called descriptions were, however, pub¬
lished that the International Congress at Budapest in 1927 adopted a pro¬
vision that in the future (after 1930) names would have to be accompanied
by a differential diagnosis. Since it is impossible to prove that a description
qualifies as a differential diagnosis, this was modified in the 1961 Code
[Art. 13a(i)] to state that the name must be “accompanied by a statement
that purports to give characters differentiating the taxon,” etc. However,
a direct comparison with another specified taxon (differential diagnosis)
is not required.
Respectable taxonomists go well beyond this minimal requirement and
describe and illustrate carefully the diagnostic differences between a newly
proposed taxon and its closest relative(s). If this relative is a little-known
species, the characters which separate the new species from a well-known
or common species should also be stated. The same provision is true for
new genera and new subspecies. Thus, direct comparison is not a mandatory
provision of the Code but one which characterizes the conscientious worker,
the worker who has the respect of his peers.

20. INDICATION

The various conditions which satisfy the requirement of an “indica¬

tion” in the sense of Art. 12 are listed in Art. 16a, and what does not
constitute an indication in Art. 16b. It is important to emphasize that
 Interpretation of the Rules of Nomenclature 349

reference to a vernacular name or to a type-locality is not sufficient to

make a name available. Many of the indications sufficient to ensure avail¬
ability prior to 1931 (Art. 12) are now no longer sufficient (Arts. 13-15).

21. NAMES GIVEN TO HYBRIDS

Hybrids are normally individuals, not populations, and hence not

taxa. Article 1 states clearly that a name given to a hybrid as such is
not available. Article 17 (2) specifies that a name given to an animal later
found to be a hybrid remains available only for the purposes of homonymy,
not (as was clarified by the International Congress at Washington) for
the purposes of synonymy. (A Declaration is in preparation.) A name
given to a species hybrid cannot be applied to either of the parental species.
The Code does not specify what to do with names given to specimens
taken from hybrid populations. It is here recommended that Art. 17 (2)
be applied to cases of hybridization between two species, but that such
a name be considered available for the purposes of homonymy and
synonymy in the case of intergrading populations between subspecies. Such
populations are usually highly variable, and if the intermediate population
is not recognized as a separate subspecies, a name given to an individual
in this population may, if appropriate, be applied to the more similar
of the two adjacent subspecies.

22. DATE OF PUBLICATION

The relevant provisions are clearly stated in Art. 21. The date of
publication is the date on which the publication was mailed to subscribers,
placed on sale, or, where the whole edition is distributed free of charge,
mailed to institutions and individuals to whom such free copies are normally
distributed. Journals are sometimes mailed weeks or even many months
after the date printed on the covers. In these cases the mailing date is
the correct date, not the printed date. (For publication, see also Arts.
8 and 9.)
Even more confusing are serial publications, parts of which are some¬
times issued over a period of 20 or 30 years. Each part of such publications
has a separate publication date, the date on which it was actually mailed.
In tracing the actual publication date of historical periodicals and series,
the Journal of the Society for the Bibliography of Natural History (starting
with vol. 1, 1936) is invaluable.
350 Principles and Application of Zoological Nomenclature

23. VALIDITY

Validity is a term that refers to the rights of names in relation to

homonyms and synonyms. Synonyms are different names for the same thing.
The earliest published synonym is referred to as the senior synonym; all
later synonyms are junior synonyms. A name becomes a senior synonym
at the time when a second name for the same taxon is made available.
Two kinds of synonyms can be distinguished. One consists of names
that objectively refer to the same thing, such as a new name for a supposedly
preoccupied name, or names based on the same specimen or illustration.
These are called objective synonyms. The other consists of names based
on different type-material. Taxonomists may nevertheless decide that they
refer to the same taxon. These are called subjective synonyms. A generic
lumper, for instance, may consider as synonyms certain generic names which
a splitter would consider valid.
Incorrect spellings of names (Art. 33b) do not qualify as synonyms.
At a given time only one name can be the valid name of a taxon. This
is normally the oldest available name that is not preoccupied by a senior
synonym or homonym and has not been suppressed by the Commission
for the sake of nomenclatural stability. The provisions are clearly stated
in Art. 23. Suppression by the Commission converts a previously valid name
into an invalid name.
One provision which is occasionally misunderstood concerns the se¬
niority ranking of coordinate taxa [Arts. 23e(iii) and 476], If it is found
that the name of a polytypic species is invalid, it is replaced by the next
oldest name given to any subspecies of this species and not necessarily
by the replacement name for the formerly nominate subspecies. The same
is true for the replacement name of a genus consisting of several subgenera
(Art. 44b). The situation with respect to family names is different (13.36).

24. STATUTE OF LIMITATION

In order to give concreteness to the statement in the Preamble “when

stability of nomenclature is threatened in an individual case,” a statute
of limitation was adopted by the Copenhagen and London congresses, as
stated in Art. 23b. Statutes of limitation are almost universal in law. De¬
mands that a limit be set on priority had been made early in the history
of taxonomy. When the ornithologist Coues found in 1880 that the well-
known name Ectopistes migratorius of the Passenger Pigeon was threatened,
he wrote: “Is it worthwhile to make this change? Cases like this make
one wish that there were in our nomenclature some law of limitation by
 Interpretation of the Rules of Nomenclature 351

which a name which has not been challenged for, say, 50 years or a century,
might then acquire an inalienable right to recognition.” Article 23b is in¬
tended to achieve this.
The article gives an automatic protection to names that have been
in unchallenged use for a period of at least 50 years. When after the
end of such a 50-year period a senior synonym is discovered that has never
been used during that period, it is to be considered a forgotten name
(nomen oblitum).
Article 23b is not well worded, which is one of the reasons w7hy
it has been widely criticized. Instead of stressing the basic intent of the
rule, the automatic protection of well-established and universally used junior
names, it concentrates on the suppression of unused older names (which
is merely a method for achieving the basic aim). It does not clearly define
“unused” and “widely used,” and a few zoologists have even misinterpreted
the definition of the 50-year period, which actually is the 50-year period
immediately preceding the rediscovery of the senior synonym.
In view of these difficulties the Washington Zoological Congress in¬
structed the Commission “to prepare an interpretative Declaration on Arti¬
cle 23b.” The President of the Commission has appointed a committee
which is now scrutinizing each of the individual provisions in order to
determine (1) w'hether they are workable, (2) w'hether they should be
replaced by a preferable alternative, and (3) what warding to adopt in
order to exclude all conceivable misunderstandings. The report of this com¬
mittee will eventually be voted on by the Commission as a w'hole, so that
it can discharge its task. For instance, the Declaration may specify, as
was suggested at the Washington congress, that in order to deserve protec¬
tion the junior name must have been used at least ten times and by at
least two authors during the stated 50 years. Such a provision wrould greatly
help zoologists w7ho work in inactive groups, for whom the benefits of Art.
236 are questionable.
It has been quite rightly suggested that the term “used” should be
construed rigidly, so that only use in the primary zoological literature quali¬
fies. This is a term for literature in which the name is actually applied
to a definite zoological object. A report on a collection, a faunal list, any
zoological or physiological paper, any textbook—all such publications are
primary. Anything that does not refer to zoological objects but is merely
a catalog of names without diagnostic or distributional information does
not qualify as primary zoological literature. Accordingly, the mere listing
of a name in synonymy or in a bibliographic publication such as the Zoolog¬
ical Record, or Biological Abstracts, or any index of names does not qualify
as primary zoological literature. Catalogs and checklists cannot be desig¬
nated as “primary or not” on an overall basis. Some checklists like the
American Ornithologists’ Union Checklist are critical surveys of zoological
352 Principles and Application of Zoological Nomenclature

objects. Other so-called checklists are nothing but uncritical compilations

of names from the Zoological Record and other bibliographic sources. If
an author is in doubt, he should be guided by the spirit of the Preamble.
Whenever stability is not endangered, it is advisable to abide by strict
priority.
The primary purpose of Art. 23b is to prevent the revival of a long-
forgotten name which would replace a well-known one, a common practice
prior to the existence of such a statute. There are many groups of organisms
in which species names are referred to in several hundred publications
annually. Furthermore, many of these publications, such as field guides,
zoological textbooks, and literature on embryology and public health, have
a circulation of thousands of copies. In the past there was no automatic
protection for such names. When a bibliographer exhumed a long-forgotten
name, the entire burden of saving the well-known name was placed on
the general zoologist. By the time the Commission was able to get to a
ruling—and in periods of disturbed international communication there was
sometimes a lag of ten or more years—the replacement name had gained
sufficient currency to make the situation virtually insoluble. Article 23b
provides for the automatic protection of all names that have been in current
use for more than 50 years without depriving names in inactive taxa of
their rights.
The greatest difficulty the Commission faces in the preparation of
its interpretative Declaration is the need to come up with a formulation
that is equally acceptable to zoologists working on animals with a very
active literature, like mammals, birds, butterflies, and disease vectors, and
also to zoologists working in groups in which there is only a single specialist
every 30 to 50 years and names are only rarely cited in the primary litera¬
ture. Hopefully, the principles laid down in the Preamble will guide the
Commission in arriving at a workable compromise.

25. COMBINED AND DIVIDED TAXA

A taxon formed by the union of two or more taxa takes the oldest
valid name among those of its components [Arts. 23d(i), 23c] (see also
Art. 67k). In the case of family names, the strict application of priority
may be set aside [Art. 23rf(ii)] (see 13.33).
If a taxon is subdivided, its valid name must be retained for one
of the components. This rule applies equally on the family (Art. 37), generic
(Art. 44), and species levels (Art. 47). For instance, when a species is
subdivided into several subspecies, the subspecies which contains the topo-
typical population becomes the nominate subspecies, its subspecific name
being the same as that of the species.
 Interpretation of the Rules of Nomenclature 353

Linnaeus described the red-winged blackbird (Agelaius phoeniceus)

on the basis of Catesby’s drawings and descriptions from South Carolina.
South Carolina has therefore been designated as the type-locality. When
this species was subdivided into several subspecies, the subspecies of eastern
North America (including the region of South Carolina) became automati¬
cally the nominate subspecies, namely, Agelaius phoeniceus phoeniceus
(Linnaeus).
Occasionally authors create synonyms by ignoring this rule. For in¬
stance, Thienemann (1938) found that the well-known turbellarian worm
Planaria alpina consisted of two subspecies. The northern one (northern
Germany, Scandinavia) he called septentrionalis: the southern one (Alps)
he called meridionalis. Since the species had been described originally from
Scandinavia and the Alps, it is obvious that either meridionalis or septen¬
trionalis is a synonym of the nominate subspecies P. alpina alpina, the
type-locality of which needs to be restricted to one of the two areas.
If a genus is divided into several genera, one of the new genera
must bear the name of the originally undivided genus. Traditionally this
has also been considered the preferred method for higher taxa (see 13.34).

26. PAGE AND LINE PRECEDENCE

If two names for the same taxon are published in the same publication,
they are considered as “published simultaneously” (Art. 24a). One of these
names may have line or page precedence, but their priority is determined
by the action of the first reviser (13.4). The first reviser must give serious
consideration to Recommendation 24A and “select the name that will best
ensure stability and universality of nomenclature.” Almost invariably one
of the two competing names is better known, is based on a better description,
is based on better type-material, is based on an adult phenotype rather
than on an immature stage, or has some other nomenclatural advantage.
This is the name the first reviser should select. Chronological priority is
not involved in the case of simultaneous publication, it is replaced by desig¬
nated priority. The same is true for simultaneous publication in different
works (also Art. 24a).

27. WORK OF AN ANIMAL

Prior to 1931 the description of the work of an animal, for instance

its gall, was considered a valid indication [Arts. 16a(viii), 246(iii)]. After
December 31, 1930, a name does not become available unless it also satisfies
Art. 13a by being accompanied by a statement attempting “to give char¬
acters differentiating the taxon” from close relatives.
354 Principles and Application of Zoological Nomenclature

28. FORMATION OF NAMES

Mandatory provisions governing the formation of zoological names

are given in Arts. 26-30. Since such names must be either Latin or latinized
or at least so constructed that they can be treated as a Latin word (Art.
11 b), it is most important for the zoologist to familiarize himself with
the rules that govern the correct transliteration and latinization of words.
Three appendices (B, C, D) to the Code are specifically devoted to these
matters. They are not mandatory provisions but have the same status as
recommendations. Some older treatments are still useful.1 See also Brown
(1954).
Appendix B (Code, pp. 95-101) deals with the transliteration and
latinization of Greek words. Appendix C (Code, pp. 101-103) deals with
the latinization of geographical and proper names, while appendix D (Code,
pp. 105-141) deals quite generally with recommendations on the formation
of names. Every zoologist who plans to propose a new name for a taxon
should carefully read these recommendations.
A few points might be stressed particularly. Complex, long, and diffi-
cult-to-pronounce names should be avoided. It is good practice to propose
only the sort of new generic name that differs from an old, already existing,
generic name in more than a single letter or its termination. It is very
unwise in describing a new species to choose a specific name already in
use in a related genus. Such a name would become a homonym if a subse¬
quent author should combine these genera. For instance, names like afri-
canus, robustus, capensis, etc. were given repeatedly to closely related
hominids which subsequent authors might well place in the genus Homo.
Names for protists should be such as would also be valid under the botanical
rules.
A noun in opposition cannot be treated as an adjective. Words like
longicauda, melanogaster, or albipectus in which the ending is a noun
cannot be changed when transferred to a genus with different gender.
Compound names are to be united into a single word without a
hyphen (Art. 26), with one exception (Art. 26c).

29. DIACRITIC MARKS

Diacritic marks, apostrophes, or diaereses are not to be used in a

zoological name and are to be deleted from names originally published with
such marks (Art. 27). The only exception is the German umlaut sign,
which is to be deleted from a vowel, the letter “e” to be inserted after
,
the vowel [Art. 32c(i)]. Thus a, 6 u become ae, ce, ue.

1 W. Miller, 1897. Calif. Acad. Sci., (3) vol. 1, no. 3, and F. E. Clements, 1902. Univ. Studies,
Nebraska, vol. 3, no. 1, are two examples.
 Interpretation of the Rules of Nomenclature 355

30. STEM OF A FAMILY NAME

Names of the family group are formed by the addition of prescribed

endings (-idae, -inae) to the stem of the name of the type-genus (Art.
29).
The 1901 Code did not specify how the stem was to be determined,
and many family names were formed in such a way as to give euphonious
combinations. The 1961 Code specifies (Art. 29a) that “the stem is found
by deleting the case-ending of the appropriate genitive singular.” When
retroactively applied, this provision will lead to a change in the names
of some well-known families. For instance, the family of stick-insects or
phasmids, almost universally called Phasmidae in zoology textbooks, will
have to be called Phasmatidae.

31. GENDER OF GENERIC NAMES

If the specific name is an adjective in the nominative singular, it

must agree in gender with the generic name. To apply this rule correctly
the zoologist must understand the rules governing the gender of Latin
nouns, whether of Latin, Greek, or nonclassical origin. These are given
in Art. 30. Various handbooks on scientific names are helpful, and standard
Greek and Latin dictionaries are indispensable (see also Appendices B,
C, D).
Latin and Greek grammar are full of pitfalls. A Latin noun ending
in -us is by no means necessarily a noun of the second declension and
thus automatically masculine: it may be feminine like domus or neuter
like pectus. A latinized Greek word takes the gender appropriate to its
Latin termination [see examples under Art. 30a(i) (3)].
A zoologist who proposes a new name in the genus group should
give the etymology and the gender of the new name (Appendix E, 16).

32. ORIGINAL SPELLING AND EMENDATION

One of the curses of nomenclature has always been the zoologists’

habit of “correcting” names proposed by other authors. Article 32 states
clearly that the original spelling must be retained unless some exceptional
condition exists. There must be evidence in the original publication of
an inadvertent error such as a lapsus calami or a copyist’s or printer’s
error. Other kinds of errors such as incorrect transliteration, improper
latinization, and use of an inappropriate connecting vowel are not to be
considered inadvertent errors.
356 Principles and Application of Zoological Nomenclature

This provision has been severely criticized by those with a classical

education. But it must be realized that fewer and fewer zoologists have
such an education and that the meaning of zoological nomenclature has
changed since the time of Linnaeus. At that time it was a Latin language,
now scientific names are essentially arbitrary combinations of letters that
serve as labels for taxa. Many names for species are misleading, but this
is irrelevant considering the now accepted meaning of nomenclature. It
is contrary to this meaning and to the Code to change Australopithecus
to Australanthropus because one considers this fossil hominid to be closer
to man than to the anthropoids.
Although the article prohibits any corrections of grammatical mistakes
in generic and specific names, except for matters of form (Arts. 26-28,
30), it makes mandatory in Art. 32zz(iii) emendations in the stem of familv
names, which seems quite inconsistent.
An incorrect original spelling, as defined in Art. 32c, is to be corrected
wherever it is found. The incorrect spelling has no separate status in nomen¬
clature. The same is true for incorrect subsequent spellings (Art. 33b),
that is, any unintentional error in the spelling of a name.
Any demonstrably intentional change in an original spelling of a name
is an “emendation.” If this emendation is not justified by the provisions
of Art. 32, it is to be considered an “unjustified emendation” [Art. 33a(ii) ].
An unjustified emendation has the status of a new name with its own
date and author and is a junior objective synonym of the name in its
original form. It is available as a replacement name in case the name
as originally spelled is found to be invalid owing to homonymy.

33. FAMILY NAMES

The names of taxa above genus rank are always uninominal and
in the plural. Provisions relating to names in the family group are found
in Arts. 11c, 23c?, 29, 35-41, 63-65. To avoid grammatical mistakes, remem¬
ber that these names are in the plural. One can say, “The family Fringillidae
is the largest family of songbirds” but must say, “The Fringillidae are
the largest. . . .” The same is true for the names of orders, classes, and
other higher taxa.
Linnaeus did not employ the family category, but after it was intro¬
duced by French zoologists in the 1780s and 1790s, it was soon universally
adopted. The early codes, however, had relatively few provisions concerning
family names.
An author proposing a new taxon of family rank has the taxonomic
freedom to select the genus which he considers central (“most typical”)
for the new taxon. Seniority of the name of the type-genus is irrelevant.
Provisions for the formation of family names are given in Art. 29.
 Interpretation of the Rules of Nomenclature 357

The 1961 Code introduces limited priority for family names as the
result of a decision made by the Copenhagen Congress. However, since
family names are widely used in zoology, even by nonspecialists, stability
is even more important on the family level than on those of genus and
species. The Code therefore provides for two important stabilizing devices.
Article 23z/(ii) decrees that “if a zoologist observes that the strict application
of the Law of Priority to two or more synonymous family-group names
would upset general usage,” he is to request the Commission to decide
which name is to be accepted.
There are no satisfactory nomenclators that list the dates and authors
of names in the family category. Prior to 1900 (Art. lie) no family name
needed to have standardized endings, but they are now mandatory (Art.
29). In groups with little history, little published literature, and few special¬
ists it is usually not difficult to establish authorship and original date of
family-group names. In groups with a rich literature and long history,
such as mammals and birds, it is virtually impossible to establish the original
date of publication for the older, traditional names. This has created count¬
less difficulties. Most specialists will agree with Myers and Levitan (1962,
p. 290), who state: “An extension of priority to family names will take
zoologists into a maze of old group names which often cannot be clearly
recognized as of familial (or any other) hierarchical grade.” Some mitigat¬
ing provisions will have to be found, perhaps along the lines suggested
by Bradley (1962, SZ, 11:178-179), to facilitate the placing of family
names on the Official List.
The other provision favoring stability relates to name changes in the
type-genus. The name of a taxon in the family group must be formed
from the stem (see 13.30) of the name of the type-genus (Art. 29).
The type of a family taxon is the zoological object identified by the name
of the type-genus. There is, however, no need, for the sake of stability,
to change the name of a family each time the currently valid name of
the type-genus is changed. The type of the family is, as stated, a zoological
object and not a name. The validities of the name of the family and
of the name of the type-genus are independent (Art. 40). Family names
are formed by tautonymy from the name of the type-genus, and Art. 40
merely codifies what has always been true for generic names formed by
tautonymy from specific names. Likewise, they are never changed when
the specific name which served in the tautonymy loses its validity.
When a zoologist finds that the name of the type-genus of a family
is threatened owing to the availability of a senior synonym, he may well
want to bring this situation to the attention of the Commission. It is some¬
times advisable to suppress the senior name in order to retain the convenient
tautonymous relation between the name of the family and that of its
type-genus.
358 Principles and Application of Zoological Nomenclature

34. NAMES FOR HIGHER TAX A (ABOVE THE FAMILY

GROUP RANK)

The 1961 Code does not contain rules dealing with names of taxa
in categories above the family group. Attempts were made at the Copen¬
hagen and London congresses to draft some rules, but these were unsuccess¬
ful. There is still much uncertainty about the basic subdivision of some
of the phyla, like the Porifera (sponges) and Turbellaria. A premature
freezing of the names of classes and orders might stultify subsequent efforts
for a complete reclassification on the basis of new character complexes.
The worst threat to stability has been the misconception of a few
zoologists that removal of some genera from a higher taxon requires a
complete renaming (13.25). There is no excuse whatsoever for renaming
the well-known Bryozoa with more than 3,000 species as Ectoprocta because
a few genera with 75 species were removed into the separate class or
phylum Entoprocta (Mayr 19686). Even though the names of higher taxa
are not based on the type method, the removal of some taxa of lower
rank does not justify the renaming of a higher taxon. Fortunately this
principle was firmly adhered to when heterogeneous elements were removed
from the Mollusca and Insecta as defined by Linnaeus.
It is frequently proposed that uniform endings for the names of the
higher taxa should be employed. This proposal, on first sight, seems so
logical and sensible that only mature consideration brings out its grave
shortcomings. First of all, it violates the concept of priority as well as
that of stability. For instance, six of the seven orders of insects recognized
by Linnaeus, i.e. Coleoptera, Hemiptera, Lepidoptera, Neuroptera,
Hymenoptera, and Diptera are still recognized essentially in the same sense
as 200 years ago. They all use the suffix -ptera (wings). The major orders
of mammals, such as Rodentia, Carnivora, Insectivora, and Primates are
equally old and well-known. Many ordinal names in birds, such as Im-
pennes, Tubinares, Oscines, are likewise not based on a type-genus. Simpson
1952, SZ, 1:20-23) well presents the argument against uniform endings.
Furthermore, little would be gained by having uniform endings be¬
cause the ranking of higher taxa is a particularly disputed area of zoology.
Different authors may rank the same taxon as superfamily, suborder, order,
or even subclass. Standardized endings would invest a given ranking with
far greater authority and definiteness than is justified by the facts. Finally,
there are several competing systems of uniform endings.
It would seem important, however, for the zoological congresses or
specialists in the groups to stabilize the nomenclature of higher taxa in
all cases where several names compete for the same taxon, as for instance
in the case of the bivalve mollusks (Bivalvia, Pelecypoda, Lamellibranchia).
 Interpretation of the Rules of Nomenclature 359

There are a few general principles concerning the names of higher

taxa above family rank. They are single words in the plural. Although
they are not based on a type-genus, there is generally a consensus as to
the central (“most typical’’) component of a higher taxon. Zoologists main¬
tain well-established names of higher taxa even though they may find some
other word “more appropriate.” When a composite higher taxon is divided,
the original name should be retained for the “more typical” or larger group
and a new name applied to the newly recognized group. See also MLU,
276-278. The history of the names of many higher taxa is uncertain. Corliss
(1957) has established it in exemplary fashion for the ciliates.

35. COORDINATE STATUS OF CATEGORIES

Article 36 specifies that the proposal of a new taxon in the family-

group implicitly constitutes permanent authorship for that taxon, even when
changed in rank. When a given taxon, first described as a new subfamily,
is raised to family rank, “it is thereupon available with its original date
and author” for use, based on the same type-genus, in the higher ranking
category. Article 37 specifies that describing a taxon in the family-group
automatically makes the describer also the implicit author of any nominate
subtaxon. The given example of the family Tipulidae well illustrates this
principle. Article 43 confirms the same principle for the genus and Art.
46 for the species.
The creation of superfamilies creates difficulties not properly taken
care of in Art. 36 (Mayr, 1954). If a family is raised to rank of a super¬
family, the provisions of Art. 36 must be applied. On the other hand,
if an author makes an entirely new superfamily by bringing together various
families not previously associated, he is not giving a new rank and name
to a previously established taxon. It would violate the principle of taxonomic
freedom (13.2) if he had to choose the type of the new superfamily
on the basis of the oldest included family name. Quite correctly, the prin¬
ciple is specifically rejected in Art. 64 for the choice of the type-genus
of a family and implicitly in the provisions for selection of type-species
for genera (Arts. 66-70).

36. SUBORDINATE TAXA

The provisions for subordinate taxa are essentially the same for the
family group (Art. 37), for the genus group (Art. 44), and for the species
group (Art. 47). The nominate subtaxon consists in each case of that
subtaxon which contains the type. One of the included subtaxa, which
360 Principles and Application of Zoological Nomenclature

therefore is called the nominate subtaxon, must have the same name as
the taxon. The only exception is on the family level in the case of renaming
the type-genus (Art. 40). Also, in the family group the subtaxon has a
different ending.
In the case of genus and species (Arts. 44b, 47b) it is provided
that the subtaxon with the next oldest name becomes the nominate subtaxon
when it is found that the name of the original nominate subtaxon is invalid.
This provision is a slight deviation from a rigid application of the type
method (see below). For this reason it is not applied (in Art. 37) to
subtaxa on the family level. Changes in the names of families have become
rarer under the new Code owing to the provisions of Arts. 23c? and 40.

37. HOMONYMY BETWEEN FAMILY NAMES

Homonymy between such names occurs only infrequently. It is of

course irreconcilable with the principle of uniqueness of names, laid down
in the Preamble, for a family name to be based on a generic name that
is a junior homonym. Should an author discover such a situation, it is
advisable that he report it to the Commission. If the senior homonym
should be either a forgotten name (Art. 23b) or a never-used junior syn¬
onym, the Commission can exercise its plenary powers to suppress such
a name. This would result in preservation of the family name based on
the junior homonym.
Article 55 advises on situations where identical family names are based
on nonidentical generic names. Such cases must always be referred to the
Commission.

38. GENUS GROUP NAMES

Provisions dealing with generic names are found in Arts. 11/, 13b,
and 42-44. A genus group name must be a noun in the nominative singular
or be treated as such. A noun in the plural when first published takes
its date and authorship from its first publication in the singular. An author
proposing a new generic name should make certain that his proposal does
not omit any of the following five essential points:

1. That it is clearly indicated as a new genus: X-us, new genus.

2. That the proposed generic name does not violate the rules and the
recommendations concerning the formation of names (consult Code Ap¬
pendix D).
3. That the newly proposed name is neither a homonym nor a synonym.
4. That the description contains a clear statement of the characters in
 Interpretation of the Rules of Nomenclature 361

which the new genus differs from previously described genera (Art. 13a).
It is desirable that a differential diagnosis is added in which a direct
comparison is made with that genus or those genera which are believed
to be most closely related to the newly described one.
5. That the type-species is clearly and definitely stated (Art. 13b). The
generic limits may be interpreted differently by succeeding authors. But
the type-species will forever anchor the concept of the stated genus to
a clearly defined zoological object.

For additional comments on generic names see MLU, pp. 261-270.

39. COLLECTIVE GROUPS

A collective group of identifiable species of which the generic position

is uncertain may be treated, for the sake of taxonomic convenience, as
a genus (Art. 42c). Such a collective group does not require a type-species.
Assignment to a collective group is a temporary identifying label which
is particularly convenient in parasitology and paleontology.

40. NAMES OF TAX A IN THE SPECIES GROUP

The rules concerning these names are given in Arts. 5, llg, 23c(ii),
34b, and 45-49. There are only two categories in the species group, the
species and the subspecies. The specific name is the second word in the
binomen (Art. 5), and the subspecific name, when employed, is the third
word of the trinomen. Linnaeus used the term nomen triviale for the specific
name, but for various reasons this usage became essentially obsolete in
zoology after 1800. The initial letter of these names is lowercase (Art.
28). Since a scientific name in the modern interpretation is considered
a recognition symbol formed by a sequence of letters (13.32), it is not
advisable (indeed leading to confusion with generic names) to capitalize
specific names at the beginning of a sentence.
Articles 11 to 19 list the prerequisites for making a specific name
available. Particularly important is Art. 13a(i), according to which names
published after 1930 must be “accompanied by a statement that purports
to give characters differentiating the taxon.” It is good taxonomic practice
to include in the original description not only such diagnostic information
but also a differential diagnosis that consists of an actual comparison with
closely related species (see 13.19). Advice on correct endings of species-
group names formed from modern personal names is given in Recommenda¬
tion 31A of the Code. A vote of the Washington congress in 1963 deleted
Art. 31 (of 1961) and replaced it with this recommendation, thereby vali-
362 Principles and Application of Zoological Nomenclature

dating many of the practices current prior to 1961 and converting some
of the mandatory provisions of Art. 31 (of 1961) into recommendations.
Three widespread usages in particular were permitted, namely (1) treating
modern personal names as nouns in apposition like Calypte anna, (2)
employing the correct grammatical genitive of a latinized family name,
e.g. fabricii instead of fabriciusi, and (3) employing the correct masculine
genitive ending, -ae, in words derived from names like Costa, Molina,
Kuroda. When used as masculine nouns, Latin words like agricola also
have the genitive -ae, e.g. agricolae.
The frequently made suggestion “to correct” the latinized dedication
genitive -ii to -i is implicitly rejected by the wording of Art. 32a that
the original spelling of a name is to be retained as the “correct original
spelling.” The ending -ii is not listed as contravening this provision.

41. INFRASUBSPECIFIC NAMES

Linnaeus did not recognize the subspecies category. The term variety
used by him and his followers referred to a medley of deviations from
the type of the species (3.3.1; also, see Mayr, 1963, pp. 334-346).
No distinction was made by him between different kinds of infraspecific
variants. As a result of a long and tortuous history we now clearly distinguish
between two kinds of infraspecific variants. The subspecies designates a
genuine taxonomic category based on populations; variety names are names
for phena and aberrant individuals and have no standing in nomenclature
(Art. 1).
For more that 150 years zoologists did not make a clear distinction
between varieties that are genuine taxa (based on populations) and varieties
that designate merely groups of individuals or phena. It would be ritualistic
to determine the availability of infraspecific names on the basis of the
original terminology chosen by the author. Although the Code clearly states
that names given to “infrasubspecific forms as such” are excluded (Art.
1), it deals with other aspects of infrasubspecific names in Arts. 106 and
45c, d, e. Any name proposed after 1960 is unavailable if not clearly given
to a subspecies taxon. In the case of infrasubspecific names given prior
to 1961, corroborating evidence is to be used, such as the statement that
a taxon is “characteristic of a particular geographical area or geological
horizon” [Art. 45zf (ii) ]. A name first established with infrasubspecific rank
becomes an available name (Art. 106) when used for a taxon of the species
group and takes the date and authorship from the time of such change
in usage. It is taxonomic practice to give the benefit of the doubt to authors
having introduced “varieties” prior to 1961 (Art. 45c).
The indiscriminate naming of infrasubspecific forms has brought much
 Interpretation of the Rules of Nomenclature 363

discredit to taxonomy. The Latin language is traditionally used for the

names of taxa, and if one wants to designate infrasubspecific forms, it
is far better to use vernacular words from modern languages. The names
for the mutant forms of Drosophila are a good example. Linsley (1944)
has listed other kinds of infrasubspecific variants that should not be given
Latin designations. This includes sexual dimorphs, castes in social insects,
alternate generations, all kinds of morphs and polymorphs, seasonal forms,
pathological forms, and various kinds of aberrations and teratologies. Now
that zoologists clearly understand the difference between populations and
individual variants (phena), there is no longer any excuse for terminological
sloppiness.

42. AUTHORSHIP

The meaning of authorship and the rules concerning the citation

of names of authors are given in Arts. 50 and 51. Contrary to the statement
of the Code [Art. 51 b (i)], almost all zoologists customarily separate the
name of a taxon from that of a subsequent user (not its original author)
by a comma.
In view of the fact that an extraordinarily large number of species
were superficially and hurriedly described so that some vain author would
“get his name in print,” the whole institution of citing authors’ names
has been frequently attacked, not without reason. If zoologists have not
given it up, it is for the single reason that the name of an author is an
identifying label, particularly useful in the case of homonymy. The name
of the author does not form part of the name of the taxon, and its citation
is optional (Art. 51a).
The author of a name is the person who is alone responsible both
for the name and for “the conditions that make it available” (Art. 50),
that is, for the diagnostic description. Consequently, if someone other than
the author of the paper is responsible for a name and its availability, it
is this other contributor who shall be considered the author of the name
(Art. 51c). It is sometimes difficult to determine “responsibility” when
the unfinished manuscript of a zoologist is published after his death by
another zoologist. If the manuscript contains only the names, and the editor
supplied the diagnostic descriptions that “make the names available,” it
is the editor who is to be regarded as the author of these names, according
to Art. 50.
When there wrere only a few zoologists, abbreviations for the names
of authors were customary, like L. for Linnaeus or F. for Fabricius. So
little is gained by these abbreviations that they are no longer considered
good practice. Special problems arise when an author changes his name
364 Principles and Application of Zoological Nomenclature

during the period when he is actively publishing, or assumes a title (e.g.

Laporte to Comte de Castelnau). In these cases it is customary to cite
as author’s name the name under which the new taxon originally was
published. Names published anonymously after 1950 are no longer available
(Art. 14).

43. HOMONYMY

Homonyms are identical names for two or more different taxa. The
earliest of such names is the senior homonym; later ones are junior homo¬
nyms. Articles 52-60 of the Code deal with the validity of homonyms and
with replacement names for invalid homonyms. This area is one of the
most difficult in zoological nomenclature, and it is probable that future
congresses may modify some of the provisions, particularly where they seem
to be in conflict with either taxonomic freedom or stability.
A junior homonym in the genus group is always invalid (Arts. 53,
56). However even a single-letter difference prevents homonymy of generic
names.
There are some real conflicts with respect to the application of the
law of homonymy to species-group names. Homonymous specific names
originally published in the same genus (primary homonyms) necessitate
the renaming of the junior homonym. However, when two identical specific
names are secondarily brought together under one generic name, such
homonymy may be a matter of purely subjective taxonomic judgment. For
an author who rejects the lumping of the tw'o genera into one, no
homonymy exists, and there is no need whatsoever to reject the junior
name. The 1901 Code said that a junior homonym was to be rejected.
Some zoologists interpreted this to mean only primary junior homonyms,
others both primary and secondary junior homonyms. Article 59c copes
in part with this situation by stating very clearly that “a secondary homonym
is to be restored as the valid name whenever a zoologist believes that the
two species-group taxa in question are not congeneric.” Unfortunately, the
editorial committee added after the London congress that this provision
was applicable only to “a name rejected after 1960.” It did not make
clear what to do with subjective junior homonyms rejected prior to 1961
only by a minority of workers in a field. This again is an area where
different practices prevail in groups where there is only one specialist every
50 years and in those where there are simultaneously 10, 20, or 50 specialists.
In ornithology, for example, it has never been the practice to suppress
secondary homonyms permanently. For instance, in the 1950s one ornitholo-
 Interpretation of the Rules of Nomenclature 365

gist adopted an exceedingly broad generic concept for the Old World
flycatchers and lumped many well-known genera under the name Musci-
capa. As a result numerous specific names, some of them very well known
and of long standing, became for him junior homonyms, and he renamed
them. Nearly all other ornithologists rejected the sweeping lumping of the
first author and refused to accept his replacement names. Several recent
applications in BZN indicate that the custom of not accepting taxonomically
unjustified secondary homonymies is not limited to ornithology. Indeed
it would be an intolerable infringement of the taxonomic freedom guaran¬
teed in the Preamble if Art. 59c were rigidly interpreted.
On the other hand it would be in the interest of stability to adopt
a statute of limitation for the revival of names long and universally rejected
as secondary junior homonyms. Otherwise every generic split may disturb
stability on the species level.

44. ACTUALITY PRINCIPLE

The classification of all homonyms into primary and secondary

homonyms is an oversimplification. Blackwelder (1948) and MLU (p. 226)
present various examples of homonymy being complicated by changes in
the generic association of two potentially homonymous specific names. There
is a strong trend to get away from all ritualism and to sanction the renaming
of a specific name only when an author thinks that two specific names
are truly congeneric at the time of the discovery of the potential homonymy.
The principle of basing nomenclatural decisions on the actual situation
existing at the time when a potential conflict of names is discovered may
be designated as the actuality principle.
For instance, an African weaver bird was described by Cretzschmar
in 1827 as Ploceus superciliosus. Another weaver was described by Shelley
in 1873 as Hyphantornis super ciliosus. Eventually the latter species was
transferred to the genus Ploceus, but long after the original P. superciliosus
of Cretzschmar had been transferred to the genus Plocepasser. No homonymy
ever existed in the sense that any ornithologist ever thought the two species
were congeneric. As a result even the most recent monographs and catalogs
retain superciliosus as valid specific name in both respective genera.
A further brake to needless renaming is applied in Art. 57c, which
states that homonymy does not exist when two identical species-group names
are used in entirely different genera that merely happen to have homony¬
mous names, such as when the name Noctua at one time was used both
for some birds and for a group of Lepidoptera.
366 Principles and Application of Zoological Nomenclature

45. REPLACEMENT NAME (FIRST REVISER)

Articles 56c and 57c deal with the rare eventuality of the simultaneous
publication of two homonyms now placed in different categories. In order
to serve best the interests of stability, the mandatory provision is made
that if one of the two names is for a subcategory (e.g. subgenus versus
genus, or subspecies versus species), the name proposed for the full category
takes precedence over the one proposed for the subcategory. As in other
cases of renaming of homonyms, the “actuality principle” should be invoked.
When at the time of the discovery of homonymy one of the homonyms
is the name of a subgenus or of a subspecies, this is the homonym to
be renamed, in order to preserve the well-established older name for the
full genus or species.

46. HOMONYMY AMONG SUBSPECIES

All specific and subspecific names in a genus are potentially homony¬

mous (Art. 57). As the example given in this article illustrates, it is not
permissible to use the same subspecies name in different species of the
same genus.

47. REPLACEMENT OF REJECTED HOMONYMS

Provisions governing the publication of replacement names are given

in Art. 60. Before proposing a new name as a replacement name (nomen
novum) for one which is preoccupied, an author must be certain that it
meets these conditions:

1. That there is no other name available for the species (or genus). In
the past there have been a few nomenclaturists who published replacement
names for all junior homonyms whenever a catalog or nomenclator was
published. Since in most of these cases replacement names had already
been provided by specialists of the respective groups, such wholesale
renaming resulted in nothing but an added burden to synonymies and
a loss of respect for the author of these unnecessary names.
2. That the original author of the preoccupied name is no longer alive.
According to paragraph 3 of the Code of Ethics (Code, Appendix A,
p. 93) a zoologist should communicate with the author of a junior
homonym, if he is still alive, and give him a reasonable opportunity
to publish a replacement name.
3. That the new name is proposed in the form recommended in the Code.
As stated in Art. 13a(ii) and as recommended by several zoological
congresses, there must be “a full bibliographic reference” (not merely
 Interpretation of the Rules of Nomenclature 367

“Smith, 1907”) to the original citation of the preoccupied name, and,

in the case of preoccupied generic names, to the name of the type-species
(Appendix E, 15, p. 145).
4. That it is desirable to propose a replacement name. For instance, there
is no excuse for renaming preoccupied names that are invalid synonyms.
More importantly, a replacement name automatically takes the same
type and the same type-locality (in case of a species) of the preoccupied
name. If the type of the preoccupied name is no longer in existence,
or if there is even the slightest doubt as to the identity of the species
with the preoccupied name, it is sometimes preferable simply to redescribe
the taxon in question, as if entirely new, and to provide it thus with
an unambiguous type and type-locality.

Under exceptional circumstances a homonym may provide an oppor¬

tunity for shifting an originally ill-chosen type-locality of a subspecies. For
example, let us assume that there is a species with a northern and southern
subspecies, meeting in a narrow zone of intergradation. The type-locality
of the southern subspecies is far in the south, but the type-locality of the
population whose name had always been applied to the northern subspecies
is actually located in the zone of intergradation. If it is found that this
name is preoccupied, it is better not to replace it by a nomen novum
but to redescribe the northern subspecies and select a new type-locality
in the middle of its range. The number of cases where such a shift of
type-locality would be desirable is undoubtedly very small; in most cases
it would only be confusing. A genus (or species) divided into subtaxa
takes the name of the next oldest available subtaxon if the name of the
nominate subtaxon is found to be a junior homonym (see 13.36 and Arts.
446, 476).

48. THE TYPE METHOD

In taxonomic practice doubt often arises as to the identity of taxa

to which names are attached. The descriptions are frequently insufficient
for establishing such identity, particularly the rather short descriptions of
earlier authors. Sometimes a description applies equally well to several
species since the species-specific diagnostic characters were not mentioned
in the early description. In the case of higher taxa, the contents of these
taxa change as additional species are discovered. When such a higher taxon
is split, the question arises as to which of the components the name should
be attached to. It is obvious that a secure standard of reference is needed
to tie taxonomic names unequivocally to definite, objectively recognizable
taxa. These standards are the types, and the method of using types in
order to tie names to taxa is called the type method.
Few current workers in taxonomy realize how comparatively modern
the type method is. Indeed its introduction during the last 100 years was
368 Principles and Application of Zoological Nomenclature

one of the major conceptual changes in the theory of taxonomy and nomen¬
clature. Early taxonomy was dominated by the Aristotelian concept of types.
All specimens that conformed to the taxonomist’s concept of the type of
a taxon were considered typical. The types—and there were as many types
as there were typical specimens—formed the basis of the description of
the species. This was the concept of type held by Linnaeus and his con¬
temporaries. As is well described by Cain (1958) and Svenson (1945),
Linnaeus never hesitated to replace specimens in his collections which we
would now consider as types with “better ones.”
“Linnaeus never designated any specimens as type. Whether his de¬
scription was based on one single or on several specimens, it cannot even
be taken for granted that [these] were preserved in his collection. Or,
the original specimen may later have been substituted by another, in better
condition, by Linneaus himself” (Lindroth, 1957). Macan (BZN, 18:328)
remarks, “The Linnaean collection is notoriously unreliable since many
of the original specimens . . . were replaced later by other specimens,”
an observation confirmed by Lindroth and other Linnaean scholars. No
nomenclatural decision should ever be made by relying on a “Linnaean
type.” There is no such thing. Consequently, “If description and authentic
specimen disagree, the former is decisive” (Lindroth, loc. cit.). This point
must be made emphatically because there have been a few ill-advised at¬
tempts in recent years to change well-known Linnaean names because imagi¬
nary types were found to belong to species different from those to which
the name had always been applied.
Linnaeus was not alone in these practices. It was customary in several
European museums in the first half of the nineteenth century to substitute
“new” type-specimens whenever the old ones became faded or were dam¬
aged by insect pests. This was quite legitimate under the Aristotelian type-
concept. In other instances, the inadvertent transfer of labels from one
specimen to another has caused an obscuring of the identity of type-speci¬
mens. Evidence derived from old types must be treated with extreme care
and discrimination and never be used to upset stable nomenclature. In
such cases, as with Linnaean “types,” if description and so-called authentic
specimen disagree, the former is decisive.
Consistent with the Aristotelian type-concept, typification of higher
taxa was done by the process of “elimination” of atypical elements. This
was the standard method, practiced until far into the nineteenth century,
and it was one of the most upsetting decisions of the 1901 Code not to
ratify this traditional procedure in the case of old type-fixations but to
apply retroactively the modern type method.
The history of the gradual adoption of the type method after about
1850 has not yet been written. How little the importance of this change
was realized is indicated by the fact that the original 1901 Code does
not include any directives concerning types. Provisions for generic types
 Interpretation of the Rules of Nomenclature 369

were adopted at the Boston congress in 1907, and for the types of species
even later. Some authors who speak of allotypes, plesiotypes, metatypes,
homotypes, hypotypes, etc., still do not fully understand the true nomen-
clatural function of the type-specimen. Such pseudotypes have no nomen-
clatural significance, but may be helpful in identification. Even in the new
Code there are still relics of the Aristotelian type-concept, such as the
provision (Art. 72b) : “The type-series of a species consists of all the speci¬
mens on which its author bases the species,” as if it were the function
of the types to form the basis of a species.
Species consist of variable populations, and no single specimen can
represent this variablity. No single specimen can be typical in the Aristo¬
telian sense. As Simpson (1961) has discussed perceptively, the only
function of the type-specimen is to be a “name bearer.” Indeed he even sug¬
gested that one might drop the misleading term “type” and call the name¬
bearing specimen the onomatophore (Greek for name-bearer). The term
type, however, is too firmly fixed in the taxonomic tradition for such a
change to be practical. The younger generation of taxonomists now under¬
stands clearly that a type is nothing more and nothing less than a specimen
(or taxon) which tells us to which taxon a given name should be attached.
The term type-series is an anachronism with respect to the current concept.
A type is always a zoological object, never a name. The type of
a genus is a species, the type of family is a genus (Art. 61). This is important
in the case of “misidentification of the type,” that is, in a case where
an author, when designating the type of a new taxon, refers to it under
a wrong name (13.49). Once designated the type cannot be changed,
not even by the author of the taxon, except—by exercise of the plenary
powers of the Commission (Art. 79)—through the designation of a neotype
(Art. 75).
Description of a new species is based on the entire material available
to the zoologist, including the type-specimen. It is not the function of
the type to serve as the exclusive or primary basis of the description. Simp¬
son (1961, pp. 183-186) discusses this aspect fully. He also introduces the
term hypodigm for the entire sample of specimens personally known to
a given taxonomist at a given time and considered by him to consist of
unequivocal members of the taxon. Disagreements among taxonomists are
often caused by the fact that they have studied different hypodigms.

49. CORRECTING THE MISIDENTIFICATION OF TYPES

It is unfortunately true that the type-species of new genera or the

type-genera of new taxa of the family group have sometimes been misidenti-
fixed (= misnamed) by the original authors of the new taxa. Articles 41,
49, 65£>, 67;’, and 70a contain provisions about how to correct the error
370 Principles and Application of Zoological Nomenclature

of the original author. The principle on which such corrections are based
is that the type of a taxon is not a name but a zoological object. The
type (-species or -genus) is then the zoological object which the original
author had before him (when making the type designation) and not the
name which he may have erroneously attached to this object.
However, it would be an intolerable burden on the working taxonomist
if he had to make sure in each case that the original type was correctly
identified and named. It is therefore provided in the Code that a zoologist
must assume that the author correctly identified this type (Arts. 65a, 70).
If there is strong or clear evidence that a misidentification is involved,
the case is to be referred to the Commission. The Commission in such
cases tends to make a ruling that will maintain stability and continuity.
If reexamination of the type of a well-known species proves that
the type actually belongs to a different species, the Commission can by
its plenary powers (Art. 79) suppress the original type and designate a
neotype (see 13.55) which conforms to the accepted concept of the species.
Wisely administered, the various provisions on misidentification can
do much to preserve stability and continuity in nomenclature.

50. TYPE OF REPLACEMENT NAME

Articles 67i and 72d unequivocally provide that if the name of a

taxon is expressly proposed as a replacement name for a prior name, it
retains the type of the prior name.
Former generations of zoologists did not necessarily follow this rule,
and many names, seemingly replacement names, clearly have a composite
basis. They are based on the description of specimens (including a new
type) but also refer to a preoccupied name. A retroactive application of
the new Art. 72d to such composite names, when the name has been uni¬
versally applied to the newly designated type-material for 50 years or more,
would surely be in conflict with the stability concept expressed in the
Preamble.

51. KINDS OF TYPE-SPECIMENS

Since the type-specimen is the official standard of reference for the

name of a species, it can have full authority only if it is unique (Art.
456). When there are two or more type-specimens, it has all too often
been found that they actually belong to different species. Which of these
species then shall have the name? The new Code clearly specifies (Art.
72a) that a taxon of the species group (species or subspecies) can have
 Interpretation of the Rules of Nomenclature 371

only a single type, either the specimen designated or indicated as the type
by the original author at the time of publication of the original description
(holotype) or one designated from the type-series (lectotype) [misprinted
in the Code], or a neotype.
Specimens before the original describer that are neither the holotype
nor the lectotype (of a subsequent author) are conventionally referred
to as paratypes. Paratypes have no special standing under the Code and
do not qualify as types, by the exclusion clearly formulated in Art. 72a.
Recognition of a “type-series” in Art. 72b introduces a contradiction.

52. TYPE DESIGNATION

Since the information concerning the type-specimen is nomenclaturally

the most important information given on the occasion of naming a new
taxon, it has become customary in taxonomic literature to record this infor¬
mation immediately following the new name. The information concerning
the type takes the same place in the sequence as does the synonymy in
a redescription (see 11B.1.7-1 IB.2).
The definite fixation of a type-species is, after 1930, a prerequisite
for making a name in the genus group available (Art. 13b). Designation
of a type-specimen is not a prerequisite for making a name in the species
group available. In certain groups, for instance certain genera of protozoans,
it is exceedingly difficult to preserve specimens in such a way that they
can serve as a permanent standard of reference. Yet even in protozoology
the designation of type-specimens has increasingly become standard practice
(Corliss, 1963). For rules concerning types in the species group, see Arts.
71-75.
Even though the publication of information on the type-specimen
of a new species-name is not mandatory, it is nevertheless expected from
the zoologist that he should clearly designate a single specimen as the holo¬
type of the new species and that he should supply in the original description
the following information, in addition to measurements and other descrip¬
tive data, characteristic for the type-specimen:

1. Precise collecting locality and other relevant data on the labels of the
specimen
2. Sex
3. Developmental stage or form (if significant) to which the type is referable
4. In the case of parasites, name of the host species
5. Name of the collector
6. Collection in which the holotype is deposited and, when specimens are
numbered, the number assigned to it
372 Principles and Application of Zoological Nomenclature

7. Altitude of the type-locality or depth in meters below sea level at which

the holotypc was taken
8. In the case of fossil species, geological horizon

Great care should be exercised by the author, if a large hypodigm

is available, to select as the type a specimen which—owing to its state
of preservation, sex, age, or locality data—is most suitable as a name-bearer.
The diagnostic characters relevant in a given genus are sometimes better
accessible in some specimens than in others. The type should have maximal
usefulness for taxonomic discrimination.
In the case of fossil material, if the hypodigm consists of many indi¬
vidual pieces (e.g. bones), it is advisable to designate the most diagnostic
of them as type, particularly if there is even the slightest doubt as to whether
the pieces actually belong to a single individual. Many “types” of formerly
described fossil species have, on reexamination, turned out to be composites
of several different species.
The following additional advice may be offered with regard to types:

1. Type designation or fixation should always be completed before

publication.
2. Type designation should be clear and unambiguous; deposition (and
museum number) of type should always be recorded.
3. Types of undescribed species should not be distributed prior to publication.
4. Type labels should never be changed or removed.
5. Type fixation for species of older authors should be attempted only
by a specialist during revisionary work.

53. TYPE-MATERIAL

Type-specimens, as the official standards for the names of species,

“are to be regarded as the property of science,” according to Art. 72/.
Recommendations 72A, 72B, 72C, and 72D specify the responsibilities of
the caretakers of such type-material. All types should be transferred as
quickly as possible to public institutions where their safety is guaranteed
and where they are accessible to other research workers. In most museums
distinctive labels are used to indicate clearly that a given specimen is a
type.
Types should not be handled in connection with ordinary identification
work. However, monographers should have free access to types, particularly
when there is doubt as to the zoological identity of a type-specimen. Most
museums are increasingly ready to mail types to specialists. In case of
loss there is always the possibility of replacing a type by a neotype (Art.
75). For further comments on type collections see also 6.2.5.
 Interpretation of the Rules of Nomenclature 373

54. LECTOTYPES

If the name of a species was based on a series of syntypes, any zoologist

may designate one of these syntypes as the lectotype (Art. 74 and Recom¬
mendations 74A, 74B, 74C, and 74D). Selection of a lectotype should be
undertaken only by a specialist during revisionary work and ordinarily only
if it contributes to the unambiguous affixation of a name to a given taxon.
It should never be done merely in order to add a type-specimen to the
collection. If the description of a species is clearly based on a particular
specimen, that specimen should be made the lectotype. If one of the syntypes
was illustrated, it should be selected as lectotype, other things being equal.
Syntypes are often widely scattered as the result of exchanges. This
requires special considerations in lectotype selection. When possible, a lecto¬
type should be chosen from syntypes in the collection of a public institution,
preferably of the institution containing the largest number of syntypes of
the species, or containing the collection on which the author of the nominal
species worked, or containing the majority of his other types.
Many classical authors clearly designated one specimen as the type
in their collections without specifically citing such a specimen as “the type”
in the published description. The present wording of Art. lib would imply
that such a specimen does not qualify as the holotype. Such specimens
have traditionally been accepted as holotypes, in contrast to cases where
an author labeled numerous syntypes as “type.” It would seem that Art.
73b could well be modified to legitimize the traditional practice (see Fen-
nah, 1957, and Young, 1958, SZ, 7:120-122).
In cases where the syntypes are from several localities and a previous
reviser has already restricted the type-locality of the species, a responsible
zoologist will give due consideration to this fact in the selection of the
lectotype.

55. NEOTYPES

In Art. 75 the Code regulates the designation of neotypes: “If through

loss or destruction no holotype, lectotype, or syntype exists.” The provision
specifically forbids the manufacturing of neotypes as a matter of curatorial
routine or for the sake of having a type for every species. Most of the
older species of birds, for instance, have no types but are such clearly
understood taxa that no type is needed. Neotypes ought to be designated
“only in connection with revisory work, and then only in exceptional circum¬
stances, when a neotype is necessary in the interest of stability of nomencla-
374 Principles and Application of Zoological Nomenclature

ture” (Art. 75a). Even when the original type is damaged or even lost,
it is in most cases quite unnecessary to designate a neotype.
Least desirable is for a neotype to be designated for a species whose
name is not in general use either as a valid name or as a synonym, or
for any name, not in use, that is a nomen dubium. No author should
designate a neotype until he has carefully checked that his action conforms
entirely with all the provisions of Art. 75. (See 75c for neotypes designated
prior to 1961.)
The Commission has the power (Art. 79) to suppress an existing
type if this is in the interest of stability of nomenclature and to designate
a neotype to conform with the traditional usage of a name. For instance,
some years ago it was discovered that the type of the Hottentot Teal (Anas
punctata) in the Oxford University Museum actually belonged to the Mac-
coa Duck (Oxyura maccoa) and that a most confusing switching of names
was inevitable if this type-specimen were not suppressed. The Commission
designated a neotype for the species, and the stability of names that had
existed for the previous 125 years was preserved. If a zoologist finds that
the type of a well-known species has been misidentified, he can, if stability
is seriously threatened, apply to the Commission for a suppression of the
type and the designation of a neotype conforming to the traditional usage
of the name. Such action should be confined to exceptional cases.

56. TYPES AND FIRST REVISERS

There are a number of situations in which a zoologist may have

to undertake an action in order to clarify the status of a type.

A. Type of a Species
1. If a series of syntypes belongs to two or more species, a zoologist must
determine whether or not a previous first reviser restricted the name
to one of the components. If not, he has to select a lectotype from
among the syntypes in such a way that it best serves stability of nomen¬
clature and is in accordance with the provisions of Art. 74 and the
comments given in 13.54.
2. If the type is found not to belong to the species to which the name
is traditionally applied, two avenues are open. In inactive groups the
zoologist may prefer simply to shift the name to the correct species.
However, if such action would cause a serious disturbance of stability,
particularly in the case of names in active and universal use for more
than 50 years, the zoologist may request the International Commission
to use its plenary powers to suppress the original type and designate
a neotype conforming to existing usage (see 13.55).
3. The Code contains no provisions regarding what to do when the type
 Interpretation of the Rules of Nomenclature 375

lacks all diagnostic characters. In the case of a forgotten name or nomen

dubium, such a name is best transferred to the Official Index of Rejected
Names. In the case of names in general use, a neotype may be designated
by the Commission provided such an action is “essential for solving
a complex zoological problem” (Art. 75). In most cases no such neotype
designation will be needed.

B. Type of a Genus
1. If no species is included in the original naming of a genus, published
before 1931, follow the provisions of Art. 69a (ii).
2. If none of several included species has been previously designated as
type-species, the first reviser will again act according to the provisions
of Art. 69.
3. If the evidence indicates that the name used for the previously designated
type-species resulted from misidentification, the provisions of Art. 70
are to be followed. In such cases the Commission is charged “to designate
as the type-species whichever species will in its judgment best serve
stability and uniformity of nomenclature.”

57. TYPE-LOCALITIES

The type-locality is the place where the population occurs from which
the type-specimen was taken. Specimens collected at the type-locality are
called topotypes, and the population that occurs at the type-locality is called
the topotypical population. The 1961 Code does not contain any mandatory
provisions concerning type-localities, but Recommendation 72E contains
advice on their designation or restriction. The comments contained in the
ensuing paragraphs represent the best current practices, but they are not
part of the Code. As in the case of lectotypes or neotypes, the designation
or restriction of type-localities should not be done routinely but only by
a specialist in connection with revisionary work.
Species can ordinarily be identified by single specimens, subspecies
often only be adequate population samples. The type-locality, consequently,
is unimportant at the species level but frequently decisive for the determina¬
tion of the validity of subspecies. In view of the not infrequent overlap
in the characteristics of subspecies, a single specimen can be the name-bearer
(“type”) for a subspecies only to the extent that it helps to identify the
population from which it was sampled. Where it fails to do this, a knowl¬
edge of the type-locality becomes a necessity.
Original Designation of a Type-locality. When describing a new spe¬
cies or subspecies a worker often has before him material from many locali¬
ties within the range of the new taxon. It is his duty to make as prudent
a choice of the type-locality as possible. In this he should be guided by
the following considerations, among others:
376 Principles and Application of Zoological Nomenclature

1. To choose a type-locality from which are available many topotypes which

constitute a fair sample of the population and illustrate its variation.
2. In the case of a variable species or subspecies, to place the type-locality
in the area inhabited by the population that the describer considers most
typical for the new taxon.
3. In the case of a new subspecies that is part of a cline, to place the
type-locality at the other end of the cline from the location of a previously
described subspecies.
4. Never to place a type-locality in an area of intergradation or
hybridization.
5. To record, in the description of a new taxon, the location of the type-
locality as precisely as possible. Taxonomically distinct populations of
a species sometimes occur only 100 m apart from each other. In paleontol¬
ogy a few centimeters vertically may mark a change from one bed to
another. This is why type-localities must be fixed with extreme accuracy.
It is also the reason why it is so important that the collector give precise
locality data on the labels of the specimens (see Chap. 6). When such
data are lacking, it is sometimes possible to obtain them from the collector
if he is still alive. In other cases the information may be found in
published or unpublished journals or fieldbooks.

Restriction of a Designated Type-locality. Earlier authors, not appre¬

ciating the need for exact type-localities, often described new species from
“California” or “Brazil” or “Africa.” When later collections indicate that
the species from “Brazil” is geographically variable and consists of two
or more subspecies, it becomes necessary to determine the exact locality
from which the type of the nominate subspecies came.
Most workers accept the principle that the “first reviser,” the person
who first realizes the geographical variability of such a species, has the
right to designate arbitrarily a more restricted type-locality, provided that
evidence derived from a study of the type itself does not contradict his
designation. Such a fixation is followed unless it can be shown that the
action of the first reviser is erroneous. Obviously, if the first reviser restricts
to Rio de Janeiro the type-locality of a species from “Brazil,” his restriction
should not be binding if the type is still in existence and belongs to a
subspecies which is confined to the neighborhood of Cayenne. To avoid
such mistakes, the first reviser should make a careful investigation of the
probable route of the collector. Even in the absence of exact information
certain conclusions may be obvious; a type collected in China in 1775
most likely came from Canton or some part of Fukien, not from Szechuan,
Kansu, or some other place far in the interior.
In the case of a “Voyage,” it is often possible to determine the exact
locality by a study of the course of the voyage. For example, a small owl,
Ninox ocellata, collected by the Voyage au Pole Sud, was described by
Hombron and Jacquinot as having come from Chile, South America. This
 Interpretation of the Rules of Nomenclature 377

is an obvious error, since the genus does not occur in the Americas. Later
on, Mathews, believing ocellata to be an earlier name of N. roseoaxillaris
Hartert (1929, San Cristobal, Solomon Islands), restricted the type-locality
of ocellata to San Cristobal. However, it is stated in the report of the
above-mentioned voyage that the expedition landed in the Solomon Islands
only on Ysabel Island (and adjacent St. George), where no owl resembling
ocellata occurs. Mathews’ restriction of the type-locality is therefore un¬
tenable. Subsequently it was shown by Peters that the Coburg Peninsula,
Northern Territory, Australia, is the only locality touched by the voyage
where an owl occurs that agrees with the description of N. ocellata. Peters
therefore restricted the type-locality to Coburg Peninsula, and this restriction
has been universally accepted. This case clearly shows how much care must
be exercised in the correction or restriction of type-localities.
Correction of a Wrong Type-locality. There are two sets of circum¬
stances under which an error in the originally stated designation of the
type-locality can be corrected:

1. Exact type-locality given in the original description. If the author or

some subsequent worker can prove beyond doubt that the type(s) did
not come from the locality given in the original description (owing to
some error or misinformation), he can shift the type-locality to the place
from which the type really came, or, at least, where the taxon is known
to occur. Actually this is not a shift of the type-locality but only of
the “stated” type-locality, since the type never came from the originally
designated locality.
A type-locality cannot be altered because an author finds that the
population at a different locality is “more typical” or because he has re¬
ceived “better material” from a new locality. Proposals for the shift of
type-localities for these and similar reasons must be rejected.
2. Exact type-locality not given in the original description. If no type-locality
is given, or if only a vague one (“India”), the first reviser may designate
a restricted type-locality. Such a restriction may later be set aside if
it conflicts with the available evidence, but only if the case is unequivocal.
The fixation of a type-locality cannot be set aside because that locality,
at the time of the collection, was “less accessible” than some other locality,
or because the species is “rather rare” at that locality. It should be
changed, however, if it is clearly outside the range of the species.

Designation of a lectotype is equivalent to a restriction of the type-

locality if the original syntypes came from several localities. If there is
a conflict between lectotype selection and restriction of type-locality, lecto-
types take precedence. A type-locality should never be changed or restricted
in such a way that it upsets the stability of nomenclature.
When designating a lectotype an author should take into careful con¬
sideration the possibility of a prior restriction of type-locality (see 13.54).
378 Principles and Application of Zoological Nomenclature

58. NOMINAL TAXON

The Code frequently refers to “nominal species” or to “nominal”

combined with some other category (rank) designation. The expression
nominal species means a species for nomenclatural purposes without refer¬
ence to its taxonomic status. Such a species is indicated exclusively by
its name, and the name, in turn, is defined by its name-bearer (= type).
The term is therefore correctly defined in the glossary of the Code as
“a named species, objectively defined by its type-specimen.” Equivalent
definitions pertain to nominal genus and nominal family.

59. INTERNATIONAL COMMISSION ON ZOOLOGICAL

NOMENCLATURE

The duties, powers, organization, and operation of the Commission

are regulated by Arts. 76-82 of the Code, the Constitution of the Commis¬
sion {BZN, 21(3) : 181-185), and the By-laws (BZN 22(1) :3-8).
Since this is often misunderstood, it must be pointed out that the
rules of nomenclature are phrased and adopted by the Zoological Con¬
gresses. These are the legislative bodies responsible for the rules. The Com¬
mission is authorized by these International Congresses, as stated in Arts.
76-79, to interpret or suspend provisions of the Code in individual cases
and to submit to the Congresses recommendations for the clarification or
modification of the Code.
In particular the Commission is empowered to suspend the application
of any provision of the Code “if such application to a particular case
would in its judgment disturb stability or universality or cause confusion”
(Preamble and Art. 79). Under these plenary powers it may “annul or
invalidate any name, type-designation, or other published nomenclatural
act, or any publication, and validate or establish replacements,” by a two-
thirds majority.

60. APPLICATIONS TO THE COMMISSION

Any zoologist may submit cases involving nomenclatural problems

to the Commission (Art. 78). It must be remembered, however, that the
Commission is a judicial, not a fact-finding board. It is under no obligation
to supplement or verify information contained in applications. A zoologist
planning to submit an application will do well to study the style of such
 Interpretation of the Rules of Nomenclature 379

applications in recent volumes of BZN and also to circulate his application

first informally among cospecialists. The processing and publication of the
applications is described in Art. 17 of the Constitution, and the By-laws,
Sec. IIIC [BZN 21 (3) and 22(1)] respectively.

61. CASES UNDER CONSIDERATION

For the sake of stability in nomenclature it is of extreme importance

that zoologists obey the provisions of Art. 80. This states that, while an
application to the Commission is pending, “existing usage is to be main¬
tained, until the decision of the Commission is published.”

62. AMENDMENTS TO THE CODE

As stated in Art 87, amendments to the Code can be made only

by an international congress of zoology. All such amendments are first
processed by the Commission and are submitted to the Congress only if
approved by a majority of the section of nomenclature of that Congress
(Arts. 77 and 78a). In accordance with its legislative authority the Congress
can vote to add or delete articles and to review any decision of the Commis¬
sion (Art. 78g).

63. OFFICIAL LISTS AND INDEXES

Article 14 of the Constitution provides that “the Commission shall

compile and maintain” the following lists and indexes :

Official List of Family-group Names in Zoology

Official List of Generic Names in Zoology
Official List of Specific Names in Zoology
Official Index of Rejected and Invalid Family-group Names in Zoology
Official Index of Rejected and Invalid Generic Names in Zoology
Official Index of Rejected and Invalid Specific Names in Zoology
Official List of Works Approved as Available for Zoological Nomenclature
Official Index of Rejected and Invalid Works in Zoological Nomenclature

Section III (A) of the By-laws delegates to the secretariat of the Commission
responsibility for “the preparation and editing for publication of official
lists and indexes.” Provisions in the Code pertaining to these official lists
and indexes are found in Arts. 23b, 23d, 77, and 78/.
380 Principles and Application of Zoological Nomenclature

64. APPENDICES TO THE CODE

Attached to the Code are five appendices to serve as a guide to

good usage in nomenclature. They are not mandatory, as are Arts. 1-87
of the Code proper, but rather have the same status as Recommendations
in the Code.
Appendix A (Code, p. 93) is a Code of Ethics. Every zoologist should
carefully study the eight paragraphs of this appendix in order not to violate
well-established conventions and risk losing the respect of his colleagues.
For instance, no zoologist should publish a replacement name for a junior
homonym during the lifetime of its author without following the procedure
of Paragraph 3 of the Code of Ethics. A new name proposed in violation
of these provisions is, however, available if it otherwise satisfies the provi¬
sions of the Code.
Appendix B deals with the transliteration and latinization of Greek
words (Code, pp. 95-101).
Appendix C deals with the latinization of geographical and proper
names (Code, pp. 101-103).
Appendix D contains extraordinarily detailed and helpful recommen¬
dations on the formation of names (Code, pp. 105-141). Since every zoolo¬
gist must have his own copy of the Code handy at all times, we have
not included a corresponding section in this text.

65. GENERAL RECOMMENDATIONS

Appendix E (Code, pp. 143-147) contains a useful summary of recom¬

mendations to the working taxonomist. The beginner, in particular, should
carefully study the 24 recommendations, because this will surely help him
to avoid making mistakes or at least to improve the quality of his publica¬
tions. Numbers 15, 16, 22, and 24 are of special importance.

66. GLOSSARY AND INDEX

If a zoologist is in doubt as to the meaning of a word used in the

Code, he should consult the Glossary to be found on pp. 148-154 (English)
and pp.155-161 (French).
Even the experienced worker sometimes has difficulties in finding the
exact article containing provisions concerning a given case. Even though
there are only 86 articles, there are over 600 individual provisions in the
Code and in the appendices (exclusive of the tables). The index of the
Code (pp. 163-176) is an invaluable guide to these provisions. Items printed
in boldface in the index refer to the glossary.
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Wiegmann’s Archiv fur Naturgeschichte. 1835. et seq. Bericht fiber die Leistun-

gen im Gebiete der Naturgeschichte wahrend des Jahres 1834 [. . . 1922], Berlin.
Williams, C. A., Jr. 1964. Immunochemical analysis of serum proteins of the
primates: a study in molecular evolution. In J. Buettner-Janusch (ed), Evolutionary
and genetic biology of the primates. Academic Press Inc., New York, no. 2, pp.
25-74.
Wilson, E. O. 1965. A consistency test for phylogenies based on contemporaneous

species. Syst. Zool., 14:214-220.

-, and W. L. Brown. 1953. The subspecies concept and its taxonomic applica¬
tion. Syst. Zool., 2:97-111.
Wirth, M., G. F. Estabrook, and D. J. Rogers. 1966. A graph theory model for

systematic biology with an example for the Oncidiinae (Orchidaceae). Syst. Zool.,
15:59-69.
Wood, A. E. 1950. Porcupines, paleogeography, and parallelism. Evolution, 4:87-98.

Wood, C. A. 1931. An introduction to the literature of vertebrate zoology. Oxford

University Press, London, xii—xix 643 pp.
Wright, C. A. 1966. Experimental taxonomy: A review of some techniques and

their applications. Intern. Rev. Gen. Exptl. Zool., 2:1-42.

Wright, A. H., and A. A. Wright. 1949. Handbook of frogs and toads. Cornell
University Press (Comstock), Ithaca, N.Y., xii -(- 640 pp.
- and -. 1957. Handbook of snakes. Comstock Press Associates, Cornell
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Zweifel, F. W. 1961. A handbook of biological illustration. Phoenix Books, Uni¬
versity of Chicago Press, vii -f-131 pp.
Glossary

Accessory sexual characters. The structures and organs (except the gonads) of
which the genital tract is composed, including accessory glands and external
genitalia (cf. Secondary sexual characters).
Adaptation. The condition of showing fitness for a particular environment, as
applied to characteristics of a structure, function, or entire organism; also the
process by which such fitness is acquired (cf. Environment).
Adaptive radiation. Evolutionary divergence of members of a single phyletic line
into a series of rather different niches or adaptive zones.
Affinity. Relationship. Sometimes misleadingly employed as synonym for phenetic
similarity.
Agamic. A species or generation which is not reproducing sexually.
Agamospecies. A species without sexual reproduction, an asexual species.
Albinism. In zoology, the absence of pigmentation, and particularly of melanins,
in an animal (cf. Melanism).
Allele. Any of the alternative expressions (states) of a gene (locus).
Allochronic species. Species which do not occur at the same time level (cf.
Synchronic species).
Allometric growth. Growth in which the growth rate of one part of an organism
is different from that of another part or of the body as a whole.
Allopatric. Of populations or species, occupying mutually exclusive (but usually
adjacent) geographical areas.
Allopatric hybridization. Hybridization between two allopatric populations (species
or subspecies) along a well-defined contact zone (cf. Sympatric
hybridization).
Allopatric speciation. Species formation during geographical isolation (cf. Sym¬
patric speciation).
Allotype. A paratype of the opposite sex to the holotype (cf. Paratype).
Alpha taxonomy. The level of taxonomy concerned with the characterization
and naming of species (See 1.4.3).
Alternation of generations. The alternation of a bisexual with a unisexual (parthen-
ogenetic) generation.
Amphiploid. A polyploid produced by the chromosome doubling of a species
hybrid, that is, of an individual with two rather different chromosome sets.
Analogous. A similar feature in two or more taxa which cannot be traced back
to the same feature in the common ancestor of these taxa (cf. Homologous,
and 4.8.1).

397
398 Glossary

Anatomy. The science of internal morphology, as revealed by dissection.

Antibody. A serum globulin which is produced in the blood of an immunized
animal in response to the introduction of a foreign antigen (cf. Antigen,
Antiserum, Serology).
Antigen. A substance capable of inducing the formation of antibodies when intro¬
duced into the bloodstream of an animal (cf. Antibody, Precipitin reaction,
Serology).
Antigenic. With the properties of an antigen.
Antiserum. Blood serum containing specific antibodies (cf. Antibody, Precipitin
reaction).
Apomorph. A derived character.
A posteriori weighting. The weighting of taxonomic characters on the basis of
their proved contribution to the establishment of sound classifications, i.e.
of monophyletic taxa.
A priori weighting. The weighting of taxonomic characters on the basis of precon¬
ceived criteria, e.g. their physiological importance.
Archetype. A hypothetical ancestral type arrived at by the elimination of special¬
ized characters (cf. Phylogeny).
Artenkreis (Rensch). Superspecies (q.v.).
Artificial classification. Classification based on convenient and conspicuous diagnos¬
tic characters without attention to characters indicating relationship; often
a classification based on a single arbitrarily chosen character instead of
an evaluation of the totality of characters (cf. Classification, Phylogeny).
Asexual reproduction. Not involving the fusion of the nuclei of different gametes.
Atlas. In taxonomy, a method of presenting taxonomic materials primarily by
means of comparative illustrations rather than by comparative descriptions
(cf. Monograph).
Authority citation. The custom of citing the name of the author of a scientific
name or name combination [e.g., X-us Jones, X-us albus Jones, Y-us albus
(Jones)]. (See 13.42).
Autopolyploid. A polyploid originating through the doubling of a diploid chromo¬
some set.
Autosome. One of the chromosomes other than a sex chromosome.
Available name. A name published in a manner satisfying the requirements speci¬
fied in Arts. 8-20 of the Code (cf. Valid name, and 13.16).
Baculum. An ossification (bone) in the phallus of some mammals.
Beta taxonomy. The level of taxonomy concerned with the arranging of species
into a natural system of lower and higher taxa (cf. Alpha taxonomy, Gamma
taxonomy; see 1.4.3).
Bibliographical reference. For nomenclatural purposes, the citation of the name
of the author and date of publication for a scientific name; a full biblio¬
graphical reference includes, in addition, the citation of the exact place
of publication of a scientific name (i.e., title of book or journal, volume,
page, etc.). (See 11.B.2.)
Binary. Refers to designations consisting of two kinds of names (see 13.11; cf.
Binominal nomenclature).
 Glossary 399

Binomen. The scientific designation of a species, consisting of a generic and a

specific name.
Binomial nomenclature. The system of nomenclature first standardized by Lin¬
naeus and now generally referred to as binominal nomenclature.
Binominal nomenclature. The system of nomenclature adopted by the International
Congress of Zoology, by which the scientific name of an animal is designated
by both a generic and specific name (cf. Binary nomenclature).
Biological classification. The arranging of organisms into taxa on the basis of
inferences concerning their genetic relationship.
Biological races. Noninterbreeding sympatric populations, which differ in biology
but not, or scarcely, in morphology; supposedly prevented from interbreeding
by preference for different food plants or other hosts (cf. Sibling species).
Biological species concept. A concept of the species category stressing reproductive
isolation, and the possession of a genetic program effecting such isolation
(cf. Species; see 2.2.3).
Biota. The flora and fauna of a region (cf. Fauna, Flora).
Bisexual. Of a population composed of functional males and females; sometimes
also applied to an individual possessing functional male and female reproduc¬
tive organ (= hermaphrodite).
Catalog. An index to taxonomic literature arranged by taxa so as to provide
ready reference to at least the most important taxonomic and nomenclatural
references to the taxon involved (cf. Checklist).
Category. See Taxonomic category.
Character. See Taxonomic character.
Character displacement. A divergence of equivalent characters in sympatric species
resulting from the selective effects of competition.
Character gradient. See Cline.
Character index. A numerical value, compounded of the ratings of several charac¬
ters, indicating a degree of difference of related taxa; also a rating of
an individual, particularly a hybrid, in comparison with its most nearly
related species (see 9A.3).
Checklist. Usually a skeleton classification of a group listed by taxa for quick
reference and as an aid in the arrangement of collections (cf. Catalog).
Cheironym. A manuscript name (q.v.).
Chorology. The study of the geographical distribution of organisms.
Chromatophore. A pigment-bearing intracellular body.
Chromosomal inversion. Reversal of the linear order of the genes in a segment
of a chromosome.
Chromosome. A deeply staining DNA-containing body in the nucleus of the cell,
best seen during cell division.
Circular overlap. The phenomenon in which a chain of contiguous and intergrad¬
ing populations curves back until the terminal links overlap geographically
and behave like good species (noninterbreeding).
Cladism. A taxonomic theory by which organisms are ordered and ranked entirely
on the basis of “recency of common descent,” that is, on the basis of
the most recent branching point of the inferred phylogeny.
400 Glossary

Cladistic. Based on the principles of cladism.

Cladogram. A dendrogram based on the principles of cladism; a strictly genealogi¬
cal dendrogram in which rates of evolutionary divergence are ignored (see
10.7.2).
Classification. The delimitation, ordering, and ranking of taxa (cf. Taxonomy,
Systematics, Horizontal classification, Vertical classification, Artificial classi¬
fication, Biological classification).
Clinal. Varying gradually, of characters.
Cline. A gradual and nearly continuous change of a character in a series of
contiguous populations; a character gradient (cf. Subspecies).
Clone. All the offspring derived by asexual reproduction from a single sexually
produced individual.
Clustering methods. Methods of grouping related or similar species into species
groups or higher taxa (see 10.5.2).
Coccids. Scale insects.
Code. International Code of Zoological Nomenclature (see Chap. 12).
Code of Ethics. A set of recommendations on the propriety of taxonomic actions,
to guide the taxonomist. Formulated in Appendix A of the International
Code of Zoological Nomenclature (see 13.64).
Coefficient of difference. Difference of means divided by sum of standard devia¬
tions (see 9B1.1):

CD = Mi — Ma
SDa + SD b

Coefficient of variability. The standard deviation as percentage of the mean:

SD X 100
CV
M

Collective group. An aggregate of related species of which the generic position

is uncertain, used principally in paleontology and parasitology (see 13.39).
Commission. The International Commission on Zoological Nomenclature (see
13.59).
Common name. Colloquial name = vernacular name (q.v.).
Competitive exclusion. The principle that no two species can coexist at the same
locality if they have identical ecological requirements.
Complex. A neutral term for a number of related taxonomic units, most commonly
involving units in which the taxonomy is difficult or confusing (cf. Group,
Neutral term).
Congeneric. A term applied to species of the same genus (cf. Genus).
Conspecific. A term applied to individuals or populations of the same species
(cf. Species).
Continuity. In nomenclature, the principle that continuity of usage should take
precedence over priority of publication in determining which of two or
more competing scientific names should be adopted for a particular taxon
(cf. Priority).
Continuous variation. Variation in which individuals differ from each other by
 Glossary 401

infinitely small steps, as variation in quality of expression of a character

or group of characters (cf. Discontinuous variation).
Convergence. Morphological similarity in but distantly related forms (see 10.1.3,
10.4.5).
Cope’s rule. The generalization that there is a steady increase in size in phyletic
series.
Correlated characters. Characters that are associated either as manifestations of
a well-integrated ancestral gene complex (10.4.2) or because they are func¬
tionally correlated (10.4.3).
Cotype. Syntype (q.v.).
Cryptic species. Sibling species (q.v.).
Cyclomorphosis. A seasonal (and thus cyclic) nongenetic change of phenotype
in species of planktonic freshwater organisms, particularly cladocerans and
rotifers.
Cytogenetics. The comparative study of chromosomal mechanisms and behavior
in populations and taxa, and their effect on inheritance and evolution.
Cytology. The study of the structure and physiology of the cell and its parts.
Darwin principle. The taxonomic importance of characters that are not the result
of a specific ad hoc adaptation (10.4.2).
Data matrix. A tabulation of differences between species (or other taxa) in rows
(characters) and columns (taxa) (10.2.2).
Delimitation. In taxonomy, a formal statement of the characters of a taxon which
sets its limits (cf. Description, Diagnosis, Differential diagnosis).
Deme. A local population of a species; the community of potentially interbreeding
individuals at a given locality.
Dendrogram. A diagrammatic drawing in the form of a tree designed to indicate
degrees of relationship (see 10.7.2; cf. Phylogenetic tree).
Derived character. A character that differs materially from the ancestral condition
(10.3.1).
Description. In taxonomy, a more or less complete formal statement of the char¬
acters of a taxon without special emphasis on those which set limits to
the taxon or distinguish it from coordinate taxa (cf. Delimitation, Diagnosis,
Differential diagnosis).
Designated priority. In cases of simultaneous publication of several names, the
priority established by the first reviser (see 13.26).
Diagnosis. In taxonomy, a formal statement of the characters (or most important
characters) which distinguish a taxon from other similar or closely related
coordinate taxa (cf. Differential diagnosis, Description).
Dichotomous. Divided or dividing into two parts.
Differentia. Linnaeus’ polynominal species diagnosis.
Differential diagnosis. A formal statement of the characters which distinguish
a given taxon from other specifically mentioned equivalent taxa.
Dimorphism. Occurrence of two distinct morphological types (morphs, phena)
in a single population (cf. Sexually dimorphic, Polymorphism).
Diploid. Having a double set of chromosomes (2n); the normal chromosome
number of the cells (except for mature germ cells) of a particular organism
derived from a fertilized egg (cf. Haploid, Polyploidy, Chromosome).
402 Glossary

Discontinuous variation. Variation in which the individuals of a sample fall into

definite classes which do not grade into each other (cf. Continuous variation).
Discriminant function. The sum of numerical values of certain diagnostic charac¬
ters multiplied by calculated constants (9A.3).
Dollo’s rule. The principle that evolution is irreversible to the extent that structures
or functions once lost cannot be regained.
Dominant. An allele which determines the phenotype of a heterozygote (cf. Reces¬
sive, Homozygous, Heterozygous).
Eclipse plumage. Inconspicuous plumage of birds worn in alternation with a
bright nuptial plumage.
Ecological isolation. A condition in which interbreeding between two or more
otherwise sympatric populations is believed to be prevented by mating in
different ecologic niches (cf. Reproductive isolation, Geographic isolation).
Ecological race. A local race that owes its most conspicuous attributes to the
selective effect of a specific environment (cf. Ecotype).
Ecology. The study of the interactions between organisms and their environment.
Ecophenotypic variation (habitant variation). A nongenetic modification of the
phenotype by specific ecological conditions, particularly those of a habitat.
Ecospecies. “A group of populations so related that they are able to exchange
genes freely without loss of fertility or vigor in the offspring” (Turesson).
Ecotype. A descriptive term applied to plant races of varying degrees of distinctness
which owe their most conspicuous characters to the selective effects of
local environments (cf. Subspecies).
Edaphic factor. The influence of soil properties on organisms (especially plants).
Electronic data processing (EDP). The sorting and storage of data with the
help of computers.
Electrophoresis. A process of separating different molecules, particularly polypep¬
tides, owing to their differential rates of migration in an electric field.
Emendation. In nomenclature, an intentional modification of the spelling of a
previously published scientific name (cf. Error, Lapsus calami) (see 13.32).
Environment. The totality of physical, chemical, and biotic conditions surrounding
an organism.
Equal weighting. The method which treats all taxonomic characters as equally
important, a key assumption of phenetics.
Error. In nomenclature, an unintentional misspelling of a scientific name, as
a typographical error or an error of transcription (cf. Emendation, Lapsus
calami).
Essentialism. A school of philosophers, originating with Plato and Aristotle, later
maintained by the Thomists and so-called realists among the philosophers,
who believed in the reality of underlying universal or essences; in taxonomy
usually referred to as typology (see 4.3.2).
Ethology. The science of the comparative study of animal behavior.
Eucaryotes. Organisms with a well-defined nucleus and meiosis. All higher orga¬
nisms above the level of procaryotes (q.v.).
Eyepiece micrometer. A linear scale in the field of vision of the eyepiece (or
one of a pair of eyepieces) of a microscope for use as a measuring device.
 Glossary 403

Family. A taxonomic category including one genus or a group of genera or tribes

of common phylogenetic origin, which is separated from related similar
units (families) by a decided gap, the size of the gap being in inverse
ratio to the size of the family (see 5.5).
Family name. The scientific designation of a taxon of family rank, recognized
by the termination idae, which termination may not be used in names
of other taxa.
Fauna. The animal life of a region (cf. Flora, Biota).
Faunal work. A publication in which taxa are included on the basis of their
occurrence in a specified area rather than on the basis of relationship (cf.
Local list; see 11A.6).
First reviser. The first author to publish a definite choice of one among two
or more conflicting names or zoological interpretations which are equally
available under the Code; in order to qualify as first reviser an author
must give evidence of a choice between available alternatives (see 13.4).
Flora. The plant life of a region (cf. Fauna, Biota).
Form. A neutral term for a single individual, phenon, or taxon (cf. Group, Neutral
term).
Formenkreis. A collective category of allopatric subspecies or species (Klein-
schmidt); in paleontology, a group of related species or variants.
Full bibliographical synonymy. A reasonably complete list of references to a
given taxon arranged so as simultaneously to serve the needs of nomenclature
(chronology of names) and zoology (pertinent taxonomic and biological
sources) (cf. Synonymy).
Gamma taxonomy. The level of taxonomy dealing with various biological aspects
of taxa, ranging from the study of intraspecific populations to studies of
speciation and of evolutionary rates and trends (see 1.4.3).
Gause’s rule. The theory that no two species with identical ecological requirements
can coexist in the same place (cf. Competitive exclusion).
Gene. A hereditary determiner; the unit of inheritance, carried in a chromosome,
transmitted from generation to generation by the gametes, and controlling
the development of the individual (cf. Chromosome).
Gene flow. The exchange of genetic factors between populations owing to dispersal
of zygotes or gametes, e.g., pollen.
Gene frequency. The percentage of a given gene in a population (cf. Gene,
Population).
Gene pool. The totality of the genes of a given population existing at a given
time.
Genetic drift. Genetic changes in populations caused by random phenomena rather
than by selection (cf. Local population).
Genotype. The genetic constitution of an individual or taxon (cf. Phenotype).
Use of this term in nomenclature for the type-species of a genus is confusing
and contrary to the terminology of the Code (sec Art. 42b, Recommendation
67A).
Genus. A category for a taxon including one species or a group of species, presum¬
ably of common phylogenetic origin, which is separated from related similar
404 Glossary

units (genera) by a decided gap, the size of the gap being in inverse
ratio to the size of the unit (genus) (see 5.4).
Geographic isolate. A population that is separated by geographic barriers from
the main body of the species.
Geographic isolation. The separation of a gene pool by geographic barriers; the
prevention of gene exchange between a population and others by geographic
barriers.
Geographical race. Subspecies (q.v.).
Grade. A group of animals similar in level of organization; a level of anagenetic
advance.
Group. A neutral term for a number of related taxa, especially an assemblage
of closely related species within a genus (cf. Complex, Neutral term,
Section).
Gynandromorph. An individual in which one part of the body is masculine,
the other feminine; most frequent are bilateral gynandromorphs, in which
the left and right halves are of different sex.
Handbook. In taxonomy, a publication designed primarily as an aid to field
and laboratory identification rather than the presentation of new tax¬
onomic conclusions (cf. Manual, Monograph).
Haploid. Having only a single set of chromosomes; gametes are usually haploid.
Hermaphrodite. An individual having both male and female reproductive organs
(cf. Intersex).
Heterozygous. Having different alleles at homologous loci of the two parental
chromosomes (cf. Allele, Locus, Homozygous).
Hierarchy. In classification, the system of ranks which indicates the categorical
level of various taxa (i.e., kingdom to species) (cf. Taxonomic category).
Higher category. A taxonomic category of rank higher than the species (i.e.,
from subgenus to kingdom) (cf. Supraspecific).
Higher taxon. A taxon ranked in one of the higher categories.
Histogram. A set of rectangles in which the midpoints of class intervals are
plotted on the abscissa and the frequencies (usually, number of specimens)
on the ordinate (see 8C.3.1).
Holistic. Looking at wholes as more than the sums of their parts.
Hollow curve. For explanation see 10.5.3, Fig. 10-9. A curve demonstrating an
excess over expectancy of very small (e.g. monotypic) and very large higher
taxa.
Holosteans. A group of fishes ancestral to the teleost fishes.
Holotype. The single specimen designated or indicated as “the type” by the
original author at the time of the publication of the original description
(see 13.51).
Homologous. A feature in two or more taxa which can be traced back to the
same feature in the common ancestor of these taxa (cf. Analogous).
Homonym. In nomenclature, one of two or more identical but independently
proposed names for the same or different taxa (cf. Senior homonym, Junior
homonym, Primary homonym, Secondary homonym; see 13.43).
Homozygous. Having identical alleles at the two homologous loci of a diploid
chromosome set (cf. Allele, Locus, Heterozygous).
 Glossary 405

Horizontal classification. Classification which stresses grouping together species

in a similar stage of evolution, rather than location on the same phyletic
line (cf. Vertical classification).
Host races. Different genetic races of the same species in oligophagous food special¬
ists or parasites occurring on different hosts.
Hybrid belt. A zone of interbreeding between two species, subspecies, or other
unlike populations; zone of secondary intergradation (see 9B.2).
Hybrid index. See Character index (see 9A.3).
Hybridization. The crossing of individuals belonging to two unlike natural popula¬
tions, principally species (cf. Allopatric hybridization).
Hypodigm. The entire material of a species that is available to a taxonomist.
Identification. The determination of the taxonomic identity of an individual (see
4.3.1 for theory and 7.3 for practices of identification).
Indication. In nomenclature the publication of certain types of evidence or cross
references which establish the typification of a name and thus make it avail¬
able (see Art. 16 and 13.20).
Individual variation. Variation within a population (see 8A, 8B).
Industrial melanism. The evolution of a darkened population owing to selection
in the sooty surroundings of an industrial area (cf. Melanism).
Infraorder. An optional category below the suborder.
Infraspecific. Within the species; usually applied to categories (subspecies) and
phena (varieties) (cf. Subspecies, Variety, Infrasubspecific form).
Infrasubspecific form. Individual and seasonal variants in a single interbreeding
population.
Infrasubspecific name. A name given to an infrasubspecific form (see 13.41).
Intergradation. Merging gradually through a continuous series of intermediate
forms or populations.
International Code of Zoological Nomenclature. The official set of regulations
dealing with zoological nomenclature (see Chap. 12).
Intersex. An individual more or less intermediate in phenotype between male
and female (cf. Hermaphrodite).
Introgressive hybridization. The spread of one or more genes of one species
into the germ plasm of another species as a result of hybridization (cf.
Hybridization).
Irreversibility rule. See Dollo’s rule.
Isolating mechanisms. Properties of individuals that prevent successful interbreeding
with individuals that belong to different populations.
Isophene. A line connecting points of equal expression of a character; lines at
right angles to a cline on a map (cf. Cline).
Junior homonym. The more recently published of two or more identical names
for the same or different taxa (cf. Homonym, Senior homonym).
Junior synonym. The more recently published of two or more available synonyms
for the same taxon (cf. Synonym, Senior synonym).
Karyological character. A character involving chromosome structure or number.
Key. A tabulation of diagnostic characters of species (or genera, etc.) in dichoto¬
mous couplets facilitating rapid identification.
Key character. In taxonomy a character of special utility in a key.
406 Glossary

Lapsus calami. In nomenclature, a slip of the pen, especially an error in spelling

(cf. Error, Emendation; see 13.32).
Lectotype. One of a series of syntypes which, subsequent to the publication of
the original description, is selected and designated through publication to
serve as “the type” (see 13.54).
Line precedence. Occurrence of a name on an earlier line of the same page
than another name for the same taxon (see 13.26).
Linnaean hierarchy. A structure of categorical ranks for taxa where each category
except the lowest includes one or more subordinate categories (sec 5.2).
Local population. The individuals of a given locality which potentially form
a single interbreeding community (cf. Deme).
Locus. The position of a gene in a chromosome (cf. Gene, Chromosome).
Lumper. A taxonomist who emphasizes the demonstration of relationship in the
delimitation of taxa and who tends to recognize large taxa (cf. Splitter;
see 10.5.3).
Macrotaxonomy. The classification of higher taxa.
Manuscript name. In nomenclature, an unpublished scientific name (cf. Nomen
nudum).
Material. In taxonomy, the sample available for taxonomic study (cf. Series,
Hypodigm).
Melanism. An unusual darkening of color owing to increased amounts of black
pigment; sometimes a racial character, sometimes, as in cases of polymor¬
phism, restricted to a certain percentage of individuals within a population
(cf. Industrial melanism, Albinism).
Mendelian population. A population with unrestricted interbreeding of individuals
and free reassortment of genes.
Meristic variation. Variation in characters that can be counted, like number of
vertebrae, scales, fin rays, and so forth.
Metamorphosis. A drastic change of form during development, as when a tadpole
changes into a frog, or an insect larva into an imago.
Metric system. A decimal system of measures (with the meter as base) and weights
(with the gram as base); the universal system in science for reporting
measures and weights.
Microgeographic race. A local race, restricted to a very small area.
Millimeter (mm). 1/1,000 m, or 0.03937 in., approximately in. (cf. Metric
system).
Mimetic polymorphism. Polymorphism in Lepidoptera in which the various
morphs resemble other species distasteful or poisonous to a predator; often
restricted to females.
Mimicry. Resemblance in color or structure to other species that are distasteful
or poisonous to a predator.
Monogenic. Determined by a single gene (cf. Polygenic).
Monograph. In taxonomy, an exhaustive treatment of a higher taxon in terms
of all available information pertinent to taxonomic interpretation; usually
involving full systematic treatment of the comparative anatomy, biology,
ecology, and detailed distributional analyses of all included taxa (cf. Revi¬
sion, Synopsis).
 Glossary 407

Monophyly. The derivation of a taxon through one or more lineages from one
immediately ancestral taxon of the same or lower rank (see 4.3.5).
Monotypic. A taxon containing but one immediately subordinate taxon, as a
genus containing but one species, or a species containing but one (the
nominate) subspecies.
Morph. Any of the genetic forms (individual variants) that account for
polymorphism.
Morphospecies. A typological species recognized merely on the basis of morphologi¬
cal difference (cf. Phenon).
Mosaic evolution. Evolution involving unequal rates for different structures, organs,
or other components of the phenotype.
Muellerian mimicry. Similarity (usually consisting of a similar warning coloration)
of several species which are distasteful, poisonous, or otherwise harmful.
Multivariate analysis. The simultaneous analysis of several variable characters.
Mutation. In genetics, a discontinuous change of a genetic factor, usually the
replacement or loss of one or several base pairs in the DNA.
Natural selection. The unequal contribution of genotypes to the gene pool of
the next generation, through differential mortality and differences in repro¬
ductive success, caused by components of the environment.
Neontology. The science dealing with the life of Recent organisms (cf.
Paleontology).
Neoteny. Attainment of sexual maturity in an immature or larval stage.
Neotype. A specimen selected as type subsequent to the original description in
cases where the original types are known to be destroyed or were suppressed
by the Commission (see 13.55).
Neutral term. A taxonomic term of convenience, such as form or group, which
may be employed without reference to the formal taxonomic hierarchy
of categories, and which has no nomenclatural significance.
New name. A replacement name for a preoccupied name (cf. Substitute name;
see 13.45).
Niche (ecological). The precise constellation of environmental factors into which
a species fits or which is required by a species.
Nomenclator. A book containing a list of scientific names assembled for nomen¬
clatural, rather than taxonomic, purposes (cf. Catalog).
Nomenclature. A system of names (see Chaps. 12 and 13).
Nomen conservandum. A name preserved by action of the Commission and placed
on the appropriate official list (see 13.63).
Nomen dubium. The name of a nominal species for which available evidence
is insufficient to permit recognition of the zoological species to which it
was applied.
Nomen oblitum. A name losing its validity under the statute of limitation (Art.
23b) (see 13.24).
Nominal taxon (species, genus, etc.). A named taxon, objectively defined by
its type (see 13.58).
Nominalism. A school of philosophy, denying the existence of universals, and
emphasizing the importance of man-given names for the grouping of individ¬
uals (see 4.3.3).
408 Glossary

Nominate. Of a subordinate taxon (subspecies, subgenus, etc.), which contains

the type of the subdivided higher taxon and bears the same name (see
13.36).
Nondimensional species. The species concept, represented by the noninterbreeding
of species at a given place and time.
Numerical phenetics. The hypothesis that relationship of taxa can be determined
by a calculation of an overall, unweighted similarity value.
Objective synonym. Each of two or more synonyms based on the same type.
Official Index. A list of names or works suppressed or declared invalid by the
Commission (see 13.63).
Official List. A list of names or works which have been conserved or declared
to be valid by the Commission (see 13.63).
Oligogenic character. A character determined by only few genes.
Onomatophore. “Name-bearer” = type (Simpson) (see 13.48).
Ontogeny. The developmental history of an individual organism from egg to
adult.
Original description. A statement of characters accompanying the proposal of
a name for a new taxon in conformance with Arts. 12 and 13 of the
Code (see 1 IB.1.1).
Orthogenesis. Evolution of phyletic lines following a predetermined rectilinear
pathway, the direction not being determined by natural selection.
Overall similarity. A (usually numerical) value of similarity calculated by the
summation of similarities in numerous individual characters (see 10.2).
Page precedence. Occurrence of a name on an earlier page in the same publication
than a synonym or homonym of it (see 13.26).
Paleontology. The science that deals with the life of past geological periods (cf.
Neontology).
Parallelism. The independent acquisition of similar characters in related evolution¬
ary lines (cf. Convergence).
Parapatry. Of populations or species, in nonoverlapping geographical contact with¬
out interbreeding.
Parasitoid. Wasps and flies, the larvae of which parasitize (and usually kill)
individuals of the host species.
Paratype. A specimen other than the holotype which was before the author at
the time of preparation of the original description and was so designated
or indicated by the original author.
Parthenogenesis. The production of offspring from unfertilized eggs.
Patronymic. In nomenclature, a dedicatory name, a name based on that of a
person or persons.
Pheneticist. See Numerical phenetics.
Phenetic ranking. Ranking into categories, strictly based on degree of overall
similarity (see 10.2 and 10.5.2).
Phenogram. A diagram indicating degree of similarity among taxa (see 10.7.2).
Phenon. A sample of phenotypically similar specimens; a phenotypically reasonably
uniform sample (see 1.2).
Phenotype. The totality of characteristics of an individual (its appearance) as
a result of the interaction between genotype and environment.
 Glossary 409

Phyletic. Pertaining to a line of descent (cf. Phylogeny).

Phyletic correlation. Correlation of characters that are phenotypic manifestations
of a well-integrated ancestral gene complex (see 10.4.2).
Phyletic weighting. Assessing the taxonomic importance of a character on the
basis of its phyletic information content (see 10.4).
Phylogenetic tree. A diagrammatic presentation of inferred lines of descent, based
on paleontological, morphological, or other evidence.
Phylogeny. The study of the history of the lines of evolution in a group of
organisms; the origin and evolution of higher taxa (cf. Classification).
Phylogram. A tree-like diagram indicating degree of relationship among taxa
(see 10.7.2).
Pleiotropy. The capacity of a gene to affect several characters, that is, several
aspects of the phenotype.
Plenary powers. Special powers granted to the Commission (see 13.59 and Art.
< 79).
Plesiomorph. Of characters, primitive or as found in the ancestor (Hennig).
Polygenic. Of a character, controlled by several or numerous genes.
Polymorphism. The simultaneous occurrence of several discontinuous phenotypes
or genes in a population, with the frequency even of the rarest type higher
than can be maintained by recurrent mutation.
Polynominal nomenclature. A system of nomenclature consisting of a scientific
designation of a species through more than two words; the antecedent of
the Linnaean “binomial” system.
Polyphyletic. A term applied to a composite taxon derived from two or more
ancestral sources; not of a single immediate line of descent (cf.
Monophyletic).
Polyploidy. A condition in which the nuclear complement of chromosomes is
an integral multiple (greater than 2) of the haploid number.
Polythetic. Of taxa, in which each member has a majority of a set of characters
(see 4.7; also Simpson, 1961, p. 42, under polytypic).
Polytopic. Occurring in different places as, for instance, a subspecies composed
of widely separated populations.
Polytypic. A taxon containing two or more taxa in the immediately subordinate
category, as a genus with several species or a species with several subspecies
(cf. Monotypic).
Population. See Local population.
Precipitin reaction. The formation of a visible precipitate at the interface when
an antigen and the corresponding antiserum are brought together (cf. Anti¬
gen, Antiserum, Antibody).
Predictive value. The capacity of a classification to make predictions on newly
employed characters or newly discovered taxa (see 4.5).
Pre-Linnaean name. A name published prior to Jan. 1, 1758, the starting point
of zoological nomenclature (see 13.8).
Primary homonym. Each of two or more identical species-group names which,
at the time of original publication, were proposed in combination with
the same generic name (e.g., X-ut albus Smith, 1910, and X-us albus Jones,
(1920) (see 13.43).
410 Glossary

Primary intergradation. A zone of intermediacy between two phenotypically differ¬

ent populations, having developed in situ as a result of selection (cf. Secon¬
dary intergradation; see 9B.2).
Primary zoological literature. Literature dealing with animals or zoological phe¬
nomena, not merely a listing of names (see 13.24).
Priority. The principle that of two competing names for the same taxon (below
the rank of an infraorder) ordinarily that is valid which was published
first (see 13.3).
Procaryotes. Those microorganisms (viruses, bacteria, blue-green algae) that lack
well-defined nuclei and meiosis (cf. Eucaryotes).
Pseudogamy (= Gynogenesis). Parthenogenetic development of the egg cell after
the egg membrane has been penetrated by a male gamete.
Q technique. An analysis of association of pairs of taxa in a data matrix (see
10.2.2).
Race. Subspecies (q.v.).
Ranking. The placement of a taxon in the appropriate category in the hierarchy
of categories (see 10.5.2).
Rassenkreis (Rensch). Synonym for polytypic species (cf. Subspecies, Polytypic;
see 3.2).
Recapitulation. The theory that ontogeny recapitulates phylogeny (cf. Ontogeny,
Phytogeny).
Recent. Of taxa which still exist, antonym of Fossil.
Recessive. Of a gene, not affecting the phenotype of the heterozygote.
Reductionism. The erroneous belief that complex phenomena can be entirely
explained by reducing them to the smallest possible component parts and
by explaining these.
Redundant characters. Characters so closely correlated with other, already used
characters that they do not contribute new information to the analysis (see
10.4.3).
Regression analysis. A form of multivariate analysis (see 8C.2.5).
Regressive character. A character which is being reduced or lost in the course
of phylogeny, sometimes independently in several related lines (see 10.4.3).
Relationship. For meaning in classification, see 10.1.
Replacement name. Substitute name (q.v.).
Reproductive isolation. A condition in which interbreeding between two or more
populations is prevented by intrinsic factors (cf. Isolating mechanism).
Reticulate evolution. Evolution “dependent on repeated intercrossing between a
number of lines, and thus both convergent and divergent at once” (Huxley).
Revision. In taxonomy, the presentation of new material or new interpretations
integrated with previous knowledge through summary and reevaluation (cf.
Synopsis, Monograph).
R technique. An analysis of association of characters in a data matrix (see 10.2).
Saltation. Discontinuous variation produced in a single step by major mutation
(cf. Mutation).
Sample. That portion of a true population which is actually available to the
taxonomist.
 Glossary 411

Scatter diagram. A bivariate or multivariate graphic method of population analysis

(see 8C.3.3.).
Scientific name. The binominal or trinominal designation of an animal; the formal
nomenclatural designation of a taxon (cf. Vernacular name).
Secondary homonym. Each of two or more identical specific names which, at
the time of original publication, were proposed in combination with different
generic names but which, through subsequent transference, reclassification,
or combination of genera have come to bear the same (or an identical)
combination of a generic and specific name (see 13.43).
Secondary intergradation. A zone of hybridization or strong steepening of character
gradients where two separately differentiated populations have reestablished
contact (see 9B.2).
Secondary sexual characters. Characters which distinguish the two sexes of the
same species but which do not (like gonads or accessory sexual characters)
function directly in reproduction (cf. Sexually dimorphism).
Section. A neutral term usually employed with reference to a subdivision of a
taxon or a series of related elements in one portion of a higher taxon (cf.
Higher category, Neutral term, Group).
Selection. See Natural selection.
Semispecies. The component species of superspecies (Mayr); also, populations
that have acquired some, but not yet all, attributes of species rank; borderline
cases between species and subspecies.
Senior homonym. The earliest published of two or more identical names for
the same or different taxa (cf. Homonym, Junior homonym).
Senior synonym. The earliest published of two or more available synonyms for
the same taxon (cf. Synonym, Junior synonym).
Series. In taxonomy, the sample which the collector takes in the field or the
sample available for taxonomic study (cf. Material, Hypodigm).
Serology. The study of the nature and interactions of antigens and antibodies
(cf. Antigen, Antibody).
75-percent rule. The rule that population A can be considered subspecifically
distinct from population B if at least 75 percent of the individuals of
A are different from “all” the individuals of population B (see 9B.1.2).
Sex Chromosome. A special chromosome, not occurring in identical number or
structure in the two sexes and usually concerned with sex determination;
the X chromosome or Y chromosome (cf. Chromosome, Autosome).
Sex-limited character. A character occurring in one sex only (cf. Secondary sexual
character, Sex-linked character).
Sex-linked character. A character controlled by a gene located in the sex chromo¬
some (cf. Sex chromosome).
Sexual dimorphism. The phenotypic difference between the two sexes of a species.
Sexual reproduction. Reproduction resulting in a diploid zygote with a maternal
and a paternal chromosome set.
Sibling species. Pairs or groups of closely related species which are reproductively
isolated but morphologically identical or nearly so (cf. Species; see 9A.2).
Sonagram. A graphic representation of the vocalization of an animal (see 7.4.10;
Fig. 7-6).
412 Glossary

Speciation. The splitting of a phyletic line; the process of the multiplication

of species; the origin of discontinuities between populations caused by the
development of reproductive isolating mechanisms (cf. Allopatric speciation,
Sympatric speciation).
Species. Groups of actually (or potentially) interbreeding natural populations
which are reproductively isolated from other such groups (cf. Subspecies,
Population, Reproductive isolation).
Specific name. The second component of the binominal name of a species (see
13.40).
Splitter. In taxonomy, one who divides taxa very finely, to express every shade
of difference and relationship, through the formal recognition of separate
taxa and their elaborate categorical ranking (see 10.5.3).
Standard deviation. (SD) The square root of the sum 2 of the squared deviations
d from the mean, divided by N:

Standard error (of the mean). Standard deviation divided by the square root
of the sample size, N:

Statute of limitation. A provision in the Code (Art. 23b) to protect universally

adopted junior names against the revival of forgotten senior synonyms (see
13.24).
Strickland Code. A code of nomenclature prepared by a committee of the British
Association for the Advancement of Science under the secretaryship of H. E.
Strickland and first published in 1842.
Subfamily. A category of the family-group subordinate to the family; an individual
taxon ranked in the category subfamily (see 5.5).
Subgeneric name. The name of an optional category between the genus and
the species, enclosed in parentheses when cited in connection with a binomi¬
nal or trinominal combination and therefore excluded from consideration
when determining the number of words of which a specific or subspecific
name is composed [eg., X-us (Y-us) albus rufus is a trinominal].
Subjective synonym. Each of two or more synonyms based on different types,
but regarded as referring to the same taxon by those zoologists who hold
them to be synonyms.
Subspecies. A geographically defined aggregate of local populations which differs
taxonomically from other such subdivisions of the species (see 3.3.2).
Substitute name. A name proposed to replace a preoccupied name and auto¬
matically taking the same type and type-locality (=New name; see 13.45).
Superfamily. The taxonomic category immediately above the family and below
the order; an individual taxon ranked in this category.
Superspecies. A monophyletic group of entirely or largely allopatric species (cf.
Allopatric, Semispecies; see 3.7).
 Glossary 413

Supraspecific. A term applied to a category or evolutionary phenomenon above

the species level.
Synipatric hybridization. The occasional production of hybrid individuals between
two otherwise well-defined sympatric species.
Synipatric speciation. Speciation without geographic isolation; the acquisition of
isolating mechanisms within a deme.
Sympatry. The occurrence of two or more populations in the same area; more
precisely, the existence of a population in breeding condition within the
cruising range of individuals of another population.
Symplesiomorphy. The sharing of ancestral characters by different species (Hen-
nig) (see 10.1.3).
Synapomorphy. The sharing of derived characters by several species (see 10.1.3).
Synchronic species. Species which occur at the same time level (cf. Allochronic
species).
Synonym. In nomenclature, each of two or more different names for the same
taxon (cf. Senior synonym, Junior synonym, Objective synonym, Subjective
synonym).
Synonymy. A chronological list of the scientific .names which have been applied
to a given taxon, including the dates of publication and the authors of
the names.
Synopsis. In taxonomy, a brief summary of current knowledge of a group (see
11A.2).
Syntype. Every specimen in a type-series in which no holotype was designated
(see 13.54).
Systematics. The science dealing with the diversity of organisms (see 1.1).
Taxon (pi. taxa). A taxonomic group that is sufficiently distinct to be worthy
of being distinguished by name and to be ranked in a definite category
(see 1.2).
Taxonomic category. Designates rank or level in a hierarchic classification. It
is a class, the members of which are all taxa assigned a given rank (see 1.2).
Taxonomic character. Any attribute of a member of a taxon by which it differs
or may differ from a member of a different taxon (see 7.1).
Taxonomy. The theory and practice of classifying organisms (cf. Classification,
Systematics).
Teratology. The study of structural abnormalities, especially monstrosities and
malformations.
Therapsid reptiles. The reptilian group from which the mammals evolved.
Topotype. A specimen collected at the type-locality (see 13.57).
Tribe. A taxonomic category intermediate between the genus and the subfamily.
Trinominal nomenclature. An extension of the binominal system of nomenclature
to permit the designation of subspecies by a three-word name (see 13.40).
Triploid. A cell or individual with three haploid chromosome sets, one of the
forms of polyploidy.
Trivial name. An obsolete designation by Linnaeus for the specific name; also
a synonym for “vernacular name.”
Type. A zoological object which serves as the base for the name of a taxon
(sec 13.48-13.56).
414 Glossary

Type designation. Determination of the type of a genus under Arts. 67-69 of

the Code (see 13.52).
Type-locality. The locality at which a holotype, lectotype, or ncotype was collected
(cf. Topotype) (see 13.57).
Type method. The method by which the name for a taxon is unambiguously
associated with a definite zoological object belonging to the taxon (see
13.48).
Type selection. Type designation (q.v.).
Type-species. The species which was designated as type of a nominal genus.
Typological thinking. A concept in which variation is disregarded and the members
of a population are considered as replicas of the “type,” the Platonic eidos.
Uninominal nomenclature. The designation of a taxon by a scientific name consist¬
ing of a single word; required for taxa above species rank.
Univariate analysis. A biometric analysis of a single character.
Valid name. An available name that is not preoccupied by a valid senior synonym
or homonym (see 13.23).
Variance. The square of the standard deviation.
Variation, ecophenotypic. Variation caused by nongenetic responses of the pheno¬
type to local conditions of habitat, climate, etc.
Variety. An ambiguous term of classical (Linnaean) taxonomy for a heterogeneous
group of phenomena including nongenetic variations of the phenotype,
morphs, domestic breeds, and geographic races.
Vernacular name. The colloquial designation of a taxon (cf. Scientific name).
Vertical classification. Classification which stresses common descent and tends
to unite ancestral and descendant groups of a phyletic line in a single
higher taxon, separating them from contemporaneous taxa having reached
a similar grade of evolutionary change (cf. Horizontal classification, 10.5.4).
Weighting. A method for determining the phyletic information content of a charac¬
ter; the evaluation of the probable contribution of a character to a sound
classification (see 10.4.).
Index
Index

Page references in italics indicate Figures.

Abbreviations, 363 Anderson, E., 185

Aceros plicatus, 150 Andrena, 156
Acknowledgments, 289 Anodonta, 153
Actuality principle, 365, 366 Anonymous names, 305
Adanson, M., 59, 83, 210, 218 Anopheles maculipennis, 7, 138, 184
Adaptation, 125 Aphid(s), 160
Adaptive zone, 234 Apomorph, 398
Aedes, 206 Appendices to the Code, 380
Affinity, 82 Applications to the Commission, 378
Agamospecies, 31 Applied biology, 7-8
Age variation, 148 Arenicola, 141
Aristotle, 56, 66, 92, 276
Aix galericulata, 131
Aristotelian type-concept, 368
Aix sponsa, 131
Aristotelianism, 91
Allometry, 154-155
Arkell, W. J., 241, 243
Allopatric, 194
Artemia salina, 31
Allopatry, kinds of, 194
Artenkreis, 52, 398
Alpha taxonomy, 15
Asexual entities, 46
Amadon, D., 53
Asexual species, 31
Amateur, role of, in systematics, 17
Astrapia, 53
Ambystoma jeffersonianum, 185 Atlases, 261
Amendments to the Code, 333, 379 Australopithecus, 234
Amphiploidy, 32 Author of a name (Art. 50), 317, 363
Analogous, definition, 85 Author’s name, 288
Anas, dendrogram of, 240 Availability, criteria of, 303, 347
Anchoviella mitchilli, 175 Ax, P., 245
418 Index

Baculum, 130, 205, 398 Cailleux, A., 10

Bader, R. S., 202 Cain, A. J., 24, 55, 62, 65, 70, 92, 203,
Baer, J. G, 139, 216 217, 219, 269, 343, 368
Bailey, R. M., 94 Calidris alba, 223
Balance, 241 Camin, J. H., 84, 212
Barber, H. S., 135 Carson, H. L., 133
Bdelloid rotifers, 31 Cases under consideration, 379
Beckner, M., 55, 65, 67, 83 Castes, insect, 151
Bees, 88 Catalog, 116, 263
classification of, 206, 207 Cataloging, 107
Behavior, 135 by lots, 108
Bessey, C. E., 25 Categories, coordinate, 359
Beta taxonomy, 15, 398 hierarchy of, 87-98
Bias, 165 infraspecific, 41
Bibliographies, references to current, infrasubspecific, 46
116-118 Category, adaptive significance, 93
Bibliography, references and, 285—286 definition, 5
Bill length, 150 higher, definition, 91
Binary, 343 Cavalli-Sforza, L. L., 212
Binomen, 29, 302, 343 Cercaria, 149
Binomial, 343 Cerion, 32, 33
Binominal nomenclature, 56 Cestodes, 242
Biochemistry, 6, 81 Character, derived, 213, 220
Biological control, 7 single, 205
Biological races, 399 taxonomic, definition, 121
Biology, theoretical, 8 Character complexes, functional, 225
Birds, 241, 251 Character displacement, 125
Blackwelder, R. E., 19, 66, 365 Character gradient, 44
Blair, W. F., 132 Character index, 185
Bloch, K., 65 Character state, 122
Bock, W. J., 85, 210, 243 Characters, 80
Bombus agrorum, 49 and adaptation, 125
Borgmeier, T., 66 ancestral, 213, 215
Boyce, A. J., 205, 209 biochemical, 133
Bracket key, 279 in common, 67
Bradley, C., 357 correlated, 93, 221
Bradley draft, 300 ecological, 138
Branching, 235 generic, 93, 142
Branching patterns, 211 genetic basis, 123, 131
Branching points, 71 geographical, 140
Brown, W. L., 42 good, 219
Bryozoa, 358 with high weight, 220-221
Bryson, V., 18, 134 kinds of, 126—142
Buchner, P., 140 larval, 131
Buck, R. C., 67, 90, 236 with low weight, 222—225
Bufo, 132 meristic, 150, 166, 171
Biihler, P., 187 monogenic, 222
Bullis, H. R., Jr., 346 morphological, 126
Burma, B. H., 176 new, 208
Butterflies, 88 number of, 205
 Index 419

Characters, physiological, 133 Code (see International Code of Zoologi¬

and rank, 142 cal Nomenclature)
redundant, 123, 225 Coefficient, of difference, 189
regressive, 222 table of, 190
scarcity of, 209 of variability, 169
sexual, 158 Coefficients of similarity, 205
table of, 127 Cohort, 89, 90
taxonomic, 121-143 Colaptes, 195, 197
unreliable, 124 Coleman, W., 59
variable, 222 Collecting, preparations for, 104
Checklists, 260, 263 Collecting techniques, 104
Chi-square, 175 Collections, 101-112
Chromosomal phylogeny, 215 arrangement of, 109
Chromosomes, 132 consolidation of, 112
Circular overlaps, 35 purpose of, 102
Circularity, 78 sorting of, 112
Cisticola, 103 Collective group, 361
Citation, sample, 286-287 Color keys, 270
Cladism, 70, 230 Color plates, 285
Cladistic approaches, 211—217 Color records, 158
Cladistic methods, 212-213 Coloration, 129, 270
Cladogram, 255, 256 Combined taxa, 352
Classes, orders, and phyla, 97 Commission on Zoological Nomenclature,
Classification, 124 300, 330, 378
biological, 54-86 Communication, system of, 297
construction of, 97, 198 Comparative method, 77
definition, 4 Comparison, 101, 119
evolutionary, 76 Complex, 47
history of, 55-65 Complexity, 220
horizontal, 244 Computer, 14, 205, 208, 211, 219, 232,
improvement of, 245—249 277, 346
Conklin, H. C., 56
and information retrieval, 229-245
objective of, 78, 79 Consistency, 220
presentation of, 249 Constance, L., 3
principles of, 77 Constancy, 220
procedure of, 198-257 Continuity evolutionary, 244
Conus, 138
provisional, 80
stages in, 15 Convergence, 202, 226
Coordinate status of categories, 359
theories of, 65-77, 79
Cope’s rule, 216
zoological, definition, 55
Corliss, J. O., 80, 107, 279, 336, 346,
Classificatory activities, 247
359, 371
Classifying, 198-257
Correlated characters, 401
Clay, T., 36, 139
Correlation, functional, 225
Clerck, C., 342
phyletic, 221
Cline, 44 Cotype, 328
Clustering, 232 Coues, E., 350
Clusters, 235 Crane, J., 103
Cnemidophorus, 185 Creodontia, 244
Cochran, W. G., 165 Crickets, 183
420 Index

Crocodilians, 70, 231 Dollo’s rule, 216

Crowson, R. A., 241 Drawing, 282
Curating, 107-112 Drongos, 248
Cuvier, G. L. C. F., 59, 251 Drosophila, 18, 138, 203, 215, 238
Cyclomorphosis, 151 number of new species, 13
phylogeny of, 72—73
Duplicates, 110
Dali, W. H., 152
Daphnia, 151
Darwin, C., 60-62, 68, 76, 87, 91, 93, Eades, D. C., 209
220, 223, 231 Echinoderms, 245
Data matrix, 204 table of, 246
Dates, important, 342 Eclectus, 159
Daughter lines, 74 Eclipse plumage, 150
de Candolle, Augustin, 2 Ecological races, 152
Declarations, 331 Ecological shifts, 221
Define, 6 Ecophenotypes, 152
Definitions, 3-6, 28 Editors, 346
Deme, 46 Edmondson, W. T., 279
Dendrogram, 252—257 Ehrlich, P. R., 3, 88
Description (s), 265—272 Eidos of Plato, 24
contents of, 269 Electronic data processing, 108
of higher taxon, 272 Electrophoresis, 134
of new taxa, 260 Elton, C., 6
original, 118, 266 Embryology, 131
Determination labels, 113 Emendations, 355-356
Dethier, V. G., 139 Empirical period, 59
Diacritic marks, 354 Empiricism, 69
Diagnosis, 265 Endings of taxa names, 90, 313, 358
differential, 266, 305, 348 Enzymes, 133
Diamond, J. M., 24 Eohippus, 244
Dice-Leraas diagram, 176 Equidae, 253
Dicrurus, 173, 275 Essentialism, 24, 66
Dicrurus hottentottus, 50 Ethology, 8
Difference, degree of, 234 Evans, H. E., 131, 136, 216
Differentia, 343 Evidence, 29, 76, 201-202
Difficulties, species concept, 30-35 Evolution, mosaic, 84
Dipodomys merriami, 193 parallel, 243
Dipodomys ordii, 39 Evolutionism, basis for classification, 79
Diptera, 243 Exchange of material, 110
Discontinuity, 233 Exclusion, competitive, 125, 138
Discriminant functions, 186 Ex-group problem, 231
Discrimination of entities, 9
Discrimination grid, 181
table of, 146 Fabricius, 338
Distinctness, 233 Family, 94-97
Divergence, 235 definition, 94
Diversity, 2 Family group names, 313, 356, 360
Divided taxa, 352 Family name, stem of, 355
DNA matching, 134 Farris, J. S., 220
 Index 421

Fasciola hepatica, 36 Gould, S. J., 155, 225

Faunal works, 262 Grade, 75, 244
Fell, H. B., 246 Graph clustering, 232
Ferris, G. F., 267, 271, 282 Graphic presentation, 175-180, 251
Field guides, 262 Gregg, J. R., 65, 67
Field number, 108 Group, 47
Field station, 105 Grouping, 14
Files, 108 Guides, 114
Finches, 205 Gunther, K., 65
First-reviser principle, 340 Gynandromorphs, 159
Fisher, R. A., 186
Flamingos, 139
Food niche, 223 Habitat variation, 152
Footnotes, 285 Haeckel, E., 62, 251
Ford, E. B., 162 Hagmeier, E. M., 193
Form, 47 Halcyon chloris pealei, table of, 171
Formenkreis, 40 Halftones, 284
Fossils, 6, 10, 35, 43, 89, 145, 272 Halkka, O., 132
Freedom of taxonomic thought, 336 Handbooks, 262
Friedmann, H., 273 Handler, P., 18, 134
Frogs, 213 Harrison, G. A., 70, 153, 203, 219
Fulton, B. B., 136 Hatheway, W. H., 186
Fusion, chromosomal, 133 Hennig, W., 65, 70, 71, 84, 86, 202,
212-217, 219, 220, 243
Herrera, A. L., 345
Galapagos finches, 94, 95, 139 Heslop-Harrison, J. W., 24
Gall insects, 128 Heterogeneity, 165
Gamma taxonomy, 15 Heteromi, 131
Gap, size of, 233 Hierarchy, Linnaean, 89
Gam, S. M., 127, 166 Hippocrates, 56
Gender of generic names, 311, 355 Hirundo, 137
Gene pool, 26 Histograms, 176
Generations, alternating, 159 History of taxonomy, periods, 55—65
Genetic program, 77 Holland, G. P., 83, 139
Genitalic structures, 129 Hollow curve, 236, 237
Genotype, 218 Holotype, 270, 327
Genus, definition, 92 Hominidae, 239
Genus group names, 315, 360 Hominoids, 72
Geographic distribution, 216 Homology, 84-85
Geography, 140 definition, 85
Geolycosa xera, 192 Homonyms, secondary, 320, 364
G4ry, J., 189 Homonymy, 318, 360, 364-365
Gesner, C., 57 subjective, 364
Ghiselin, M. T., 6, 209, 231 of subspecies, 366
Gilmour, J. S. L., 68 Hopkins, G. H. E., 139
quoted, 249 Host specificity, 138-139
Gisin, H., 55, 69, 70, 234 Hoyer, H. B., 134
Gnathostomulida, 12 Hubbs, C., 175
Goshawk, 148 Hull, D. L., 67, 90, 236, 346
Gosline, W. A., 15 Huxley, J. S., 44, 50, 64, 244
422 Index

Hybrid index, 185 Key, indented, 278

Hybridization, 32, 195 pictorial, 279
Hybrids, 302, 349 Key characters, 58, 124, 278
Hymenolepis, 36 Keyl, H. G., 132
Hypodigm, 270, 369 Keys, 67, 276-281
Kinsey, A. C., 159
Kohn, A. J., 138, 184
Identification, 4, 66, 112-115, 124 Kosmoceras, 35
Illustrations, 119, 271, 282-285
Imbrie, J., 35
Labeling, 106
I mmunogene tics, 134
Lacerta, 185
Indication, 305, 348
Lack, D., 93, 139
Inference, 29, 81, 202
Lagopus, 150
Information retrieval, 229-245, 259
Lamarck, J. B. de, 59
Information storage, 14, 229
Larval stages, 148
Infraorder, 242
Latimeria, 10
Infrasubspecific forms, 302
Latin, 304, 343, 354
(See also Names, infrasubspecific)
Laws, phylogenetic, 85
Inger, R. F., 42, 77, 94, 131, 192, 199,
Lectotypes, 328, 340, 373
213
Legends, 291
Integration, level of, 3
Length, total, 167
Intergradation, 49
Leone, C. A., 18, 134
primary, 194
Leptocephalus, 148
secondary, 194
Levi, C., 131
International Code of Zoological Nomen¬
Lewontin, R. C., 170
clature, 298, 301—333
Lidicker, W. Z., Jr., 193
history of, 299
Limitation, 350
range of authority, 341
statute of, 308
text of, 301
Lindroth, C. H., 254, 368
Intersexes, 159
Line precedence, 353
Irreversibility, 226
Linnaean types, 368
Irwin, M. R., 134
Linnaeus, C., 57, 93, 96, 241, 265, 299,
Isolate, geographical, 49
337, 343, 345, 361, 362, 368
peripheral, 49
Linsley, E. G., 156, 363
Isolated populations, classification of, 196
Literature, 116
Isolating mechanisms, 26, 125
taxonomic, 117
Isophene, 45
Loans, 111
Locust a migratoria, 154
Logic, 67
Jepsen, G. L., 251
Lovejoy, A. O., 56
Johnsgard, P. A., 136, 240
Lumpers, 239-241
Johnston, R. F., 161
Lumping, 241
Jolicoeur, P., 193
Jordan, K., 26
MacIntyre, G. T., 244
McKinney, F., 137
Keast, A., 50 Macrotaxonomy, 198
Kendrick, W. B., 208, 209 Malaria mosquito (Anopheles maculipen-
Kennedy, J. S., 154 nis), 7, 138, 184
Key, dichotomous, 277 Mallophaga, 36, 139
 Index 423

Mammalian phylogeny, 75 Mustela, 150

Manuscript, preparation of, 290-293 Mutillid wasps, 159
revision of, 291 Myers, G. G., 357
Manuscript names, 273
Manwell, R. D., 182
Name, in synonymy, 337
Map diagrams, 179-180
trivial, 343
Maps, 282
Name changing, 344
Margoliash, E., 212
Names, 341
Maslin, T. P., 84, 213, 216, 219
formation of, 310, 354
Material, 115
generic, 360
biological, 102
for higher taxa, 358
Matthey, R., 132
of hybrids, 349
Mean, 169
infrasubspecific, 304, 341, 362
Measurements, 166
kinds of, 347
Measuring tools, 168
published simultaneously, 309
Meise, W., 186
rejected, 347
Melania, 161
of species, 29
Mergus, 221, 223
for specimens, 341
Meristic variation, 166
validity of, 308
Metamorphosis, 131
vernacular, 342
Metcalf, Z. P., 276
Natural system, 59, 67
Michener, C. D., 3, 55, 88, 93, 199,
Neave, 117
203, 207, 209, 223-224, 231, 233,
Necturus, 132
234, 243, 339, 344, 345
Nematodes, 232
Microscopic techniques, 107
Neopilina, 10
Micro-species, 31
Neositta, 51
Milne, L., 252
Neotypes, 329, 373
Mining insects, 128
Neutral terms, 47
Minkoff, E. C., 205, 209
New systematics (see Systematics)
Misidentification of types, 369—370
Newell, N. D., 44
Misidentifications, 317, 326
Nichols, D., 140
Mites, 211
Noctua, 365
MLU, 334
Nomen dubium, 347
Mobius, K., 10
Nomen nudum, 347
Mollusks, 272
Nomen oblitum, 308, 351
Monographs, 261
Nomen triviale, 361
Monophyly, 75, 141
Nomenclature, alternate systems, 345
Monothetic, 83
binominal, 304
Moody, P. A., 133
Linnaean, 344
Moreau, R. E., 236
scientific, 343
Morphospecies, 5
theory of, 300
Morphs, 46
zoological, 297-380
Mosaic evolution, 209, 228, 251
Nominal, 378
Mosquitoes, key to larve, 281
Nominalism, 68
Motacilla alba, 40
Nominate, 359
Moynihan, M., 136
Numerical schemes, 68
Multivariate analysis, 187
Multivariate methods, 185
Muscicapidae, 96 Official indexes, 379
Musophagidae, 236, 238 Official lists, 331, 379
424 Index

Onomatophore, 369 Phyla, 97

Operationalism, 69 Phylogenetic trees, 62, 63, 251-253
Opiliones, 239 Phylogeny, 70-74, 77-79
Opinions, 299, 331 chromosomal, 215
Orders, 97 Phylogram, 256
Orians, 184 Phytomyza, 128
Orthoptera, 242 Pictorial scattergram, 177-178
Orton, G. L., 131 Pie graph, 179
Orycteropus, 236 Pieris napi, 45
Osche, G., 231-232 Pipilo, 32, 34
Ostrea knorri, table of, 36 Planaria alpina, 353
Overall similarity, 203 Plato, 24, 66
Overlap, geographical, 42 Platypus, 214
linear, 188 Plenary powers, 331, 339
variational, 185 Plethodon jordani, 191
Owen, R., 85 Pleurocrypta, 157
Owls, 205 Pogonophora, 12
Oyster, 152 Polymorphism, mimetic, 162
Polyphyly, 75
Polyploidy, 132, 185
Page precedence, 353 Polythetic, 82-83, 88
Paleontology (see Fossils) Polytopic, 193
Pan, 70, 135, 140, 216, 234 Polytypic species (see Species)
Papilio, 162, 163 Pongidae, 70
Papilionidae, 242 Popper, K. R., 66
Parallelism, 82 Population, asexual, 31
definition, 243 Population continuum, 48
Paramecium, 183 Population sample, 169
Parapatry, 53 Population structure, 48
Parasites, 36, 139, 182, 216, 231, 301 Population systematics, 64
Parasitism, 156 Population taxonomy, 47
Paratypes, 328, 371 Population thinking, 8
Parthenogenesis, 46 Porcupines, 141
Parus carolinensis, 172 Post-mortem changes, 157
Passer, 137 Preamble, 298, 301
Pericrocotus, 182 Prediction, 7
Periphyllus, 160 Predictive value of classification, 79
Petersen, A., 45 Preparation of material, 107
Phena, 46, 144, 182 Preservation, 105
assignment of, 145 Primary zoological literature, 351
classification of, 29, 228 Primates, 213
Phenetic method, weaknesses, 208 Primitive, 213
Pheneticists, 69 Primitive ancestors, 62
Phenetics, 231 Priority, 302, 308, 337-340
Pheidole, 155 designated, 353
Phenograms, 254, 255 of usage, 338
Phenon, 10 Procaryotes, 133
definition, 5 Profession of systematist, 16
Photography, 285 Proofreading, 291
Photuris, 135 Propinquity of descent, 76
 Index 425

Proteus, 132 Reproductive isolation, 146

Protozoans, 107, 127, 140, 371 Reptilian phylogeny, 71
Pseudacraea eurytus, 162 Retroactivity, 342
Publication, criteria of, 303, 346 Reviews, 260
date of, 307, 349 Reviser, first, 309, 353
simultaneous, 353 Reviser principle, first, 340
in synonymy, 304, 347 Revisions, 260, 263
taxonomic, 258-293 Reyment, R. A., 164, 189
Publications, evolutionary, 264 Rhabdocoela, 245
kinds of, 259-265 Ride, D., 189
Publishing, strategy of, 258 Ridgway, R., 270
Rohlf, F. J., 204, 206
Role, evolutionary, 234
Q technique, 204 Rollins, R. C., 3, 69
Rosen, D. E., 94
Rotifers, 151
R technique, 204 Rozeboom, L. E., 17
Race, 44 Rules, phylogenetic, 216
ecological, 44
microgeographic, 153
Rank, 142 Sabrosky, C. W., 8, 13
Ranking, 14 Salt, G., 156
cladistic, 230 Sample, adequate, 166
criteria, 233—242 size of, 102
equivalence, 241 Samples, allopatric, 187—197
phenetic, 231 unbiased, 105
weighted, 232 Sampling, 165
Rassenkreis, 40 Saturniidae, 88, 224
Rates of change, 71 Scala naturae, 56, 251
Ratios, 171 Scatter diagram, 172, 177, 185
Ray, J-, 57 Schmidt, K. P., quoted, 339
Reclassification, 245 Seal, H. L., 164, 189, 205
Recommendations, 342 Selander, R. K., 161, 270
Redescriptions, 271 Semispecies, 33, 53
References, 285 Sequence, of characters, 268
Regression analysis, 173 linear, 249, 250
Reinig, W. F., 49 Serology, 133
Relationship, 83-85, 200 Seventy-five percent rule, 190
diagrams of, 254 Sex, 106
genealogical, 70 Sex differences, 158
genetic, 70 Sharov, A. G., 73
Relative, nearest, 200 Sheals, J. G., 211
Relict types, 216 Sherborn, C. D., 117
Remane, A., 13, 65, 84, 85, 213, 219 Short, L. L., Jr., 197
Rensch, B., 52, 70, 77, 85, 216 Sibley, C. G., 32, 134, 195, 208
Replacement name, 366 Sibling species, 138, 183-185
type of, 370 Similarity, 68, 84, 144, 201-205
Reprints, 292 causes of, 202, 227
Reproduction, 283 overall, 124
uniparental, 30 phenetic, 203
426 Index

Similarity, quantification of, 203 Spelling(s), original, 312, 355

Simpson, G. G., 2, 4, 14, 19, 28, 31, Spermathecae, 129
35, 43, 44, 52, 55, 62, 67-69, 72, Spermatozoa, 127
75-77, 82, 85, 135, 140, 199, 213, Spider nomenclature, 342
216, 219, 341, 358, 369 Spieth, H. T., 136
Simpson, Roe, and Lewontin (SRL), 164 Splitters, 238-240
Sims, R., 211 and lumpers, 230, 238-241
Siphonaptera, 83 Sponges, 97, 246
Sister groups, 73 Squires, D. F., 108
Size, 123 SRL (Simpson, Roe, and Lewontin), 164
of taxon, 235 Stability, 248, 298, 301, 335
Slides, microscopic, 107 Stafleu, F. A., 210
Smart, J., 115 Standard deviation, 169
Sneath, P. H. A., 83, 203 Standards, 230
Sokal, R. R., 164, 203, 209 Star-type phenogram, 254
Sonagraphs, 136 Statistical analysis, 163-180
Sorting centers, 113 Statute of limitation, 350-352
Sotavalta, O., 124, 222 S teatoda, 187
Sound analysis, 136 Stenzel, H. B., 76
Specializations, narrow, 223 Sterility, 131
Species, allopatric, 193 Stirton, R. A., 253
asexual, 31 Stoll, Norman R., 299
chromosomal, 31 Strategy, 15, 258
definition, 26 Stresemann, E., 68
fossil, 10 Strickland, H. E., 299
grouping of, 199 Strickland Code, 338
intermediate, 242 Sturtevant, A. H., 203
inventory of, 10 Style, telegraphic, 267, 277
monotypic, 37 Style manual, 287-290
multidimensional, 31 Stylo ps, 156
nominal, 118, 378 Subgenus, 303
nondimensional, 27, 37 Subspecies, 41, 193-197
number of, table, 11-12 definition, 41
parapatric, 195 difficulties, 42
polytypic, 37-41, 64 nomenclature of, 43
difficulties, 38 or not, 188
primitive, 214 polytopic, 42, 193
relational definition, 26 temporal, 43
Species category, 23-36 Subspecies borders, 193
Species concept, biological, 25-27 Subspecies recognition, 189
difficulties, 30-35 Subspecific name, 361
nominalistic, 25 Superspecies, definition, 52
typological, 24-25 Svenson, H. K., 265, 343, 368
Species group, 47, 238 Syagrius fulvitarsis, 8
Species group names, 316, 361 Sylvester-Bradley, P. C., 35, 36, 44
Species problem, 23 Symbionts, 140
Specific name, 361 Sympatric samples, analysis, 181-187
Specimens, authentic, 119 Symplesiomorph, 202
Spectrophotometer, 270 Synapomorph, 202
Spelling(s), homonymous, 320 Synonymy, 273-276, 350
 Index 427

Synonymy, generic, 275 Topotypes, 375

Synopses, 260 Transliteration, 354
Syntypes, 327, 373 Triangular graph, 176
Systema naturae, 57, 342, 343 Trinomen, 302, 343
Systematic Zoology, 19, 43, 265 Triturus, 344
Systematics, biological, 14 Tubinares, 248
contribution of, 6-9 Turbellarians, 97, 245
definition, 2 Twins, 28, 68
future of, 19 Type by elimination, 368
new, 50-52, 64 Type-collection, 110, 372
as profession, 16-19 Type-concept, 321
theory of, 265 Type-designation, 332, 371-372
Systematist, training of, 9 Type-localities, 327, 375
Type-locality, correction, 377
restriction of, 376
t test, 174 Type method, 367—369
Tautonymy, 306, 357 Type-species, misidentified, 315
Taxa, adaptive significance, 139 Type-specimens, 120, 370
formal recognition, 229 Types, curating of, 109
higher, 87-98, 263 family-group, 321
monotypic, 236 and first revisers, 374
polythetic, 83 genus-group, 322
subordinate, 359 species-group, 326
Taxon, definition, 4 Typing, 290
higher, definition, 88 Typology, 67
qualifications, 92
Taxonomic character, 121
Taxonomic publication, title, 287 Umlaut, 354
Taxonomist, task of, 9-16 Uninominal nomenclature, 345
training of, 18 Uniqueness, 298
Taxonomy, chemical, 134 Universality, 298
definition, 2 Uvarov, B. P., 154
experimental, 64
history of, 55-65
numerical, 69 Vacelet, J., 246
role of, 8-9 Vachon, M., 126
Teratology, 156 Validity, 350
Termites, 243 Van Emden, F. I., 131
Tetrahymena, 336 Van Tyne, J., 106
Tetrapod phylogeny, 74 Variation, allometric, 154
Tetrodotoxin, 81 analysis of, 144-180
Thamnophis ordinoides, 174 continuous, 161
Theory, scientific, 79, 94 density-dependent, 154
Therapsid reptiles, 75 genetic, 158-162
Thielcke, G., 137 host-determined, 153
Thomomys, 130 individual, 146, 161
Thompson, W. R., 66 neurogenic, 155
Throckmorton, L. H., 72, 129, 134, 208, nongenetic, 147—158
219 seasonal, 150
Tinbergen, N., 136 social, 151
428 Index

Variation teratological, 156 Weighting, a posteriori, 60, 217-226

traumatic, 155 a priori, 217
Variety, 41, 362 Wells, G. P., 141
Vaurie, C., 50 Wheeler, W. M., 155
Vogel, H. J., 18 White, M. J. D., 132
Voss, E. G., 276 Whitney South Sea Expedition, 104
Wickler, W., 85, 136
Wagner, R. P., 138 Wilson, E. O., 42, 84, 212
Wagner, W., 84, 212 Wood, A. E., 141
Walker, T. J., 183 Woodpeckers, ivory-billed, 210
Weevils, 237 Work of an animal, 127, 309, 353
Weight, 106
Weighting, definition, 218
difficulties in, 226-228 Zone of secondary intergradation, 49
importance of, 211 Zoological Record, 116
phyletic, 226 Zweifel, F. W., 282
 DATE DUE / DATE DE RETOUR

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