An Introduction to Plant Taxonomy

AN INTRODUCTION TO

Plant Taxonomy

GEORGE H. M. LAWRENCE
DIRECTOR, BAILEY HORTQRIUM

c01lNELL UNIVEllSJTY, ITHACA, N. Y.

ILLUSTRATIONS BY
Marion Ruff Sheehan

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PREFACE

THIS BOOK is written for the adult amateur botanist and the student of
a J.x:al 1I0ra course at the college level. It is restricted in scope to the
vascular plants-fems, gymnosperms, and 1I0wering plants. It is intended
to provide sufficient information for an understanding of plant structures,
a comprehension of why we have different contemporary systems of
classification, an understanding of fundamental principles of plant
nomenclattlre, and an appreciation of the mechanics of plant identifica-
tion, including the preparation and pre,ervation of herbarium specimens.
Taxonomy is a complex subject. It is based on a knOWledge and
understanding of plant morphology and anatomy, cytology, and genetics,
as well as paleontology and plant geography. It is not presumed that the
user of this book has knowledge or training in these subjects, but it is
presumed that he has had the equivalent of an introductory course in
biology or botany. This volume is intended to serve as an aid to the
non-taxonomist in the use of a manual or 1I0ra that accounts for the
plants of any region of North America.
In writing this book, I have drawn freely from pertinent parts of my
earlier work, Taxonomy of Vascular Plants (1951). No bibliographies
have been included, and persons desiring references to original and
background papers are invited to consult that earlier work. In addition
to the more general taxonomic situations, there has been included a
selection of about fifty of the larger or more dominant families of
vascwar plants; lor eacn inere nas Deen !iJven inilf &siIDguiSning cnar-
acters, by which members of these families may be recognized at this
classification level. A chapter on the history of taxonomic studies in the
United States of America, as indicated by biographical sketches of the
makers of this history, has been included to provide background and
perspective.
•
 PIlEF.4Ui
The glossary provided in Taxonomy of Vascu/ar Plants, corrected and
amplified, is repeated in combination With an index. The figures used in
the earlier glossary have been transferred to appropriate chapters of this
new text. A study of glossaries in most ftoras and manuals demonstrated
the need of bringing the definitions of many terms into harmony with
modem concepts of morphological and anatomical situations. This is
especially true of terms pertaining to the organs of the ftower. Persons
requiring a more complete glossary !han provided here, or wisbing to
consult a listing of English equivalents of most specific epithets, are
invited to consult Featherly, H. 1., Ttuonomic Terminology of Higher
Plants (1954).
All illustrations in this book have been drawn by Mrs. Marion Ruft
Sheehan, and many were prepared especially for this introductory work.
Except as noted, these figures have been prepared from the plants
themselves.
GEORGE H. M. LAWRENCE
Bailey H orlorium
Cornell University, Ill=a, New York
CONTENTS

CHAPTER I. Introduction 1
Taxonomy, what it is, its impoJ"blDCO

CIfAPTER n. Plant Classification 6

A review of Pre-Linnaean taxonolllY. the wystemo of LlmI-. de
lussieu, Engler. Bessey, Hutchinson, Tippo

CIfA~TER In. Evolution and Units of Classification 19

CIfAPTER IV. Plant Structures 26

A review of the organography and terminology common to
the ferns, gymnosperms, and angiosperms

CHAPTER v. Collecting and Identifying Techniques 84

CHAPTER VI. Nomenclature 88

The role of nomenclature in taxonomy. the importance of Latin
names, and a review of the saliellt features of the International
Code of Botanical Nomenclature

CHAPTER Vll. Phylogeny and BioaysterruJtics 98

CHAPTER Vlll. Taxonomy i1l North America 103

As retlected by the lives and accomplishmellts of about thirty
of its leading scholars
0111
CONTENTS
CIl.\~TBR IX. Important Families and Their Characters 119
Presented with a view towards aiding the student to recognize
important families on sight, giving special emphasis to diagnostic
and distinguishing characters

147
 CHAPTER 1. Introduction

WHAT IS taxonomy?
Taxonomy is a science that treats of the identification, nomenclature,
and classification of objects. When concerned with plants, it is often
referred to as systematic botany, and in this book it is restricted to the
systematics of ferns, of conifers and other gymnosperms, and of the
flowering plants. These comprise the vascular plants. Taxonomy may be
considered the mother of biological Sciences. Before mall could study
plant structures, the way plants grow, or could accurately record the
plants about him, he had to know the names and characteristics of those
plants. In gaining this knowledge he has tried to group plants together
in accordance with their presumed affinities. Today, anyone dealing
with plants in any way depends on the labors of the taxonomist.
To understand the scope and functions of taxonomy, it is necessary to
understand what is meant by identification, nomenclature, and cIassili-
cation. It is necessary also to understand how each of these differs from
the others and how they are interdependent.
Identification is what one does when keying out an unknown, when
determining the kind of a plant by comparing it with a plant of known
( identity, or with a description of sucb a plant. If someone tells you only
the common name of a plant, he has identified it. The name given by
this identification may not be correct, but the function or process re-
mains the same. A more precise and formal definition of identification
would be:

the determination of a plant as being identical with or aimilar to III1Othet'

and already known element; or, as being unlike any previl>llsly known do-
ment and therefore one that i. new to science.
1
z INTRODUCTION TO PLANT T.4XONOMY
A plant may be identified by the aid of books or papers on the sub-
ject (such as manuals, fioras, monographs, or revisions), or by com-
paring it with plants, the identity of which is already known (such as
living plants in collections, or pressed and dried herbarium specimens).
One must remember that identification, in the strict sense, has nothing
to do with the correct name of the plant Or with determining that name,
for the latter-as we soon shall see-is a function of nomenclature.
Suppose you are given an armful of all the known kinds of lilies, and
that you spread them out on a table. Sorting them out, you place like
kinds in separate piles. For purpose of easy reference, you then assign
each pile a number. Having done this, assume that a neighbor hands
you a lily from his garden. Immediately, you recognize it to be like
those placed by you, for example, in pile number 8. By this recognition,
you have identified the neighbor'. lily. No name has been involved nor
is the assigned number a part of that identification. Hence, it is seen
that identification is a distinct function of taxonomy. Nomenclature is a
very different function. In practice, one usually finds that a specialist or
an author has handled the nomenclatural part so that, when one comes
to the end of the identification, there also is the correct name for the
plant.
Nomenclature has reference to the correct naming of the plant that
has been identified. It is· the part of taxonomy that tells us how to go
about the determination of what name is correct, whether a particular
name is only a synonym, or whether it has no standing at all. Botanical
nomenclature deals only with the Latin names of plants. It is not con-
cerned with common or English names (sometimes referred to as ver-
nacular names). The naming of plants is controlled by rather rigid
rules that are explained in the chopter on Nomenclature.
Classification is the placing of a plant (or group of plants) in cate-
gories according to a particular system, and in conformity with a nomen-
clatural system. A very simple system of classification is that whicb di-
vide. plants into such groups as trees, shrubs, and herbs. Another might
divide them into ferns, conifers, dicots, and monocot.. Classification Is
a grouping together of those plants whose similarities are greater than
their differences. In practice, species of plants having many characters
in common are united under the name of a genus, as the lilies are all
treated as species of the genus Lilium. Related genera, such as the
tulips, lilies, and fritillaries, for example, are classified as belonging to
In.roduction B
a single family, the Liliaceae. Modem systems of classification attempt
to align the various groups of plants in accordance with their presumed
relationships.
Taxonomy-it! Significance
Any understanding of the natural resources of the earth requires an
appreciation and knowledge of its plants. The role of taxonomy is all-
pervading and fundamental; too often it is taken for granted. A primary
objective of taxonomy, prerequisite to all studies and uses of plants, is
to provide an accounting of the kinds growing on the earth. For the last
two hundred years, and longer, botanists have worked toward the com-
pilation of a list of the kinds of plants. This work has progressed at
about the same rate as has man's scope and spread of communications.
The development of steamship lines, of railroads, highways, and air-
ways has enabled the botanist to penetrate regions never before explored.
Despite these aids to travel, many significant parts of the earth remain
to be thoroughly explored for their plants. The inventory, as it stands
today, is far from complete. Probably we know of no more than three-
fifths of the flowering plants of the world, and many of these are known
imperfectly.
The plants of the earth compose its flora, just as the animals compose
its fauna. One may refer to the flora of the world, to that of a continent,
or to the flora of a political unit as a country, state, or county. Some-
times a book that merely lists the plants (with or without their descrip-
tions) of a particular area is also referred to as a flora. The taxonomist's
record of the flora is the herbarium. The specimens collected, pressed,
and dried, compose the herbarium. In the 1700's and even later, the
snn.<Lw.<l nf .nru' n r .m<lW .<;"!:.,nrumhlikt> "",b.1.rnl'S..<ll' ~
.hr.r.ha<illD'
pages the dried specimens were pasted or sewed. Later, as collections
increased in size, and allowance had to be made for continued expan-
sion, the sheets were placed loose in folders and arranged in the pigeon-
holes of specially constructed cases. A collection of dried, pressed plants
is an herbarium, no matter if occupying only a notebook or if contained
in hundreds of cases. Each herbarium specimen is accompanied by a
label giving the name of the plant, the place where collected, by whom
and when. Other pertinent information should also be added. Each
specimen serves as a permanent record that the plant grew at a given
place, and that it was collected there at a particular season and year. It
4 INTRODUCTION TO PL..uIT T A.XONOMY
is only by assembling the plants of a major area in herbaria that the
plants can be studied comparatively and that those of a wide area can
be brought together to determine their similarities and differences.
These data about the kinds of plants are assembled in books or
articles known as floras or manuals. While a flora lists or describes
briefly the plants of a region, a manual assists the user in identifying a
plant of that area. This is done usually by means of keys. Another form
of presenting the results of taxonomic study is a monograph or revision.
This is a comprehensive treatise on the taxonomy of a natural group,
such as the genus Rosa (the rose) or the family Rosaceae (of which, for
example, Rosa, Spiraea, Prunus, and Amelanehier are members). In
general, the monograph differs from a revision in being more exhaustive
and comprehensive. Studies by the taxonomist enable others not only to
identify and name plants but also to handle intelligently the plant mate-
rial and products with which they work. Clearly, taxonomic studies are
fundamental to any subsequent study of the natural resources of an
area or of land potentials, and to evaluations of the raw materials requi-
site to allied activities such as forestry, medicinal work; horticultural and
agricultural pursuits, and pharmaceutical and biological industries.
Taxonomy demonstrates the great diversity of plants in nature and
their relationships. In this way it aids in academic studies relating to the
genesis, evolution, and heredity of plants. As documented knowledge of
the earth's flora accumulates, the taxonomist can begin to understand
better the inter-relationships of plants. From this knowledge he may
form a better system of plant classification. Other botanists-the phy-
siologist, ecologist, anatomist, cytogeneticist and morphologist-<lepend
on taxonomic findings to identify and name the plants with which they
work, as do others who work with plants and plant products. Here is
one of the major functions of taxonomy.
Opportunities In Taxonomy
Floras have been written to account for the plants of most areas of
the world. Many of these are over a century old and were writtell on the
basis of scant collections made at the fringes of a country before rail-
roads or roads existed or before modem concepts of adequate 1I0ristic
coverage were developed. Many of these same lIreas are being explored
anew, by trained men, and with collecting techniques unknown even a
half-century ago.
These areas whose plants are inadequately known are not all tropical
Inrf'Odudion 5
rain-for..ts, inaccessible arid regions, or Oriental lands difficult of ac-
cess. Some are here in the United States, in Mexico, and in Canada. The
fiowering plants of the southeastern United States are not adequately
known, those of northern and western Canada are only beginning to be
known, and, despite existence of 1I0ras published during the present cen-
tury, the lands of Central America and of Mexico have not been well
explored for their plants. The problems are many and open to any stu-
dent trained in taxonomy. In this country there is an acute need for
more complete county fioras, even of states presumed to be well bot-
anized. No Hora for Virginia, settled in 1607, has been written. It may
be noted that many of our best county 1I0ras are the resnlt of intensive
field studies and collections by amateur botanists.
At the same time, other problems await resolution by the trained
taxonomist who is most interested in monographic work. The need for
revisions and monographs of plant groups is of increasing importance.
Recently accelerated research in cortisone and allied products has fo-
cused attention on how little is known of the large genus Dioscorea.
This is an isolated example. There are a thousand and more other
genera equally in need of modem and thorough taxonomic study. Many
of these are large genera, of a hundred or more species; others of equal
importance are small and may have no more than a dozen species.
From the above, it should be clear that plant taxonomy is funda-
mental to all biological sciences and that its Objectives have only begun
to be achieved. With this perception of its underlying significance and
scope, there should be the realization that taxonomy is not an end in
itself, but that it is a tool by the use of which the student of other
biological sciences~)f facets of them---can bring his problems closer to
fmition.
CHAPTER ,II. Plant Classification

PLANT CLASSIFICATION is the arrangement of plants -into an

orderly sequence. It may be based on the form of plants-whether trees,
shrubs, or herbs. It may be based on numerical distribution of parts of
the flowers-the number of stameJls, pistils, or ovary locules. Or, it may
be based on real or presumed phylogenetic relationships. Most classifica-
tions of plants from the time of Aristotle to the present bave been of
one or another of these three fundamental types.
Plant classifications follow a system of arrangement set forth by a
specialist in matters of phylogeny. No less than fifty different systems
have been proposed, with two or three holding positions of dominance
in recent years. The differences in them often have been those of de-
gree. During the centuries nothing has exerted greater influence than the
introduction and establishment of the doctrine of evolution, the recogni-
tion of mutations arising in nature, and an understanding of the me-
chanics and principles of inheritance.
Changes in concept of taxonomic units and in taxonomic procedures
reflect the influence of the same factors that have brought about changes
in systems of classification. For this reason, a better appreciation of the
problems involved is achieved if selected classificatory systems are briefly
recounted and their significance described. A knowledge of them is es-
sential to a comprehension of the objectives of taxonomy as a whole.
Early man classified plants before he had a written language. Cer-
tainly he ate plants, or parts of them, used them for sbelter, and from
them fashioned weapons for slaughter and delense. For each use, some
sorts were found to be superior and others inferior. Man talked about
these plants. Names for them were a prerequisite to communication,
and, since many kinds of plants were involved, he must have classified
them as well as identified and narned them.
6
PIa.,. Claui/ic4tfo" 7
The first deliberate attempt to cla.sify all of the plaots known was
that of Theophrastus (370-285 B.C.), noted student of Aristotle. His
was a classification based on habit or form-trees, shrubs, undershrubs,
aod herbs-aod he distinguished those of annual, biennial, or perenniaJ
duration. In addition, he distinguished between determinate and inde-
tenninate infiorescences, between superior and inferior ovary position,
aod between polypetalous aod gamopetalous corollas.
Although several classifications were published during the period
from the time of Theophrastus to the seventeenth century, none was
predominant until that set forth by Joseph Pitton Toumefort in 1700.
Toumefort (1656-1708), a professor of botany aod medicine at the
University of Montpellier, France, followed Theophrastus in dividing
flowering plants into two categories of trees and herbs. Each of these
was subdivided into groups based on such characters as flowers petaI-
bearing or non-petal-bearing, flowers simple or compound (groups akin
to the polypetalous and gamopetalous subdivisions of later authors),
aod flowers regular or irregular. His system was the first to group !o-
gether as genera the plants we know as oaks, maples, peas, verbenas,
and willows. The system was widely adopted in Europe aod remained
dominant for nearly a century.
A contemporary of Toumefort was John Ray (1628-1705), re-
nowned English naturalist, who devised a classification, based to a large
extent on precepts of several earlier botaoists, accounting for nearly
18,000 species. It was superior to that of Toumefort in that Ray rec-
ognized the monocots as distinct from dicots, separated taxa on the
basis of fruit-type (cone-bearing, nut-bearing, berry-producing, pomif-
erous, pruniferous, etc.) aod subdivided these on the basis of leaf aod
flower characters. It was a system based on the form and gross morphol-
ogy of plant structures. Unfortunately, it was overshadowed by other
systems of the era, and its superior features were unrecognized for
nearly a century.
CAROLUS UNNAEUS AND ms CLASSIFICATION
The great Swedish naturalist and physician, Carolus Linnaeus (1707-
1778), was the dominant figure in systematic botany during his time--
a dominance that persisted for much of the half-century after his death.
At the age of thirty-two he was the author of fourteen botanical works.
Among them was his Genera Plantarum which was based on his se-
called sexual system. It was an artificial system by which all flowering
B INTRODUCTION TO PUNT TA.XONOldf
plants were subdivided among twenty-three classes, the first ten con-
taining respectively those plants with flowers having one, two, three, four
stamens, etc. Other classes were those whose flowers bad stamens that
were didynamous, tetradynamous, monadelphous, syngenesiolls, etc.
Each class was subdivided into orders on the basis of the nu'mber of
styles in each flower.
In addition to providing a numerical basis for classification, the Lin-
naean system offered a device wher,~by anyone knowing the funda-
mental flower structures, and having the ability to count, could identify
as well as classify an unknown plant. The system is artificial, because it
does not bring related genera-and often related species of the same
genus-together. It was not a natural classification nor was it intended
to be so. .
Linnaeus is remembered also for his famous work, Species Plantarum
(1753), in which he classified, described, and named every species
known to him. It represented the first time that a binomial (the Latin
name of a species, composed of two words-the generic name [such as
Quercus] and the specific epithet [such as alba]) was assigned to prac-
tically every species of plant known to the author. The importance of
this work is made clear in the chapter on Nomenclature.
DE 1USSIEU AND THE PERIOD OF THE NATURAL SYSTEMS
Two members of a family of famo,", French botanists were Bernard
de Jussieu (1699-1766) and his nephew, Antoine Laurent de Jussieu
(1748-1836). The first devised and the other improved and published
a classification system representing an improvement over that of John
Ray, for it utilized such additional characters as ovary position, fusion
of floral parts, and types of ovule arrangement. Their thirteen classes
of angiosperms have continued to the present to be accepted as valid
taxa, sufficient evidence of the keenness of perception evidenced by de
Jussieu.
It was Augustin Pyrame de Candolle (1778-1841) who amplified
and publicized the basic tenets of the de Jussieu system. This he did in
several works, but notably in his monumental Prodromus systematis
naturalis regni vegetabilis, a seventeen-volume work proposing to clas-
sify, describe, and name every species of plant then known. It was
never completed for all flowering plants.
The de lussieu system served as the basis for one that was improved
and publicized in English-speaking countries by John Lindley (1799-
"'_1 Cla.i/icadon 9
1865) in his Vegetable Kingdom. It was the first to be widely accepted
in Great Britain and America as a successor to that of Linoaeus.
I! was not until the British botanists George Bentham (1800-1884)
and Sir Joseph D. Hooker (1817-1911) published in three volumes
their famous Genera Plantarum that the basic concepts of de Jussieu
received their highest development. Their Genera Plantarum continues
to be a standard and mnch-consulted work, and one whose value is en-
hanced by Our knowledge that the detailed descriptions in it are based
on the authors' own meticulous observations of the plant structures and
are not copied from previous writings.
These systems, from that of de J ussieu to that of Bentham and
Hooker, are called natural systems, because in each case the authors
tried to group obviously related taxa together. All of these systems were
predicated on the dogma of the constancy and immutability of species,
and one represented improvement over another only in degree. All were
published in the era prior to the pUblicaiion of Wallace's and of
Darwin's theories of evolution and the origin of species, which auto-
matically closed this period in the history of classification systems and
introduced the next.

SYSTEMS BASED ON PHYLOGENY

The publication of Darwin's Origin of Species opened a new era in
biological thinking. I! refuted the dogmas of constancy of species and
of the origin of existing species by special creation, and displaced these
and related views with theories of descent and evolution. By these
theories most biologists hold that existing forms of life are the products
of evolutionary processes, and that some combination of characters are
the result of gene exchange, and others of mutations.
Recognition of evolutionary forces brought the realization that pres-
ent-day species of a given genus have a common ancestor, that present-
day genera of a family probably are descended from a common ancestor,
and that the plants of today are derived from plants long extinct (of
which a few are known from the fossil record). This stimulated paleobo-
tanical studies of fossils and allied relics of plants that existed millions
of years ago. These studies confirmed some earlier views and gave bases
for rejection of others.
A new direction was given to botanical thinking with regard to plant
relationships. Botanists were asking, "How are these plants related,
. . . from what are they descended, . • . which characters are those
JO INTRODUCTION TO PL4NT TnONOMY
of ancestrally more primitive plants and which ones are derived from
them, . . . which characters have longest remained unchanged, . . .
have characters arisen only once or have some arisen many times inde-
pendent of one another?" The term phylogeny (meaning "the evolution
of • 'genetically related group of organisms" [Webster]) came into being,
and botanists began to rearrange plant families in an effort to place to-
gether those families believed to be more closely related to one another,
with derivative groups following those presumed to be ancestral. None
of these efforts, not even those of the present day, has developed a truly
phylogenetic system. One reason is that not enough is known of prehis-
toric forms to enable the botanist to know, from the evolutionary stand-
point, which characters reflect primitive and which reflect advanced
conditions. However, this revolution of thinking has resulted in a score
or so of different classification systems for higher plants. A few are well
known and adopted in various countries, others are known locally, and
some have been accepted by few except their authors.
American floras and manuals are based, for the most part, on one or
another of three systems of classification-<Jr of recent modifications of
them. They are the systems of Engier, of Bessey, and of Hutchinson.
Contemporary systems in some European countries are those of Pro-
fessor Pulle of Utrecht, and of Professor Skottsberg of Stockhoim. Most
British floras continue to follow the Bentham and Hooker system of
classification.
The Engler Syotem
The dominant system of classification in North America today con-
tinues to be that proposed over sixty years ago by Adolph Engler. It is
the basis ior arrangement oi iamilies of higber plant. in manuals ano
floras such as those by Fernald, Gleason, Small, Jones, Rydberg,
Abrams, Jepson, and Rehder, and for the arrangement of specimens in
the larger herbaria of the country.
The widespread acceptance of the Engler system has not resulted from
its correctness-it was not an original work, nor are its major tenets ac-
cepted today, to even a substantial extent-but rather from the system-
atic thoroughness with which Engler and his colleagues applied the sys-
tem to the flora of the world. In his first rna jar work, using this system,
he applied it to the classification of all the families of plants and their
genera. This work, called Die natiirlichen Pflanzenfamilien, was pub·
Iished under the joint editorship of Engier and his colleague, A. Prantl.
Plan. Clauifi,cation 11
It is in twenty volumes and accounted for all the genera of algae, fungi,
bryophytes, and higher vascular plants then known for the world. Each
genus is described, accounted for in keys, and many are well illustrated.
This is the last work of its scope to account for the world's flora at the
generic level. The system has been revised from time to time by the

i ACHLAMVDIE I HOMOIOCHLAMVDIE

--
HETEROCHLAMVDlf

SCITAMINAE

9A
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~E
R
M
~

A.CHLAMYDIE HOMOIQCHLAMVDI£
ARCHICHLAMVDEAE

FIg. 1. Classification of the angiosperms, after Bnaler.

 INTRODUCTION TO PUNT TAXONOMY
publication of a Syllabus der tUltlirlichen Pflanzenfamilien, a single
volume accounting for the families of all these major groups but lacking
detailed review of the genera and their components; Each revision of
the Syllabus has held to the major tenets of the Engler system, and no
recognition has been given to evidence in support of changes of the basic
framework. .
Engler's system, based largely on an earlier and little-known system
by Eichler (1883), divided plants among four divisions: the algae and
fungi, the mosses and liverworts, the ferns, and the seed plants. The last
was subdivided into the gymnosperms (cycads, ginkgo, conifers) and
the dowering plants (composed of the monocotyledoneae and dicotyle-
doneae). In the classification of the dicot families, the most primitive
were considered to be those families whose dowers had no petals (the
Apetalae, e.g., willows, birches, oaks), secondly those families wbose
dowers had separate petals and sepals (the Choripetaleae, e.g., butter-
cups, chickweeds, roses, violets, cacti), and lastly those families whose
dowers had the petals and the sepals united to varying degrees in two
separate perianth units (the Metachlamydeae, e.g., primuJas, borages,
mints, cucurbits and daisies). Basically, Engler followed the belief that
dowers now appearing simple had always been simple, a view rejected
by most present-day botanists.
Much has been learned about plants since the time of Engler, and it
is now generally accepted that his system is not based on the best evi-
dence. Its present position spring. from the reluctance of taxonomists to
change today for a system that may be subject to displacement tomor-
row. Nonetheless, such a change is overdue and is to be expected. By
modem views the Engler system is archaic. The peak of its utility is past.

The Beuey System

Charles Bessey, for many years head of the botany department at the
University of Nebraska, was the first American to devise a system of
cJass.ification. It was, in its latest revision (1915), a modification of the
Bentham and Hooker system, but it rellected to a marked degree the in-
lIuence of evolution on the thinking of that day. The system, despite its
obvious merits, has suffered from lack of publicity.
Bessey took many of his basic ideas from de Jussieu, from de Cau-
done, from Lindley, and perhaps some of them from his contemporary
German botanist, Hallier. In his system he attempted to collate what he
thought to be the beSt of these systems (including adoption of some
EngJerian principles). The arrangement does not differ so much fn;m
14 INTRODUCTION TO Pl.A.NT TA.XONOMY
that of Bentham and Hooker or from that of EngIer, as the latter two
differ from each other.
According to Bessey's system, the seed plants are composed of three
phyla: angiosperms, gymnosperms, and the extinct seed-ferns (Pterido-
spermae). Only the angiosperms were treated by Bessey, a taxon 1 he
considered to be derived from the cycads. He interpreted the strobiloid
type of flow"r, with many spirally arranged parts (e.g., that of magnolia)
to be primitive and flowers showing fusion of unlike parts (as corolla
and calyx to ovary, in most kinds with an inferior ovary) to be more
advanced; those showing fusion of like parts (e.g., petals fused to form
a one-piece corolla) and those whose flowers show both types of fusion
he considered to be the most highly advanced of all.
The Bessey system is little known outside the United States, and for
several reasons: it has never been applied to the world's flora; it has not
b<>en championed in recent time by an influential school of taxonomic
study; and it has not been revised in the light of recent knowledge. This
third factor is probably responsible for the limited acceptance of the
system today in this country. While many botanists recognize its good
points and accept it as being closer to a realistic phyletic classification
than any other system, they also know that many of the family relation-
ships embodied in it are contrary to present evidence. A critical and
documented revision of this system, accompanied by its association with
the classification of other segments of the plant kingdom and applica-
tion to the world's flora may produce a more acceptable arrangement
than any now available.

The Hutchinson System

John Hutchinson, now retired from the staff of Britain's Royal Botanic
Garden at Kew has published a system of classification (1926-34)
based on many of the same prin'ciples adopted earlier by Bessey but
differing in two fundamental respects. One point of divergence is his be-
lief that the dicotyledonous plants represent two distinct subgroups: the
Herbaceae, composed of families whose members are fundamentally
and predominantly herbaceous, and the Lignosae, composed of families,
whose members are fundamentally and predominantly woody. '}:he

1 The term taxon (plural, taxa) is of recent widespread usage in botanical literature.
although coined and introduced about a quarter-century ago, It is used to designate any
taxonomic entity or group without indkating the category to which it roay belong. It
is the plant, the population, or any number of populations that is the taxon-not the
unit of species, nor of genus, or any other category. For example, the taxo~ Gleason
called Arabis Shortii was treated by Fernald as Arabis perstell(Jta var. Shorti,.
15
16 INTRODUCTION TO PLANT TAXONOMY
second departure from the Bessey system is Hutchinson's defendable
view that families with corollas of fused petals have not had one or a few
common origins but have evolved independently of one another and thus
do not represent a single advanced levef.
The Hutchinson classification of the monocots has been viewed more
favorably in America than has his treatment of the dicots. For the dicot.
and the monocots, he has treated the families of the world, but" in neither
case has he presented reasons for his alignments, nor is there evidence
that he gave consideration to available morphological, anatomical, and
cytOlogical findings. This system, by its critical appraisal of family re-
lationships and by the excellent keys published with it, has provided more
grounds for discussion and greater stimulation of thinking along phyletic
lines during the last two decades than has any other.
No floras, except those by Hutchinson, have been based on this system
of arrangement, although one book devoted In American wild flowers
has adopted it.
Other Systems
The systems encountered in American herbaria, manuals, and floras
have been described briefly. There are other systems used currently to a
limited extent in other countries. These include the earlier system by
Wettstein of Vienna (a modification of the Engler system and in many
respects an improvement over it), that by Pulle of Utrecht (a con-
temporary system little known in America but deserving of greater con-
sideration), that by Skottsberg of Goteborg (a modernization of Wett-
stein's system), and a few less well known systems.
In 1942, Professor Oswald Tippo, of the University of Illinois, pro-
posed in very broad outline a classification of the plant kingdom based·
largely on anatomical and morphological criteria. By this system the
vascular plants were subdivided into four subdivisions of the phylum
Tracheophyta, with the last composed of the three classes indicated:

Psilopsida
Sphenopsida
Lycopsida
Pteropsida
Filicinae
Gymnospermae
Angiospermae
 /
""------
""I'"
......
,
I --- I
\~,I,__,
\

, i1t';J.... I
..... _---- '"

__ ------.
_/

___ _____ ~ TRACH_~_EOPHYT ___A_____ . _ . <;z:~---

CHLDr"'A

FIg. 4_ Classification of the angiospertllll, adapted from Tippo (1942). Extinct

taxa are marked by an asterisk. The Bryophyta and Chlorophyta (algae) are non-
vascular cryptogams.
17
III INTRODUCTION TO PLANT TAXONOMY
Unfortunately for taxonomic usage, Tippo,did not elaborate his system
below the level of the monocots and dicots, No classification was indi-
cated for the families of these groups.
From this brief review, the query may be raised, "What is the correct
system of classification?" The answer, and one so common to many
biological situations, is that there is none. Oassifications are embodied
in the taxonomy books one uses-the floras, manuals, monographs, and
revisions. Current teaching practices in the United States reflect the
adoption to an increasing extent of the Tippo classification, with many
authors filling in the gaps for the flowering plants with adaptations from
the Bessey system. This is especially true in current books and courses
dealing with elementary botany, where a trend set by morpbologis1s
seems certain to exert an influence in taxonomic works of the future.
CHAPTER Ill. Evolution and Units
of Classification

EVOLUTION, in the field of biology, is the doctrine which holds that

present species of living organisms have descended from more primitive
ancestors and that through successive generations--perhaps tens of
thousands of them-a series of modifications has occurred. These modifi-
cations, or changes in form and function, have been brought about by
various means, including natural selection, hybridization, and mutations.
The mechanism of natural selection may be illustrated in its simplest
form by the following sequence of events. Assume that a number of
individuals of the same parentage occupy a given area and that thoy pro-
duce offspring by sexual reproduction. Assume forther that there are
some genetical differences among tbese individuals; no two sexually re-
producing individuals are exactly alike. Nata",1 selection acts on this
variability. It works in tbe direction of adaptation, for some of the varia-
tions may be toward qualities or cbaracters that better enable the in-
dividual to compete and succeed over others or may enable it to grow,
reproduce, and invade areas where conditions are favorable or unfavor-
able with respect to members of the initial population.
Hybridization is the crossing of two genetically different individuals
by the fusion of gametes. It is considered that some present-day species
originated-probably thousands of years ago-by the hybridization of
two other species now extant. Products of this kind of evolution perhaps
include such well-known plants as timothy (Phleum protense), annual
bluegrass (Poa annual, tobacco (Nicotiana Tabacum), the common
plum (Prunus domestica), and Spartina Townsendii.
Mutations result from changes in the molecular structure of the gene,
19
 INTRODUCTION TO PLANT TAXONOMY
and these changes are reflected in the characters which the genes control.
Mutations may alIee! major or minor characteristics. These changes are
heritable and are transmittable from generation to generation.
It was once believed· that evolutionary changes were caused directly
by changes in environment, as for example, when a habitat gradually be-
came more desert-like, the structures of SOme of the plants growing in it
were modified, and their descendants became adapted to the new en-
vironment, or that organs or structures were changed by use or disuse
on the part of the individual. This belief, known as Lamarckism, is not
now generally accepted.
Following the work of Lamarck, and as a product of studies by Dar-
win, there came into mOre general acceptance the belief that evolution
results from the transmission of variations from parent to ollspring.
Mendel demonstrated that characters among hybrids are inherited in cer-
tain predictable ratios.
Following the reasoning presented above, it is now generally held that
existing species have been produced through evolutionary processes.
Severo related species may have had a common ancestor. These several
species may compose a genus, in which case this common ancestor was
that of the genus. Perhaps another allied genus also had a common an-
cestor. If the two ancestors of the genera in question came from the
same stock, it could be said that those genera and their species had a
monophyletic origin (at least back to the level under discussion). This
implies a forking or branching type of ancestry. Some botanists carry
this concept to the extent of believing that the dicots as a taxon, or the
monocots similarly, had a monophyletic origin; others extend it to the
angiosperms, while there are authors who have postulated that all plant
life has arisen from a single unicellular individual. Few botanists today
hold these extreme views of development by monophyletic lines. The
greater number of botanists, while accepting monophyleticism for groups
of specie:;, genera, and even for some families and orders, consider the
major taxa to have had polyphyletic origins; that is, groups now con-
sidered a part of the same taxon are presumed to have evolved from
several ancestral forms. The features in common of taxa within these
major units are believed to represent the result of convergent evolution.
In other instances, selected characters (such as the inferior ovary, or the
gamopetalous corolla) have arisen in dillerent genera and families (and
even in dillerent orders) by what is termed as parallel evolution. That is,
characters resulting from parallel evolution <l!,d not have a common an-
EtJOIutlo" cmd URI.. 01 Ciaui/i<..lo.i -J
cestor responsible for that character; they do not express a monophyletiC
origin.
Regardless of the type of origin (monophyletic or polyphyletic), it is
accepted that some groups of species are more closely related to one an-
other than to species within other groups. Several species, comprising a
genus, may have had a common ancestor (possibly extinct). These units
of classification-species. genu~, family, etc.-are assumed to be natural
units in that they are genetically related. Because of this genetical re-
lationship, some genera are more distinct than are others; sometimes
they are distinct, but in other instances a group of allied genera may
lack fixed limits or boundaries (in which cases, authors may not concur
on the number of genera involved).
To a lesser extent the same situations prevail at the lower level of the
species. There is no universally accepted definition of what a species is.
Despite the convictions and efforts of some geneticists, the determination
of species limits or circumscription is a matter of opinion-an opinion
based on knowledge of the components of a group, on judgment, and on
taxonomic competence. These differences in interpretation account for
the fact that the late Professor Fernald recognized nineteen species of
Juneberry (Amelanchier) for northeastern North America, while Glea-
son recognized eight species of the same genus for approximately the
same area. This exemplifies the difference in judgment of the same data
by two eminent taxonomists and illustrates the extent of individual
opinion in taxonomic studies. It cannot be said that one author is right
and the other wrong. Each is entitled to his own opinion. The student
may accept one or the other or reject both in favor of still other views.
Such is the freedom requisite to aU scientific work in the field of biology.
The units of classification and their sequence of relatIOnship are
established by the International Code of Botanical Nomenclature (cf.
Chapter VI, pp. 90-91). The plants of the world compose the vege-
table kingdom (as opposed to the animal kingdom). This, the largest
unit, is subdivided into successively smaller units, as follows (examples
based on the Tippo system of classification):
Division (sometimes, but not legitimately, designated Phylum)--example: Tra-
cheophyta (the vascular plants),
Subdivision--example: Pteropsida (the fems and seed plants)
Class--example: Angiospermae (the flowering plants)
Subclass--example: Dicotyledoneae
Order--example: Rosales
Family--example: Rosaceae
zz INTRODUCTION TO PL4IVT TAXONOMY
Subdivisions of family include: subfamily and tribe, each composed
of one or More genera
Genus---example: Rosa. Subdivisions of genus include: subgenus,
section, subsection and series. in that order. These are only of
taxonomic- significance to show affinities of included species.
Specics--exampJc: Rosa .retigera
Varietas
Forma
Clone

The Latin names given to taxa in the highest four units of classification
vary with the system followed, but the names and sequence of the units
themselves (as division, class, etc.) are always the same. This is illus-
trated in Table I. From this table, it will be noted that, except for Tippo's
system, the taxa are retained in the same categories but sometimes are

Table I. Names ,pven t& major unite of dicotyledonous plante by various

authors.

..umoa op DIVIStON
BY""'" (P~yl...) SUBDIVIIION CLASS SUBCLAlIS

Ben""'" ODd
Hooker
Phanero'lDlae Angiospermae Dicotyledoneae Polypetalae

~ EmbzyopiJyta A:JsWopermae DlcotyledODeae Archichlamydeae

SiphoD()JJInara
IIeaoJ PhanerollDlae Anthophyta Altemifoliae Strobiloideae
TIppo Tncllcophyt. Pteropaida Angiospermac Dicotyledoneae

given different names. The system of Tippo, in which the vascular plants
[the ferns and the seed ,plants] compose a single category (a division),
shifts the taxon Dicotyledoneae from the category of class to subclass.
Note, however, that the sequence of the units remains the same. The
shift in taxonomic emphasis reflected by this system is an expression of
the author's opinion. Many contemporary authors believe that there are
no discrete taxa intermediate between the Dicotyledoneae and the orders
that compose them (a current view in opposition to that held a half-
century and more ago). Certainly there is little evidence to support the
taxonomic validity of ~ subclasses accepted by Bentham and Hooker
and by Engler (shown in Table I).
The category of order stands below that of class (and subclass) and
above that of family. Names of orders end in -ales, such as Ranales and
Rosales. Generally, an order is composed of several families, such as the
Evolution and Unit. oj Cla,,'jicarloft 23
Rosaceae, Saxifragaceae, and Crassulaceae, included among those placed
in the Rosales. The components of an order usually are bound together,
or have in common, one or more phyletically important characters. Some
orders contain only a single genus, one that is so distinct from all others
as to comprise not only a family by itself, but a separate order (e.g.,
Ginkgo of the Ginkgoales, Welwitschia of the Welwitschiales, and Cas-
uarina of the Casuarinales).
The family is the largest category commonly encountered in routine
taxonomic work. It is usually a readily recognized taxon composed of
one or more genera whose similarities are greater than their differences.
Names of families end in -aceae. with the exception of eight families
whose names end in -ae. However it is admissible to use alternative
names ending in -aceae for those eight exceptions and an increasing num-
ber of authors have adopted the alternative names. These eight families
and their a1temative names are:
Gramineae - Poaceae Guttiferae Clusiaceae
Palmae - Arecaceae Umbelliferae Ammiaceae
Cruciferae - Brassicaceae Labiatae Lamiaceae
Leguminosae - Fabaceae Compositae Asteraceae
A family is often distinguished by characters of reproductive structures
(e.g., the fruit in such families as the Aceraceae [samara], Umbelliferae
[schizocarp], Leguminosae [legume, loment], and Cucurbitaceae [pepo],
or the inflorescence in the Gramineae [spikelet]. Fagaceae lament],
Amaryllidaceae [umbel], and Compositae [head]),' or by combinations
of these and other characters.
The genus is often recognizable by one or more characters of gross
morphology. The names <.f genera are nouns from any source whatso-
ever. They may be masculine, feminine, or neuter in gender and may be
in Latin or Latinized derivatives from the Greek.
The species (abbreviated .p. when singular and spp. when plural) is
the basic unit of classification. A species name is composed of two words,
the generic name plus a specific epithet, such as Rosa gallica. The species
name is sometimes designated the binary name or the binomial, and the
species epithet (e.g., gallica) the binary or specific epithet. Specific
epithets are ordinarily spelled with a small initial, but in much of the
literature certain specific epithets may commence with a capital initial
(when named for a person, when commemorating a generic name, when
a noun in apposition, and when commemorating a barbaric name). The
24 INTRODVCTION TO PLANT TAXONOMY

practice of capitalizing certain specific epithets is now optior~l, with an

increasing number of authors decapitalizing all specific epithets.
lnfraspeci{ic units of classification arc those whose rank is below that
of specie,. A wbspecies (abbreviated ssp. or subsp.) is that category
immediately below species. It is variously defined; by some authors it is
a "baby species" or an "incipient species" (one whose evolutionary de~
velopment has not progressed to the' extent of its being mar~edly and
consistently distinct from its parent species)) by others the sUQspecies is
a subdivision of a species having its own distribution but not sbfficiently
distinct (morphologically or genetically) to deserve elevation to the rank
of species. A varielaS (more commonly known in this country by its
English name varin),. and abbreviated var.) is a unit subordinate to the
subspecies when the latter category is used (otherwise it is subordinate
to the sl'ecies) and like the subspecies is subject to varying definitions.
The varietas of some authors is treated as subspecies by others. A forma
(abbreviated t.) is a minor variant (the variation perhaps often caused
by a single gene difference, or distinguished by a single character) of a
species or higher infraspecific unit, and one lacking constancy and usu-
ally occurring sporadically in random populations (such as albino
forms, or forms differing in vestiture, or minor Jeaf character). The clone
(abbreviated c/.) is a vcgetative propagule, an individual; generally the
term is reserved for individuals of horticultural value (e.g., the so-called
garden "varieties" of rose, iris, gladiolus, etc.). The apomict is genetically
the same as a clone. It is a plant grown from a seed which has developed
from an ovule that was not fertilized. It contains no paternal germ plasm
and is the cytogenetic eu"uivalcnt of its "mother."
In considering these units of classification, it is important to remember
their sequence of arrangement, to remember that a class is composed of
one or more orders) an order of one or more families, a family of one
or more genera, and a genus of one or more species. The student un-
famili~r with Latin must remember that the plural form of the word
species is no different fwm the singular form but that the plural form of
the word genus is always genera. It should also be remembered that the
ending for family names is -aceae and for that of order is -ales. The
botanically trained ear becomes tuned to spot the name of a family im-
mediately by association of that ending -aceae, and of the next higher
category, oKier, by association of the ending -ales.
The study of evolution soon becomes complex and technical and,
when pursued, leads one to the related fields of genetics and cytogenetics.
Evolution and Unit. 0/ t:la6&ijication 25
The student should not be disturbed or unduly confused over matters of
phylogeny and the evolution of present-day plants, but he should find
solace in the knowledge that even specialists are no more than intelligent
speculators, arriving at conclusions with a minimum of evidence. Be-
cause of the lack of knowledge with regard to the sequence of events in
the far distant past, the evidence is not now at hand on which to base
dependably any phylogenetic system of classification. The student should
view any new system of classification with an open mind, knowing it is
unlikely to be the ultimate, and should be prepared to accept its later re-
jection for another. These systems may reflect stages or levels in the in-
crease of knowledge, and as such each plays a contributory role to be
welcomed and not decried.
 CHAPTER lV. Plant Structures

PLANT IDENTIFICATION is based on one's ability to recognize the

characters by which one kind of plant may be distinguished from an-
other. This requires a knowledge of plant structures and a familiarity
with the names by which they and their components are known. The
terminology employed in any manual, flora, or plant dictionary is ex-
tensive. The names or terms used may seem unduly technical, alien to
one's vocabulary, and in the aggregate may (and sometimes do) con-
tribute to a jargon that is almost meaningless to the non-botanist. None-
theless, these terms do give precision to expression, they are concise, and
they are a part of botanical literature.
The greater the scope and detail of a Bora or manual, the more techni-
ca! it is likely to be. One of the ways to comprehend this technica! jargon
is to take the time to study and comprehend the gross morphology of a
plant, to understand the relationships of its parts, and to know the terms
applied to them. These terms are often in polysyllables and, like the
scientific names of most plants, are generally taken from the Latin or
Greek. A knowledge of one or both of these languages may facilitate
understanding the terminology, but it is not a prerequisite. A technical
terminology becomes essential when precision and exactness are re-
quired. It enables a writer to express much with a single word. Botani-
ca! terminology has a heritage and has developed over the centuries.
When provided with a knowledge of botanical terms and the barest
minimum of Latin grammar, anyone can read or comprehend a Latin
description of a plant.
This book treats the vascular plants: the ferns and lycopods, the
gymnospel'll'o, aud the angiosperms. The gross structures of each are
markedly different, and, in general, each has its own terminology. FOI
%6
Plant Structure. %7
this reason the morphology and terminology peculiar to each major
group is taken up separately.

VASCULAR CRYPTOGAMS
The vascular cryptogams include the three subdivisions Psiiopsid,
(Psi/o/um), Sphenopsida (horse-tails), Lycopsida (Club-mosses), an'
the class Filicinae (true ferns) of the Pteropsida. These are spore-pro
dueing vascular plants. Tbey do not produce lIowers, fruits, or seeds.

Fig. 5. SELAGINELLACEAE. A, Selaginella pallescens: Aa, habit of plant; Ab,

branch tip; Ac, fertile branch, vertical section; Ad, same, showing megasporangium
(left) and microsporangium (right). B, Selaginella Kraussiana: Ba, habit of plant;
Bb, branch tip. (From L. H. Bailey. Manual of cultivated plants. The Macmillan
Company, 1949. Copyright 1925 and 1949 by Liberty H. Bailey.)

A spore is a simple reproductive body, usually composed of a single

detached cell, and contains a nucleated mass of protoplasm (but no
embryo) that is capable of germinating and developing into a new in-
dividual. In the vascular cryptogams, the spore germinates and produces
a tiny, independent, usually green, fiattened tballus, the gametophyte.
The gametophyte is sO small that most persons never notice it in nature.
It bears sex organs (of one or both sexes), and hom the fertilized egg
of the archegonium (the female organ) is produced the familiar con-
spicuous plant. In contrast to the gametophyte, it is called tbe sNro-
phyte. The sporophyte is the usually foliaceous spore-producing plant.
In most ferns the spores are alike, and the species is said to be homos-
porous. In other vascular cryptogams (e.g., Selaginella, Fig. 5; lsoeles,
Fig. 6; and Saivinia, Fig. 7), two kinds of spores are produced, and the
species is said to be heterosporous. These two kinds are microspores that
on germination produce antheridia-bearing (male) gametophytes, and
megaspores that on germination produce archegonia-bearing (female)
gametophytes. The surface of spores may be variously pitted, net-like,
 28 INTRODUCTION TO PLA.NT TA.XONOMr

knobby, or spiny-features that may prnvide characters for identifica-

tion.
Spores are borne in cases called sporangia. These are of many types
and shapes (Figs. 8 f, g, h). In the Filicinae, or true ferns, the spo-
rangium may bear a ring or cluster of thick-walled cells, the annulus, and
is then said to be annulate (Fig. 8 f showing an incomplete and vertical

Fig, 6, lsoETAcEAE. I,metes Engelmannii: a, plant, habit; b, corm, with attached

leaf bases; c, leaf, cross~section; d, leaf base, ventral side; e, leaf base, vertical
section~ f, corm with portion of leaf bases, vertical section; g, megaspore. (i,
velum; I, ligule; me, megaspores; mi. microspores; s. sporangium.)

annulus, Fig. 8 h showing a complete and apical annulus) or no annulus

may be present (Fig. 8 g). The sporangium may be stalked (Fig. 8 f)
or sessile (Fig. 8 g). Its mode of dehiscence or opening may be trans-
verse (Fig. 8 f) or by a terminal slit (Fig. 8 i).
Fern sporangia may be scattered over or may densely cover the under

FIg. 7. SALVINIACEAE. A. Salvinia rotundi/olia: Aa, habit of plant; Ab, single

plant with sporocarp and submerged pinnatisect leaves; ACt sporocarp~ Ad, sporo~
carps, vertical section with megasporangia (left) and microsporangia (right).
B, Azolla filiculoides: Ba, habit; Bb, sterile branch; Be, leaf. (From L. H. Bailey,
Manual oj cultivated plants, The Macmillan Company, 1949. Copyright 1924 and
1949 by Liberty H. p.i1ey.)
 ~ ,/-:;;:;
--~

"II

Fig. 8. FILICINAE: a, simple leaf-blade; b. one-pinnate leaf: c, one-pinnate leaf,

the pinnae one-pinnatifid; d, naked rl1izome; e, scaly rhizome; f, sporangium with
incomplete annulus; g. sporangium; h, sacus, single sporangium; i, segment of
fertile frond; j, naked SOrllS; k. sorus subtended by lacerated indusium; I, sarus,
indusium pcltate; m. same, vertical section; n, sorus, indusium cup-shaped; 0,
same, vertical section; P. sarus a marginal flap; q. same, vertical section; r, sarus
marginal, contiguous; s, sa.me, much enlarged; t, veins dichotomous and free; u,
veins anastomosing and reticulate.
29
30 INTRODUCTION TO PLANT TAXONOMY

(dorsal) side of a leaf. More often they are arranged in tiny distinct
clusters called sari (sorus, the singular form). The sorus may be sub-
tended, surrounded, or more or less covered by an indusium. This in-
dusium may be superior and umbrella-like (peltate) (Figs. 81, m), or it
may become slit or opened towards the central stalk and is more or less
horseshoe-like. It may be inferior and cuplike (Figs. 8 n, 0), or it may
be inferior and shield-like (the shield may be segmented into ribbons as
in Fig. 8 k). In some sori the indusium is a marginal flap of the leaflet
(Figs. 8 p, q, s). When the sorus has no indusium it is said to be naked.
However, by the time the induslate sporangium is fully developed, the
indusium is often shrivelled and may have lost its characteristic form.
For this reason, whenever possible, examination should be made of a
series of sori of varying degrees of development to determine more ac-
curately whether or not an indusium is present and what are its charac-
teristics.
The position and arrangement of sori are of taxonomic importance.
Sori may be distributed over the dorsal surface of all leaflets or only on
the more terminal leaflets (Fig. 8 b). In some taxa they are in a row, and
when sO close together as to touch one another and appear as if one
"long sorus" are said to be confluent or contiguous. These rows may be
marginal (Figs. 8 r, s) or may extend along the mid-vein or a lateral
vein.
The vegetative structures of ferns are of equal importance to identifi-
cation. This is more true when engaged in field studies than in critical
revisionary studies. The habit of growth is important. Except for the
exotic and subtropical tree-ferns, the stems of most ferns are rhizom-
atous-although some ferns are climbers by the production of slender
stem-like leaves. These rhizomes may be scaly (Fig. 8 e) or naked (Fig.
8 d). Sometimes when scaly, or woolly, the vestiture (covering) extends
to the leaves. Often the rhizomes are creeping.
Fern leaves provide many taxonomic characters. They are usually
circinate in vernation. That is, they are usually rolled coil-wise in bud
from the'top downward, with the apex nearest the center of the coil.
Young Circinate fern leaves just beginning to expand are sometimes
called fiddleheads or crosiers, from their resemblance to those objects.
In much of the literature, fern leaves are designated fronds, a term ap-
plied also to the compound leaves of cycads and palms. There is no
structural difference between a leaf and a frond.
A fern leaf may be simple or compound. When simple, it consists of a
Plant Srructure$ 31
blade and a stalk or petiole (sometimes termed the stipe). The blade of
a simple leaf may be entire (as is the leaf-blade of lilac or grass) or it
may be variously toothed, lobed, or dissected. When a portion of the
tissue of adjoining lobes or segments of a deeply divided leaf extends
along the mid-vein and connects one segment of the blade with another,
that leaf is considered simple.
A fern or other leaf is compound when the segments are compl~tely
separate (Figs. 8 b, c). Each segment is a leaf/et. In fern literature the
leafiet is commonly called a pinna. That portion of the axis of a com-
pound leaf from the lowest pair of pinnae upwards is termed the rachis.
A leaf with one rachis and a single series of pinnae is one-pinnate (Fig.
8 b). In some ferns each pinna is compound and its mid-vein bears pairs
of lateral secondary leafiets called pinnules. Such a leaf is two-pinnate or
decompound (Fig. 13 f) and some decompound leaves are three-pinnate
or more (Fig. 13 g).
In most ferns the leaves of a given plant are more or less alike. When
sporangia are present the leaf is said to be tertile, when not present it is
said to be sterile. In a few genera or families tbe plants produce some
leaves that remain sterile and others that become fertile. When the fertile
leaves are dissimilar to the sterile ones, the leaves of that plant are called
dimorphic. Leaf dimorphism may be represented by the fertile leaves 01
segments conspicuously narrower, bearing abundant and often densely
congested sporangia. In the more extreme examples, one part of the leaf
may be foliaceous and another reduced to an apparent axis covered with
sporangia.
The venation (arrangement of veins) of fern leaves is of taxonomic
significance. In some ferns, the veins branch by forking successively in
pairs, a type of branching termed dichotomous (Figs. 8 p, r). The veins
may extend to the margins, not uniting with others, and are free (Fig.
8 t), or they may anastomose (coonect with one another) to form a net-
work, and the venation is reticulate (Fig. 8 u). The meshes of such
a network are called areoles.
Explanations and illustrations of terms of vestiture, leaf form, margin,
and texture are to be sought in the section of this chapter devoted to the
angiosperms.
GYMNOSPERMS
The gymnosperms, together with the angiosperms, compose the
present-day group known as the seed plants. The seed plants (sometimes
32 INTRODUCTION TO PLANT TAXONOMY

designated as phanerogams or spermatophytes) differ from the vascular

cryptogams in that the gametophyte is reduced to a comparatively few
cells, always enclosed within sporophytic tissue, and is never an inde-
pendent body. Two other distinguishing characteristics are the forma-
tion of pollen tubes (from the pollen grains) and the production of seeds.
In the gymnosperms, represented by the cycads, ginkgo, conifers, and
ephedra, the female element is the megasporangium (ovule) and the
male element is the microsporangium (pollen grain). Each type is borne
on the surface of a megasporophyll and a microsporophy/l respectively.
In the gymnosperm megasporangium, the outer tissues are those of the
sporophyte and consist of the integument and nuce/lus. At the tip is a
minute opening, the micropyle. Within the nucellus is the female game-
tophyte. At the upper end of this gametophyte, and directly below the
micropyle, two or more archegonia are produced. Each archegonium
contains one egg-cell which, after fertilization, develops into the embryo
within the new seed. Because of their minuteness, none of these charac-
ters is of everyday use in identification of the gymnosperms. They are
reviewed here to point out the fundamental differences between gymno-
sperms and angiosperms and are of vital importance when considering
phyletic relationships of families and orders cf gymnosperms.
In ordinary taxonomic studies, the gymnosperms may be distinguished
from the angiosperms by the absence of flowers, by the ovules borne
naked on sporophylls, and by the seed not enclosed within a fruiting
structure of ovarian origin. There are also differences in vegetative
anatomy that collectively separate one taxon from the other, notably,
in the gymnosperms, the absence of vessels in the secondary xylem and
in the presence of resin canals.
The gymnosperms are composed of at least four, and probably six
orders: Cycadales, Ginkgoales, Coniferales, and Gnetales (the com-
ponents of the latter are so divergently different as to be considered by
more recent authors to represent three orders-Ephedrales, Gnetales,
and Welwitschiales). Each order of gymnosperm is so different from an-
other as to have almost separate terminologies for its distinguishing
characters.
The Cycadales contain the one family Cycadaceae, of which only one
genus (Zamia) is native to the United States. All members of the order
have pinnately or bipinnately compound leaves that are persistent (the
plants are evergreen), remaining on the plant for three to ten years.
The sexes of reproductive organs are grouped separately (the species
Pia'" S,,.UtIUre~ 33
are dioecious, an individual plant being either male or female). The male
organs are borne on a strobilus (pI. strobili), a cone-like structure (Fig.
9 c) consisting of a central axis about which are many closely packed
spirally arranged fertile sporophylls termed microsporophylls (Fig. 9 d).
Each microsporophyll bears an abundance of microsporangia or pollen-
sacs on its lower (dorsal) surface (Fig. 9 e). Polten grains produced
from these microsporangia are carried by air currents. After a pollen
grain comes in contact with the ovule it germinates and produces the
motile sperm by which fertilization is effected.
In Cycas the megasporophylls or ovulate scales are several in number,
pinnately lobed, and are arranged in a whorl in the center of the crown
of leaves, at the top of the stem. Each bears four to eight erect naked
ovules that are about the size of duck's eggs (Figs. 9 a, b). In other
genera the ovulate-scales are much more numerous and are spirally ar-
ranged on an axis (as are the microsporophylls) and form an ovulate
strobilus called the female cone.
The Ginkgoales contain a single family (Ginkgoaceae), which is com-
posed of a single genus (Ginkgo). The one species (G. bi/oba) is a tree
unlike any other gymnosperm. Its leaves are deciduous, falling in au-
tumn, and are fan-shaped with dichotomous venation (Fig. 9 f). The
species is dioecious (sexes on separate plants, one plant male, the other
female). It belongs to the gymnosperms because it produces naked
ovules (Fig. 9 f), and its seed is not endosed by any ovarian structure
(Fig. 9 h). The drupelike seed consists of a fleshy outer integument, a
hard bony inner integument, and an embryo (Fig. 9 h). As in the cycads,
the Ginkgo egg is fertilized by a motile sperm.
The Coniferales are probably the best-known North American repre-
senlatives of Ihe gymnosperm>. They are Ihe cone-bearing e~etgreens
and are represented by Ihe pines, firs, cedars, cypresses, and araucarias
plus those not producing cones as the podocarps and taxads (the last two
placed in a separate order, Taxales, by some authors). The conifers
differ from the cycads and ginkgo in the leaves needle- or scalelike (at
least in North American natives), in the pollen not producing motile
sperm, and in most genera being monoecious (the sexes separate but on
the same plant). In most members the scales of the ovulate strubilus
(each usually subtended by a bract) become woody at time of seed
maturity (leathery in Junipers). The male strobilus is much smaller
than the female and is herbaceous in texture. In the printitive families of
the order (as in Taxaceae), the male strobilus consists of an axis bearing
 Fig. 9. GYMNOSPERMAE: a, Cycas, ovulate sporophyll; b, same, bearing seeds;
c, staminate strobilus; d, fertile scale of staminate strobilus, lower surface; e, same,
enlarged; f, branch of Ginkgo bearing one drupe-like seed; g, Ginkgo, ovulate
branch; h. Ginkgo seed, vertical section; i. Taxus. fruiting branch; i. Taxus seed
and arit (ariI in vertical section); k. Taxus. ovulate branch; I, same, vertical sec-
tion; m, Taxus, staminate strobilus; n, Taxus, peltate microsporophyll; o. Pinus
branch with one- and two-year-old cones; p. Pinus, staminate strobilus; .q. Pinus,
microsporophyll, lower (abaxial) surface; r, same, three..quarter view of adaxial
surface; s, Pinus, ovulate strobilus; t, Pinus, ovulate scale, abaxial surfacc; u,
same:'""end view.

34
Plant Structure. 35
three to fourteen peltate fertile microsporopbylls subtended by several
fiattened sterile bracts (Figs. 9 m, n), and the ovulate "strobilus" of
an axis tenninated by a single naked ovule and subtended by sterile
scales (Figs. 9 j, k). The seed is solitary but enveloped by a red or pur-
ple fieshy aril (Figs. 9 i, I). In the more advanced Pinaceae, the bract
subtending each ovulate scale is more or less completely fused to the
scale and at maturity the strobilus is a woody cone (Fig. 9 0). In the
genus Pinus most of the leaves are horne on very short shoots and in
clusters (fascicles) of usually two to five, depending on the species (Fig.
9 0). The microsporophylls of the male strobilus (Fig. 9 p) contain two
pollen-sacs on the lower side (Figs. 9 q, r). The seeds are often winged.
In the Cuppressaceae the leaves and cone scales are often whorled or
opposite, and are decussate in arrangement (each successive pair at
right angles to the preceding pair), with the leaves often small and scale-
like.

ANGIOSPERMS
The angiospenns, like the gymnospenns are seed plants, but differ
from the gymnospenns in the production of flowers, in the ericlosure of
the ovule within an ovary (and the seeds enclosed within a fruit), and
in vegetative aspects of stem anatomy where the wood, in contrast to
that of gymnosperms, usually contains vessels and lacks resin ducts. Un-
like the gymnosperms, no archegonia are produced within the female
gametophyte which, in the angiosperm, consists of an embryo sac con-
taining a megaspore nucleus. This nucleus divides three times to form
eight nuclei, one of which becomes the egg cell. Following fertilization, '
by one, sperm nucleus from the genninated poUen grain, an embryo is
developed.
The angiosperms are composed of two subclasses, the Monocotyle-,
doneae and the Dicotyledoneae. The basic reproductive structures o(,i'
each of these taxa are similar. In the treallnent that follows, the vegeta-., '
tive structures are treated separately from the reproductive. Vegetativ~ "
structures of stems and leaves are used extensively in plant identification", '
and in the characterization of many taxa. They may be classified as'
stems, leaves, and systems of inflorescences.

STEMS
Stems provide many characters and these may be related to (1) axis
and habit, (2) direction of growth, and (3) types of modified stems.
IltJ INTRODUCTION TO PLA.NT T,jXONOMY

The axis of a plant is its main stem, as the trunk of a tree or stalk of an
herb. The point of origin on that axis of a leaf or bud is a node. The span
between two adjoining nodes is an internode. The angle formed by the
axis and leaf is the axil.
Any plant producing a conspicuous stem above the ground is said to
be cau[escent, whereas one in which the stem is inconspicuous or seem-
ingly absent (as in dandelion or plantain) is termed acauiescent. Many
acaulescent plants bear flowers singly or in terminal heads on long leaf-
less stalks known as scapes and are called scapose (Fig. 26 a). Plants
whose stems are very short, but much branched and usually covered
with leaves, often form cushion-like tufts or mounds ",nd are cespitose.
Those havirig several stems, or branches, growing close together and up-
ward to give :\ columnar effect are described as strict and in the extreme
fOfill are called fastigiate. Stems may bend abruptly at the nodes (as in
some grasses) and be termed geniculate, or may fork successively in
equal pairs (as in club-mosses) and be termed dichotomous.
In some groups of plants, stems are given special terms, such as culm
for the stems of grasses, boles for the unbranched stems of forest trees,
and caudex for the stems of woody monocotyledonous plants, such as
palms and some aroids.
DIRECTION of stem growth provides a character of aid in field identifi-
cation. Many of the terms used are common to the ordinary vocabulary
but here may have more precise meaning. A stem is erect when growing
straight upwards, a flower is erect when opening skyw~rds. Ascending
means arched upward and approaching erect (Fig. 10 a); a decumbent
habit is when the stem lies Hattened immediately above but not on the
ground, the branch tips usually ascending, such as in some prostrate
junipers (Fig. lOb); procumbent differs from decumbent in that the
stem lies on the ground, but does not root at the nodes, e.g., garden weed
Purslane (Fig. 10 c); stoloniferous applies when the plant sends out leaf-
less runners whose tips root and these produce new plants, e.g., in straw-
berry (Fig. 10 d) (the term is applied to underground stolons function-
ing similarly) ; repent is the term for the usual creeping condition, where
a stem grows along the surface, rooting at the nodes, such as in Checker-
berry or chickweed (Fig. 10 e); soboUlerous is a growth condition pro-
duced when plants (usually trees or shrubs) spread and form clumps by
underground rhizomes, e.g., sumac or sassafras (Fig. 10 f).
Plants whose stems are vinelike are called Uanes (pronounced lie-
a )1-n5). and .stems of this type also may be termed scandent. Those that
Plan' ~trudure. 37
grow over obstructions, and are without tendrils or other me.ns of self-
support, are said to be clambering, while others that grow upwards to
whatever height the available support or growth limit permits are said to
be climbing. Climbing lianes may derive their support by the stems
twining about an axis, such as wisteria; by means of modified structures

~~~-~
D\_.
d

Fig. 10. Stem habit types: a, ascending; b. branches decumbent; c, procumbent;

d, stoloniferous-; e, repent; f, soboliferous.

such as tendrils, e.g., in the grape or pea; by twisting leaf-stalks, e.g., in

clematis; by cirrhous leaf tips, such as are present in the Gloriosa-Iily; or
by production of aerial rootlets, such as in the Poison Ivy or English Ivy.
DURATION AND TEXTURE of stems (and plants) provide such terms as
annual, meaning living for one year; biennial (for stems and plants) liv-
ing for two seasons, the stem for the first year often acaulescent and
rosette-bearing the first season and caulescent the second; perennial liv-
ing several to many seasons (used mostly for plants rather than stems).
Plants living two to many years and dying after flowering and fruiting
the first time are termed monocarpic, such as many bamboos, some
palms. Herbaceous stems die to the ground after blossoming or at the
end of the growing season; woody stems live for a successive number of
years; suDrutescent stems are woody at the base, where they ;urvive
from year to year and with herbaceous distal portions (such a plant is
suDruticose). The arborescent habit is that of trees, or of plants that are
treelike" m appearance, growth, "or size.
TYPES OF MODIFIED STEMS, other than described above, are mostly
subterranean. The more common of these mclude rhizome (sometimes
38 INTRODUCTION TO PL,4NT TAXONOMY

termed rootstock, but a rhizome is properly a stem and not a root), a

creeping stem growing beneath the surface, consisting of a series of
nodes and internodes with roots often produced from the nodes, and
producing buds in the leafaxils (Figs. 11 a, f); tuber, a stem (usually
but not necessarily underground) much enlarged and modified as a food
storage organ with very minute scale-like leaves and buds or "eyes," such
as in the Irish potato (Fig. 11 j); corm, a very short thick firm fleshy

Fig. 11. Underground stem types: a. stoloniferous rhizome; b, corm, with

membranous tuni,c; c, same, vertical section; d, corm ~ith fibrous tunic;, e, same,
vertical section; f, rhizome; e, bulb, tunicated; h, sam~~ vertical section; i, bulb,
scaly; j. tuber. ' .

subterranean stem, usually broader than high, producing stems from the
base, and leaves and flowering stems from the top, such as in the jack-in-
the pulpit, crocus, and gladiolus (Figs. 11 b, c, d, e); cormel, the minia-
ture corm produced annually and vegetatively in leafaxils at the top and
upper side of a mature corm; bulb, a modified underground stem that is
very short, usually flattened, and crowned by a main body composed of
usually fleshy, more or less imbricated, non-green, scale-like leaves;
bulbs may be covered with a coat or tunic which is thin and membranous
as in onion or tulip (Figs. 11 g, h), fibrous and reticulated, or the scales
may be naked, e.g., in Lilium (Fig. 11 i); bulbel, the miniature bulb
asexually produced about the base of scales of a mature bulb; bulbil. the
miniature bulb produced in leafaxils of some bulbous plants or in leaf
sinuses; bulblet. diminutive for bulb, irrespective of point of origin on
the parent plant.
A cladophyl/ is a modified stem resembling a leaf in form and ap-
Plant S'r~cture. 39
pearance, but arising in the axil of a minute, bractlike, often caducous
true leaf, as in asparagus, or Ruscus. The so-called leaves of some cacti,
as Opuntia and the Christmas-cactus, are true stems flattened and
superficially leaf-like but are not cladophylls since they do not resemble
leaves in form or venation.

BUDS
Buds are much-condensed, undeveloped shoots. Each is a stem with
exceedingly short internodes, the leaves and floral parts enveloping one
another, and often a bud is covered with leaves modified to scalelike
bracts. The buds of woody plants are more conspicuous than those of
herbaceous plants and hence are more frequently used for identification
purposes. Most buds are mixed. That is, they are nascent ("embryonic")
branches bearing both leaves and flowers, but in some plants, such as
pear or apple, some buds are flower-producing and others produce only
vegetative shoots.
Buds may be in one of several positions-lateral, or axillary, on the
side of the stem (Fig. 12 a); terminal, at the stem apex (Fig. 12 g), or

Fig. 12. Bud types and bud-scale arrangements: a, axillary bud with imbricate
scales; b. naked bud; c, buds (three) superposed; d, stalked bud with valvate scales;
e, subpetioiar bud; f, pseudo-terminal bud; g, terminal bud. (b.s., bundle scar;
a·p., growing point abscission scar; 1.s., leaf scar; p, pseudo-terminal bud; s.s.,
.tipule scar.)

p.reudo-termirtlll (Fig. 120. A pseudo-terminal bud, such as in Cas-

tanea, the chestnut, is a lateral bud distinguished from a terminal bud by
the presence of the abscission scar of a twig that failed to mature (Le.,
40 INTRODUCTION TO PI,ANT TAXONOMY

one that never developed beyond the primordial bud stage) on Oll(O side
close to the apex and that of the leaf scar on the opposing side of the
conspicuous pseudo-terminal bud; the last being a lateral axillary bud
that developed so close to the stem apex as to appear to be terminal.
Buds not covered by scales are termed naked (Fig. 12 b). Those
covered by scales are termed scaly. There are two arrangements of bud
scales-imbricate, with usually several scales whose edges overlap (Fig,
12 a), and valvate, with one or few scales whose edges meet but do not
overlap (Figs. 12 c, d). Buds may be stalked, e.g., in witch-hazel (Fig.
12 d), or more commonly are sessile. In some genera, such as the hick-
ories, lateral buds may be superposed (Fig. 12 c), and in others, such as
Jycamore, the buds are in/rapetiolar ~nd are enveloped by the petiole
base of the "sublending" leaf (Fig. 12 e).
LEAVES
Leaves generally provide more characters for identification purposes
than all other vegetative parts combined. The terms applied to them are
used also to describe parts of the flowers, and sometimes of fruits, The
descriptive value of leaves is found in characters of (1) structure, (2)
venation, (3) form or general outline, (4) the apex, (5) the base, (6)
margin, (7) position and arrangement, (8) vesliture, (9) surface, (10)
texture.
STRUCTURE. In its simplest form a leaf consists of a blade and its leaf-
stalk or petiole, When there is no petiole the leaf is sessile. There is often
a small earlike appendage at each side of the base of a petiole called a
stipule. A leaf with one blade is generally a simple leaf (Figs. 13 b, c).
A leaf of two or more apparent blades is a compound leaj and each ap-
parent blade is a leaflet (Figs. 13 d, e, h, i, j), The stalk of a leaflet,
when present, is a petiolule (pronounced petty-o-lool), and the tiny
earlike appendage sometimes present on each side of the petiolule is a
stip/e.
A compound leaf has a petiole and two or more leaflets. When these
leatlets all originate from an apparent common point at the end of the
peliole the leaf is palmately compound (Fig, 13 h). The leatlets may be
sessile, as in clover (Fig. 13 i) or each may be borne on a petiolul.
(Fig. 13 h). When the leaflets are borne on a rachis, an apparent con·
tinuation of the petiole, the leaf is pinnately compound. If such a leaf
is terminated by a leatlet it is described as odd-pinnate (Fig. 13 d), and
when not so it is even-pinnate (Fig. 13 e). A leaf that is twice pinnate
Plant Structure. 41
(also termed bipinnate or two-pinnate) is decompollnd (Fig. 13 f) and
if the decompounding is always in threes, the leaf is said to be temately
decompound (~uch a leaf is also palmately decompound).

Fig. 1~. Leaf parts ar..d types: a, leaf sessile, parallel venation (bl, blade; sh,
sheath; st, stem); b, leaf alternate and petioled, pinnate venation; c, leaves oppo-
site and petioled, palmate venation; d, leaf compound, odd-pinnate; e, leaf com-
pound, even-pinnate; f, leaf decompound (2-pinnate); g, leaf decompound, tern ate;
h, leaf compound, palmate; i, leaf trifoliolate and palmate; j, leaf trifoliolate and
pinnate.

A leaf of three leaflets is trifoliolate (pronounced try-fo-Iie-o-Iate), a

term not to be confused with trifoliate, for tIie latter means three-leaved
(descriptive of the situation in Trillium). A trifoliolate leaf may be piJl-
nately or palmately compound, depending in part on its origin, for most
trifoliolate leaves represent reductions from multi-foliol.te ancestral
types. If an odd-pinnate five-leaflet leaf has lost the two lowest leaflets
il becomes trifoJiolate and the terminal leaflet will be stalked. This
petiolule of the terminal leaflet (which is composed in part of the rachis
top) is longer than those of the lateral leaflets and serves to identify the
leaf as pinnately compound (Fig. 13 j). When the terminal leaflet of ..
trifoliolate leaf is sessile, the leaf is assumed to be palmately compound
(Fig. 13i).
VENATION. Leaf venation is a pattern in the leaf blade representing
the major vascular strands. The so-called mid-rib of a leaf is the main
strand extending from tbe petiole to the leaf apex. In much of the litera-
ture the veins are called nerves and, wbile one term is as correct as the
other, these vascular strands are in no sense homologues of veins or
 42 INTRODUCTION 10 PLANT T.-lXONOMY

nerves of animals. There are three basic venation types: parallel, pal-
mate; and pinnate. Combinations of these types also occur.
Parallel venation is that in which the veins are more or less parallel to
the leaf margins. There mayor may not be a definite mid-vein. Two
forms of parallel venation may be noted: the simple parallel form in
which all veins extend from base to apex, such as in grasses and irises
(Fig. 13 a); and the penni-parallel form in which the veins are parallel
but arise from a mid-vein such as in banana, calla-lily, or pickerel-weed.
Palmate-venation is a type of reticulate or netted arrangement in
which three or more primary veins diverge from the point of petiole at-
tachment (Fig. 13 c); the pattern formed is often similar to toes of a
bird and is sometimes designated digitate-venation or pedate venation.
Pinnate-venation is the most common reticulate or netted type, some-
times termed penni-veined, and is distinguished by the presence of one
.entral mid-vein and many secondary veins arranged along the mid-
'vein, like the plume of a feather on each side of the shaft.
FORM. The terms given to leaf form or leaf outline, as described in
botanical and more general accounts, reflect man's effort to give preci-
~on where exactness and conformance to definition may not exist. For
centuries botanists have assigned geometrical terms such as oblong, ellip-
tic, or rhombic to forms of leaves. Some leaf forms are equivalents of
these terms, others approach equivalency and more represent intermedi-
ates between them. However, no more convenient system of leaf-form
classification has been devised and botanists continue to use these terms,
arid hyphenated combinations of them, to express situations. These
, terms are used also to describe forms of floral parts.
1m' Subulate. Awl-shaped, as a juvenile leaf of Juniper, tapering from
>!!f.ise to apex and usually sharpcpointed (Fig. 14 a). :
<1(' • Acicular. Needle-shaped, as a pine leaf, very slender, usually round-
P.'&h in cross-section and not flattened (Fig. 14 b). , .
. ,I, Filiform. Threadlike, often flexuous; very slender and cylindrical, as
the divisions or segments of a leaf of fennel'(Fig. 14 cr ".>,.
Linear. Long and narrow, flattened, the sides parallel' or nearly so,
as blades of many grasses (Fig. 14 e) . .
Lorate. Strap-shaped, flattened and flexuous, as in many Arnarylli-
daceae, the tip obtuse or bluntly acute but not tapering to a pOmt (Fig.
14 f)_
Lanceolate. Shaped as the head of a lance, broadened althe base ""d
tapering toward the apex, as some Eucalyptus or willows (Fig. 14 g).
Ovate. Egg-shaped, broadest below the middle, usually but not nec-
Plan' Structure.

essarily rounded at each end (Fig. 14 h). Some authors differentiate oval
from ovate, defining the former as "broadly elliptical with the breadth
considerably more than half the length" and with the broadest point at
or about the middle. More often, oval and ovate are used interchange-
ably.

Fig. 14. Leaf form or outline: a, subulate; b, acicular; c, filiform; d, filiform

segments; e, linear; f, lorate; g, lanceolate; h, ovate; i, elliptic; j, oblong; k,
oblanceolate; I, spatulate; m, runcinate; n, pandurate; 0, oboy-ate; p, deltoid; q,
rhombate (rhomboid); r, reniwform; s, orbicular. ('

Elliptical. Similar to ovate but the broadest point midway between

the ends and the width about one-half the length (Fig. 14 i). The term
may be qualined, when necessary, as narrowlY'-elliptical or broadly-
elliptical, depending On the ratio of width to length.
Oblong. The sides parallel or nearly so, and the length two to three
times the breadth (Fig. 14 j).
Oblanceolate. The reverse of lanceolate, with the broadest half above
the middle and tapering somewhat toward the base, the apex acute or
obtuse or otherwise (Fig. 14 k).
Spatulate. Similar to oblanceolate but tapering to a very narrow base
and the apex usually obtuse (Fig. 14 I).
Cuneiform. Wedge-shaped, widest at or near the apex (which is
more or less flattened) and tapering to a narrow base.
Runcinate. Generally oblaneeolate or spatulate and the sides very
coarsely cut or saw-toothed with the teeth pointing toward the base,
such as in dandelion (Fig. 14 m).
Obovate. The reverse of ovate, with the broadest half above the mid-
dle and the narrower end toward the base (Fig. 140).
 INTRODUCTION TO PLANT TAXONOMY
Pandurate. Fiddle-shaped, a modification of obovate with concave
to varying degrees, such as in some Ficus (Fig. 14 n).
Delloid. More or less equilateraUy triangular, such as the leaves· of
some poplars.
Rhomboidal. About as long as wide, broadest at the middle and
tapering to base and apex, shaped like a rhomboid (Fig. 14 (I).
Reniform. Kidney-shaped, broad as long or broader, the ape.•
broadly obtuse and the base somewhat cordate (Fig. 14 r).
Orbicular. Circular or nearly so, e.g., as in Nelumbo or the garden
nasturtium (Fig. 14 s).
THE APEX (pI. apices) of an organ is its terminal end, that which is
farthest from the point of attachment. Thirteen variations of apex form
are distinguished below and the terms are applicable to any appropriate
organ of the plant.
Cirrho.e (cirro.e). Filiform and coiling as a tendril as in the leaf-
tip of Gloriosa-Iily. It is also used for the tendril-like petioles of some
Clematis (Fig. 15 a).

FIg. I!. Leaf apices: a, cirrhosc; h, aristate; c, caudate; d, acuminate; e, acute;

f, cuspidate; g, mucronate; h, mucronulate; i. apiculate; j, obtuse; k, retuse; I,
emarginate; m, obcordate.

Aristate. Terminated by "slender often bristlelike appendage, usually

the continuation of the mid-vein, as in oats (Fig. 15 b).
Caudate. Tailed or bearing a tail-like appendage, as in some aroids
(Fig. 15 c). .
Acuminate. Sharp-pointed, tapering gradually or abruptly with the
. sides of the apex somewhat concave (Fig. 15 d).
Plan' Structure. 45
Acute. Sharp-pointed, tapering to a point, the sides straight or some-,
what convex (Fig, 15e),
Cuapidate. Somewhat abruptly and sharply concave and constricted
into an elongated sharp-pointed tip, this tip usually rigid (Fig. 150. '
Mucronate. Abruptly tipped with a shoot projection of the mid-vei",
(a mucro), accompanied Or not by a small amount of leaf tissue (Fig.
15 g).
Apiculate. Tipped with a small sharp point not representing an ob-
vious extension of the mid-vein or mucro, but harsh or sharp to toucl!
(Fig. 15 i).
Obt ...e. With a rounded apex (Figs. 15 j, 17 h).
Retu.e. With a rounded apex very slightly notched at the terminus of
the mid-vein (Fig. 15 k).
Emarginate. A condition more extreme than retuse but less so than
obcordate, the apex markedly notched but not lobed (Fig. 15 I).
Obcordate. With the apex two-lobed, inversely heart-shaped, such
as the leallet of Oxalis or of most clovers (Fig. 15 m).
THE SASE of an organ is usually the end of attachment. Twelve varia,
tions of base type< are distinguished below and while all apply to leaf
bases, many are equally applicable to other organs, '
Attenuate. With the lower sides constricting concavely and graduall1
into a somewhat winglike petiole, the base "drawn out" (Fig. 16 a). .'
Cuneate. With a Darrow to broad wedgelike taper, acute, the sides
straight (Figs, 16 b, 17 a, h).

PIa. 16. Leaf bases: a, attenuate; b, cuneate; c, oblique; d, obtuse; e, truncate;

f, cordate; I, auriculate; h. aagittate; i, bastate: j, peltate; k, perfoliatej ~ connate.
perfoliate.
46 INTRODUCTION TO PLANT TAXONOMY

Oblique. With the lowermost sides markedly unequal, such as in

Ulmus or Celtis (Figs. 16 c; 17 O.
Obtuse. Rounded, constripling abruptly to the petiole (Figs. 16 d,
17 e).
Truncate" With a nearly!straight line across the bottom, as if cut
across (Fig. 16 e). . s:
Cordate. With two lobes giving a heart-shaped appearance, such ali
in most leaves of common lilac (Fig. 16f).
A.uriculate. With a small earlike lobe (auricle) on either side of the
petiole and the two auricles separated by a narrow sinus (Fig. 16 g);
sometimes applied to stipules when enlarged and earlike, such as in sorne
Salix.
Sagillate. With a pair of basal lobes (or ears) turned downwards and
inwards, their apices acute or obtuse, arrow-shaped (Fig. 16 h).
HaSlate. With a pair of basal lobes flaring outward (Fig. 16 i).
Peltate. With the petiole attached at or near the center of the lower
surface of a usually orbicular blade, such as in Nelumbo or garden nl\S-<,.
turtium (Fig. 16 j); a falsely pellate situation may occur, such as iil
Nymphaea or Nuphar, where a cut or sinus extends from one edge of aml]
orbicular or elliptical blade to the center where the petiole is attached.
Per/oliate. With a base that extends around the stem (the leaf bi:ing
sessile), such as in Uvularia (Fig. 16 k).
Connate.per/oliate. With two opposite sessile leaves having their
bases fused (connate), such as in terminal leaves of spme Lonicera
species (Fig. 161).
THE MARGIN of leaves and other foliar organs'varies widely. Many
situations will be found to be intermediate between the conditions ac-
counted hy the terms given below and a compound hyphenated term is
then used. Some of the terms given below are encountered more com-
monly in margins of petals or bracts than of leaves (as fringed or
laciniate) .
Entire. Uncut, without indentation from the margin (Fig. 17 a); an
entire margin may however be variously wavy in a vertical plane (as un-
dulate) or may be provided with a row of hairs (ciliate).
Undulale. With an edge wavy in a vertical plane (Fig. 17 b); the
margin may be weakly or strongly undulate.
Sinuate. With the margin sthmgly wavy by turning inwards and out-
wards but too shallow to be lobed.
 47,
Crenale. Withpl!l~,Qr rounde(Lbroad teeth that may be direcled,
forward or at right angles to the mid-vein, scalloped (Fig, 17 c). f, ,
$errale. With,~!mrP.: Illther c~se .sawlike teeth that point fOIWan!
(Fig. l,",d). ,
Serrulate. A margin representing the diminutive of serrate, finely
serrate (Fig. 17 e).

Fig~ i7. Leaf margins: 8. entire; b, undulate; c, crenate; d. serrate; e, serrulate; 1

f, double-serrate; g, dentate; h, denticulate; i, ciliate.

DQ~ble-.errale_ With the teeth of serrate margins them'selves ser-

rulate or bearing teeth (Fig. 17 f). ' ,
Deniale. With sharp rather coarse teeth that point outwards from
the mid-vein (Fi~.,V g). , . : .... • .',.. <, '" ,"
Denticulate. A" IIlargin
._
represellting_th'e
, ",-" J,
diminlldve
. ,
:of dentate, finely
,-
dentate (Fig, 17 h). . . , • . ,: . .'.
Ciliate. With a row of fine hairs; sometimes so minute as sCltr~elY
to be discernible by the naked eYe (Fig. 17 i). .,,',
r';cised. When the margin is cut jaggedly into very deep teeth, ('Pi,.
18 a ) . ' , ", , 0 0 '0 ,

Lacerate. With the margin irregularly cut about one-half to tWQ:.

thirds the distancetbthe mid-vein; the IIliugins of segmen\s so(orrnell.
maybe entire or oiIierwis~ (Fig. }8 b). ' '"'' . ' " ',. ". ",',
Laciniate. With the blade Cut into narrow more or less 'ribbon-like
segments (Fig, 18c). ',.0 .' . '.'0.' .' . '

Lobed. With sinlises (inCision$) not more tlUui hilIfway f~m margin
:_ -, ,; "t, . '

d Q]
48 INTRODUCTION TO Pl.ANT TAXONOMY

to mid-vein and usually the lobes and sinuses more or less obtuse (Fig.
18 d).
Clefl. With sinuses extending somewhat more than halfway from
margin to mid-vein and usually the segment and sinus sharp or acute
(Fig. 18 e).

FIg. 18. Leaf margins: a, incised; b, lacerate; c, laciniate; d, lobed; e, cleft;

f, parted; g. pinnatifid; h. palmatifid; i, crispate.

Parted. With sinuses nearly but not quite reaching the mid-vein or
base of the blade, with the terms two-parted, three-parted, etc., express-
ing the number of segments produced (Figs. 18 f, a five-parted leaf,
18 g, h).
Pinnalifid. With the margin pinnately cleft or parted (Figs. 18 g, j).
Palmatifid. With the margin palmately cleft or parted (Fig. 18 h).
Cri.pate. With the margin curled in a vertical plane in minute waves,
as in parsley (Fig. 18 i).
POSITION AND ARRANGEMENT of leaves are sources of fundamental
characters, often more constant than most others. Special terms descrip-
tive of the situation include:
A.lternale. With one leaf at a node (Fig. 19 a); for critical ap~isal
of this and kindred terms as applicable to deciduous woody species, it
is best to examine the young shoot or twig that is the current season's
growth.
Di.,icho .... An alternate arrangement with adjoining leaves on op-
posite sides of the twig, the third leaf directly above the first in a ~
phyllotaxy (Fig. 19 b).
P1ant Structures 49
<"
Opposite. With two leaves at a node, one on the opposite side of the
twig from the other (Figs. 19 c, h); in some plants, one leaf of a pair
may be slightly below or above the other, a condition called subopposite.

Fig. 19. Leaf arrangements: a, a\ternate~ b, alternate, distichous; c, opposit~

d, opposite, decussate; e, whOJ;led; f. fascicled; g, imbricated; h, cauline; i. rosulate
(basal); j, equitant; k, cross~section through equitanl arra.uq:ement.

DecuSlale. A special arrangement of opposite leaves, with the leaves

of one pair emerging and disposed at right angles to the pair above and
below (Figs, 19 d, g).
Whorled. With three or more leaves at a node (Fig. 19 e).
Fasciculate. With leaves in clusters, often subtended by a bracteate
sheath, such as in Pinus (Fig. 19 h) where the leaves of each fascicle
are borne on a short, very slow-growing shoot and actually are alternate
in a high phyllotaxy and with exceedingly short internodes.
Imbricale. With the leaves overlapping, shingle-like, such as in some
Selaginella (Fig. 19 g).
Cauline. Borne on the stem, as opposed to basal or rosulate (Fig.
19 h).
Rosulale. Borne in a basal rosette, usually but not always on or
close to ihe ground, generally alternate in a high phyllotaxy (Fig. 19 i).
Equilanl. Of upward growth, the sides at right angles to the ground
and each leaf basally folded together lengthwise and enveloping the next
younger leaf, such as in iris (Fig. 19 j with sectional view in 19 k).
so INTRODUCTION TO PLANT TAXONOMY

VESTITUR};
Vestiture (or vesture) implies covering. In botanical writings it refers
to the condition of hairiness, scaF:--ess, giandularity, or other covering
that may be on the surface of any external part of a plant or organ.
When hairiness in general is to be distinguished from other types of
vestiture, the term indumentum (meaning garment or covering) may be
used. Terms of vestiture are commonly applied to stems, leaves, floral
components, and fruits. The conditions represented intergrade imper-
ceptibly from one to another and the terms employed cannot be defined
within sharp limits, but are used in a relative sense. The distinction be-
tween some of these terms is so difficult that botanists of long experi-
ence may not diagnose a particular state of vestiture with the same terms,
and botanists have been known to use two or more terms for the same
situation when faced with identifying it on separate occasions. Because
of their relativeness, terms of vestiture and especially of hairiness are
difficult to illustrate. In an effort to explain these terms as adequately as
possible, they are here described, many are illustrated, and the more
common ones then arranged in the form of an analytical dichotomous
key.
Terms applied to hairs (trichomes) and hairy coverings:
Glabrou•• Said of surfaces devoid of vestiture, not properly a term
of vestiture but the antonym of those terms.
Glal,rate. Essentially glabrous but found, especially by examination
with a lens, to be provided with some vestiture, usually very sparsely
or minutely so.
Pubescent. A general term lacking any uniformly accepted definition
and meaning hairiness of any type, as oppo'ed to glabrous.
Glandular-pube.cent. A general term indicating presence of hairs
and glands (usually trichome-like glands) intermixed on a given surface.
PuberuloUl (puberulent). Covered with exceedingly short, fine,
rather dense straight hairs at right angles to the surface, scarcely per-
ceptible to the naked eye except when viewed by transmitted light
(Fig. 20 a).
Jlelulinoru. Velvety, pilelike, covered with erect straight dense hairs,
much longer than puberulous (Fig. 20 b).
Tomentole. Covered with more or less matted woolly hairs, curled.
and 'appressed to the surfac,\ (Fig. 20 c) .
Tomentulo.... Diminutive of tomentose.
Plnnl S,rudure. 51
Floccose. Bearing more or less scattered tufts of appressed woolly
or long and soft hairs that rub off easily, such as in some species of
be\onia.
Arachnose. Bearing a cobweb-like indumentum of long entangled
hairs.
Woolly (Ianate). Covered with dense, long, soft, entangled, curled
hairs, not appressed close to surface (Fig. 20 d).

; ~ ,r,
,
~ )
J"I ~ ~
-' -, l
Fig. 20. Vestiture types (surface view at left. sectional view at right): a, pu~
berulous; b, velutinous; c, tomentose; d, woolly; e. villous; f, pilose (a and c with
enlarged inserts of sectional view, aU others drawn at same scale).

Cri.p-hairy. Bearing long kinky hairs, obtus< ')/ zigzagged.

Villous. Bearing moderately dense, long, soft, often curly hairs, more
or less erect but not necessarily straight (Fig, 20 e),
Pilose. Clothed with soft, very long, rather straight hairs, not dense
but somewhat shaggy (Fig. 20 f).
Sericeou•. Silky, with straight, soft, long hairs, appressed (Fig. 21 f),
often producing a satin-like sheen, especially when dense.
Strigose. With harsh, straight, stiff, short hairs, mostly appressed or
weakly ascending, bristle-like in harshness as in some species of Ulmus
(Fig. 21 b).
Himae. Covered with short erect stiff (but not harsh) hairs (Fig.
21 d).
Hirtellous. Minutely hirsute.
A..perou•• Rough to the touch, especially when rubbed "the wrong
way," a general tenn without reference to trichome character.
 52 INTRODUCTION TO PLANT TAXONOMl'
Hispid. Bristly, bearing dense, erect, straight, harshly stiff (but some-
what flexuous) hairs (Fig. 21 c).
Hispidulous. Diminutive of hispid.
Setose (setaceous). Similar to hispid, but not so dense.
Setulose. Minutely setose and probably not different from sparsely
or weakly hispidulose.
Scabrous. Covered with scattered harsh hairs, not erect, usually with
dilated (or bulbous) bases (Fig. 21 a).

,. " ...- -..-",..._,;

Fig. 21. Vestiture types (surface view at left, sectional view at right); a, scabrous;
b. strigose; c. hispid; d. hirsute; e. stellate; f, sericeous.

Scabrescent. Minutely scabrous.

Ciliate. Fringed with a row of hairs (Fig. 17 i).
Plumo.e. Provided with feather-like compound hairs.
Stellate. Bearing forked or multiple-branched hairs, each on a short
central stalk or sessile (Fig. 21 e).
SquamelaeJorm (squamose). Scaly, provided with fiattened usu-
ally plate-like sessile hairs (sometimes fringed), or short-stalked and
peltate.
Lepidote. Covered with scurfy scales, usually easily removed.
Barbed. Having hairs whose tips are bent back acutely or whose
sides provided with teeth or minute retrorse barbs (Fig. 22 h).
Uncinate. Having hairs with obtusely hooked tips (Fig. 22 i).
Clavellate. Club-shaped hairs or glands, broadest at apex (Fig. 22 f).
Echinate. Beset with stiff sharp long spines, as the burr of a chestnut.
Moniliform. A multicellular hair or gland with constrictions be-
tween the component cells (Fig. 22 g).
Plant. Structure. 53

Spin~:. Bearing one or more modified stems, leaves or stipules re-

duced to spines (hard, sharp, needle-like) (Fig. 22 b).
Spinescent. Covere~ith spines, usually minute ones.
Prickly. Covered with prickles, spine like excrescences of bark, as in
roses (Fig. 22 a).
A.culeate. Furnished with prickles.

Fig. 21. Armature and special hair types: a, prickles; b, stipular spines; c,
areote of spines; d, arcole of one spine and many glochids (single glochid shown
enlarged at right); e, gland~tipped hairs; f, clavate hairs (or glands); g, stamen
bearing moniliform hairs, single hair at right; h, an achene with retrorsely barbed
awns, single awn at right; i, uncinate-tipped achenes.

Key to terms of common types of indumentum (adapted from manu-

script of Prof. K. M. Wiegand, Cornell Univ.) :
[. Hairs compound
2. Forked, or branched radiatingly .... ,.,., .•... ,.,.,.,., ....... stellate
2. Pinnate or feather-like ................ ' ..................... plumose
I. Hairs simple
3. Surface with some or all hairs knob-tipped (pin-headed)
4. Trichomcs all knob·tipped .............. , ............. ,. glandular
4. Trichomes mixed, some with knob-tips and others
being simple taper~tipped hairs ................. . glafldular-pubescent
3. Surface with none of the hairs tipped by glandular knob,
5. Hairs curved or bent, at least at tips, not straight
6. Tips hooked or bent
7. The tip hooked over obtusely ... ' .... ,' ...... , ....... uncinate
7. The tip, or teeth at tip, bent back acutely .. , ........... barbed
6. Tips straight
8. Each hair kinky. obtusely zigza.gged ,., ... , .. ' ..•.. . crisp-hairy
8. Each hair curved or bent, not kinky
54 INTRODUCTION TO I'L1NT TAXONOMY
9. Indumentum close and dense, but not apfJtCssed. . . ... woolly
9. Indumentum close and tightly appre~"cd to surface
10. Hairs distributed evenlY, not rubbing off easily .. . /ontentose
10. Hairs scattered in bunches, rubing off easily .. "" floccose
S. Hairs straight
11. Each hair bent at base and pointing toward base of
organ bearing same .................... retrorse
11. Each hair not bent basally, or if so pointing forward
12. Hairs appressed against surface
13. The hairs very short, in bunches, rubbing oft' ..... . floccose
13. The haiTs not in bunches nor rubbing off easily;
usually more than 1 mm. long
14. Each hair very short, harsh, stiff..... . ... . strigose
14. Each hair long, the indumentum usually dense
and soft, silky-appearing ...................... sericeous
12. Hairs erect or nearly so
15. The hairs standing out from margin in a row, like
eyelashes ... , .....,.................. . ... , .. , , . ciliate
15. The hairs borne over the surface of the organ
16. Hairs soft to touch, at least not harsh
17. Trichomes 0.5 mm. long, very minute
and rather dense ..... , ................ . puberulent
17. Trichomes mostly 2 mm. long or more,
dense or scattered
18. Hairs ntedium stiff ...................... hirsute
18. Hairs soft, not stiff
19. Trichomes very long ................. . pilose
19. Trichomes moderately long, very soft
20. Hairs very dense, pile!ike .. relutinous
20. Hairs not dense nor pilelike ..... villous
16. Hairs harsh and very stiff
21. The hairs minute, less than 0.5 mm. long ... . hispidulous
21. The hairs 1 mm. long or longer
22. Trichomes quite dense ...•............... . hispid
22. TJ'ichomes scattered ..................... . selose

SURFACE
Surfaces of major plant parts, as stems, leaves, flowers and fruits,
whether devoid of vestiture or not, provide many characters for identifi-
cation purpose.s. Some of the more common terms associated with sur-
face types are:
Glabrous. Devoid of all vestiture.
Reticulate. Bearing a net-like pattern represented by weak grooves
or color variegation outlining the veinlets beneath; or slight wrinkling,
as of a seed-coat, to produce a weakly netted effect.
Rugose. With the reticulation more deeply grooved over the net-
work of veinlets, as in Rosa rugosa (Fig. 23 e).
Pla"t Strudu.... 55
Bullaw. A surface appearing as if much puckered or blistered, as
in Savoy cabbage (Fig. 23 d).
Papillate. Bearing minute nipple-like projections (Fig. 23 a).
M uricate. Roughened by minute firm epidermal proliferations
scarcely classifiable as hairs, squamellose Or other types of vestiture
(Fig. 23 b).

-d)"'" ,

~ ~"!~.
~.

Fig. 23. Surface types (each with highly magnified insert): a, papillatej b, muci-
cate; c. rugose; d, bullate.

Foveolate. Pitted with shallow distinct depressions.

Alveolale. Honey-combed or resembling such by its geometrical uni-
formity.
Punctate. Covered by minute impressions, scarcely possessing depth,
and appeanng as if made by a pinpoint, such as seed coats of Anagallis
or of Geranium pratense.
Glandular-punclale. The epidermis provided with minute glands,
translucent when pale or\amber-colored (as viewed by transmitted
light) or blackish, such as in leaves of Hypericum.
Striate (lilleate). Marked (often by color differences) by longi-
tudinal lines.
Sulcale. Furrowed with longitudinal grooves, such as stems of many
members of Umbelliferae.
Annulate. Ringed by circumferential grooves or ridges.
Areolate. Divided into a number of irregular, squarish, or angular
spaces.
F ene,trale. Perforated by obtuse window-like openings.
56 INTRODUCTION TO PLANT TAXONOMY

Yi.cid. Covered with a sticky exudate.

Mucous. Covered with a slimy exudate.
Roridulate. Covered with transparent parenchymatous tissue as ele-
vations or over fenestrations.
Pruinose. F rasted, having an opaque dewy appearance.
Glaucous. Covered with very fine bloom of wax, as on grapes.
Pulrerulent. Covered with a very fine powder, representing a waxy
exudate.
Farino,e. As in pulverulent, but the powder granules coarser.
Glaulcent (cae,iow). Weakly glaucous.
TEXTURE
In addition to such common conditions of texture as herbaceous and
woody, there are a score or more of other types, of which tne common
includes: hyaline, thin and almost wholly transparent; membranous,
thin and semi-transparent; chartaceous or papery, opaque but thin;
coriaceous, leathery and thickish; cartilaginous. hard, tough, and often
thin; scarious, thin and dry, appearing as if shrivelled, such as the
phyllaries (involucral bracts) of many species of Centaurea; suberous,
corky; other terms of texture of equivalent meaning as in English in-
clude: spongy, horny, bony, fleshy, succulent, waxy, fibrous, and pithy.

DURATION
In addition to such terms of duration as annual, biennial, and peren-
nial which concern longevity of plants as a whole, there are others used
for various parts of a plant, including: diurnal, opening during the day;
nocturnal, opening at night; ephemeral, open for only a day and then
perishing; deciduous, falling off after maturity (as leaves that fall in
autumn); caducous, falling off early, as sepals of Papaver; marceseent,
withering or fading but not falling off; fugacious, perishing rapidly and
falling: monocarpie, bearing fruit but once and dying after fruiting; and
poiyearpic, bearing fruit many times; protantherous, when the leaves
appear before the flowers; synantherous, when flowers and leaves appear
at the same time; and hysteranthous when leaves appear after the flowers.

THE INFLORESCENCE
An inflorescence is now generally accepted to be the arrangement of
flowers on the floral axis and is a branch system, but by early botanists
it was considered to be the mode of flowering. Present systems of in-
Plan' "Crudures 57
fiorescence classification are descriptive and have little bearing on
phyletic origins (for discussion of latter, cf. Lawrence, Taxonomy of
Vascular Plants, 1951, pp. 59-64). In much taxonomic literature in-
florescences are classified as determinate, the terminal or central flower
developing first and thereby arresting growth by elongation of the pri-
mary axis, or indeterminate, with the terminal or central flower open-
ing last and the inflorescence often elongating as the flowers develop
from lower to upper portions of the primary axis. It is now recognized
that the division into these two types is of little significance, since for
example the umbel of an onion (Allium) is determinate whereas the um-
bel of the carrot is indeterminate. The sequence of inflorescence types
given below is descriptive and arbitrary and makes no pretense to reflect
or suggest phyletic relationships.
When the flowers appear singly in the axils of ordinary foliage leaves
they are said to be axillary and solitary, and no conventional inflores-
cence type is present. However, when the foliage leaves are much re-
duced in size (or are of different form or character) they are designated
bracts and the mode of flowering is then an inflorescence of one of the
types given below. If the bracts are very small, as compared to the size
of the flower, they may be termed bracteoles.
The stalk supporting a single flower is a pedicel. If the flower has no
pedicel it is termed sessile. A common leafless axis bearing several
flowers (sessile or pedicellate) is sometimes called a peduncle and that
te.rm may be used also for single flowered "inflorescences" believed to
represent a phyletic reduction from a multiflowered ancestral condition
(such as in uniflowered species of Narcissus). The peduncle in this last
type is better known as a scape.
A dichasium, in its simplest form, is a determinate cluster of three
flowers arising from a common pedunde by a dichotomous branching
immediately beneath a terminal flower (Fig. 24 q); more common is
the compound dichasium (Fig. 24 a) such as found in Gypsophila or
some Anemone spp. (Fig. 25 b).
A cyme is a determinate inflorescence representing a reduction from
a compound dichasium by loss of secondary axes to produce a more or
less flat-topped or convex cluster (Fig. 24 m), such as in Comus alterni-
folia or Viburnum spp. A modification of the cyme, and represented by
a loss of flowers from the same side of each dichotomy (forking) is the
helicoid cyme (often incorrectly referred to as a scorpioid cyme) and
common among many members of the Boraginaceae, such as Heliotrope,
 i/\ t
I f

g
____y

Fig. 24. Schematic diagrams of hypothetical evolution of inflorescence types:

a, compound dichasium; b, helicoid cyme; c, cincinnus; d, scorpioid cyme; e,
thyrse; f. panicle; g. compound corymb; h. simple corymb; i, raceme; j, indeter~
minate umbel; k, spike; 1, indeterminate hea.d; m, cyme; 0, determinate umbel;
0, determinate head; p, verticillate inflorescence; q. simple dichasium.
58
Plant Structure. 59
Symphytum or Baraga (Figs. 24 b. 25 c). The cincinnus (sometimes
termed glomerule) is a modified helicoid cyme in which the peduncles
and pedicels are shortened or absent. such as is to be found within the
head of Armeria (Fig. 24 c). The scorpioid cyme (often confused with
helicoid cyme) is a nearly straight but determinate inflorescence and
resembles a raceme. An example of this is the lily-of-the-valley (Fig.
24 d). In some plants the flowers of the cyme are much congested and
said to be fascicled (in a bundle) as in Sweet William. In the Labiate
fhe so-called verticillate inflorescence is a primary indeterminate axis
(sometimes much condensed and capitulate. e.g.• in Monarda) bearing
pairs of cymes or cymules (Figs: 24 p. 25 h. and 60 a).

Fig. 15. Inflorescence types (somewhat schematic): a, cOTymb~ b, cymes (di-

chasia) (Anemone); c, helicoid cyme or cincinnus; d, umbel (Hoya); e, capitulum
(Trifolium); f, capitulum, vertical section, to show conical torus (Rudbeckia);
gl inflorescence of verticillate cymes (Salvia); h, "verticel" (Salvia).

An umbel is a descriptive term applied to inflorescences whose flowers

are pedicelled and the pedicels seemingly arising from a COmmon point
at the stem or peduncle apex; often subtended by a number of bracts
forming an involucre. The umbel may be determinate (Fig. 24 n) or
indeterminate (Figs. 24 j. 25 d). It may be simple or compound. as in
most Umbelliferae (Fig. 56 a). and when compound each peduncle.
which is termed a primary ray bears an umbel/et Or miniature umbel
(Fig. 62 b) whose lIower pedicels are termed secondary rays and whose
bractlets form an involucel.
A head Or capitulum is an inIIorescence of sessile or nearly sessile
60 INTRODUCTION TO PLANT TAXONOMY
flowers on a very short or flattened axis (when flattened as in the daisy
it is termed the receptacle) producing a globose to flat-topped cluster
(see Figs. 69 a, b). It may be derived from an umbel (by loss of lower
pedicels) or from a spike (by compression of the axis and loss of in-
ternodes between the flowers (Figs. 24 j, n). The head may be naked
as in Cephalanthus or clover (Fig. 25 e) or subtended by an involucre
of bracts (phy//aries) as in the sunflower or aster.
A thyrse is a branch system of compound dicbasia, arising from a
primary axis of indeterminate growth; this type somewhat r~embles
and is often confused (as in lilac) with a panicle (Figs. 24 e, 26 f).
A panicle is an indeterminate branch system whose primary axis
bears branched secondary axes and pedicellate flowers (Figs. 24 f, 26 d),
such as in many grasses.
A raceme is an indeterminate single axis bearing pedicellate flowers
(Figs. 24 i, 26 c). Many so-called spikes are properly termed racemes,
e.g., in Gladiolus.

Fig. 26. Inflorescence t)-pes: a, scapose (flower solitary); b, spike; c, raceme;

. d, panicle; e, dichasium; f, thyrse.

A spike is an indeterminate single axis bearing sessile flowers, such

as in Plantago (Figs. 24 k, 26 b). Many so-called spikes are not this
inflorescence but only superficially resemble it and should then be termed
spicate or spike-like; as, for example, the catkin or ament, a flexuous
spike-like assemblage of bracteate dichasia, in the willow, oak or birch;
or the spadix of members of the Araceae (usually subtended or envel-
oped by ? large leafy bract or spathe).
Plant Structure. 61
A corymb is an indeterminate Jlat- or convex-topped inflorescence, a
modified panicle (Figs. 24 g and h), such as in some goldenrods.
When the flower, Or head of flowers, is solitary and borne on a scape
(naked pedicel or peduncle) the inflorescence is termed scapose (Fig.
26 a), e.g., in Cyclamen, dandelion, and most tulips.

THE FLOWER
A flower is homologous with a leafy stem in that it consists typically
of an axis (the pedicel and receptacle) and four whorls of leaf-like com-
ponents, arranged from the bottom upwards and known as the calyx,
corolla, androecium (stamens), and gynoecium (pistils or carpels).
These floral components are homologues of leaves in that they have a
position and origin comparable to leaves. This does not mean necessarily
that they are modified foliage leaves or that they have been derived
from them-but they and the leaves may have had a common ancestor.
Each of these four sets of floral components is often referred to as a
floral whorl, or either of the outer two may be designated as a floral
envelope, even though the actual arrangement may be in a spiral or the
units reduced or fused into one. Any understanding of the descriptive
terms applied to flowers is predicated on a familiarity of the components
typical of most flowers.
FLOWER TYPES AND ARRANGEMENT. A complete flower is one com-
posed of all four whorls. An incomplete flower lacks anyone of them. A
per/ect flower is bisexual (hermaphroditic) and contains both androe-
cium and gynoecium (Fig. 32 c), but need not possess calyx or corolla.
An imperject flower lacks the elements of one sex or the other. A neutral
flower possesses neither an androecium nor gynoecium, and a nude
flower has one or both sexes but lacks both calyx and corolla. A flower
whose reproductive parts consist only of an androecium (one or more
stamens) is a staminate flower (Fig. 27 a), and one in which they con-
sist only of a gynoecium is a pistillate flower (Fig. 27 b). When a plant
bears flowers of only one sex (i.e., the flowers are either staminate or
pistillate) it is called dioecious (di- from the Greek for two, oecious
meaning households, in allusion to two plants required to house both
sexes). An example of a dioecious species is the willow. When a plant
bears unisexual flowers, but with flowers of both sexes on the same
plant (as in maize Or in many cucurbits), it is called monoedous (one
plant to house both sexes) (see Fig. 26 d).
A flower that is radially symmetrical in form is said to be actina-
62 INTRODUCTION TO PLA.NT TAXONOMY
morphic (Fig. 27 e). If it is bilaterally symmetrical, i.e., divisible longi-
tudinally through the center into two equal halves, as are orchid flowers,
it is termed zygomorphic (Fig. 27 f); but if it is not possible to divide
it into two equal halves, as is true of the flower of the canna, it is called
irregular (Fig. 27 g). In some works, the term zygomorphic is used to

Ilg. 27. Flower types and arrangement: a, flower staminate (periaoth uni-
seriate); b, flower pistillate (perianth uniseriate}j c~ ftower bisexual (hermaphro-
ditic) (perianth biseriate aDd corolla polypetalous); d, flowers unisexual (plant
'f
monoecious); e, ftower actinomorp)"Olc or reaular; f, flower zygcmorphic; flower
irregular; b, corolla gamopetalous.

include also irregular. Frequently authors apply one or the other of

these terms to flowers, when only perianth or corolla is intended.
PERIANTH is a collective term for the calyx and corolla. When the
components of calyx and corolla are similar in size, form, and coloration
they are termed tepais, e.g., in Liliur. or Tulipa. Terms used to describe
parts of the perianth are those used to describe leaves.
Plant Structure. 63
Calyx is a descriptive term for the outer whorl of usually green leaf-
or bract-like structures called sepals. They may be distinct 1 and the
calyx then said to be polysepalous (aposepalous), or connate 2 and the
calyx then gamosepalous (synsepalous), or in some families be reduced
to modified hairs or scales and termed a pappus. In some genera the
sepals are highly colored and the calyx resembles a corolla (such as in
Mirabilis, Delphinium, Helleborus).
The corolla is tl _ >'cond or inner envelope of the perianth and is
composed of petals. ,\ r rolla of distinct petals is termed polypetalous
(apopetalous), while one in which the petals are connate in any degree
(as at the base only or throughout their length) is termed gamopetalous
(synpetalous). Usually the corolla is colored other than green and con-
tributes to the showiness of the flower.
COROLLA TYPES. The terms actinomorphic, zygomorphic. and irreg·
ular, explained above under flower types, are especially pertinent for
corolla classification. Polypetalous corollas are commonly actinomor-
phic and do not vary so much in form as do the gamopetalous corollas.
However, among the former there are a few types, as:
Cruciform corollas, such as are found in the members of the mustard
family, generally are actinomorphic (zygomorphic in lb ...is. the cdndy·
tuft) and consist of four petals disposed in an X or cross-form (Fig.
55 a).
Papilionaceous corollas (the sweet-pea type) are found in the Legume
family and are zygomorphic (Figs. 29 e, f, 57 a). There are five petals
(Fig. 57 b): a large standard or banner which is the most posterior
petal, two lateral wing petals, and two keel petals that are usually con-
nate along the lower edge; this limited extent of connation in no wny
causes the corolla to be termed gamopetalous. since the petals rail
separately and the corolla does not fall as a single piece. A modification
i& the cesalpinaceous corolla in which the standard is in a position an-
terior to the wings.
There are several types of gamopetalous corollas, whose general out-
line is also applied to types of polypetalous corollas. The actinolllorphic
1 Distinct is used in botanical expressions to indicate cexnplete separation of like parts
(as one petal distinct from another), whereas free indicates freedom from union of
adjoining, but unlike. parts (as the stamens frec from the petals).
2 Connate means the fusion of like parts (as petal to petal); coherent means the
meeting in close proximity by cohesion (by viscidity, vestiture, etc.) or otherwise of
like parts but lacking fusion of their tissues (as pollen grains sometimes coherent);
adnau refers to the fusion of unlike parts (as stamens to petals); adhesion refers to
the coming into close contact of unlike parts but lackina: fusion of their tissues (a..
the indehiscent husk of luglan~ adherent to the nul).
 INTRODUCTION TO PLANT TAXONOMY
types compose one grouping and the zygomorphic another. Those of the
former include:
Rotate, (wheel-shaped) with a very short tube and a broad limb at
right angles to it, such as in borage or forget-me-not (Fig. 28 e);

Fig. 28. CorolJa types: a, urceolate (Pieris); b, campanulate (Campanula);

c, funnel-form (infundibular) (Salpig/ossis); d. salverform (Quamoclil); e, rotate
(Brunnera).

Salver/arm, with a long corolla-type, and a shorter limb that is at

right angles to it, such as in tobacco (Fig. 28 d) ;
Funnel/arm (infundibular), with a tube gradually expanding upward
a. doe. a funnel, the limb may be lIaring or .omewhat at right angle. to
the lIower axis (the type intergrades with salverform) such as in petunia
or morning-glory (Fig. 28 c);
Campanulate (bell-shaped), with a broad tube about as long or longer
than broad and a lIarin~ limb or lobes, as in canterbury-bells (Fi~.
28 b);
Urceolate, with a large urn-shaped tube, usually somewhat constricted
above the middle, and small more or less flaring lobes, such as in Vac-
cinium or Monotropa (Fig. 28 a).
Tubular coral/a, with a more or less cylindrical tube composing the
conspicuous portion; the limb small by comparison, e.g., the disk
lIower of a daisy (Figs. 27 h, 69 d).
The zygomorphic type. indude, in addition to the papilionaceous
corolla described above, the following:
Bilabiale corolla, in which zygomorphy is due to a two-lipped ar-
rangement of the lobes (in many mints, two lobes form one lip. and
 6S
three lobes form the other, as shown in Fig. 66 f). The bilabiate type
may have the throat open, such as in salvia (Fig. 66 a), in penstemon
(Fig. 29 b) or in catalpa, or closed by a palate and then termed per-
sonate, such as in toad-flax (Fig. 29 d) or snapdragon.

FIe. '9. Corolla types: af Ugulate or ray type (Helianthus); b, tubular or

disc type (also bilabiate) (Penslemon) (see also Fig. 69d); c, biJabiate, the base
scniculate (ScutelJaria); d, bitabiate and penonate, the corolla-base spurred or
calcarate (Linaria); e, papilionaceous, side view (Lotus); t, same, petals spread
(I<, keel petal; w, wing petal; 51, standard or banner).

Ligulate (or ray) corolla characteristic of most of the aster family is

a three- .to five-lobed gamopetalous zygomorphic corolla that is split
along one side and having a large lIattened limb and a small tube (Figs.
29 a, 69 c).
Zygomorphy may be due also to a protrusion from the base of the
<:orolia in. the tQ=' Q{ a £!'w, eJ~,., in toad-flax o~ Lirwria (Fi¥, 29 d),
a large in.IIation on one side, termed a gibbosity (the corolla being gib-
bous) , in snapdragon; and when less so the swollen condition is termed
ventricose, such as on the ventral side.of Scutellaria (Fig. 29 c).
THE ANDROECIUM is the male reproductive component of the 1I0wer
and is composed of one or more stamens. The stamen is usually com-
posed of an anther (the pollen-producing element) and a filament (the
stalk), but the filament may be so reduced that only th<;..anther is ap-
parent. The stamens may be very numerous, as in Magnoliaeeae or
Nymphaea, or more commonly are few in number. In some 1I0wers there
is only a single stamen (e.g., in Euphorbia, Typho). The stamens may
be spirally arranged (e.g., Ranunculaceae) or more commonly in two
66 INTRODUCTION TO PLANT TAXONOMY

whorls. When in two whorls, those of the outer whorl are generally op-
posite the petals, a condition called diplostemony. When in one whorl,
a reduction is believed to have taken place, and usually only those of
the outer whorl remain. Sometimes the remnants of the inner whorl are
represented by staminodes or nectaries.

Fig. 30. Aestivation and corona types; a, valvate aestivation (Clematis panic-
ulala), habit of bud and diagram of perianth arrangements; b, imbricate aesti-
vation (Tulipa), habit of flower and diagram; c, convolute aestivation (Phlox),
habit of opening bud and diagram; d, plicate aestivaticn (Nicotiana), habit of
opening bud and diagram; e, corona from perianth, vertical section (Narcissus);
f, corona from stamen filaments, vertical section (Hymenocallis).

The anther is composed of two anther-sacs or thecae. Sometimes

these thecae are called anther-cells. The two sacs are separated by a
connective, which in reality is the continuation of the filament. Funda-
mentally each anther is of four thecae (representing four sporangia), but
the septum (partition) separating the two on one side of the connective
usually disappears during anther development and prior to dehiscence
(Fig. 31 Bb), although this is not always true (e.g., in Lauraceae). In
the flowers of some families a mature anther is com posed of a single
theca formed from two sporangia (e.g., Malvaeeae) and are sometimes
described as one-celled.
When the anther is terminal on the filament (or seemingly so), it is
said to be basifixed (Fig. 31 Da), and when the filament appears to be
affixed to the side of the anther it is dorsifixed (Fig. 31 Dc), or some-
times versatile (if free-moving on the filament, such as in grasses or lilies
[Fig. 31 Db]).
Plan' S,rudure. 67
Sometimes, as in Malvaceae, the stamens are connate by their fila-
ments and termed monadelphous (Fig. 32 d, e, f). In other families, .s
in many Legumes, they may be connate in two groups and are then
diadelphous (Fig. 57 cJ. When stamens are connate by lheir anthers
(e.g., in Compositae) they are termed syngenesious (Fig. 69 cJ. In
some flowers, as in those of the mint family, the stamens are four in

FiR. 31. Androecium and ,mIne-r types. A, Androecium types: Aa. Slamen~
spirally arranged; Ab. stamens in three whorls; Ac. Siamcns in two Whorls. those
of the outer whorl the shorter (ojiplos.ten1l1n(1l1s): Ad. stamens in a single whorl
(dehhccnce introrsc 1. n, Anther-cdl number: Ba, ceUs four at anthes:is~ Bb, same
cros~·scction; Bc. cells two at anthes.is; Bd. cell one at anthesis. C. Anther de-
hiscence types: Ca, longitudinaL Cb. transv!!rse: Ce. valvular flaps; Cd, apical
pores. D, anther position: Da. bas.ifixed: Do. dorsilixed and versalile; De, dorsi~
flxeJ, not vcr5atilc (Jchiscence extrorse).

number and in two pairs, on~ pair longer than the other, a condition
termed didynamolls (Fig. 32 a); in those of the mustard family the
stamens number six, with two shorter than the other four, a condition
termed lelradynamolls (Fig. 32 b).
Stamens provide other characters of taxonomic value. Ih the aster
family they are frequently appendaged. The appendages may be distal
at the anthor tips, usually by projection and expansion of the connective,
or basal and formed by tail-like projections from the anther sacs (Fig.
32 h). In the heath family the illament may be appendaged and be pro-
68 INTRODUCTION TO PLANT TAXONOMr
vided with glabrous or variously hairy sac-like swellings (Fig. 32 i). In
other taxa stamens may be provided with various types of vestiture,
such as moniliform hairs in C ommelina (Fig. 22 g).

Fla. :n. Androecilll types: I, stamens didynamous (Origanum); b, stamens

tetradynamous (Thlaspi); c, stamens synieDesious (Aster); d, stamens mono-
delpbous (Hibiscus); e, stamens syngeDesious (Sinningia); f, stamens mona-
delphous (D~smosium); I, stamens diadelphous (Lathyrus); h, stamens with apical
appendage (lnula); i. stamen with basal and ventricose appendage (Chimaphila).
I
Staminodes are sterile nonfunctional and often anth_tless stamens,
present in some flowers as accessory structures. They are not properly a
part of the andro_cium, since they are not a reproductive organ. These
are usually the vestigial remains of once-functional stamens. They may
be distinct and arise from the receptacle, or be adnate to the corolla.
Sometimes they are petaloid in form and color.
A nther dehiscence may be one of several types. When the thecae split
along the surface facing outward the dehiscence is termed extrorse (Fig.
31 Dc), when along the surface facing an adjoining anther or inward
toward the flower axis it is introrse (Fig. 31 Ad). In some genera the
dehiocence may be transverse (Fig. 31 Cb) or at right angles to the
vertical axis (e.g., in Hibiscus, Elatine); in others it may be by a pore
(e.g., in Polygalaceae, most Ericaceae), a dehiscence type that is termed
poricidal (Fig. 31 Cd). A modification, where the pores are covered
by flap-like valves, is found in the Lauraceae.
Pollen grains, produced from most stamens at anther maturity, result
from two successive divisions of a pollen mother cell within the anther.
Plant Structure. 69
The outer membranous coat of a pollen grain is the exine. The surface
of the exine may be smooth or variously sculptured (furrowed, papillose,
honeycombed or areolate, echinate, etc.). Most pollen grains bear three
furrows and are termed tricolpai}; others have a single furrow down one
side and are called monocolpate. Distinct pollen grains are usually
granular; those coherent or connate in clusters of four are in tetrads
(e.g., most Ericaceae); and when the pollen from each theca is aggluti-
nated into a single often hard and horny mass it is termed a pollinium
(pl., pollinia) and is a condition characteristic of most Asclepiadaceae
and Orchidaceae (Figs. 49 Bf, 64 d).
Plants pollinated almost exclusively by wind are called anemophilous,
and those pollinated by insects are termed entomophilous. Plants in
which the pollen is shed before the maturity of the stigma of the same
flower are termed protandrous (or proterandrous), and those whose
flowers produce mature stigmas before the pollen is shed from stamens
of the same flower are termed proterogynous.
GYNOEClUM means literally the female household, and in flowers the
term is applied to the female reproductive element. The basic unit of
the female element of a flower is the carpel (Figs. 33 a, b). y. is an
ovule-bearing structure believed to have been evolved from a non-green
leaf-like appendage (homologous with a leaf but not derived from a
foliage leaf), and as such was a megasporophyll. In ancient flowering
plants this megasporophyll is believed to have been palmately three-
veined, fiat, and to have borne ovules on or near its margins. This pre-
historic sporophyU is further believed to have folded lengthwise, with
the ovules inside and margins fused. The resultant structure is a unicar-
pellate, one-IDculed ovary (Figs. 35 a, 33 c). This ovary is the product
of a single carpel and is termed a simple ovary. The space within such
an ovary (or any ovary) is termed a locule or chamber (Fig. 35 e). In
many works it is called a cell, and in them a simple ovary'may be de-
scribed as one-celled. This usage of cell is far removed from the more
precise cytological meaning and the terms locule or chamber are pre-
ferred.
The upper end of an ovary is terminated by a stigma (Fig. 35 d).
Pollen grains germinate on the stigma reior to fertilization of the ovules.
The stigma may be directly at the ovary apex or, more commonly, it is
separated from the ovary by a narrow constricted "neck" termed the
style (Fig. 35 d). These three structures, ovary, style, and stigma com-
prise the pistil (Fig. 35). The gynoecium may be composed of many
 70 INTRODUCTION TO PL.' , r4XONOMY

distinct pistils (e.g., in the rose, Fig. 39 Ca, or the butte" p, Fib'
of a single unicarpellate pistil (e.g., in the cherry, Fig. 39 Ba), or, n.
commonly, of a single multicarpellate pistil (e.g., in the lily, apple, or
cactus) .
Placentation is the mode of arrangement of the ovules within an ovary.
The placenta in the ovary of a flower is not a distinct tissue, as is the
placenta in the uterus of a mammal, but is a zone or area occupied by

,,,
,,
,,
,,,
,
,,,
,
~,
1I ,-
... ~{/ ...
Q
,
~

Fig. 33. Hypothetical evolution of simple and compound ovary: a, three-

lobed carpel with submarginal ovules (dotted lines indicate v:;'ScuIar strands);
b. same, somewhat involute; c, simple ovary derived from "b" by infolding of
ovules and connation of ventral margin~; d, axis bearing three involute open
carpels; e, compound ovary derived from "d" by connation of edges of adjoining
carpels; f. axis with three open carpels with adjoining sides more or Jess parallcl;
g, compound ovary derived from "r' by connation of adjoining sides and mar-
gins~ h. cross-section of "g" (hypothetical); i. cross-section of "g" (actual) show-
ing loss of carpellary demarcation in the three septa. (Note: vascular strands
shown with xylem elements blackened.)
Plm" Structures 71
the ovules. For descriptive purposes, the intrusion of the ovary wall into
the lacule is often designated the placenta (Fig. 35 c, d), and while it
usually lacks morphologic;'l significance it may provide characters of
considerable taxonomic value. In the simple unicarpellate ovary de-
scribed above the ovules are in a double row along the inner abaxial 8

Fla. 34. Gynoecial types: a. pistils many and spiralled, the gynoccium apo-
CfU'POUS; b, pistils many and whorled (cyclic), the SYDoecium apocarpous; c, pistil
solitary, the gynoccium syncarpous; d, pistil one, ovary compound; e, same, ovary
vertical section; f, same, ovary cross--section; g, same, style apex and three-lobed
,tigma; h, pistil one, ovary simple; i, same, vertical section; j, same, ovary crosa-
section; k, same, style-tip and simple stigma.

wall of the ovary (Fig. 35 a). The placentation in a simple ovary is

termed marginal or ventral,' although sometimes the number of ovules
may be reduced to one whose position may be basal or pendulow from
the apex of the loeule.
In a majority of unipistillate (apocarpous) gynoecia, a compound
f7Vary is present (Fig. 35 b, d). That is, the ovary represents the fusion

• The terms abaxial and ventral are synonymous and mean "toward the axis" or more
specifically, toward or OD the side nearest the center of the axis. The antonym of eacb
is adaxial and dorsal. respectively. 'The ventral side of a leaf (or its abaxial side) iJ
that aide facina: the axis of the leaf bud before unfolding, the upper side. The axis
of an open dower is that imaginary zone that would be occupied by the center of th4J
pedicel if it were projected on through the flOWCl.
to In Taxonomy of Vascular Plants (p. 75) this type of placentation was termed
pamtal. However, ,cod phyletic reasons han been advanced for rcatricting ~ term
parietal to a placental situation of the compound ovary.
72 INTROnUCTION TO PLANT TAXONOMY
of two or more carpels. This may have been brought about by anyone
of several mechanisms and may be represented by different types of
placentation. Partially opened ovaries within the same flower standing
in close proximity to one another may have become fused. This fusion

Fig. 35, Placental and carpellary types: a, pistil with simple ovary and mar-
ginal placentation (unicarpellate); b, compound ovary with parietal placentation
(tricarpellate); c, compound ovary with axile placentation (bicarpellate); d, com-
pound ovary with parietal placentation, each placenta intruded and spreading
(bicarpellate) .

may have occurred along adjacent ovary margins (Fig. 33 d). It re-
sulted in a single compound ovary that has a single locule and as many
placentae as there are carpels (Fig. 33 e). Here the placentae (rows of
Plont Stru.ctu.re. 73
ovules) are on the inner sides of the ovary wall and the placentation is
termed parietal (Latin for walls). Usually the number of placentae indi-
cates the number of carpels comprising the pistil (Fig. 35 b). In some
situations there may be an intrusion of the ovary wall at the point of
-union of each adjoining carpel margin. In some extreme conditions the
edges within the locule may flare away from one another (Fig. 35 d).
These modifications are also considered to be parietal placentation.
When two or more simple pistils (eacn unicarpellate and unilocular)
become fused by their abaxial surfaces, a single compound pistil results.

Fig. 36. Placental and carpellary types: a, ovary with axile placentation (S-car-
pellate); h, ovary with basal placentation (carpel Dumber not determinable); c,
ovary with lameIJate placentation (9~arpellate); d, ovary with free-central-placen-
lation (5-carpcllag)
74 INTRODUCTION TO PLANT TAXONOMY
If two such simple pistils were involved the resultant compound ovary is
two-carpellate. two-Ioculed, and its placental shuation is termed axile
(Fig. 35 c). If five simple pistils were involved the resultant ovary is
quinquelocular (Fig. 36 a). The term axile owes its name to the ovules
seemingly in the axils of the partitions. The point of common union of
these carpels may produce a central axis, but this axis does not repre-
sent a continuation of the flower pedicel or receptacle into the ovary.
In the ovary of some kinds of flowers that once had axile placentation,
there has occurred a loss of the partitions (the fused walls of :wo adjoin-
ing carpels) leaving a central column bLaring as many double rows of
ovules as there were original carpels. Such an ovary has a single locule.
The placentation in this type is termed free-central (Fig. 36 d) and is
characteristic of flowers in the primula family and of most members of
the pink family (Fig. 53). When, by reduction, the central column and
most of its ovules are lost and the few remaining ovules appear to "sit"
on the bottom of the ovary locule, the placentation is termed basal (Fig.
36 b).
In a few families the compound ovary is composed of many carpels
and the partitions have the ovules scattered over their surfaces. This
situation occurs in Nymphaea, the common water~lily, and the placenta-
tion type is termed lamellate (Fig. 36 c). In the poppy there may be
many incomplete partitions converging from the inner surface of the
ovary wall toward the center of the loculc (but not meeting there).
These intrusions bear an abundance of ovules over their surfaces. This
place"tal type is classed as parietal since all placentae appear to be in-
trusions of the ovary wall.
Styles and stigmas provide important taxonomic characters. Ordinarily
their number is a guide to the carpellary number of the ovary below.
Styles may be simple (Fig. 37 b) or branched (Fig. 37 a, d), or some-
times petaloid (Figs. 37 h, i). In the grasses they are often plumose (Fig.
37 a), whereas in the asters they are simple and forked (Fig. 37 O. The
stigma may be obvious because of its enlargement, or it may scarcely be
differentiable from the style. When enlarged it may be bluntly lobed as
in lily (Fig. 46 d) or tomato (Fig. 35 c); linear and papillate as in many
mallows (Fig. 37 e), or markedly discoid as in hibiscus (Fig. 37 c). In
Iris and related genera the stigma appears as a narrow transverse linc
acrOss the lower side of the petaloid style-branch (Fig. 37 h). Often the
stigma is recognizable by the presence of a glossy viscid exudate present
when receptive to pollination.
Plant Structure. 75
The ovule is the egg-containing organ within the ovary (Fig. 35 a).
After fertilization it develops into a seed. Ovules vary in position and
attachment. These distinctions, while of considerable taxonomic value,
are difficult to recognize in the field, or in the laboratory, except with
a magnification of X 60 or higher. Typically the ovule is borne on a
stalk or funiculus (Fig. 38 Ab), "tho"gh it may be sessile. The point
where the funiculus meets the ovuh: b termed the hilum (the hilum is

Fig. 31. Style and stigma types: a, stigmas plumose (Gramineae); h, stigma
capitate (A Ichemilla); c, stigmas discoid (Hibiscus); d, style branches filiform
(Armerja); e, stigmas linear (Ki/aihelia); f. style bifurcate (A mica); g, stigma
branches radiate (Papaver) j h, style branches petaloid (Iris); i, stigma transverse
(Iris) (s.b., style branch; st, stigma).

also the scar left when a seed breaks from its "stalk" as in a bean). The
opening or "mouth" of the ovule is the micropyle (Fig. 38 Ad) or fora-
men. The body of the ovule is the nlleelllls and it is typicaIJy enveloped
by two coats or imeguments (Fig. 38 Ad).
Ovule positions of taxonomic importance include:
Orthotropous, a condition present when the ovule stands straight and
the micropyle is at the end away from the funiculus, as in buckwheat
(Fig. 38 Aa);
Campylotropous (incurved), when (he ovule is curved and the micro-
pyle nearly meets the funiculus, as in chickweed (Fig. 38 Ad);
Amphitropous (half-inverted) when the developing ovule retains an
essentially straight axis but turns 90' on its funiculus with the latter
fused to its exterior for that part of the tum, as in Mallow (Fig. 38 Ab);
and
 76 INTRODUCTION TO PL:4NT TA.XONOMY
Analropous (inverted), when an ovule has turned 180 0 during its
development and stands with the funiculus fused along on I side and the
micropyle close to the placental surface, as in most taxa (Fig. 38 Ac).
OVARY POSITION. The position of an ovary may be superior, inferior,
Or half-inferior. A superior ovary is situated above the zone of attacb-
ment of perianth and androecium (Figs. 39 A, B, C), and when t~

Fig. 38. Ovule and cotyledon types: Aa, ovule orthotropous; Ab. ovule aDl¥
phitropous; Ac, ovule anatropous; Ad, ovule campylotropous. B, Cotyledons
incumbent: Ba, embryo with cotyledons folded against radicle; Be, same, sepa-
rated; Bd, same, cross-section. C, Cotyledons condupIicate: Ca, cross-section Deaf
distal end of embryo; Cb, same, medium section. D, Cotyledons accumbent: Da.
embryo, habit; Db, same, cross-section (co chalaze; co, cotyledon; e, egg; f, funic~
UillS; Ii, inner integument; Ic.., outer integument; ro, micropyle; n, nuceIlus; r,
radicle).

stamens are adnate to the perianth, it is the area of attachment of the

latter that determines the ovary position (i.e., if the ovary is above that
point of attachment [superior] Or if it is below it). An interior ovary is
below the apparent area of attachment of perianth and androecium
(Figs. 39 f, g). An ovary whose position is mOfe or less intermediate be-
tween the two is termed halt-interior or subinferior (Fig. 39 D).
Two theories have been advanced to explain the derivation of the in-
ferior ovary from a superior ovary: the receptacular theory and the ap-
pendicular theory. Most older works and some current elementary works
explain the inferior position of the ovary solely by the receptacular
theory, but a preponderance of anatomical data have brought about a
Plan' Siructurea 17
rejection of the validity of this theory for most situations, and the ap-
pendicular theory is now the more widely accepted.
The appendicular theory of ovary position is predicated on the ac-
ceptance of the flower being. morphologically speaking. a determinate

FIg. 39. Ovary position. A, ovary superior, perianth and stamens hypogynous;
B. ovary superior, perianth segments and stamens perigynous; Ba, vertical section;
Rb, detail of Ba to show composition of hypanthium; hc, same, minus lines of
adnation of perianth and androecium; C, ovary superior, perianth and stamens.
perigynous~ hypanthium of "Rosa" type; Ca~ vertical section; Cb, detail to show
relation of receptacular cup to hypanthium and composition of latter; D, ovary
haIf·inferior, stamens perigynous; E. ovary inferior, perianth and stamens epi~
gyoous; Ea. vertical section of flowers; Eb, detail of same to show adnation of
hypanthiwn to ovary; F, ovary inferior, hypanthium present, perianth and stamens
epigynous; Fa, vertical section of flowers; Fb, detail to show componel)ts of
hypanthiumj Fe. section of hypanthium; G, ovary inferior, stamens exserted,
hypanthium present. (h, hypanthium; pe, petal; r, receptacle; se, sepal; 5t, stamen.)
78 INTRODUCTION TO PLANT TAXONOMY

stem with appendages tbat are homologous with leaves. By this view it
is held that some of those appendages were carpellary and contributed to
the pistil, that others bore microsporangia and developed into stamens,
and that still others below were accessory to the reproductive functions
and became the parts of the perianth. In some kinds of plants, as Prunus
(the plum), the ovary is surrounded by a cup-like structure on which the
sepals, petals, and stamens appear to be borne (Fig. 39 Ba). This struc-
ture is termed the hypanthium or lIoral-cup). Anatomical studies show
this hypanthium to have been formed by the adnation (fusion) of the
sepals, petals, and stamens. The unfused parts of the calyx, corolla, and
androecium which appear above the hypanthium are designated as
sepals, petals (not calyx-lobes, corolla-lobes), and stamens (Fig. 39
Bb). Inasmuch as the hypanthium is composed of these three elements,
it is incorrect to refer to it as a calyx-tube, for, strictly speaking, the
calyx-tube is the tubular portion of a gamosepalous calyx.
In most plants, the inferior ovary differs morphologically from the
superior ovary by the adnation to it of the lower portions of calyx,
corolla, and stamens (Figs. 39 Eb, Fb). It represents a stage of phyletic
advancement over the more primitive superior ovary. These adnate
sepals, petals, and stamens are appendicular (appendage) parts of the
flower and do not comprise the distal end of the receptacle or torus'
By the receptacular theory it was believed that the inferior ovary was
embedded in or surrounded by a tube or cup of receptacular tissue (as
if the ovary were pushed into the receptacle), and that the hypanthium
of the superior ovary likewise was a tube of receptacular tissue. Although
most older works, and some contemporary semi-popular books, present
the receptacular theory as the explanation for the inferior ovary, this
theory is nowhere accepted by scientists today as representing the situa-
tion for the large majority of plants with inferior ovaries.
THE FRUIT
A fruit may be defined as the product of a ripened ovary, pistil, or
gynoecium of a lIower and may be composed in part of accessory floral
or vegetative parts. It is the seed-bearing (or seed-containing organ of a
plant). Fruits are important in the classification and identification of seed
plants, because generally they provide characters very reliable in the de-
II This is the prevailing situation in flowers with an inferior ovary. There is anatomical
evidence that in some plants the basal portion of the hypanthium (or oC tissues adnate
to the ovary) is composed in part of receptacuiar tissue, e.g., in the hypanthium of
Rosa, and the inferior ovary of lOme members of the Santaltceae (set Fig. 39 Cb).
Plant Structures 79
limitation of genera and of families. Many kinds of fruits exist. Various
classifications have been proposed but all in common usage are based on
descriptive features of the fruit rather than on HJ'!ir comparative mor-
phology and anatomy. For this reason the classifications now in use. and
inherent in taxonomic writings. are artificial and do not reflect basic
morphological situations. The types of fruit currently recognized in most
of the literature are differentiated grossly in the following synopsis (re-
produced from Taxonomy of Vascular Plants):
Fruits si~ple, the product of a single pistiL
Fleshy and usually indehiscent.
Texture homogeneous, fleshy throughoul .... ,................ .Berry
Texture heterogeneous.
Fruit exterior a firm, hard, or leather rind.
Septate present, several to many ..................... Hesperidiltm
Septate absent , ......................................... Pepo
Fruit t:xterior soft.
Center of fruit with a single "stone" which contains
the seed .............................................. Drupe
Center of fruit with paper or cartilaginous carpels ............ Pome
Dry fruits.
Fruit indehiscent, usually one~ to two-seeded.
Winged ................................................ Sanlara
Wingless.
Pericarp thin.
The pericarp adnate to the seed--........................ A chene
The pericarp loose and free from seed .................. Utricle
Pericarp thick and hard, sometimes bony.
Fruit small, from a one~loculed ovary ........... : ...... Achene
Fruit usually large, from a two~ to more~loculed ovary ...... Nut
Fruit usually dehiscent, one~ to many~seeded.
Product of a unicarpelJate ovary.
Dehiscing by ventral suture only ........................ Follicle
Dehiscing by two longitudinal or transverse sutures.
Sutures longitudinal ................................ . Legume
Sutures transverse ................................... Loment
Product of a bi- or multicarpellate ovary.
Fruit splitting into one~seeded halves ................. . Schizocarp
Fruit splitting and releasing seeds.
Dehiscence circumscissle ............................... Pyxis
Dehiscence longitudinal .............................. Capsule
(Silicles and siliques are specialized types of capsules
characteristic of the Cruciferae.)
Fruits compound, the product of two or more pistils.
The product of several pistils of a single gynoecium connate or
coherent, and usually flesby ........................... Aggregate fruit
(An accessory fruit is a type of aggregate fruit in which the
conspicuous and often fleshy part of the fruit is of nonovarian
origin.)
The product of several gynoecia aa:gregated in ODe mass ..... . Mulliple fruit
80 lNTIWDUCTION TO PLANT TAXONOMY
From the above it may be noted that some fruit types n{ore or less
parallel family limits. The hesperidium is found only in the citrus family,
Rutaceae, but is one of several types produced in that family. The pepo
is characteristic to a large extent of members of the gourd family,
Cucurbitaceae. The pome (Fig. 41 f) is represented by the fruit of the
apple, pear, cotoneaster, etc., in the Rosaceae, and the legume (Fig.
40 j) and loment are common only to the legume family.

FIg. 40. Fruit types (dry): 8, follicle (Helleborus); h, achene (Potentilla);

c, achene (CompQsitae); d, pyxis (circumscissile); e, capsule (septicidal); f, cap-
sule (loculicidal); g, capsule (poricidal); h, nut; i, lament; j, legume; k, schiZo~
carp; 1, samara (Acer); m, samara (Pulea); n, samara (Fraxinus); (ca, calpo·
phorej me, mericarp).

Aggregate fruits Occur in several families but are well known by the
blackberry-raspberry alliance (Rubus) of the Rosaceae (Fig. 41 g), to
which family belongs also the strawberry, whose gross fruit is of the ac-
cessory type but in which the true fruit is an achene embedded in the en-
larged fleshy receptacle (Fig. 41 h, k, I).
Multipk fruits are those in which the gynoecia of severaillowers form
a single structure. These are common to the mulberry family, Moraceae.
f:I=, as in the fig, the conspicuous fleshy part develops from the re-
ceptacle. In the mulberry the gynoecia retain their individuality and re-
semble tiny drupelets aggregated into one mass.
The dehiscence of dry fruits refers to the mode of the splitting or open-
ing of a fruit for release of seeds and provides valuable diagnostic
characters. There are several types of dehiscence of dry fruit, of which
the follicle, legume, loment, capsule, nut, and schizocarp are examples.
A follicle is the product of a one-carvelled unilocular ovary (Fig. 40 a)
Plan' Structure. 81
and dehisces by a single suture (line). It may be many-seeded or one-
seeded. A legume is a product of a unicarpellate ovary, but differs from
a follicle in dehiscing by two lines or sutures, as in the common beau
(Fig. 40 j). A loment has a similar morphological origin but is strongly
contracted between the seeds, falling apart (when mature) at the con-
strictions into one-seeded segments or joints. A capsule is the product of
a multicarpellate ovary. A loculicidal capsule Is one in which the line of
dehiscence (the split) opens directly through the capsule wall into the
locule (Fig. 400. A septicidal capsule is one in which the line of dehis-
cence is from the capsule wall at the point where the septum (partition)
jam. it, and often the septum is'split longitudinally (Fig. 40e). A
poricidal capsule dehisces by pores. Each'pore may be provided by a lid
or operculum or not. The pores may be at the top of the capsule, as in
the poppy (Fig. 40 g) or at the base as in some species of Campauula.
A circumscissile capsule, known also as a pyxis, dehisces by a circum-
ferential line whereby the top comes off as a single piece, as in portu-
lacca (Fig. 40 d). A nut is a one-seeded fruit enclosed in an involucre 6
and its diminutive form is nutlet. It may be dehiscent or indehiscent. A
schizocarp is a dry fruit dehiscing into two equal one-seeded halves
(each a mericarp) and usually suspended or supported from a short
central axis (stylopodium) by a wiry carpophore. This type of fruit is
common to the carrot family and to the samarous fruits of maple. Dry
indehiscent fruits include the nut (discussed above) samara, utricle, and
achene. The samara is a one-seeded winged fruit. The wing may be on
one side as in the ash (Fig. 40 n) or may surround the seed and be disci-
form, as in the wafer-ash (Fig. 40 m). The utricle is a one-seeded usually
small, bladdery fruit as in Atriplex. An achene is a one-seeded indehis-
cent fruit in which the pericarp (outer layer) is firmly adoate to the seed.
Achenes are the fruit-type encountered in the aster family, and here
there are many modifications in form and appearance of tax6nomic
value. The achene may be attenuated into a beak which, in lettuce and
dandelion, is terminated by a whorl of plumes (Fig. 42 a); it may be
winged and have terminal awns in the form of bristles, spines (Fig.
42 c), or be terminated by a double pappus (Fig. 42 h, i). The achene
surface may be smooth or variously ribbed, glandular, echinate, or
papillate.
1;1 The involucte is an accessory l'art of the fruit and may be cup-like and represent
the fusion of abundant bracts as in the oak. or bur~like and of similar origin as in the
thestnut, or somewhat capsular as in the beech. The acorn of the oak is the nut plus the
cup-like involucre and the term. is Dot synonymous with Dut. or with the nut of a
particular kind of tree.
82 INTRODUCTION TO PLANT TAXONOMY
Fleshy fruits are more difficult to classify but in general are one of
basic types. The berry is any fleshy fruit containing one or more seeds as
the grape, date, and tomato. The drupe is a fleshy one-seeded fruit in

Fig. 41. Fruit types (fleshy); a, drupe (Prunus); h, same, vertical section;
C, pyrene or "pit" of drupe; d, berry (Vilis); e, same, vertical section; f, pome
(Malus), vertical section; g, aggregate fruit of drupelets (Rubus); h, same, ver-
tical section; i, pyrene from Rubus drupelet; j, accessory fruit (Fragaria); k,
same, -vertical section; I, achene from Fragaria fruit. (ca, calyx; end, endocarp
[bony wall of pyrene]; ex, exocarp; mes, mesocarp; P. pyrcne [the so-called "pit"
or seed]; r, receptacle; s, seed.)

which the seed is enveloped by a stony endocarp (formed from part· of

the ovary wall). Here, as in the plum, peach, or olive, the ellocarp is the
"skin," the mesocarp is the pulpy "flesh," and the endocarp, the bony
structure enveloping the seed; the bony endocarp plus the seed com-
prise the "stone" or "pit." Sometimes the "pit" of a drupe ot more com-
monly of a drupelet is termed a pyrene, as in the raspberry (Fig. 41 h, i).
Seeds are fertilized mature ovules. Each seed is composed of one or
two seed coats, the endosperm (sometimes lacking) and the embryo.
The embryo is made up of a plumule, one to two cotyledons (rarely
more), a hypocotyle, and radicle. The external features, aside from the
seed coat, are: the hilum, a scar that marks the place of funiculus (stalk)
attachment; the raphe, a ridge formed by the funiculus (and produced
only by anatropous ovules); and micropyle, a minute scar (not always
present) indicative of the opening into the ovule. Taxonomically, these
features are not used extensively, except for the surface characters of
the seed coat and (in the mustard family) the arrangement of the cotyle-
Plant Structure. 85
dons and radicle. In some taxa the location and form of the embryo pro-
vide important characters for delineation of a genera and family; in
others the presence (or absence) and character of endosperm are of
equal diagnostic value. However, while these are characters of phyletic
significance, they rarely are suited for field use or general identification
purposes.
The shape, surface, and "appendages" of seeds often are of value as
field characters. Seeds may be plumose, as in milkweed (Fig. 42 q), or
winged, as in the trumpet-vine (Fig. 42 n) or the surfaces variously

Fig. 41. Achene and seed types: a, achene, beaked (TmgopogorJ); b, achene,
plumose pappus; c, achene, winged and awned; d, achcne, capillary pappus; e.
same, detail of capillary trichome; f. achene with double pappus, of scales and
plumose trichomes; g, same, detail of plumose trichome; h. same, detail of outer
pappus scales (trichomes excised); i, achene with pappus of scales; j, seed with
reticulate coat \te~ta)~ k., ~eed with mur\ca\e iUt\'ace', \, tT~quetTon'!. '!.ffi()oth ~eed
(Fagopyrum); m, same, cross-section, embryo curved; n, seed, winged; 0, seed
with rows of tubercles; p. seed with alveolate or honeycombed surface, pitted;
q, seed winged and with coma.

sculptured or papillate (Figs. 42 j, k, 0, p). The number of seeds within

a locule or fruit often provides reliable distinguishing characters, as may
the size, color, and form.
The position of the cotyledons and radicle within the seed may be
of taxonomic value, especially in the crucifer family (see Fig. 49). When
the two cotyledons have their edges against the radicle they are said to
be accumbent; when one side of the cotyledon rests on the radicle, they
are said to be incumbent; and when the two cotyledons fold over the
radicle, they are said to be conduplicate.
CHAPTER v. Collecting and Identifying
Techniques

APART of the training of every taxonomist and professional botanist

includes the preparation of herbarium specimens-the pressing, drying,
and mounting of plants. This chapter is designed to meet the needs, not
of the professional, but of the amateur botanist, the naturalist, and the
person whose curiosity has been whetted to the point of wishing to pre-
pare an "unknown" for personal use or to send to an authority for
identification and naming. Very often one sees flowers in woods Or fields
and wishes to preserve material for future identification. It is good to
know and practice the right ways of doing this.
For the beginner, it is more simple to sit with some freshly collected
flowers and "work out" their identity with a flora or manual than to
work over flattened and dried specimens later. Frequently it is easier to
study and understand the structures and floral organs from fresh flowers
than from pressed ones. With experience, the latter present no diffi-
culties. Aside from the requisite flora or manual-<lesirably one with
keys-it is only necessary to obtain a good hand lens (preferably one
with a Hastings triplet lens, or at least a Coddington type lens, of X 7
or X 10 magnifying power), a pair of forceps, and a sharp scalpel or
razor-blade, plus, of course, flowers to dissect.
The inexperienced person should select, as a "starter," a plant having
large flowers, and avoid those such as of the aster family or others
equally minute which are more complex than is apparent. It is folly to
attempt to identify an unknown in a hurry. Study the plant and especially
the flower before attempting to identify it with aid of a book. Observe
the nature of the perianth and be able to name its parts. Count the parts.
84
Collecting and Identifying Technique. 85
Note whether sepals and petals are fused or distinct. Note the number of
stamens, where they are attached, and the positioning of the anthers. In-
vestigate the gynoecium, count the number of pistils, of styles and
stigmas. Slice a flower lengthwise through its center and note the point
of attachment of tbe calyx and corolla and observe the ovary position
(whether superior, inferior, half-inferior). Remove the perianth from
another flower, and the stamens, too. With a sharp blade, slice crosswise
through the ovary, and identify the carpellary and placental situation.
Observe the number of locules of the ovary, the approximate number of
ovules, and then decide whether the placentatiou may be marginal, axile,
parietal, or free-central. Sometimes the ovule is solitary, or the number
is few, and the position basal or penduious. If these characters are de-
termined from a preliminary study, the process of identification becomes
more sure, and one is more confident of step-by-step decisions.

IDENTIFICATION
After the plant has been examined, and familiarity has been gained
from study of its flowers, the next steps are to identify and name it.
This calls for the use of a book. If available, a simple local flora or guide
written for use in the particular region should be sought. This, in prefer-
ence to the larger, more technically written manuals that cover a major
region of the country but which are more difficult for the beginner.
Simple works, such as Muenscher's Keys to Woody Plants, Jones' Flora
of Illinois, or Styermark's Spring Flora of Missouri (and there are many
others for other areas), are to be commended to the beginner. Spring
flowers are well suited to "first tries" and come at a season when field
collecting is most inviting.
A. k.e~, for plant identi1ication purposes, is a device whercb~ com-
binations of characters not present in the unknown plant at hand are
eliminated and by means of those characters that are present, one "comes
out" to a tinal answer. The keys in most modern works are called dichot-
omous keys. In these, contrasting conditions are combined in paired
couplets, with each of the two opposing situations in the couplet termed
a lead. The following is an example:
1. Flowers short~pedicelled, in racemes; periantb showy.
2. Ovary superior, stamens six ............................. Liliaceae
2. Ovary inferior, stamens three ........................... /ridaceae
1. Flowers sessile. in spikes; perianth inconspicuous, or none.
3. Plants herbaceous; inflorescence with one herbaceous spathe .. Araceae
3. Plants woody; inflorescence with one to many woody spathes ... Palmae
86 INTRODUCTION TO PL-INT T-IXONOMY

The first couplet is composed of the two leads preceded by the figure
one. Some authors prefer to designate couplets with letters, using here
for example A in place of the first figure one and AA in place of the
second. In the key above there is a total of three couplets. Note that the
leads of anyone couplet provide opposing statements. One is supposed
to fit the situation, and the other does not. If, for example, the situation
in the second lead of the first couplet prevails, then determine if it is the
first or second lead of the third couplet that matches the condition in the
unknown. Ultimately, one arrives at a name of the family, genus, or
species of the plant under study.
Often a book will contain one key by which to identify the plant with
its family, and a second key (or series of keys) by wbich to determine
the particular genus of that family. The third and last step is to turn to
the treatment of the genus, where (if more than one species is involved)
a key (0 the species will usually be given.
Usually, it is not sufficient to accept the species name determined by
the key as final. Confirmation is to be sought by comparing the plant in
hand with the description of the species whose name was arrived at by
the key. If there i, not reasonably close agreement, the plant may have
been misidentified and should be keyed again in a recheck for possible
error. Experience gained in the use of the less complex keys, and simi-
larly restricted floristic treatments of small areas, should provide a back-
ground and confidence to encourage one to use a more technical work
like Fernald's Gray's Manual or Abram's llIustrated Flora of the Pacific
Coast States.

PRESERVING MATERIAL FOR FUTURE IDENTIFICATION

As enthusiasm increases for xnowing more kinds of plants, it will be-
come evident that often it is not possible that each plant be studied as a
freshly-picked specimen. Furthermore, there develops the desire to ac-
cuinulate a record of the plants encountered, identified, and learned.
This record takes one form or another of an herbarium. In its simplest
form it may be that of cards on which are pasted fragments adequate to
associate a plant with its name, or full specimens on sheets of regulation-
sized herbarium mounting paper. These become recognition specimens.
The other extreme is the preparation of conventional herbarium speci-
mens pasted, complete with a label bearing collection data, on a
11 ~ X 16~-in. herbarium paper.
Co1l6crln6 and Identilrilt6 Technlque$ 87
The basic principles are the same, whether preparing specimens for
"recognition cards" or conventional herbarium specimens:
1. flatten the specimen under pressure, selecting a plant that is typ-
ical and undamaged, and pruning or folding vegetative parts as
needed;
2. Dry the specimen by pressure between sheets of newsprint and
blotters, changing the blotters as they absorb moisture from the
plant, or drying with heat.'
Equally important with preserving the specimen, is the preparation of
!lotes of characters and conclltions not evident from the dried specimens.
These include:
(1) Place where collected, and date;
(2) Habit and size of plant (when the plant is too large to comprise
the pressed specimen);
(3) Color of flowers and fragrance, if any;
, (4) Color of fruit, when present;
(5) Type of habitat where the plant was growing.
Occasionally the non-botanist desires to know the identity of an un-
known or to obtain verification of his own identification. This usually re-
quires sending a portion of the plant to a botanical center, such as a
botanic garden or college department of botany. The same procedures
described above for preparing and preserving the specimen should be
followed. Collection notes should accompany the specimen as an aid to
the person identifying the plant. Elaborate pressing facilities are un-
necessary. Most flowering material can be pressed and prepared for
mailing by placing between several thicknesses of newspaper, and leav-
ing under heavy weigh! (as a few books) overnight, and (leaving i( m
the newspaper) wrapping between two pieces of corrugated board (avail-
able from any carton of adequate size).
A second, and perhaps easier, method of sending material for identifi-
cation is to place the specimen (folded in accordion-plaits if necessary)
m an airtight polyethylene bag, such as is used for homefreezing of
foods, and mailmg it m a box to prevent undue crushing. Most plant
material will keep well for three to eight days if so packaged. It is not
desirable to moisten the specimen, nor wrap cut ends in wet material,
when sending fresh material m moisture-proof envelopes of this type.
1 For detaila on this and related techniquea and procedures. see Lawrence. Taxonomy
01 VQSculm Plant", pp. 234-248.
CHAPTER VI. Nomenclature

NOMENCLATURE is a function of taxonomy that deals with the de-

termination of the correct name of a plant. In its more precise applica-
tion it is associated with scientific or Latin names, but nomenclatural
studies may also involve common or vernacular names. A plant must be
identified and classified before it can be named, and it is important to
distinguish clearly between identification and classification and nomen-
clature.
It is not uncommon to find that a plant may have different names in
different floras or manuals. For example, one author may treat a plant as
a species, and assign it a name. Another may believe the same plant to
be only a variety of another species having an older name, and give it a
varietal name. The determination of the category to which a plant be-
longs is a matter of classification. The correct name of the plant in one
category (as that of species) may be different from the correct name it
would receive in another category. The differences between some genera
are not always sharp and clear cut, and it becomes a matter of opinion
as to which genus a plant belongs; some authors may believe the plant
to belong to one genus while sOme would place it in another-for this
reason the plant would have one name by some persons and another
name by others. However, when there is agreement on the correctness of
the identification and the classification, there should be only one correct
name for a plant.
Modem plant nomenclature, for the most part, dates from the eight-
eenth century. As the number of kinds of plants known to man has in-
creased, the procedures for naming them have become more technical
and legalistic. Today, many persons are familiar with the general mech-
anism of nomenclatural procedures, but so complex are some situa-
88
Nomenclature 89
tions that only experts-persons who have devoted years of study to the
subject-are held competent to resolve them. For this reason, this cha?-
ter makes no pretense to explore or explain the intricacies of nomen-
clatural problems. For them one is referred to the literature cited in
Taxonomy of Vascular Plants, pp. 220-222.
Plants are not named haphazardly. For the last two hundred years
the naming of plants has followed definite procedures. During this time,
in response to new situations, these procedures have been modified, re-
vised, and amplified. Linnaeus, the great Swedish botanist. laid down the
first rules of modern nomenclature, doing so in 1737. For a century they
remained relatively stable, but by 1867 communications had opened up
areas of the world unknown to Linnaeus and his students, and resulted
in the discovery, COllecting, naming, and describing of tens of thousands
of plants unknown to science in the mid-1700's. The revision of nomen-
clatural rules in 1867 was largely the work of Alphonse de Candolle, a
Swiss botanist of great repute. As more areas became centers of western
civilization, botanists came to be more conscious of their own inter-
national position and responsibility. In 1905 they held their third inter-
national congress in Vienna, when a new revision of a code of nomen-
clature was adopted. During this time a "splinter-group" of some Ameri-
can botanists attempted unsuccessfully to displace the international c~e
and their differences of view were not reconciled until 1930. The present
International Code of Botanical Nomenclature (hereinafter referred to
as the ICBN) was adopted at the Seventh International Botanical Con-
gress, convened in Stockholm, 1956, and is followed by botanists of all
nations. It is to be expected that subsequent congresses will legislate
changes in this Code.
One of the best ways to comprehend the scope of plant nomenclature
and to appreciate its modus operandi is to study the more important pro-
visions of the Code. This Code is divided into chapters, articles, and
recommendations. For ease of reference the original numbers of the
artides are used below, although not all comprising the Code appear in
this brief recounting. The articles of the Code are considered binding on
all who accept it. The recommendations are provided for guidance. In
the Code, almost all articles are accompanied by examples illustrating
the application of the provision. This Code sets forth the principles of
plant nomenclature (Articles 1-21) and the specific rules by which they
are activated (Arts. 27-83).
The principles provide that botanical progress is predicated on a
90 INTRODUCTION TO PL~NT T~XONOMf

universally accepted system of nomenclature that has for its objectives

(1) the fixity of names, (2) nomenclatural clarity and freedom from
ambiguity, and (3) the avoidance of the useless creation of names (Arts.
1-4). It deals with the categories of plant classification and the names
applied to the taxa assigned to them (Art. 8). Botanical nomenclature is
independent of zoological nomenclature, and there are many instances of
plant genera and animal genera having identical scientific names. The
scientific name of a plant is its Latin name, one taken from Latin (e.g.,
Digitalis, meaning glove.finger) or from Greek (e.g., Lithospermum,
meaning stone·seed), 01' is treated as if Latin (e.g., Lindera, latinization
of Linder, for John Linder, a Swedish physician).
The purpose of giving a name to a plant, or taxon, is solely to supply
a means of referring to it and not to indicate its characters or history.
For example, the name of the common milkweed is Asclepias syriaca, a
native of eastern North America, although its name would imply a
Syrian origin. The Latin name, Asclepias syriaca, is the species name. It
is a binomial. For explanation of species names and binomial nomen·
clature, see Chapter Ill, p. 23.
The names legitimate, illegitimate, valid, and invalid are constantly
encountered in nomenclatural discussions. A legitimate name is one that
is in accordance with the Code. It is the correct name. An illegitimate
name is onc that is contrary to the Code, an incorrect name. It is pro·
vided that illegitimate names, such as nomina nuda (naked names,
those not provided with descriptions) may not be included in nomen·
clatural considerations. A valid name is one that has been properly pub·
lished. One may be obliged to consider several valid names to determine
which is the legitimate name for a plant. An invalid name is one that
docs not meet all requirements for valid publication and as such is also
an illegitimate name. The terms valid and legitimate are sometimes but
improperly used interchangeably and thereby cause confusion. It is im·
portant to recognize these distinctions between them.
The Code is specific in designating the categories or units of classifi·
cation and the sequence in which they are related to One another. The
plant kingdom is divided into divisions and it depends on whose classifi·
cation is followed as to what name is employed (e.g., SOme authors treat
the vascular plants as composed of two divisions, the Pteridophyta and
Spermatophyta, others may accept the view that taxonomically these rep·
resent but a single division, and call it the Tracheophy/a). The practice
of designating these taxa as phyla; as is allowed for divisions of the
Nomenclature 91
animal kingdom, is contrary to the Code. The units of classification have
been treated in Chapter III and only the nomenclatural provisions for
genera and lower units will be reviewed here.
The genus is composed of species, but for clarity in classification the
Code provides for a number of infrageneric categories in the following
sequence of descending magnitude: subgenus, section, subsection, series,
subseries.
The name of a species is a binary combination consisting of the name
of the genus (as Quercus for the oak) followed 'by the single specific
epithet as the epithet alba, forming the binomial Quercus alba (the
species .name of the white oak). Quercus alba is the species name,
known also as a binomial, a name of two words.
Provision is made for three infraspecific categories of the species: the
subspecies, variety, and form.
From the above and the aspects covered in Chapter III, it should be
clear that every species belongs to a genus, every genus to a family,
every family to an order, every order to a class, and every class to a
division. This sequence of categories is specified by the Code and cannot
be altered.
The principle of priority is a cornerstone of the Code. It provides that
each taxon from the rank of order to genus has only one correct name,
the earliest published with the same rank. "For any taxon below the rank
of genus the correct name is the combination of the generic name witj!
the earliest available legitimate epithet or epithets validly published with
the same rank" (Art. 16). There is one exception to this principle of
priority for generic, family, and ordinal names. It is the provision (Art.
24) to avoid disadvantageous changes of names, wneIeb1 the Code pIG-
vides lists of generic names retained as exceptions to the applicati')n of
the rule of priority. The retained name is a conserved name (nomen con-
servandum) and any other name for that taxon is a rejected name (nomen
rejieciendum). Efforts to extend the provisions of this rule to cover the
conservation of species names have been defeated.
Names of all taxa are based on types. This is known as the applica-
tion of the type method. The nomenclatural type of a species is a speci-
men, usually a pressed and dried herbarium specimen. If it is the one
designated as the type by the author of the species, it is termed a holo-
type; if it is one from original material from which the species was
described it may, if the author failed to designate a holotype or if the
latter has been lost or destroyed, be designated as the lectotype; and if
92 INTRODUCTION TO PLANT TAXONOMY

it is a specimen that was not known to the author of the name but (in
the absence of any holotype) is selected by another person as typifying
the species, it is designated as the neotype. The type of a genus is a
particular species of that genus. Usually it is the first species to have
been described in the genus or it may be the species on which the genus
was based. The type of a family is a genus, that of an order is a family,
etc.
The publication of names is a subject that is strictly legislated by the
Code. There are two aspects to the publication of scientific names:
effective publication and valid publication. Effective publication refer[
to the place and the scope of publication. Until January 1, 1953, a
name could be published anywhere-a book, a magazine, on a printed
herbarium label of a widely distributed collection, in a newspaper, or a
nursery or seed list. The rules now reject names published since January
1, 1953, in newspapers or tradesmen's catalogues. To be effective, th"
name must also be published in a book or periodical that is distributed
to botanical centers throughout the world in some indelible form. Publi-
cation by communication of new names at a public meeting, or by an-
notation of collections, or by issue of microfilm of otherwise unpublished
manuscripts, does not comprise effective publication. On and after
January I, 1953, a name is not effectively published if printed on a label
and accompanied by a description and distributed l"idely with specimens
of that name.
Valid publication requires that in addition to being effectively pub-
lished, the name must meet one or the other of the following two con-
ditions: (a) it must be accompanied by a description of the taxon (in
Latin if published on or after January 1, 1953); or (b) it must be ac-
companied by a reference (direct or indirect) to a previously validly
published description. The first condition should be self-explanatory.
The second may be less obvious and requires illustration:
The species Rhododendron candense was so named by John Torrey in
1839, but it was first given a binomial by Linnaeus in 1762 who named
it Rhodora canadense and accompanied the name with a description. Ac-
cording to the Code it was correct for Torrey to validate his transfer of
the plant from the genus Rhodora to the genus Rhododendron by fol-
lowing the new name with a reference to the binomial used by Linnaeus
(the basic name or basonym) and to Linnaeus' description. On and
after January I, 1953, new transfers, such as this, are validly published
Nomenclature 93
"only when the basonym is clearly indicated with its author and
the place and date of publication" (Art. 42).
The citation of authors' names is provided for in the Code (Arts.
55-60). In floras, manuals, and technical works, the scientific name of a
taxon is followed by the name (often abbreviated) of the person who
first described it. This increases the precision and accuracy of the treat-
ment. For example, in 1753 Linnaeus named tbe , red maple Acer
rubrum. To distinguish this from a different species of maple, but given
the same name by Lamarck in 1788, one writes it Acer rubrum L. It is
known now that Lamarck's maple is the same species as that named
Acer saccharinum by Linnaeus in 1753 and one would treat Lamarck's
name as a synonym of Acer saccharinum and write it "Acer rubrum
Lamarck (1788), not L. (1753)." By citation of these authors' names,
clarity and accuracy is added.
Article 58 of the Code reads, "When a name has been proposed by
one author . . . and is subsequently validly published and ascribed to
him by another author who supplied the description, the name of the
latter author must be appended to the citation with the connecting word
ex." The Oak, Quercus hemispha!rica was named, and perhaps described,
by John Bartram of Philadelphia, but the name was published by Kar'
Willdenow of Berlin in 1805, not by Bartram. To distinguish between
the author of the name (Bartram), and that of the publisher (Will-
denow), the binomial and its author citation is Quercus hemisphrerica
Bartram ex Willdenow. For brevity this may be condensed to Quercus
hemisphtl!rica Bartram.
For situations where a genus, species, or taxon of lower rank "is
altered in rank but retains its name or epithet, the author who first
published this name or epithet must be cited in parentheses, followed by
the name of the author who effected the alteration" (Art. 59). This is
sometimes called double author citation. For example, in 1814 Frederic
Pursh named a California shrub Arbutus tomentosa. Twenty-two years
later John Lindley, of London, recognized it as a member of the ge~s
Arctostaphylos and made the new combination of Arctostaphylos tom n-
tosa (Pursh) Lindley. Pursh's name in parentheses shows he had gi en
the taxon the epithet tomentosa in another genus or category, and Lind-
ley's name following shows that he was responsible for the transfer. A
second example shows the same application if an epithet is retained
when a taxon is transferred from one category to another: A shrub was
94 INTRODUCTION TO PLANT TAXONOMY
named Ceanothus Integerrimus var. parvifolius by Sereno Watson in
1875. Later, William Trelease recognized it as a distinct species and
named it Ceanothus parvifolius. Since it is based on Watson's type, and
since Trelease retained Watson's epithet, the binomial and its full cita-
tion is Ceanolhus parvifolius (S. Watson) Trelease.
Retention of epithets of taxa usually prevails when names are trans-
ferred wilhoul change of rank. When a species is transferred from one
genus to another, the epithet must be retained unless the resulting
binomial is (I) a later homonym (duplication of an earlier name for a
different plant), (2) a tautonym (duplication of generic name as the
specific epithet, as Pinus Pinus), or (3) unless there is available in the
genus an earlier validly published specific epithet. These same rules
apply to the transfer of taxa below the rank of species, provided no
change of rank takes place.
The uniting of two or more taxa of the same rank results in retention
of the oldest name or epithet. When two or more taxa of different rank
are united, the earliest legitimate name or epithet in the new (accepted)
category is correct. In no case does a name or epithet have priority out-
side its own rank. To illustrate, the name of the variety of ash now
known as Fraxinus pennsylvanica var. subinlegerrima (Vahl) Fernald
was long treated as F. pennsylvaniea var. laneeolala (Borkh) Sargent.
However, the epithet laneeolala had its origin in the species name or
binomial Fraxinus laneeolala Borkhausen (1800). This taxon bearing
the epithet "Ianceolata" was first treated in the rank of variety by Sargent
in 1894. If the plant is accepted as a variety, the oldest epithet assigned
it in the category of variety must be used: that epithet is subinlegerrima,
given by Vahl in 1804. VaW's name is placed in parentheses because he
mistook the plant to be a variety of Fraxinus juglantiifolia. Fernald
made the transfer to F. pennsylvaniea, hence his name follows Vahl's.
The Code recognizes numerous situations under which names or
epithets must be rejected. When rejected, the name is replaced by the
next oldest legitimate name or, if there is none, it is replaced by a new
name. In this connection it must be remembered that a name may not
be rejected "merely because it is inappropriate, or disagreeable, or be-
cause another is preferable or better known, or because it has lost its
original meaning" (Art. 72).
A name must be rejected if it is illegitimate, and nomenclatural illegiti-
macy exists if:
1. it was superfluous when published (that is, if another and legiti-
mate name already existed in the accepted rank);
Nomenclature 95
2. the earliest legitimate epithet in the particular circumscription,
position, and rank had not been adopted;
3. it is a later homonym (that is, if it duplicates a name previously
published for a taxon of the same rank but based on a different
type, even if the earlier homonym was illegitimate);
4. it is a tautonym;
5. it is a binomial published in works in which the Linnaean system
of binomial nomenclature was not consistently used for species.
The Code makes other provisions by way of its Recommendations.
Those most commonly encountered deal with the subject of capitaliza-
tion of specific and ternary epithets. On this subject the present Code is
essentially noncommittal, Recommendation 82 g providing:
All specific and infraspecific epithets should be written with a small initial
letter, though authors desiring to use capital initial letters may do so when
the epithets are directly derived from the names of persons (whether actual
or mythical), or are vernacular (or barbaric) names, or are former generic
names.

The use of the single and double "i" is another source of confusion
and is covered by Recommendation 82 c (1),
When the name ends in a consonant the letters ii are added (Ramondii
from Raymond), except when the name ends in -er, when i is added (thus
Kerner; from Kerner). Those who follow this Recommendation may treat
the termination -i as an orthographic error and correct it.

There is another nomenclatural situation that is inextricably inter-

locked with taxonomy and which is a source of confusion to many bud-
ding taxonomists. That is the recognition and naming of the typical ele-
ment of a species. The typical element of a species is that which is in-
distinguishable from the type specimen of the species. If the species is so
constant and so uniform in all characteristics that all plants appear to be
like the type specimen then no problem of recognizing variants of the
species is at hand. However, suppose you were the first to discover the
species of maple known today as Acer palmaturn and that you collect,
prepare, and document an herbarium specimen of the species and pub-
lish the name and description in accordance with the rules. That speci-
men is the type of the species. As an aid to visualizing this type, assume
it to be represented by Fig. 43 a.
Let us exte"d the supposition further and assume that a year later
on a Japanese mountain side you collected another maple. From im
96 INTRODUCTION TO PLANT TAXONOMY
fruits and other characters you recognize it as Acer palmatum, yet its
foliage is so different as to be distinct from the specimen first col-
lected. Perhaps you consider it a variety of the first species and
name it Acer palmatum var. aconitifolium. The type specimen of this
new variety is the one you collected. Assume it to be represented by
Fig. 43 b.

FJg.. 43. Acer palma/urn: a. var. palmatum; b. var. aconiti/olium; c, var. dis-
sec/urn.

The species, Ace, palmalum, is now composed of two elements. The

first, on which the species was based, is known as the typical element.
As soon as the second was added, the concept of the species as a unit
had to be amplified to include both the typical element (Fig. 43 a) and
the aconitifolium element (Fig. 42 h). Since the species is composed of
two variants, each must have a varietal name. One is no more the species
than the other. It was by mere chance that one was discovered and
named before the other. It becomes desirable to differentiate nomen-
ciaturaIly, between the two. The Code provides for this by stipulating
that the typical element of a taxon be designated by repeating the epithet
of the taxon in the next higher category which is based on the same type
specimen. By application of this rule, the name of the typical element
of this maple becomes Ace, palmatum var. palmatum. Because the type
specimen of var. palmalum is that of the species (a species, no matter
the number of its varieties, can have only one type specimen), it is not
necessary to follow it with an author's citation. Its author is always that
of the binomial. Prior to legislation of the present Code there was no
uniformity in the naming of typical elements and authors designated
Nomenclature 97
them by such epithets as typicus, genuinus, or by prefixing eu- to the
repeated specific epithet. Botanists are now directed to replace these
epithets, as necessary, with that repeating the epithet of the next highest
taxon based on the same type specimen.
In most works written more than a half-century ago, and some of
more recent time, authors described the species in the sense of the typ-
ical element and treated varieties as if they were appendages of the
species. These authors did not conceive the species as a biological unit
composed of variable populations. They did not treat the first-named
population as a typical element. For this reason, in a majority of cur-
rent floras and manuals the typical element of a highly variable species
may not be accorded nomenclatural recogrution.
The nomenclature for cultivated plants must conform to that provided
for native plants insofar as the same classificatory situations prevail.
However, the many specialized situations encountered \\ ith culti\"ated
plants have resulted in the formation of an Appendix \,111 of the ICBN.
This Appendix, known also as the Intematiollal Cod" of ,\'omellciatllre
for Cultivated Plants, was presented at the Stockholm Congress in 1950.
It was later adopted, with some revision, by the international committee
on horticultural nomenclature and r~gistration at the Thirteenth Inter-
national Horticultural Congress, London, 1952.
This review of plant nomenclature, and of the major provisions of
the ICBN, increases one's appreciation of the principles involved and
should help one to understand that there are usually good reasons for
occasional changes of plant names. In this connection, it is imperative
to distinguish between changes resulting from processes of classification
and identiJlcation and those due to application of the rules. If the name
change results from a change of rank of a taxon, it is one of classifica-
tion; if from a change from affiliation with one genus to another it may
be one of identification. An analysis made by me in 1949 of changes
of about 120 plant names showed that about one-third represented the
application of the "Rule of Priority," the displacement of an existing
name by an earlier legitimate name; a fourth represented the applica-
tion of the "Homonym Rule" (Art. 72), the displacement of a name be-
cause it had been used previously for a different kind of plant; and the
balance of the changes represented a variety of taxonomic rather than
nomenclatural reasons, including misidentification by earlier authors,
the subdivision of genera into two Or more genera, and the uniting of
two or more genera under one name
CHAPTER VII. Phylogeny and
Biosystematics

PHYLOGENY is the evolutionary history of a taxon, and by the prin-

ciple, of phylogeny the attempt is made to account for the origin and
development of the taxon. In much of the literature the term phylogeny
has become associated more with studies of major units of classifica-
tion (as the family and units above it) than with the genus and in-
frageneric units. This distinction is artificial, because studies of the
evolutionary history of the species and its subdivisions may be as truly
phylogenetic as those dealing with higher units. Biosystematics, on the
other hand, is a study of living popUlations with the objective of recog-
nizing and circumscribing natural biotic units (quite different from con-
ventional taxonomic units), and to classify them objectively as taxa of
different orders of magnitude. The casual reader may think these two
topics as strange bed-fellows to be combined in the same chapter but
this is not the case. They represent two different approaches to the same
goal: the formulation of a classification based on genetic or natural rela-
tiollships.
The primary objective of phylogenetic studies is to determine the
origins and relationships of all taxa of extinct and extant plants, and
from these data to prepare a classification reflecting these relationships.
This is a goal. Such a classification does not exist today, and there :_
scant probability that it ever will exist. Why not? Because much, in fact
most, of the evidence establishing the identity of ancestors of present-
day plants is believed to be nonexistent, and it is not likely that sub-
stantial portions that may be buried beneath the earth's surface ever will
be recovered or past history reconstructed. Without these data, present
and future systems of classification must be largely conjectural.
98
Phylolleny and Bio.y.temo.tic. 99

PRIMITIVE VS. ADVANCED CHARACTERS

Phylogenetic studies have attempted to determine, among other things,
which characters of present-day vascular plants have remained substan-
tially unchanged during the geologic ages and epochs of evolution. A
character that has persisted for long periods is said to be conservative
or primitive. The terminal position of the ovary on a branch or axis is
an example of a conservative character. One of the best sources of
evidence for primitiveness of characters (or of taxa) is in the fossil rec-
ord, for here there are paleontological data of plants of former times.
Unfortunately these data are very few, and for flowering planls, fossils
have not produced much conclusive evidence. Contemporary taxonomic
writings may designate certain characters as primitive and others as ad-
vanced (of later evolutionary development), but with few exceptions
these are only allegedly or presumably primitive or advanced. The evi-
dence in this regard is more often circumstantial, or even only specula-
tive, than factual. Very often the evidence that is available serves a, the
basis for certain deductions compatible with the general theory of evolu-
tion. The synthesis of data from many sources often contributes to a
series of ded,!ctions relating to possible and plausible phyletic views that
seem to be in greater harmony with known facts than are earlier views.
By these additive processes our ideas of phylogeny take on a three-
dimension.l character, and there follows the recognition that plant rela-
tionships are not simply tree-like in their composition but are further
complicated by having the branches interlocked in a reticulate fashion.
A trend is now gaining momentum to place greater confidence in the
conservative nature of anatomical characters, especially of the vascular
anatomy. The type of stele within a stem, the presence and nature of
leaf-gaps, of orientation of vascular strands, and the vestigial remains
(or presumption of such) are now subject to greater use than ever be-
fore in determining allegedly primitive conditions. Acceptance of these
views has led to the belief that organs or plants that today may appear
simple are in fact seemingly simple by reduction or loss of parts.
Anatomical evidence is usually presented in support of such views. If
accepted, it holds that these seemingly simple structures or plants have
come from ancestors that were more complex as concerns number of
parts, hence the seemingly simple situation represents an advancement
over the earlier and more primitive condition. An illustration of this is
to be found in the catkins of birch, or alder, or hornbeam; inflorescences
100 INTRODUCTION TO PLANT TAXONOMY

now seeming simple, but adduced to be simple by reduction and fusion

of parts from ancestral types-inflorescences not simple but highly com-
plex and advanced.
A classification may be phylogenetic or it may be strictly artificial
No truly phylogenetic classification exists today for the vascular plants
or the flowering plants, or the monocots, or for any order of this taxon.
Undoubtedly some monographic works of some genera have been so
thorough as to be able to classify the species of a genus In as nearly a
phylogenetic classification as possible. Other things being equal, the
lower the classification unit (as species is lower than genus) and the
smaller the number of taxa composing it, the more likely it is that a
phylogenetic classification of its taxa can be closely approached.
In the absence of paleobotanical data, the phylogenist dealing with
higher units of classification seeks to piece together an evolutionary pat-
tern of ancestry by using as many other data as possible. These are ob-
tained from studies of anatomy, morphology, cytology, genetics, and
physiology of the plants concerned. Structures and organs too minute to
be of use in identification procedures are often very important, especially
those associated with the embryo, pollen, chromosomes, floral vascular-
anatomy, and the seed.
By application of these techniques, it is now held that the inferior
position of the ovary is derived from ancestors in which the ovary was
superior, that flowers possessing a gynoecium of one or a few pistils may
have been derived from those whose gynoecia were of many pistils, that
the simple ovary with marginal placentation preceded the compound
ovary with axile, parietal, or free-central placentation, etc. These studies
have also led to the belief that possession of certain characters of two
or more major taxa may be evidence of parallel evolution and not nec-
essarily of close or common relationship. To illustrate, a polypetalous
type {Jf corolla is generally held to be more primitive than the gamo-
petalous type. By the Engler or the Bessey system of classification all
taxa with gamopetalous corollas belong in one sector of each system,
on the assumption that this condition arose once long ago by convergent
evolution, that is, by convergence of evolutionary products and the
"fanning out" from that convergence of subsequent taxa possessing the
character in common. Evidence is continulng to accumulate to refute
this view, and to support the opposing view-that by parallel evolution
the condition of gamopetalous corollas has arisen many times among
flowering plants and among taxa phyletically far removed from one
another.
Phylogeny and Bio.y.temal;a 101
Phylogenetic studies at the level of the family and above result in
modifications of existing systems or in the formulation of new systems
of classification. That is why it is now believed by many botanists that
the systems of Engler and of Bessey are so "out of date" that neither
reftects a synthesis of the most acceptable data available. New system.
have been presented since these, but none has been adopted widely;
more new systems and revisions of systems are inevitable and will be-
come subjects of biological importance. The new and successive systems
have their place; each is to be studied, tested, and analyzed. None may
displace existing systems in our major herbaria or floras, because the
taxonomist can identify, name, and catalogue (i.e., classify) his plants
without phylogeny. Phylogeny leads to a better understanding of rela-
tionships at all levels and thereby is a significant contribution to knowl-
edge, but it is not essential to the demands on taxonomy as a functional
science.
Biosystematics has the same goal as does phylogeny, but is concerned
primarily with units of genus and below. Furthermore, it is a facet of
taxonomic endeavor that is predicated on data obtained from the be-
havior of living individuals and populations. The biosystematist rejects
most conventional taxonomic definitions of a genus or a species and
tries to determine the circumscription of these taxa (or populations)
more objectively. In doing so, emphasis is placed on cytogenetics and
cytotaxonomy supplemented by the classical approaches of morphol-
ogy, ecology, and phytogeography. Cytogenetics deals with the analysis
of breeding behavior of individuals in the light of their chromosome
complements, contents, and activity at meiosis. Cytotaxonomy is the
integration of cytology-more properly termed karyology-and taxon-
omy in the effort better to understand and to resolve problems of plant
relationships.
Biosystematic investigations include:
1. sampling of the taxon and the cytological study of its populations,
2. studies of the genetic compatibility of the populations and of the
vigor and fertility of resultant hybrids in the determination of pres-
ence or absence of breeding barriers,
3. studies of the homologies of the chromosome6 in the hybrids at
time (Jf meiosis,
4. integration of above data with those obtained from comparative
morphology and geographical distribution,
5. subjection of individuals or popUlations thus studied to criteria of
a system of classification into biosystematic units.
102 INTRODUCTION TO ~NT ~4XONOMY

The biosystematic categories have been devised in the effort to pro-

duce a better understanding of natural relationships at the rank of genus
and below. They are categories that may be counterparts of taxonomic
categories but are never to be used as substitutes or equivalents of them.
They have no status in the formal nomenclature of plants. Each pro-
vides a single-word term for a biosystematic situation. The categories
(and their approximate phyletic level) are:
comparium--comparable to the genus
ecospecies-an approximation of the conventional and conserva-
tive species, and
ecotype-sometimes parallel with the geographic variety or sub-
species of taxonomisl$.
In no case should anyone of these terms be applied to a plant or popula-
tion unless thf> situation for which the term stands has been proved ex-
perimentally to exist for the particular taxon. These biosystematic
categories represent evolutionary nodes in the sense that each category
represents a step or level in the evolutionary scale of differentiation from
that of a local population to that of a genus.
The student requiring a more complete treatment of the subject of
biosystematics is referred to Chapter VIII in Taxonomy of Vascular
Plants. where the various biosystematic categories are described in de-
tail (pp. 176-179) and the advantages and limitations of the techniques
and results are discussed.
Biosystematic studies require facilities for field experimentation, for
research studies of clonal separations of individuals under different en-
vironmental conditions, and for cytogenetic experiment. They are time-
consuming and costly. Results obtained have contributed materially
toward the solution of numerous taxonomic problems and "riddles"
not otherwise resolved. They may provide one avenue of approach to
some difficult taxonomic groups. For other groups they have been
found to be of little help--as in solving problems of taxonomic rela-
tionship and speciation in such genera as Lathyrus or Astragalus.
These experimental channels provide the taxonomist with one more
and very valuable tool with which to work, but it must be recognized
that biosystematic studies of themselves are not likely to provide the
answer to a majority of the taxonomist's problems of speciation and
determination of generic limits.
CHAPTER VlIl. Taxonomy in
North America

WHENEVER a student of North American plants makes use of floras,

manuals, or technical papers he encounters the names of the men who
described these plants, or of men whose names are associated with
binomials or important floristic works. It is not enough to know the
plants and their biology; the taxonomist profits from knowing something
of the past that has contributed to the heritage of the science-a heritage
built around the lives and accomplishments of the men concerned.
The beginnings of taxonomic botany in North America cannot be
separated from botanical studies in Europe of the same period. In the
,earliest days 'Of New World exploration and colonization, tropical Amer-
ican plants were known to Europeans to a greater extent than were
.those of temperate regions. Sir Francis Drake sent plants back to Eng-
land from tropical America in the sixteenth century. Robert Morrison
(1620--1683) of England and Botanist to the king, wrote three volumes
on plants in the Royal Gardens. Many of these plants were of American
origin.
The first book devoted to North American plants was Flora virginien-
sis written hy John Frederick Gronovius and published in 1739-43 in
Holland. Gronovius, friend and onetime counselor of Linnaeus, based
this fiora on extensive notes and a collection of pressed plants received
in 1730 from his American correspondent John Clayton of Virginia. A
second and much revised edition, based on the Linnaean system, but
using polynomial nomenclature was published in 1762. The work is of
historical interest, for the first edition was cited regularly by Linnaeus
in Species Plantarum. Some of Clayton's plants are now at the British
103
104 INTRODUCTION TO PLA.NT T4XONOMY

Museum (Natural History) in London, and not a few are the types of
Linnaean species. Clayton is commemorated by the genus Claytonia
(Portulacaceae) .
John Bartram (1699-1777), a Philadelphia Quaker, laid out and
planted one of the first arboreta in this country. He was a correspond-
ent of many European botanists, notably Peter Collinson of London,
and to whom he sent specimens, seeds, and plants of a large number of
American species and genera. His travels for plants took him as far
south as Florida and as far north as Lake Ontario, New York. The moss
genus, Bartramia, was named in his honor.
Another Philadelphia Quaker and contemporary of Bartram's was
Humphrey Marshall (1722-180 I ), also a dendrologist, who published
(1785) a small book on American trees entitled Arbustum Americanum.
Marshall named many species but by modern standards his descriptions
are neither ample nor critical. He followed the binomial system of
nomenclature and arranged the plants in alphabetical sequence. Marshall
was for many years a correspondent of Sir Joseph Banks of London and
sent to him seeds and roots of many American plants. The genus
Marshallia (Compositae), often credited as having been named in honor
of Humphrey Marshall, commemorates his nephew, Dr. Moses Marshall
(1758-1813), who was more important as a collector and exporter of
plant materials than as a scientist.
Peter Kalm (1716-1779), Swedish naturalist, student and associate
of Linnaeus, spent three years in eastern North America and, perhaps
more than any other person, provided Linnaeus with valuable collections
of plants from this region. These specimens are now to be seen in the
Linnaean Herbarium in London. His account of travels from Philadel-
phia south through Virginia and north to Canada have been translated
into Engli.sh. In identifying and naming Kalm's collections, Linnaeus
used the epithets "canadensis" and "virginiana" frequently and loosely
as the two principal localities in North America, without regard for the
limits of the political subdivisions. The genus Kalmia (Erieaceae) named
by Linnaeus commemorates Peter Kalm.
Thomas Walter (c. 1740-1789) emigrated from England to eastern
South Carolina, but little is known about his background or life. Remote
from cultural centers, libraries, or associates in science, he wrote a
flora of the region in Latin and named it Flora Caroliniana. It was based
On his Own collections, augmented by some of John Fraser's, and the
manuscript, together with Walter's herbarium, was taken by Fraser (0
Ta.;t"onomy in North America 105
London in 1788. Fraser published Walter's flora at his own expense,
and it was the second floristic work of its kind devoted to a region of
this country. It treats about 1,000 species and 435 genera; 200 species
were described as new. Whenever Walter had a plant whose generic
position was unknown to him, he resorted to the curious device of
placing it in his genus Anonymous and made many binomials therein
(e.g., Anonymous plicatus, A. rigidus). As the generic position of these
species became known to later authors, many of the epithets were trans-
ferred to the correct genus, with Walter acknowledged as the paren-
thetical author. The International Code of Botanical Nomenclature
(1952) rejects these basonyms. Subsequent combinations based on the
Walter epithets in Anonymous can no longer be treated as legitimate if
antedated by another legitimate name for the same plant. Walter's
herbarium, a bound volume of snips and scraps, was given by Fraser's
descendants to the Linnaean Society of London in 1848, and by the
latter in 1863 to the British Museum (Natural History), where it is now
available for study.

Period of 1800-1840
This period was one of active exploration along the Atlantic coast
and Allegheny Mountains and westward into the Mississippi drainage
basin. The three most notable botartists of the period were Pursh,
Rafinesque, and Nuttall.
Benjamin Smith Barton (1766-1815) was more important as a
teacher and benefactor of botany (at the College of Philadelphia, an-
tecedant of the University of Pennsylvania) and physician than as II
pUblishing taxonomist. His Elements of Botany (1803) was one of the
first American works of its kind. Much of his research dealt with prob-
lems of pharmacognacy and materia medica. He was responsible for
some of the work of Pursh and Nuttall. The genus Bartonia (Loasaeeae)
commemorates his narne.
Jacob Bigelow (1786-1879), a Boston physician, published the first
good flora of the New England area, Florula Bostonensis. The first edi-
tion (1814), based on the Linnaean system, was restricted to the plants
growing within ten miles of Boston, but the next (1824), based on the
de Candollean system, accounted for most of those of New England.
Bigelow's most valued contribution is considered to be his three-volume
work American Medical Botany (1817-20). Bigelowia (Compositae)
was named in his honor by de Candolle.
106 INTRODUCTION 1'0 PLANT TAXONOMY

William Darlington (1782-1863), another of the Philadelphia bota-

nists, and pupil of Barton, was a physician of note, As a botanist he
published a Florula Cestrica (1826), a catalogue of plants in the region
of West Chester, Pennsylvania, of which a revised, enlarged and an-
notated edition was later published under the name of Flora Cestria
( 1837) ; in the first edition he adopted the Linnrean system of classifica-
tion, and in the second the de Candolle system, His Agricultural Botany
(1847) was one of the first American works to name and describe culti-
vated plants. Torrey named the genus Darlingtonia (Sarraceniaceae) of
California in his honor.
GOlthilf H, E, Muhlenberg (1753-1815), a Lutheran clergyman, was
a teacher and botanist of Lancaster, Pennsylvania, and was a specialist
in the taxonomy of grasses, sedges, and fungi, His Catalogue Plantarum
Americae Septentrionalis (1813) was the first comprehensive account
of the then known native and naturalized plants of North America (ar-
ranged according to the Linnaean system). Previous works were of more
restricted scope. He collected assiduously, accumulated a large private
herbarium that was especially strong in sedges and grasses, and was the
author of scores of new species names in these families, of which many
binomials were published for him by de Candolle, Muhlenbergia, a genus
of grasses so named by Schreberer, commemorates his name. Probably
no American botanist of the period had a wider circle of foreign botan-
ical correspondents than did Muhlenberg, Unfortunately, a major part
of his herbarium was ravaged by insects, and many types were lost. The
remains of this collection are at the Academy of Natural Sciences,
Philadelphia,
Andre Michaux (1746-1802), a French botanist and student of
Bernard de Iussieu, was sent by his government to this country in 1785
to collect material for the botanic gardens in Paris. He settled in
Charleston, South Carolina, in 1787, and stayed there for ten years, He
traveled during this time over much of the eastern part of the country.
During this period he introduced about 60,000 American plants to
French and Austrian gardens. He published (1801) the first work on
the oaks of North America and in 1803 there was published under his
authorship the first flora of North America, Flora BoreaU-americana,
Michauxia, a campanulaceous genus, was named for Michaux by
L'Heritier.
Fran<;ois Andre Michaux (1770--1885), son of Andre, and a more
talented botanist than his father, published his North American sylva
TaS-OllOrnr in North America 107
Histoire des arb res forestiers de I'Amerique septentrionale . in three
volumes in 1810-13. Here, many species of forest trees are named and
described for the first time. Fran~ois Andre was in this country for
three different, but short periods, returning to France for.the last time
in 1809. He was better known in the United States than in France and
bequeathed a part of his fortune to the American Philosophical Society,
Philadelphia.
Frederick Pursh (1774-1820), of Saxony, Germany (not Siberia, as
often recorded), emigrated to the Philadelphia area in 1799 as a
gardener and profited from the patronage of Humphrey Marshall and
Benjamin Smith Barton. He lived in poverty most of his life and died at
the home of a friend in Montreal while working on a 1I0ra of Canada.
He travelled some in eastern North America, collected extensively, is
alleged to have pirated collections of others and to have published their
new species as his own. In 1811 he went to England where (in 1814)
he arranged for the publication of his two volume Flora Americae Sep-
tentrionales, the second comprehensive 1I0ra of North American plants.
It contains no keys and the descriptions are very brief. Many new
species were described; it nearly doubled the number of species known
to Michaux. Pursh included here also the plant records of the Lewis
and Clark expedition to the Pacific Northwest. Many of Pursh's speci-
mens are in the herbarium of the Academy of Natural Sciences, Phila-
delphia.
Thomas Nuttall (1786-1859) came to Philadelphia from Yorkshire,
England, in 1808, as a journeyman printer. During his first year in
America, and with counsel from Benjamin Smith Barton, he became
avidly interested in knowing the local 1I0ra and travelled in adjoining
!1.a.te~ ~ rot plaru&.. He =~ R~, a Scotch u.aIJIDII.-
ist, to the upper limits of the Missouri River and brought extensive
collections of plants back to Philadelphia. For the next eight years he
travelled over much of the eastern half of the country making Phila-
delphia his winter headquarters. It was during this period that he com-
pleted his two-volume work The Genera of North American Plants, and
a catalogue of the species. to the year 1817 (published in 1818). This
was the first work of its kind and is noted for its meticulousness and
accuracy. Nuttall himself set the type for the two volumes. From 1822-
1833, Nuttall was curator of the Botanic Garden at Harvard but was not
happy with so sedentary a life. In 1834, in company with John K.
Townsend, he went overland to Oregon, down the Columbia River to
108 INTRODUCTION TO PLANT TAXONOMY
Fort Vancouver, on to the Hawaiian islands, and back to Philadelphia
late in 1835 by way of Cape Hom. In 1841, he returned to England
to claim a legacy granted on the condition that he maintain his residence
there for nine months of every year. Prior to leaving Philadelphia, he
published a three-volume supplement to Michaux's Silva which he titled
The North American Silva (1842-54). Elias Durand summarized
Nuttall's work by writing, "No other explorer of the botany of North
America has personally made more discoveries; no writer on American
plants, except perhaps Asa Gray, has described more new genera and
species." The genus Nuttallia (Rosaceae) commemorates his name. Nut-
tall's plant collections contain a large number of types, and, while
many have been lost, those extant are at the Academy of Natural
Sciences, Philadelphia, and the British Museum (Natural History),
London.
Constantine Samuel Ralinesque (Schmaltz), was born in 1784 in
Constantinople of French and German parents and spent much of his
early life in southern France and Sicily. Rafinesque, as he is best known,
is one of the world's most controversial botanists. He was a genius,
yet an erratic eccentric, an egoist who was vain, ambitious, and con-
tentious to a fault. His egoism, fired by an inexhaustible energy, engen-
dered in him a passion that he be recognized as the supreme naturalist,
a conviction that drove him to write voluminously, not only in the fields
of geology, zoology, botany, and the physical sciences, but also in
philosophy, religion, and metaphysics. Dr. E. D. Merrill has written of
him, ". . . he lived to publish." The opinion that he was a psychopath
is not restricted to his contemporaries, but it may be unduly severe.
Certainly he has not always been treated fairly. It is unfortunate that
his publications in descriptive biology were so consistently ignored by
his contemporaries and immediate successOrs. Much of his botanical
work was of merit. He published several ftoristic works of nomencla-
tural importance. One of these, F10rula Ludoviciana or Flora of the
State of Louisiana (1817), was of a locality he never visited nor whose
plants had he studied. In this work he based his descriptions on the
scant notations of a French author's Flore Louisianaise (1807). Many
of his contemporaries, and botanists of the succeeding generation (in-
cluding Asa Gray), ignored his writings. Of, recent time greater study
has been given to Ratioesque', work, notably by E. D. Merrill of the
Arnold Arboretum, and although Ratinesque left no herbarium, many
of his descriptions and name, have been identified. Merrill's Index
Taxonomr ira Nor'" America 109
Ra(mesquenorum (1949) has become indispensable to all students of
Ralinesque names. The genus Ra(inesquia of Compositae was named in
his honor by Nuttall, and on one occasion, perhaps in fear his name
might not be so perpetuated, Rafinesque himself renamed a species of
Jacaranda (Bignoniaceae) as Rafinesquia!

Period of 1840-1880
Two American potanists tower over all others for this period: John
Torrey and Asa Gray. Each was trained as a physician, but Gray did
not practice the profession. This period was one of great exploration
throughout the central and western parts of North America and ooe
that was paralleled by colonial exploration in the Old World by the
British (Australia, India, South Africa). It was the period of rapid
ascendancy of taxonomic work at the Royal Botanic Garden at Kew,
England, and the British Museum, London. At Kew, such men as John
Lindley, William J. Hooker and his son, Sir Joseph D. Hooker, and
George Bentham, made their mark with monographic and floristic
studies. Robert Brown was a leader of similar research at the British
Museum. The personalities and accomplishments of these British bot-
anists provided a background of influence to their American counter-
parts. Exchanges of ideas, publications, and specimens were a continu-
ing process.
John Torrey (1796-1873), state geologist for New Jersey, chemist,
Professor of Medicine at Columbia University for thirty years, and
botanist outstanding, published two floras, e.ach of two volumes, on the
higher plants of eastern United States. One was A Flora of the State of
New York and the other, in collaboration with Asa Gray, A Flora of
North America (1838-43). Torrey was a critical taxonomist, more
thoroughly trained in science than any American predecessor. He rec-
ognized the value of type specimens, studied them in European as well
as in American herbaria, and reviewed the literature with meticulous
care. He early discarded the Linnaean system for that by de Candolle,
was one of the first to provide ample descriptions of accepted taxa,
supplementing them often with ecological notes, and keys as aids to
identification and diagnosis. Torrey was the dominant figure in Amer-
ican botany. He was a man whose views and help were sought by others
working with the lower groups as well as the fel]l$ ""d seed plant•. His
name is commemorated by the genus Torreytl (Taxaceae), so named by
Arnott of Scotland.
110 INTRODUCTION TO PLANT TAXONOMY

Asa Gray (1810-1888) born near Utica, New York, was a preco-
cious student, trained as a physician but more interested in plants than
in medicine. While a student he corresponded with Amos Eaton, Rens-
selaer Polytechnic Institute, Troy, New York (author of Manual of
Botany for the Northern States, 1817). Eaton referred him to Dr.
Torrey whom Gray visited in 1832. As a result of this visit, Gray ac-
cepted Torrey's invitation to be joint author in the preparation of the
Flora of North America. From 1840 to 1880, Gray was Fisher Profes-
sor of Natural History at Harvard. Here he specialized in taxonomic
studies on the plants of eastern North America, of western North
America, and of Japan. Evidence of the quality of his work is found in
the large number of the many species described by him as new and
which are currently accepted both as to name and to rank. Gray was
also an outstanding teacher, and his books First Lessons with Plants
and Structural Botany continue to be consulted, although long since
out of print. In 1880 he commenced publication of his Synaptical
Flora of North America, in which he proposed to include, with synop-
tical keys, descriptions of every species of flowering plant then known
for the region. Gray died in 1888, before the completion of this work.
The genus Grayia (Chenopodiaceae) of the southwestern part of the
United States commemorates his name.
With his death in 1888 Harvard's position as the dominant American
center of taxonomic thought came to an end. During Gray's (and thus
Harvard's) era of dominance, this country was expanding and students
of Gray and others were establishing new centers of taxonomic activity.
These successors, in point of time, may be treated as representing two
basic schools of thought: the Harvard School and the New York School.

The Harvard School, 1886-

Asa Gray was influenced considerably by Sir Joseph D. Hooker and
other British taxonomists. Their views on classification, nomenclature,
and concepts of classificatory units (especially of species and genera)
were accepted in principle by him. Many students profited from Gray's
teachings, and other young botanists worked in association with him if
not as students, e.g., Charles E. Bessey. These colleagues of a succeed-
ing generation continued the Grayian ideas.
At Harvard, Asa Gray had Sereno Watson (1826-1892) first as his
assistant (in 1873) and later as curator of the herbarium (from 1874
to Watson's death). Watson, a graduate of Yale, was in California in
Ta.yonomy in Nor,h America 11
1867 and there joined the King;exploring expedition. It was the exce
lence of his specimens and of his five-hundred-page report, Botany (
the King Expedition, that brought Watson to Gray's attention. This n
port by Watson became in effect the first flora of the Great Basin. Hi
two-volume Botany of California (1876-80) stood for several decade
as the most complete flora of that region, and his Bibliographical Inde.
to North American Botany (1878), a never-completed labor of love
was another "first" by Watson. With John M. Coulter he prepared th,
revised fifth edition of Gray's Manual of Botany. Watson w~s a quie
retiring man of dignity, a rapid worker, but not one to attract students
Benjamin Lincoln Robinson (1864-1935), a graduate of Williams
Harvard, and Strassburg, became an assistant to Watson in 1890 an(
succeeded-{)n the death of Watson-to the post of curator of the GraJ
Herbarium in 1892. Robinson "inherited" an institution wealthy ir
specimens and books but lacking endowments, operating funds, or ade·
quate housing. His was the responsibility of acquiring financial il!de·
pendence for the Gray Herbarium and for designing and financing the
fireproof building that held its collections until 1954. During this time
he worked over the collections and wrote the Flora of the Galapagoo
Islands (1902) and, with Fernald. prepared the entirely rewritten seventh
edition of Gray's Manual of Botany (1908). Robinson's taxonomic
work is best represented by his many papers on members of the Com-
positae. Unlike his predecessors and successors, he was not a field
botanist and was known to tell persons bringing in a fresh specimen for
name, to "press it, dry it, bring it back, and I will name it for you."
Weatherby has written of him, "His balancing and restraining influence
at a time of considerable, and too often ill-considered, innovation was
not the least of his contributions to taxonomy." With other duties, he
was also editor for over thirty years, of the New England Botanical
Club's publication, Rhodora.
A colleague of Robinson at HarVard was George Lincoln Goodale
(1839-1923), a physician by training, brought to Harvard in 1872 by
Gray as instructor in botany, and successor to Gray in 1888 as Fisher
:'rofessor of Natural History. Goodale was curator of the Botanical
Museum and of the Botanical Garden from 1879-1909 and, more than
anyone else, was responsible for the initial development of economic
botany at Harvard. Although first s taxonomist, his interests became
stronger in morphology and physiology.
Successor to Robinson as director of the Gray Herbarium and to
 112 INTRODUCTION TO PLANT TAXONOMY

Goodale as Fisher Professor was Merritt Lyndon Fernald (1873-1950).

Born at Orono, Maine, educated there and at Harvard, he became an
assistant to Sereno Watson in 1891 and remained at the Gray Herbar-
ium. Fernald was a specialist on the fiora and phytogeography of
northeastern North America, and, while a majority of his publications
were floristic in character, they contained many revisions of difficult
genera of that region. His principal monographic work was with the
genus Potamogeton, but his complete revision and publication of the
eighth edition of Gray's Manual of Botany (1950) was his crowning
achjevement.

The New York School, 1895-

The New York school of taxonomic thought radiated from the ac-
complishments and personality of Nathaniel Lord Britton. Where the
Harvard school reflected a European conservatism, a thoroughness im-
bued with caution, and a consideration tempered with respect, the New
York school, unshackled by any heritage of a past or ties with mentors
at other centers, reflected fresh and zestful thinking, recognition of new
areas to conquer, and the determination to be a center of aggressive
leadership.
Nathaniel Lord Britton (1859-1934), a geologist with a degree in
mining engineering from Columbia, was appointed botanist and assist-
ant geologist on the New Jersey Geological Survey. From this introduc-
tion to botany and a natural love for plants he became a member and
later head of the botany staff at Columhia University, New York. He
was the organizer and developer of the New York Botanical Garden,
became its first director 'in 1896, and proved to be a most capable ad-
ministrator. As a taxonomist he was an ardent and energetic field man.
He was author of the Flora of Bermuda (1918); coauthor with Addison
Brown of An [/Iustrated Flora of the Northern States and Canada (three
volumes, 1901); with C. F. Millspaugh of The Bahama Flora (1920);
and, with J. N. Rose, The Cactaceae (four volumes, 1919-23). Britton
was a brilliant person, quick thinking, a rapid worker, and possessed of
a peppery, volatile disposition. The genera Brit/oniastrum (Labiatae)
by Briquet, and Brit/onella (Malpighiaceae) by Rusby, commemorate
his name.
He attracted students and brought to the botanic garden an assem-
blage of keen minds, of rugged individualists, and of whom none was a
"Harvard school" man. Most of these men accepted the Brittonian ideas
Taxonomy in North America 113
of taxonomy. NOlie was noted for taxonomic conservatism, and none of
those now deceased could be called a "Iumper," while some have been
and are considered "splitters." Britton led one school of American
thought on matters of nomenclature, wherein he initiated many innova-
tions that were the backbone of the Rochester and American Codes
(now superseded), and he and his associates sought unsuccessfully to
have them adopted universally. A few of these innovations were ulti-
mately accepted (1930) and incorporated into the international regula-
tions. Britton encouraged his colleagues to "adopt" individually differ-
ent parts of the country for intensive floristic study and to provide floras
and manuals. By this method the ideology of the New York school blan-
keted a substantial part of the country. The initiation, in 1905, of the
serially published North American Flora, edited at first by Britton and
later by his successors, contributed toward this end .. This flora, pro-
jected to occupy thirty-four volumes, is yet far from complete. A major
portion has been written by Britton and staff members of the New York
Botanical Garden. Some of these colleagues of Britton deserve mention
for the siguificance of their taxonomic work.
Per Axel Rydberg (1860-1931) came from Sweden to this country to
study mining engineering, but a partially crippling accident in Michigan
ended these plans and he became a mathematics teacher in Nebraska.
While there he spent his summers as a botanical exporer for the U.S.
Department of Agriculture, a work that led to his Flora of the Sand
Hills of Nebraska (1895). This encouraged him to go to Columbia,
where his Ph.D. thesis was A Monograph of the North American Poten-
tillae (1898). On completion he became a member of the New York
Botanical Garden staff, remaining there until his death. During this
thirty-year period he was an herbarium botanist, engaging in little or no
field work in the areas of whose plants he published. His important and
valued works included much of the Rosaceae for the North American
Flora, the Flora of Colorado (1906), Flora of the Rocky Mountains
(1917; ed. 2, 1922), and Flora of the Prairies and Plains (published
posthumously, 1932). Rydberg was extremely liberal in his concepts
of genera and species, describing during his lifetime over 100 genera and
1,700 species as new. E. L. Greene sought to commemorate his name
by the genus Rydbergill (Compositae), but by most authors this genus
is now merged in Actinella.
Marshall Avery Howe (1867-1936), a Vermonter with his Ph.D.
from Columbia in 1898 and a member of the garden staff from 1901 to
114 INTRODUCTION TO PLAlVT TAXONOMY

his death, was a leading mycologist, who carried into cryptogamic bot-
any the Brittonian views applied by other members of the staff to the
vascular plants. He succeeded E. D. Merrill as director of the New York
Botanical Garden.
John Kunkel SmaIl (1869-1938), whose Ph.D. thesis from Columbia
in 1895 resulted in A Monograph of the North American Species of Po-
lygonum. contributed significantly to the Britton program of expansion
by his lifetime work as a specialist on the flora of southeastern United
States. This was an area that Small knew well, especially Florida. His
floras included the Flora of the Southeastern States (1930; ed. 2, 1913)
and the Manual of Southeastern Flora (1932).
John Hendley Barnhart (1871-1951) had a medical degree from
Columbia but from 1903 to his retirement was a staff member of the
Garden, serving variously as editor, librarian, and bibliographer. His
proficiency in these fields was of international repute, and he was largely
responsible for the early development of the library. Barnhart was a
competent taxonomist and the study of the Lentibulariaceae was his
specialty. He was an ardent proponent and supporter of Britton's nomen-
clatural views and defended them at every opportunity, including every
international botanical congress from that in Vienna in 1905 to that in
Cambridge, England, in 1935.
The exponents of the New York school include also Professor LeRoy
Abrams (born 1874) who received his Ph.D. from Columbia in 1910
and soon afterwards became established at Stanford University.
Abrams' major contribution, aside from that of teacher, has been "An
Illustrated Flora of the Pacific States, Washington. Oregon and Cali-
fornia," an incomplete work of which the first three of four volumes
have been published.
Other American Taxonomic Centen
Botany in the southeast, after the time of Walter, was best repre-
sented by the floristic work of Alvan Wentworth Chapman (1809-
1899), a graduate of Amherst, who settled in Georgia and later moved
to western Florida. Chapman, a physician, was long a devotee and cor-
respondent of both Torrey and Gray. His Flora of the Southern States
( 1860) went through three editions and teflected his acceptance of
Gray's species concepts and was patterned after Gray's Manual. Chap-
man, color-blind to reds, frequently described pink-flowered species as
having white flowers and not a few of his alba's and albif/ora's have pink
Tcrxonomr in Nord. A....rlca 115
to red flowers. The genus Chapmannja of Leguminosae was named in
his honor by Torrey and Gray.
The Missouri Botanical Garden, St. Louis, established in 1859 by
Henry Shaw, ardent horticulturist and philanthropist, was later endowed
by him. It operates in close cooperation with the Henry Shaw School of
Botany of Washington University, SI. Louis, with members of the
scientific staff of the garden serving on the school faculty. The garden
has been affiliated with the university since 1899, when William Trelease
became its first director. Although never a member of its staff, George
Engelmann (1809-1884) long a friend of Henry Shaw, was a dominant
and central figure in early SI. Louis botanical activity. Engelmann emi-
grated here from Germany in 1832, and, although a noted physician, he
was also an intimate and life-long friend of Asa Gray and an indisputed
authority on the taxonomy of the cacti, conifers, gentians, yuccas and
agaves, North American grapes, dodders, euphorbias, and evening prim-
roses. Plants of these taxa were sent to him for naming from sponsors
and botanists of most North American explorations. His large herbarium
of Over 100,000 specimens was given by his son to the Missouri Botan·
ical Garden.
William Trelease (1857-1945), a graduate of Cornell and later a
student of Farlow (mycology) and Gray at Harvard, was the first Gray
professor of botany at the Shaw School. He retired in 1912 to become
professor of botany and head of that department at the University of
Illinois (1913-26). Trelease, a taxonomist of liberal species concepts,
is best known for The Genus Phoradendron (1916) and The American
Oaks (1925). His small book Winter Botany (1918) deserves to be
better kn\)wn for its excellent keys to a wide range of woody plants in
winter condition. The genus Treleasia (Commelinaceae) named in his
honor by J. N. Rose is now generally included in Setcreasia.
Jesse Moore Greenman (1867-1951), a graduate of the University of
Pennsylvania (1893) and of Harvard (1899), was an assistant there to
Robinson and a colleague of Fernald. He left Harvard to study under
Adolf Engler at the University of Berlin, where he received his Ph.D.
degree in 1901. After returning to the Gray Herbarium, where he hoped
to remain but was bypassed for promotion in favor of Fernald, he went
in 1905 to the Field Columbian Museum of Chicago. He left there for
SI. Louis in 1913 to become curator of the herbarium under Dr. George
T. Moore, also a Harvard man. Greenman was a scholarly man, genteel,
quiet, and outstanding as a teacher and developer of taxonomists.
1I6 INTRODUCTION TO PLANT T AXONOMr
Taxonomically, he was more of a monographer than a floristic specialist,
although he did considerable work on Mexican collections in his earlier
years. The genus Senecio was his special interest, although the Com-
positae as a whole held much appeal for him. Greenmania, named for
him, is a genus of the Compositae.
Taxonomy on the Pacific coast centered in California where the early
leaders included E. L. Greene and W. L. Jepson. Prior to the entrance
of either into California, the fust flora of the region was VoIoey Rattan's
A Popular California Flora (1879; ed. 2, 1880; ed. 3, 1882). Rattan, a
California school teacher and correspondent of Asa Gray, was more of
an explorer than botanist.
Edward L. Greene (1893-1915), a native of Rhode Island, educated
in Wisconsin and Illinois and ordained an Episcopal minister in Col-
orado (1873), was active as a taxonomic botanist for half a century.
He was a controversial, fearless figure, an egocentric by nature, quarrel-
some, and one of America's greatest "splitters" among botanists. Greene
was the fust professor of botany at the University of California. He was
an ardent field man and knew Californian plants better than his predeces·
sors. He was a reformist in nomenclatural matters, antedating Britton
and the New York school in his, insistence on absolute priority of names
and epithets. While in California he published the periodical Pittonia
(1887-1905), Flora Frandscana (1891-97, incomplete), and a Man-
ualof the Botany of the Region of San Francisco Bay (1894). In 1885
Greene declared invalid his ordination as an Episcopal clergyman and
became a Roman Catholic layman. Ten years later he left California to
become professor of botany at the Catholic University of America,
Washington, D.C., remaining until 1904 when he went to the Smith-
sonian Institution. In 1914 he transferred his library and herbarium t<>
Notre Dame University, Greene considered species to be immutable and
this philosophy was largely responsible for his having described over
3,000 plants as new species. In early middle-life he feuded openly with
Asa Gray, insisting on the right of a botanist to publish his results with-
out fust submitting a manuscript or specimen to Gray for approval.
Despite this, Gray named the Compositae genus Greenella in his honor.
Willis Linn Jepson (1867-1946), a student of Greene, liked to think
of himself as the Hooker of California botany; this from his high esteem
of Sir Joseph D. Hooker of Kew. Jepson was a conservative taxonomist,
a man who knew California flora from field experience, and whose train-
ing under E. L. Greene instilled in him the recogniti<>n of the importance
 Taxonomy in North A.merica 117
of knowing plants in their native habitats as well as in the herbarium.
Unlike Greene, however, Jepson recognized the importance of popula-
tions, accepted the principles of organic evolution, and respected these
dicta in arriving at judgments regarding species limits. Jepson was the
first Pacific coast botanist to accept the Engler system of classification,
but persisted in following the British in adhering to the English system
<)f measurement. His best known publications are probably A Manual
'of Flowering Plants of California (1923-25) and A Flora of California
(1909-43, incomplete). Jepson never married and was a rugged indi-
vidualist, who, in his later years, became somewhat anti-social and often
difficult to approach. He became a person of very strong likes and dis-
likes, and often only those persons in his favor had access to his collec-
tions and benefit of his counsel. The genus Jepsonia, named for him by
J. K. Small, is a member of the Saxifragaceae.
The National Herbarium, now at the Smithsonian Institution, Wash-
ington, D.C., had iits beginnings with collections from various American
expeditions and explorations to the far West. Its first botanist was George
Vasey (1822-1893), who emigrated as a youth from England to up-
state New York. Later he moved to Illinois where he was a practicing
physician for many years. Independent means came to him by his sec-
ond marriage and enabled him to participate in an exploring expedition
to Colorado, which led to his becoming Botanist of the U. S. National
Herbarium in 1872. Primarily an agrostologist, Vasey published many
works on the grasses, among which are Agricultural Grasses of the
United States (1884) and his Monograph of the Grasses of the United
States and British America (1892). Successor to Vasey was Frederick
Vernon Coville (1867-1937), graduate of Cornell (1887) and botanist
On Branner's Geological Survey of Arkansas (1888). Coville was in
many ways more of an administrator than a taxonomist, his principal
contribution in the latter field being his Botany of the Death Va!ley Ex-
pedition (1893). He was important in building up the staff of the Na-
tional Herbarium and in encouraging and championing taxonomic ac-
tivity. William R. Maxon (1877-1948) succeeded Coville as head of
the National Herbarium; he was a graduate of Syracuse University and
after graduation studied under L. M. Underwood (noted authority on
American ferns). From 1898 to 1946 Maxon was curator of plants at
the U. S. National Museum, where he developed the finest fern herbar-
ium in the country, one which contains about 150,000 specimens.
Maxon was a pteridologist who had nO interest in nomenclatural squab-
118 INTRODUCTION TO PLANT TAXONOMY
bles nor in phylogenetic hypothesizing. On several occasions he studied
fern types in European herbaria and confined his researches to floristic
studies of the ferns and to careful revisions of their local groups. The
bibliography of his writings includes Over two hundred titles on ferns.
The genus Maxonia (Polypodiaceae) commemorates his name.
There are many persons who have been omitted from this brief re-
view and, with one exception, those included are not now living. Stu-
dents desiring to study the subject further should become familiar with
the following biographical works by Andrew Denny Rodgers III: Noble
Fellow, William Starling Sul/ivant (1940); American Botany, 1873-
1892 (Asa Gray and contemporaries [1944]); John Merle Coulter,
missionary in science (1944); Liberty Hyde Bailey: a story 0/ American
Plant Sciences (1949); Bernhard Eduard Fernow, a story 0/ North
American Forestry (1951); Erwin Frink Smith; a story 0/ North Amer-
ican Plant Pathology (1952).
CHAPTER IX. Important Families
-and Their Characters

AS INDICATED in a previous chapter, the process of plant identifica-

tion involves placing the "unknown" in successively smaller units of
classification. The characteristics of the larger of these units are usually
more obvious than those of the smaller. Obviously, it requires little ex-
perience to distinguish a fern from a seed-plant, a conifer from a flower-
ing plant, or a lily (a member of the Monocotyledoneae) from a butter-
cu)' (a member of the Dicotyledoneae). The person seeking further
identification of an "unknown," usually with the aid of a flora Or man-
ual, finds ft necessary to "key the specimen out" to the family, then to
the genus, and ultimately to the species to which it belongs. In most
publications, the key to the family is the most difficult to use because
the characters of family differentiation often are highly technical, minute
in size, Or are to be determined only by careful and critical dissection of
the reproductive structures. Furthermore, the key to the families repre-
sented in a particular flora Or manual must be complete and include
every family known to be represented in the area treated. Invariably,
this means the inclusion of many families that are rare or infrequent,
and the more families included the more difficult and complicated the
key becomes.
Experience has shown the advantage of being able to recognize on
sight the families to which belong the common plants of an area. Know-
ing the family, one step in the procedure of identification is by-passed.
Some families are very easily "spotted," and the amateur and beginner
should have little difficulty in knowing the distinguishing characters of
about fifty of the families of fems and seed plants encountered in most
JIll
no INTRODUCTION TO PLANT TAXONOMY
parts of the United States and Canada. It is here estimated <hat fully
tbree-fOlJI1!ls of the plants composing those groups known as spring
40wers, wild flowers, and woodland trees are to be found in the families
that are treated here briefly. Technical terms are avoided when possible.
The effort is made to focus attention on the priJ!cipal distinguishing
characters and on the special terminology common to members of these
families.
The morphology and terminolOgy peculiar to the ferns have been
treated in Chapter N (p. 27). By conservative concepts, most of our
native terrestrial ferns belong to the Polypodiaceae. These ferns, for the
most part, are clump-forming plants or produce creeping rhizomes or
stolons, and their leaves seemingly arise from the ground. Other families
discussed previously include the Selaginellaceae (Fig. 5) to which be-
long the club-mosses and Iycopodiums, the Isoetaceae (Fig. 6) known
also as quillworts, and the aquatic !loating ferns of the Salviniaceae
(Fig. 7). In addition, there are the MarsUeaceae or water-clovers, which
grow in sballow water and bave leaves of four leallets that resemble
those of oxalis or a four-leaf clover; bere the sporangia are in sori tbat
are contained within a hard pea-sbaped sporocarp situated near the base
of the leaf petiole. The filmy-fern family, Scbizaeaceae, produce pear-
shaped sporangia with an apical annulus (Fig. 8). In the Osmundaceae,
to which belong the Cinnamon, Royal, and Interrupted ferns, the
sporangium do not have the incomplete annulus characteristic of the
Polypodiaceae (Fig. 18 f) but has a small lateral annulus not readily
apparent wben examined with a hand lens (Fig. 18 i).
The gymnosperms, like the ferns, were accounted for in Chapter IV
(p. 31) and are not treated bere. The principal families of North Amer-
ica and their characteristics are to be sought in the previous account,
and for further details see Lawrence Taxonomy of Vascular Plants
(pp. 355-370). The pages that follow provide brief synopses of the
distinguishing characters of the more important families of Angiosper-
mae, the !lowering plants. These are composed of two subclasses, the
Monocotyledoneae and the Dicotyledoneae.

THE MONOCOTYLEDONEAE
For brevity the members of this group often are termed monocots and
those of the Dicotyledoneae, dicots. The monocots are not distinguisheo
by any single character but the following characters in combination,
with certain exceptions, serve to characterize the group:
1m""...."' Famllie. and T1IeIr CIoanJ..... 1Z1
Stems with vascular suands scattered through a large pit11;
Rootstocks often rhizomatous, bulbous, or cormous;
Leaves with parallel venation; and
Flowers three-merous.
The more dominant families include the following:
Cyperaceae Graminea. Iridaceae
Araceae Liliaceae Orchidaceae
Amaryllidaceae
The Gramlneae-Grau Famlly (Fig. 44)
Grass stems are often called culms, and are jointed at the nodes. The
leaves are arranged in a \>2 phyllotaxy and are composed of three parts,
the sheath, a ligule (Fig. 44 c) and a blade. The llowers are tiny, much

Fla. 44. G1<A~. !'hl.urn pratl!1lSe: a, ha'olt at plattt (cu\ms 'oent); '0.
spic:ate inftorescence; c, nodular portion of leaf. showing ligule; d. spikelet; e,
same, "exploded" tQ show orientation of organs; f, fruit oontaining seed.

reduced by loss of the perianth (no petals or sepals are present), and
are much congested into spikelets (Fig. 44 d). The grass llower is often
termed a floret. The lloret may be bisexual or unisexual, and typically it
is composed of three stamens (six in rice and most bamboos) and a
superior one-celled ovary with a single ovule, terminated by two plumose
styles or style-branches. The spikelet may contain one to many llorets.
Usually the florets are so short-stalked as to appear sessile. In a one-
llowered spikelet (see Fig. 44 e) there are two dry bracts called glumes.
Above these, and often of thinner texture and enveloping the lloret, are
Jft INTRODUCTION TO PL.4NT T..aONOMf
two more bracts, the lower is the lemmtJ and the upper the palt!4. In Fig.
44 e the relative position of these is shown in an expanded (or ex-
ploded) diagram, the dotted lines show the axis as if elongated. Actually,
it is very short and the spikelet is much condensed as in Fig. 44 d. The
fruit is a modified achene, termed a caryopsls and differing from an
achene in the coat being loose from the seed.

Cyperaeea-Sedge F8JDlI,.
Vegetatively this family is distinguished from the grasses in the stems
usually solid and triangular in cross-section, the leaves arranged in a
y., phyUotaxy, and in the absence (usually) of any ligule. The liowers
are bisexual or unisexual and arranged in spikelets that are much more
simple than the grass spikelet for here the liower is subtended by one
or two chaffy bracts, not differentiated into glumes, lemma, and palea
as in grasses. A perianth represented by bristles or scales is frequently
present. The fruit is an achene. In the large genus Carex, the ovary is
enclosed in a thin sac termed a per/gymum which persists in fruit.

Araeeae-Arum F8JDlI,. (Fig. 45)

Members of this family are commonly referred to as aroids. The
family is primarily tropical, but its range extends 'northward to the

FJi. 45. AlL\CEAB. ArisQema triphyllum: a, plant in JIowcrj b, pistillate spadix;

pistillate flowers; d, staminate flowers (staminate spadix from another plant not
shown).
arctiC. The leaf venation may be reticulate, pinnate, or parallel. Vegeta-
tive portions often contain bundles of crystals of calcium oxalate
(raphides) that impart a pungent "!lavor" and may be highly irritating
to the mouth if eaten uncooked. Members of the family are best dis-
tinguished by the spike-like inliorescence, termed a spadix, that is sub-
Importa,., l'amUiu alUl' Tlurir Claaradar. 12$

tended or enveloped by a fleshy or herbaceous spathe (Fig. S6 a). The

spadix may be fertile throughout or the flowers restricted to the basal
portion and the distal part is then sterile (Fig. 45 b). The tlowers may
be bisexual or unisexual, with or without a perianth of four to six
minute scale-like tepals not differentiated into petals or sepals. The
ovary is superior, one- to many-Ioculed, and the plac.entation basal,
parietal, axile, or pendulous. The stigma is usually· capitate. The fruit
is a berry.

LWaeeae-lJIy Famlly (Fig. 46)

The leaves are mostly parallel-veined, the flowers typically ani of
six tepals, six stamens, and a single tricarpellate pistil whose ovary is
usually superior, three-IocuJed, with aJIile placentation (Fig. 46 b, c).
The stigma is generally three-lobed or branched (Fig. 46 d) and a style

~
JI'Ie. oM. LD...u.CEAE. Lilium canadense:
~, c
~ inflorescence; b, flower. vertical sec·
tion; c, ovary, cross«:c:tion; d, stigmaS, each. bilobed; e, versatile anther (appc:at'-
in, as if basifixed in b, before anthesis).

is usually present. The inflorescence type is variable and not diagnostic.

The fruit may be a berry or a capsule. In most. members ot the family
an underground scaly bulb is present.
AmaryWda _ _Amaryrn. Famlly (Fig. 47)
Opinion is divided on the bases for distinguishing this family from the
Liliaceae. By older views, and followed in most contemporary (IorfU,
those plants whose flowers produced a superior ovary were placed in the
Liliaceae, and those with an inferior ovary were placed in the Amaryl-
1idaceae. The number of genera representing intermediates between these
two situations invalidate the distinction.
More recently, data from studies of morphology, anatomy, embryol-
lJ4 INTRODUCTION TO PLANT TAXONOMY
ogy, and karyology have supported the view that a better criterion for
distinction of AmaryJlidaeeae from Liliaeeae is provided by the infiores-
eenee. By this view, those genera whose inflorescence is an umbel sub-
tended by one or mOre papery spathes or bracts are placed in the

Ftc. 47. AMARYLUDACEAE. A. Hymenocallis narcissif/ora: Aa, plant in flower,

with bulb; Ab, flower, fllCe view; Ac. S~, vertical section; Ad, periantb section,
showing corona; Ae, style tip, with three-lobed stigma; Af, ovary, vertical section;
AI. ovary, cross-section. B, Zephyranthes grandi/iora: Ba, plant in flower, Jess
bulb; Bb, flower, habit; Be, same, vertical section; Bd, style tip and stigma; Be,
ovary, vertical section; Bf. ovary, cross-section. C, Cooperia pedunculata: Ca,
plant in llower; Ch, bulb; Cc, flower. habit, with spathe valve; Cd, same, vertical
section; Ce, ovary, cross-section. (c, corona; p.i., perianth segment.)

Amaryllidaceae. This results in the transfer to this family from the

Liliaeeae of some genera with a superior ovary, as Allium. Agapanthus.
and Brodiaea. In using this criterion for separation. it must be remem-
bered that in advanced situations the umbel may be reduced to a single
flower (as in some species of Narcissus). but even so, that amaryl-
lidaceous flower is subtendcd by one or more papery spathe-like bracts.
._. in Liliaceae, the stamens are six in number.
Impor'an' Farrailie. and Their Charader. 125
In some members of the family a COrona may be present. This may
be a part of the perianth (Fig. 42 Ad), such as the cup of Narcissus, or
it may be a thin tissue coooecting the stamen filaments.
Irldaceae--Irla FamUy (Fig. 48)
This is a distinctive family, whose members have flowers with a
trilocular inferior ovary with axile placentation, and three stamens. The
rootstock is more commouly rhizomatous than in the lily or amaryllis

F1a. 48. IRmACEAE. lrLr germanica: a, flower with subtendin, spathe-valves; b,

same, spatho-valves removedj c, same. less perianth; d, style-branch and stigma;
e, ovary croas section.

family, although bulbous and cormous members occur. In the iris tribe
the style branches are winged and petaloid (Fig. 48 c), and each of
the three stigmas is reduced to a transverse line on the lower (dorsal)
side (Fig. 48 d).
Orchldaceae--Orchid FamUy (Fig. 49)
This is a dislinctive and highly complex family of monocots that is
characterized by the flowers zygomorphic, the perianth in two series
with the outer composed of two to three green sepals and the inner of
three petals, one of which is enlarged and highly modified into a seg-
ment termed a labellum (Fig. 49 Ac). The ovary is inferior, usually
resupinate (twisted 180 0 , as in Fig. 49 Bb), one- to three-loculed with
parietal or axile placentation, and the ovules abundant but exceedingly
minute. Two subfamilies are present: Diandrae (Fig. 49 A) and Monan-
drae (Fig. 49 B). In each the androecium and style and stigmas are
fused to form a column or gynandrium. In the Diandrae, there are two
lZ6 INTRODUCTION TO l'L4JIIT T AXONOMJf
functional sessile anthers (Fig. 49 Af) producing granular tetrads of
pollen; and a third nonfunctional anther consists of a conspicuous
glandular staminode (Fig. 49 AF, Ag). In this subfamily there are
three stigma lobes, situated beneath the staminode. In the Mooandrae,

fII8. 49. ORCHlDACEAl!. A, Cyprlpedium Calc.olus: Aa, plant in ftower; Ab,

lower, habit; Ac, lame, vertical section; Ad, gynandrium, vertical section; ~,
ovary, cross section; Al, column, viewed from below; AI. same, side view. B,
HabelUJrla: Ba, plant in flower; Bb. flower, habit; Be, gynandrium. vertical section;
Bd, coltJIDn. viewed from below; Be, same, side \liewj Bf, pollinium..

there is a single terminal anther, whose pollen grains are in tetrads that
are generally agglutinated into fum to bony bodies termed pol/inio
(Fig. 49 Bf). These pollinia are generally attached to the apical ends
of the anth"" sac by slender stalks called caudicles. The number of
pollinia in an anther may be two, four, or eight depending on the genus.
In this subfamily the stigma is situated in a usually depressed cavity
below and in back of the anther. For a more detailed discussion of the
Botal morphology, see Lawrence, Taxonomy of Vascular Plants, pp.
433-437.
 127

Dleotyledolleae
The dicots may be diatinguished from the monocots by the combina-
tion of the following characters, recognizing that exceptions may occur
in one or some of those cited:
1. Stem with a true cambium cylinder, the pith usually small and no
vascular strands scattered through it;
2. Roots usually fibrous; no scaly bulbs produced;
3. Leaves mostly pinnately or palmately veined; and
4. Flowers not commonly three-merous throughout, although the
pistil is not infrequently tricarpellate.

The catkin-producing families are characterized by the flowers of

one or both sexes in flexuous aments or catkins. Here are included the
(mostly) unrelated families Salicaceae, Juglandll<\Cae, Betulaceae, and
Fagaceae.
SalIeaoea_Willow Famlly
The willows are dioecious trees or shrubs with the flowers of both
sexes In flexuous catkins. Each flower is subtended by a finger- or cup-
like gland, t1ie ovary is superior, two-carpelled, unilocular, and the
placentation is parietal. The stigmas are two in number, and each may
be lobed or fringed. The fruit is a capsule. The seeds are comose.
JuglaDdaeea_Walnut or Blekory Famlly
.This is a family of monoecious trees or shrubs with pinnately com-
pound leaves, the staminate flowers in catkins, but the pistillate flo ....rs
otten not so. The stamens are furee to one 'nundred in number, wi\'n
erect basifixed anthers. The ovary is inferior, mostly two-carpelled, and
contains a solitary ovule. The styles are usually two. The fruit. is a
dehiscent or indehiscent drupe-like nut.
Betulaceae (Corylaeeael-Blreh Family (Fig. SOl
These are monoecious trees or shrubs with simple leaves. The stami-
illite flowers, and frequently the pistillate, are in catkins. The flowers are
in much condensed cymules, with one to three bracts (sometimes fused
and then seemingly three-lobed). The staminate flowers contain many
stamens. The pistillate flower has an inferior or nude (position not then
identifiable) ovary, subtended by one large and two small bracts, which
lU INTRODUCTION TO PUNT T..aONOM~
is two-loculed and two-carpelled with basal placentation. The style is
ODe or two. The fruit is a small nut or short-winged samara.

..... so. BBT'Ul..A.CEAB. Betula peru/uta: I, flowering branchj b, staminate

.ftowersi c.. pistillate Bowers; d, schematic diagram of theoretical throe-flowered
pistillate cym.ule believed ancestral to modem types; e,. schematic diagram of
pistillate cymule of Betula Medewiewii; f, "ftoral" diagrams of inflorescence of
hypothoticol cymule shown in "dO; .. pistillate cymule, B. M.d.wl.wlt. (d·, re-
drawn lnIni Abbe, 1935.)

F..,.eeae-Beeeb Family
A family of mostly monoecious trees and shrubs, the beecb's stami-
nate Bowers are usually in pendulous catkins, and the pi.tillate ones are
solitary or clustered. The pistillate Bower is mostly within an involucre
of generally many adnate and imbricated bracteoles. The ovary is in-
ferior, three- to six-Ioculed and carpelled, the placentation axile, and the
styles as many as the locules. The perianth is composed of four to six
tepals. The fruit is a one-seeded nut, subtended or enveloped by a
cupule or involucre (sometimes two to five within the involucre).

A group of unilocular one-ovuled families mostly lacking a showy

perianth is composed of the Polygonaceae, Cbenopodiaceae, and Ama-
ranthaceae.

Polygon.ceae-Buckwheat Family
The stems usually have swollen nodes and the leaves are subtended
by a sheathing stipular growth called tbe ochrea (absent in tbe tribe _.
Eriogoneae, in which the flowers are in involucrate heads or umbels).
The !lowers are usually bisexual with perianth of three to six distinct
I",porlan' FomiliQ _Gnd T/u!ir Claarttder. 1:9
tepals in two whorls, with six to nine stamens and a superior com-
pressed or three-angled ovary which is unilocular with a basal ovule and
is terminated by two to four stigmas. The fruit is an achene whose
seed has a curved or ,-shaped embryo.
Chenopodlaeeae--Goooefoot FamUy (Fig. 51)
This is a family of mostly salt-loving plants with often fleshy or
granularly farinos .. herbage (Le., herbaceous vegetative parts). The
flowers are mostly bisexual with a perianth of two to five connate .tpals

F1I. Sl. CHENOPODlACEAE. Beta vulgaris: a, garden beet, foliage and root; b,
Bowedng stem; c, flower; d, same, vertical section; e, anther, dorsal side; I, fruit;
I, same, vertical section; h, seed. (c-e after LeMaout and Decaisne.) (From L. H.
Bailey, Manual of cultivated plants, The Macmillan Company, 1949. Copyright
1924 and 1949 by Liberty H. Bailey.)

that usually persist in fruit. No petals are present. The stamens are as
many as the sepals and opposite them. The ovary is mostly superior and
two- to three-carpelled. The fruit is a nutlet, whose seed contains a
coiled or peripheral embryo.
Amaranthaeeae--Amaranth FamUy (Fig. 52)

FIg.. 52. AMARANTHACBAB. A, Amaranthus caudatus; Bowering branch. B,

Gomphrena globosa: ba, flowering branch; Bb, flower with involucre; Be, flower,
vertical section. C, Celosia argentea var. cristata: flowering branch. (From L. H.
Bailey, Manual of cultivated plants. The Macmillan Company, 1949. Copyright
1924 and 1949 by Liberty H. Bailey.)
1M INTRODUCTION TO PLANT TAXONOMY
rhe herbage is not fleshy nor farinose, but often is red-pigmented ill
part. The flowers are bisexual, and are subtended by a membranous or
scarious persistent bract and two similar bractlets. The perianth consists
of three to five scarious sepals. The stamens are generally five in num-
ber and the filaments are basally to wholly connate. The ovary is su-
perior, two- to three-carpelled, and one-loculed. The fruit is a nutlet,
or rarely a circumscissile capsule.
For the most pari this next group of families have a showy poly-
petalous corolla (the petals falling one by one).
Caryophyllaeea_PInk Famny (Fig. 53)
This is a family of herbs that is characterized by the leaves opposite,
often linear and parallel-veined, and the nodes swollen. The flowers have
a superior, unilocular, three- to five-carpelled ovary whose placentation
is free-centraI (rarely basally trilocular with axile placentation). The
fruit is a capsule.

Fig. 53. Cn.YOPHYLLACEAE. Agrost~mma Githogo: a, ftowering branch; b.

flower. face view; c, ,same, vertical SectiOD; d. ovary, cross-section; e, capsule with
persistent calyx; f. seed.

Ranunculacea_Buttercup Famny (Fig. 54)

These are mostly herbaceous plants, often with acrid sap, but which
are characterized by the stamens many and spirally arranged, and the
gynoecium of three to many spirally arranged pistils. Each pistil con-
sists of a one-loculed, one-carpelled, superior ovary with marginal
(parietal) placentation (Fig. 65 d). The fruit is typically a follicle, but
may be an achene, berry, or capsule.
Im""._ ,.mllIe, """ rltel, c".,_,... 1111

J'Ii. st. ItANuNcut.ACB,I.E. Ranunculu.t sp.: a, flower; b. same, vertical section;

C, pistil; d, ,.,ne, vertical section; e, lame, crosa-sectioD.

Papaveraeea_Poppy Famlly
These are mostly herbs, often with mi1ky latex or colored sap, charac-
terized by a calyx that usually falls when the flower opens, and petals
that generally appear crumpled on opening. The stamens are usually
many, in several whorls. The pistil is solitary, with a superior, unilocular,
multicarpellate ovary whose placentation is parietal. The style is short
or absent and the Stignl8 is radiate or lobed. The fruit is usually a capsule
dehiscing by apical pores or by valves.
eru.lfera_M...tard FamUy (Fig. 55)
these are herbaceoU$ plants, often with stellate hairs, whose flowers
have four distinct sepaIs and four petals (Fig. 55 a). The stamens are
six and are termed tetradynamous, because one pair is longer than the
other two pairs (Fig. 55 c). There is a single pistil whose ovary
is superior, two-loculed, four-<:arpelled, and has parietal placentation
(Fig. 55 e). The ~t)'les ate short or ab...nt and the stigma!< two. The
fruit is a modified capsule, usually dehiscing by two lateral valves (hav-
ing a central septum) and when long and slender is termed a silique
(Fig. 55 j), or when short and squat is termed a silicle (Fig. 55 k). The
arrangement of cotyledons and radicle within the seed provide charac-
ters of taxonomic importance. The cotyledons may be acumbent (Fig.
5S i), incumbent (Fig. SS k), conduplicate (Fig. SS n), or double-
acumbent (Fig. SS q).
Cr...ulacea~rpln~ FamU"
Th~e are herbs or shrubs, with succulent leaves that often are in
rosettes when young. The flowers are distinctive because the sepals,
petals, stamens, and pistils are of the same number (usually four or
five), or the stamens are twice as many as the components of other
162 INTRODUCTION TO PLANT T ,4XONO/llY
whorls. Each pistil usually has a conspicuous gland at base and on the
adaxial side. The ovaries are superior, unilocular, unicarpellate, with
marginal (parietal) placentation, style and stigma one. The fruit is a
follicle.

FIg. 55. CRuCIPBIlAE. Thlaspi arvense: a, flower. face view; b, same, side view;
c, same, perianth removed; d, ovary, vertical section; e, same, cross-section; f.
silicIc, side view; g, same, cros!I:-section; h, seed; i, same, cross-section. Hesperu
matrcnalis: j, silique. dehiscing; k, seed. cross-section. Alyssum saxatile: I, winged
seed; m, same, cross-section. Brassica arvensis: n, seed; 0, same, cross-section.
Cakile maritima: P. seed; q. same, cross-section. (Redrawn from L. H. Bailey.
Manual of cultivated plants, The Macmillan Company. 1949.)

Saxifragaceae--Saxifrage Family
As conservatively defined in most floras, this is a difficult family to
distinguish. There is no single field character by which it can be sepa-
rated from some members of the Rosaceae. It differs from that family
in the more abundant endosperm of the seeds, and in its estipufate or
rarely stipulate leaves. From the Crassulaceae, it differs in the pistiho
(when two or more) not subtended by a gland.
 18$

ROlaeeae-Rooe Family (Fig. 56)

This is a large and diverse family whose members have leaves that
are usually stipulate. Its bisexual dowers are of one of three types: (1)

b
'

.
Gb
F1a:. 56. ROSACEAE. A, Spiraea Vanhouttei: Aa, fiowering branch; Ab. flower,
face vic\y; Ac, same, vertical section; Ad, single pistil, vertical section; Ae, cluster
of follicles. B, Prunus sp.: Ba, flower, vertical section: Bb, fruit. C, Malus sylves--
tris: Ca, flower, vertical section; Cb, fruit, vertical section. D, Cotoneastel' horizon-
taUs: D, flower, vertical section. E, Rubus occidentalis: Ea, flower, vertical section;
Eb, fruit, vertical section. F. Rosa canina: Fa, flower, face view; Fb, same, verti-
cal section; Fe, hip; Fd, same, vertical section showing aehenes within. G,
Fragaria chi/oense: Ga. flower, vertical section; Ob, accessory fruit, vertical sec-
tion; Gc, achene. (Redrawn from L. H. Bailey, Manual 01 cultivated plants, The
Macmillan Company, 1949.)
1114 INTRODUCTION TO PLANT TAXONOMY

the gynoecium of few to many pistils, partially to completely enclosed

in a hypanthium and whose ovaries are superior, unilocular, unicarpel-
late, and uniovulate, and the stamens several whorls; (2) the gynoecium
of a single pistil whose ovary is inferior, multicarpellate, multilocu1ar,
with axile placentation and many ovules; and (3) the gynoecium of a
single pistil situated in a hypanthium, the ovary superior, rnulticarpel-
late, unilocUlar with a single parietal ovule. In the first type the fruit is
usually an achene or follicle; in the second a pome; and in the third a
drupe.
Lepminooe-Pea FamUy (Fig. 57)
The family is composed of three subfamilies as distinguished below,
but bonded together by a common type of fruit, the legume, a dry dehis.>
cent fruit that is the product of a simple ovary and which differs from
a follicle in dehiscing by two sutures instead of by one (Fig. 57). In
some members of the family the fruit is a loment, a modified legume
which dehisces by transverse joints (Fig. 40 i). In each subfamily the-
ovary is superior, unilocular, unicarpellate, and its few to several ovules
are arranged in a marginal (parietal) placentation type.

FIe- 57. LEGUMINOSAE. Lalhyrus latifolius: a, flower, face view; b, corolla; c,

flower, less periantb; d, pistil; e, ovary, vertical section; f, legume, dehisced.

Mimosoideae: flowers actinomorpbic, the calyx and corolla valvate in bud.

Caesalpinioideae: corollas zygomorpbic, the po.c;terior petal innennost. the petals
five and distinct.
Lotoideae: the corolla zygomorphic and of the papilionaceous type (Fig. 57),
consisting of a standard (the Qutennost petal), two wing petals, and
two k.eel petals that are basally connate; the stamens are five to ten
in number and are usually diadelpbous (Fig. 57 c) or monadelphous
(Fig. 32 d, ., f).
 IS5

Euphorblae.a--Spurge FamUy (Fig. 58)

This is a family of monoecious or dioecious piants. often with milky
juice. The staminate flowers usually have the stamens as many or twice
as many as petals (or reduced to a single stamen in Euphorbia [Fig.
58 d]). The pistillate flowers have a single pistil whose ovary is superior,
mostly thiee-loculed and three-carpelled with axile placentation and one
to three ovules in each locule (Fig. 58 e. f). There are generally three
simple or bilohed styles. The fruit is usually a capsule. (In Euphorbia
the perianth is missing or reduced to bristles.)

FIg. 58. EUPHORBUCEAE. _ Ihorbia: a, 1Iowering branch; b, cyathium of stam-

inate and pistillate fl.owers; c. same, vertic.a! section; d, staminate flower; e, pistil-
latc flower; f. ovary, cross-scction.

Malvacea_Mallow . Family (Fig. 59)

The sap often is viscid or mucilaginous. The flowers are generally
bisexual, actinomorphic, and have a calyx of five sepals and a corolla
of five petals. Often an involucre of bracts sUbtends the calyx. The
stamens are numerous and monadelphous, with one-celled anthers. The
gynoecium is usually composed of a single pistil having a superior two-
to many-locuJ.ed and carpelled ovary and axile placentation (Fig.
59 Be). Sometimes the carpels are represented by separate pistils (Fig.
59 Cb).

Vlolaceae--Vlolet FamUy
The flowers are bisexual and usually zygomorpbic. with a calyx of
five sepals and corolla of five petals (the lowermost often spurred). The
five stamens are connivent about the pistil and the two lower are often
 INTRODUCTION TO PLANT TAXONOMY
spurred. There is a single pistil, with the ovary superior, one-Ioculed,
three to five-carpelled, and the many ovules are arranged in a parietal
type of placentation (Fig. 3S b).

F'Ia. 59. MALVACEAE. A, Hibiscus palustris: fruiting branch. B. Hibiscus

Moscheutos: Ba, Bower; Db, same, vertical section, perianth partially excised;
Be, ov.vy, cTws-.st!Ction. C. Mawpe 'rijido.' Ca, fuwer; Cb, same, gyDDt:Cium. D,
Gonypium hirsutum: Da, boll; Db, seed. E, Sidalcea candida, style enveloped by
double staminal tube. F. Anoda cristata: Fa. flower, side view; Fb. same, face view;
Fc, partial vertical section. (b. bract; c, corolla; ca, calyx; in, involucre; P. pistil;
s.c., ,taminai column; sty, style.) (From L. H. Bailey. Manual of cultillated plants.
The Macmillan Company, 1949. Copyright 1924 and 1949 by Liberty H. Bailey.)

Ca_eea-caclus Family (Fig. 60)

The plants are mostly succulents, with enlarged fte,by stems, produc-
ing small, often caducous, leaves, and spines (Fig. 60 a). The ftowe~
are mostly bisexual. and actinomorpbic, with the perianth only weak!
differentiated into sepals and petals and fused basally into an bypan-
thium. There is a single pistil, with a usually inferior unilocular, three-
 137
to many-carpelled ovary containing many ovules in a parietal type of
placentation (Fig. 60 b). There is a single style terminated by as many
branches and stigmas as there are carpels. The stamens are very numer-
ous, arising spirally or in clusters, and with two-celled anthers. The fruit
is a berry.

Fla. 60. CACTACEAE. Mam",lllo.ria: a. plant ill flower; b. ftower, vertical section;
c. ovary, cross-section.
Onagraeeae--Evenlngoprlmrooe FamUy (fig. 61)
This family is characterized by the flowers having typically the parts
in multiples of two or four, a single pistil whose ovary is inferior and
multiovulate, with axile placentation, and terminated by an hypantbium
from whose rim emerge the sepals, petals, and stamens.

a
~.
FIe- 61. ONAGRACEAE. Oenothera: 8. plant in flower; b, flower; c. sa.m.e, vertical
section; d, ovary, vertical section; e, ovary, cross-section.

Umbelliferae---<:arrot FamUy (Fig. 62)

The umbellifers are a family composed mostly of herbs, distinguished
by the presence of aromatic oils, the usually sheathing lear bases
 1811 INTRODUCTION TO PLANT TAXONOMY
(petioles), and the typically umbellate inllorescence. In most members,
the inllorescence is a compound umbel (Fig. 62 a) composed of pri-
mary peduncles or stalks termed primtJry rays that bear diminutive
~umbels termed umbellets (Fig. 62 b). The dowers of the umbellet are
. borne on pedicels termed secondlJry rays. The umbel and the umbellet

F1c-,~. UMlIELLlFER.AE: ~ umbel; b, umbellet; c. flower, face view; d, same

vertical section; e, ovary, cross-section; f, scbizocarpj If same, tross-section.

may be subtended by an involucre of bracts and an involucel of bractlets

respectively. The dower has a five-merous perianth of calyx and corolla
with the calyx often caducous. The stamens are five in number. The
ovary is inferior, two-locular and carpelled, each locul. with a single
pendulous ovule (Fig. 62 d). The styles and stigmas are two. The fruit,
a schizocarp, is distinctive for the family. It usually dehiaces into two
equal mericarps, each borne on a wiry carpophore (Fig. 62 f). The ad-
joining faces of a pair of mericarps are termed commissural faces (or
surfaces). The mericarp is often 3-5-7-ribbed and the oil ducts situated
just beneath or between the ribs provide diagnostic characters.

Families with gamopetalous corollas include the following:

Erleaceae-Heath Famlly
This is a family of mostly woody plants, often with persistent leaves.
The dowers have a four- to seven-lobed calyx and corolla, with the
stamens mostly twice as many as corolla-lobes and ,msing from a
 139
glandular disk. The anthers often have caudal appendagei, and dehisce
by apical pores. The ovary may be superior or inferior. It is typically
four- to seven-loculed and carpelled, with axile placentation.
~rlmulacea_Prlmula Family (Fig. 63)
This family of herbaceous plants is similar in some respects to the
Caropbyllaceae, especiall} in its superior ovary with free-central placen-
tation, but differs in the gamopetalous corolla and in the stamens borne
on the corolla-tube and opposite the corolla-lobes. The fruit is a many-
seeded capsule.

}1g. 63. PRIMULACEAE. A, Primula denticulata: Aa, plant in flower; Ab. flower;
Ac, perianth, expanded; Ad, pistil; Ae, ovary, cross-section. B t Trientalis borealis:
Ba, plant in flower; Bb, flower, face view; Be, same, vertical section (perianth
partially excised); Bd, ovary, cross-section. C, Dodecatheon Medio.: Ca, plant in
flower; Cb, flower; Ce, same, vertical section (perianth partially excised).

Apqeynaeeae--Doghane FaD1Uy
Most members of this family produce a milky latex and generally
have o;:>posite decussate leaves. The dowers are actinomorphic, with the
corolla contorted in bud, usually saivedonn or funnelionn, and often is
appendaged within. The pistil consists of two superior ovaries (uniIocu-
140 INTRODUCTION· TO PL.4NT T.4XONOMY
1ar and with marginal placentation) terminated by a single style and
stigma. The stamens are borne on the corolla, alternate with the corolla-
lobes, and produce granular pollen.
Aoclepladaeeae--MlIkweed FamUy (Fig. 64)
A family with many characters similar to those of the Apocynaceae,
but it differs from them in the ovaries terminated by separate styles and
an enlarged single, usually five-lobed massive stigma (Fig. 64 e). Fur-
thermore, the five stamens are usually adoate to the stigma, with the
pollen agglutinated into po/linia united in pairs. Each pollinium bears a
translator arm (or connective) with the two adjoining arms meeting in
a gland-like body (Fig. 64 d). In Asclepias, and some other genera, the
corolla-tube is crowned by a corona that arises from the corolla, and iI
corona-hom represents sterile staminal appendages arising from the
filament or anther. The fruit is a follicle.

FIg. 64. AscLEPIADACEAE. Asclepias curassavica: a, ftower; b. same, vertical

section; c, top of gynandrium, showing five-lobed stigma and anthers (corona
excised); d, pollinia; e. pistil, vcttical section; f, pistil, cross-section through the
two ovaries.

BOl:aginaeeae--Borage FaDlUy (Fig. 65)

The borages are a family of usually scabrous or hispid hairy plants
with alternate leaves, terete stems, and whose inflorescence is a circinate
helicoid cyme or aggregation of same. The flowers are mostly actino-
morphic. The corollas are often with toothlike faucal appendages or
scales (Fig. 65 Ac). The five stamens are borne on the corolla. There is
Impor'.," Tamale, arad Their ChtJrtl(!lBr' 141
a single pistil with a superior ovary that is two-carpelled and bilocular
(but seemingly four-loculed at maturity). The placentation is seemingly
basal but actually axile. The four ovules present in the ovary develop
into four nuUets or a four-seeded drupe.
Verbenaceae--VerbeDa Family
Members of this family usually have opposite leaves and the 1I0wers
are arranged in one or another form of a cyme, but never circinnate nor

Fig. IS. B9RAOINACEAE. A, Syml'hytum asperum: Aa, flowering branch; Ab,

cymule; Ac, flower; Ad, same, vertical section; Ae, corolla expanded; Af, ovary,
habit; Ag, same. vertical section. B, Anchusa azurea: Ba, portion of flowering
branch; Bb, flower; Be, nutlet. C. Echium plantagineum: flowers. D, Heliotropium
arborescens: Da, flowering branch; Db, flower; Dc, pistil; Dd, same. vertical section.
(From L. H. Bailey, Manual of cultivated plants, The MacmiHan Company, 1949.
Copyright 1924 and 1949 by Liberty H. Bailey.)

helicoid. The stamens ar~ commonly four and didynamous. The OTary
is not lobed apically and the style is terminal to it.
141 INTRODlJL"HU/y TO PI.4lVT TAXONOMY

Lablata_Mlnt F&bllly (Fig. 66)

A family predominately of herbs, the mint has four-sided stems, 0p-
posite leaves, and the herbage characteristically aromatic when crushed.
The flowers are in axillary pairs of dichasial or circinnate cymes often
congested and spicate. The flowers are mostly zygomorphic and bila-
biate, the stamens two or four (and then didynamous), and the superior
ovary is deeply four-lobed apically with a gynobasic style (Fig. 66 d).
The fruit usually consists of four nutlets .

.... "- WIATAE. Slllvia splend.,ms: R, flowering branch; b. flower, vertical

section; c, base of pistil;~ same, vertical section; e. fruit of four nutlets. Stochy,
grandlflora: f. flower; g, same, corolla expaoded. (Adapted from L. H. Bailey,
ltIQIUlll/ of cultivated phml•• The Maani.Uan Coo!paoy, 1949.)

Solanaeeao-NlPtohade F&bllly
These are mostly glandular plants whose herbage is variously odifer-
ous when crushed. The inflorescence is cymose and the flowers are usu-
ally actinomorphic with plicate corolla. The androecium is commonly
of four didynamous stamens (a fifth is usually represented by a stam-
inode). The gynoecium consists of a single pistil with a superior usu-
ally bilocular ovary having axile placentation and a single style with •
two-lobed stigma. The fruit is a berry or capsule.
Serophularlaceae--Figwort Family (Fig. 67)
This family has many characters in common with the Solanaceae, but
differs in the corolla not plicate and usually zygomorphic. In some mem-
CmportGRI Fatnllie. and TAeir ClutraCler. 1411
bers the corolla develops posteriorly into a swollen ventricle or spur,
and in some the throat (such as in snapdragon) is closed by a fold
called the paiate.

FIe- 67. ScitOPHULARlACEAE. Linaria vulgaris: a, 1lower, side view; b, same,

vertical sectiOD; c, same, cross-section of ovary. Veronica longifolia: d, habit of
flower.

Bublaeeae--Madder Famll"
These trees and shrubs, infrequently herbs, are characterized by
leaves opposite Or whorled and provided with stipules; the inHorescence
cymose and the Howers generally are actinomorphic (corollas often
salverform). The stamens are as many as corolla-lobes and the pistil
has a usually inferior ovary that is commonly bilocular with axile
placentation.
Caprifoliacea_Boneyeuekle FamUy
This family is so closely allied to the Madder family that the two
have been united by some authors. There is no single character by which
they can be distinguished, except that in general the leaves of the
Capriloliaceae are without stipules.
Cueurbltacea--.Gourd FamUy (Fig. 68)
A family of mostly monoecious or dioecious tendril-bearing herba-
ceous vines, the gourd has leaves which are usually palmately or pin-
nately five-lobed or divided. The staminate Howers are highly variable
as to number and arrangement of stamens. The pistillate Howers have an
inferior ovary of three to five carpels with parietal placentation (some-
times axillary). The fruit is a berry with a hard or leathery rind and is
termed a pepo.
144 INTRODUCTION TO PLAIVT TAXONOMY

FIe. 68. CucUIlBrrAcEAE. A, Cucurbita Pepo var ovilera: branch with fruit and
pistillate flower. B. Cucurbita maxima: Ba. staminate flower, less perianth; Bb.
pistillate ftowcr, less perianth and ovary; Be, pistillate flower. vertical section, less
perianth; Bd, ovary, cross-.sectiOD. C, Cyclanthera explodens: Ca. pistillate llower,
vertical section; Ch, staminate flower, vertical section. D, Mormordica Balsam/nea:
DB, staminate flower; Db, same, less perianth. E, Cucurbita Pepo: fruit, much re--
duced.

Campanulaeea_Bellllower Family
The beIl1lower family is one composed mostly of herbs, often with
milky latex, whose bisexual lIowers have actinomorphic or zygomorphic
corollas. The stamens are five and may be distinct, connivent or syn-
genesious. The ovary is inferior, usually three- or five-carpelled and
-loculed, with axile placentation (unilocular with parietal placentation
in a few genera). The fruit is usually a capsule, dehiscing commonly by
apical or basal pores.
Composlta_Aster Family (Fig. 69)
This is a very large and complex family, but its members are charac-
terized by having inllorescence and lIoral characters in common. The
inflorescence is a head or capitulum of few to many lIowers borne on a
receptacle. Each lIower may have one or more chaffy or scarious bracts
at its bpse (Fig. 69 d). The outside of the receptacle bears few to many
involucral bracts or phyllaries (Fig. 69 b) that may be distinct or com-
pletely fused to form a tubo-like structure (as in marigold). The lIowers
(usually termed 1I0rets) are typically of two types: when both types are
present in a single head, the inner ones are termed disc or tubular
1I0rets, and the outer are ray or ligulate 1I0rets (Fig. 69 a). The disc
floret typically has a tubular five-lobed corolla from whose base rises
five syngenesious stamens (connate by their anthers to form a cylinder
ImporCGnI FamUie. Grad Their Charade,.. 145
about the style). The ovary is inferior and bicarpellate, but contains a
single loeule and ovule. There is a single style terminated usually by
two stigmatic branches (becoming recurved at anthesis). The calyx is
generally represented by bristles or scales (or both) arising from the
Inp of the ovary and is termed the pappus (Fig. 69 d). The ray floret
is a modified disc floret whose corolla is split down one side and become

FIe- 69. COMPOSITAE. Helianlhus annuus: a, head, face view; b, same, back
view; c, section of receptacle to show disc florets and one·ray floret (corolla
partially excised); d. disc floret; e, same, corolla and stamens expanded.

expanded. Very often the androecium is lost and the flower is unisexual
(i.e., pistillate) and sometimes it lacks both sex element and is neuter.
The fruit is an achene and the pappus is often persistent. In some genera
the plants produce heads of ouly ray flowers, termed radiate heads (as
in dandelion) in which cases the ray florets are bisexual. In others, only
disc flowers are produced, and the heads then termed discoid (as in
AntenlUlTia). The characters of the inflorescence, ovary and androecium
combined serve In separate this family from all others.
GLOSSARY AND INDEX

THE FOLLOWING comprises a combined index to this book and a glossary

to taxonomic terminology as required by current American works on tbe
identification and nomenclature of vascular plants. In it are intercalated tbose
parts of Ibe work dealing wilb biosystematics, phylogeny, and historical
background. Terms discussed, or defined in Ibe several chapters of this book
are followed by !be pertinent page reference and tbeir definition is not re-
peated here. Many of Ibe definitions have been taken witbout change from
Lawrence, TaxolWmy of Vascular Plant•.

Abaxu,z, p. 71 (in footnnte).

Abortive. Defective; barren; imperfectly developed.
Abrams, Leroy. p. 114.
Abrupt. Changing suddenly rather than gradually, as a leaf tbat is nanowecI
quickly to a poin~ not tapering; also a pinnstely compound leaf that has
no terminal leaflet.
Acaulescent. p. 36.
Acctuory fruit. p. 79.
Accr~scenl. Becoming enlarged with age, as the sepals of some flowers.
Aecumbent. Lying against and face to face, as some cotyledons (Fis. 38D), p. 83.
Acephalous. Without a head.
A.cerO$t. NcedlcooShapcd.
Aehene, p. 81.
Achlamydeous. Lacking a perianth, without calyx or corolla.
Acicle. A very slender prickle or sti1f bristle.
Acicular, p. 44.
Acropetal. Arising or developing in a longitudinal plano from a lower to a more
apical level. the opposite of basipetal. See Centripetal.
Actinomorphic, p. 61--62.
Aculeate. Covered with prickles.
Acuminate, p. 44.
Acute. p. 45.
Acyclic. Arranged in spirals. not in whom.
Adaxial, p. 71 (in footoote).
Adherent. Unlike parts in close con~ but DOt fused.
147
1411 GLOSSARY .4JIID INDEX
Adllllle. p. 63 (in footnote).
Advanced characters, pp. 99-101. •
"'d"~nliliolU Buds. Buds appearing on oc:c:aaion, rather than resident in regular
places and order, as those arisinS about wounds.
AIl,,_lIIitiYe. A nonindigenous plant that bas become cstablished in a plant com-
munity but which is not expanding its area of occupation appceciably.
Autival. Said of a plant appearing in su.nuner.
A<JStlvtllion. lbe arrangement of the perianth or its parts in the bud, p. 66 (Fic.
30). Verllation i! the leaf arrangement in the bud.
AnTI",,- fruit, p. 80.
A/ate. Winged.
AlblUPlen. Starclly or other nutritive material accompanying the embryo in the
seed; commonly used in the sense of endosperm. for the material lur-
rounding the embryo.
AlllIIf:eow. Having the smell or taste of ollions or IUlie.
A.1UnraU. Any arrangement 'ot leaves or other parts Dot opposite or whorled;
ono ])art at a node; plaeed .ina!y at dilferent heii\lts on the axi& or stem
(FiB. 19.).
A"'.olate. Appearing like a honeycomb, With lJIIII)ar depr...ions aeparated by
thin partitlona, often in hex_Bonal form.
Alveoli: Pits or depressions rescmblinB a h()neyeomb.
Amaranthaceae, descn1>ed, pp. 129-130.
Amaryllidaceae, described, pp. 123-124.
A.menl, p. 60 and Fis- 60.
AmpltktJrpoua. ProoU4'in8 two kinds of fmits.
A",phigean. Nali.., to both the New and ()ld Worlds.
Amphiploid (AmphidiplOid). A type of p<>lyploid characterized by the addition
of both sets of chromosomes from each of two species.
A.mpldtropou.r, p. 7S.
AmDlnkaul. Clasping the stem.
Ampulla. A bladder, as in Utricularia.
Anostomo.ring, pp. 29, 31.
ANJJrOPOIU, p. 76 (Fig. 3BAc).
Ancipital. Tw<HOdged.
Androecium, defined, p. 6S; types illUilrated, p. 68.
Andro/lYnophore. An axis Dr .talk beaz:intr both stamens and pistil above the
point of periantb attachment.
Androszynow. Said of an inOorescence colllposed of both staminate and pistillate
floWers, with the stam.imlte at tho apex, as in some members of the
Cyperaceae.
Anarodioecious. Havins: staminate flowers on one plant and bisexual 1Iowen OD
another.
AndromonoeciolU. Havins: male and bisexual tlowers on the same plant.
Androphore. A stalk bearing the androechun.
Anemophilow, p. 69.
AngiosperMS. characteristics of, p. 35.
Annual, p. 37.
AnnuloJe, p. 5!1i; annulate sporangia" p. 28.
AnnullU, p. 28.
Anterior. Front; on the front side; away from the axis. adaxial; toward. the
mbtending bract.
Anlezep%",. Inserted opposite the poiDt of insertion of the aepala.
GLOSSA.RY A.ND INDEX 149
A.nther. p. 65; dehiscence types, 68; -uc, 66.
Antheridium. In Cryptogams. the organ producing male gametes or sperm.
Anlhesis. Flowering; strictly. the time of expansion of a flower when pollination
takes place, but often used to designate the flowering period; the act of
flowering.
An/rorse. Directed forward or upward.
Apetalous. Lacking petals.
Apu (pl. Apices). The tip or distal end; for terms of. see pp. 44-45.
Aphyllous. Leafless.
Apiculau. p. 45 and Fi,. lSi
Apocarpous. With the pistils (carpels) separate, not united; frequently applied
to a gynoecium of separate pistils; the opposite of Syncarpous.
Apocynac•••• described. pp. 139-140.
Apomict. p. 24.
Apop.t.lollS. p. 63.
Apophysis. The 'exposed. swollen part of a cone-scale. as in Pinus.
Aposepalous, p. 63.
Appendage. An attached subsidiary or secondary part, sometimes projectin, or
banging; for appendaged anthers. see pp. 67-68.
Appendicular Theory, p. 77.
Appressed. Closely and flatly pressed against; adpressed.
Apterous. Without wings; the opposite of Alate.
Araceae, described, p. 122.
Arachnose. p. 51.
Archegonium, p. 27.
Arcuate. Curved or bowed.
Areolate. Reticulate, netted, marked out into small spaces by venation pattern,
p. 55.
Areale. see Fig. 22; of leaf venation, p. 31:
Aril. An appendage or an outer covering of a seed, ar1smg from or Dear the
hilum or funiculus; sometimes appearing as if pulpy.
Arillate. Provided with an ariI.
Arislate. p. 44 (Fig. 15b).
Aristotle, p. 7.
Armature. Any covering or occurrence of spines, barbs. hooks, or prickles .
.A.rmed. Provided with an)' strong, or shaJtl defense, as spines, thorns, or barbs.
Arrangement of leaves, terms of, pp. 48-49.
Articulate. Iointed; provided with nodes or joints or places where separ3tion
may naturally take place.
Arum family, described. p. 122.
Ascending, p. 36.
Ascidium. A cup- or pitcher-shaped organ, as in Nepenthes leaves.
Asexual. Sexless; without involvement or influence of gametes.
Asclepiadaceae, described, p. 140.
Asperous, p. 51.
Assurgent. Ascending, rising.
Aster Family, described, pp. 144-145.
Attenuate, p. 45 (Fig. 16a).
Auricle. An ear-shaped part or appendage, as the projections at the base of lOme
leaves and petals.
Auriculilte. p. 46 (Fig. J6g).
Author citation, pp. 93-94.
150 CWSSARY AND INDEX
Autoploid. A polyploid in which each of the three or more chromosome seta has
been derived from the same species. See also Amphiploid.
Autotrophic. Neither parasitic Dor saprophytic.
Awl"haped. Narrow and sharp-pointed; gradually tapering from base to • dODder
or stiff point.
Awn. A bristle-like part or appendage.
Axil, p. 36.
Axil. Platen/ation, p. 74 (FiS. 36a).
Axillary,. ln, or arising from. an axil.
AxiJ. The main line of development; the main Item, __ p. 36.

Boccate. Berry-like; pulpy or lIeahy.

Boon, Sir Joaeph, p. 104.
Banner Petal. p. 63.
Barbed, p. 52 (Fig. 22h).
Barbellate. Minutely or finely barbed.
Barbulate. Finely or minutely bearded.
Barohart, 1. H., p. 114.
Barton, B. S., p. lOS.
Bartram, John, p. 104.
Basal Placentationl pp. 71, 74.
Bases, terms of, pp. 4S-46.
Basifixed, p. 66.
Basipetal. Developing in a longitudinal plane from an apical or distal point toward
the base, the opposite of Acropetal. See Centrifugal.
Basonym. The original epithet assigned to a species (or taxon of lower rank).
which is usually retained when transferred to a new position.
Bast. The inner bark. usually fibrous in texture.
Beard. A long awn or bristle-like hair, u in some grasses; a ~ line or zone of
pubescence, as in some perianth parts.
Beech family, described, p. 128.
Bellflower family, descnbed, p. 144.
Bentham, George, p. 9.
Berry, p. 82.
Bessey, Charles E., pp. 12-14.
Betulaceae, described, pp. 127-128.
Biciliate. Provided with two cilia or claters, as in some motile sperm or spoteI.
Biennial, p. 37. .
Bifid. Two-cleft, as the apices of some petals or leaves.
Bifurcate. Porked, as in some Y-shaped hairs, stigmas, or styles.
Bigelow, Jacob, p. 105.
Bilabjote, p. 64.
Binary, name, contrasted with binary epithet, p. 23.
Binomial.. pp. 23) 91.
Biosyslematics, pp. 98-102; defined. p. 98; units of. p. 102.
Biolype. A genotypic population.
Bipinnate. Twice or doubly pinnate. p. 41 (Fi,. llf).
Sipinnalifid. Twice or doubly pinnatifid.
Birch family, described, pp. 127-128.
Biseriote. In two whorls or cycles~ as a perianth composed of a wyx and a
corolla (FiS_ 27).
Bisexual, p. 61.
GWSSA.RY AND INDEX 151
Bladdery. Inflated; the walls thin, like the distended bladder of an animal.
Blade. The expanded part of a leaf or petal.
Bloom. A whitish.. finely powder; co'lerini af the s.url-ace, ofttn W?J..1. fA 01\
some grapes.
Bale, p. 36.
Boraginaceae. described. pp. 140-141.
Boslryx. A helicoid cyme.
Botulifol'm. Sausage-shaped.
Brachiate. With spreading acmlik:e branches.
Bract, p. 57.
Brae/eale. Bearing bracts.
BTtJCleo/e. p. 57.
Bractlet. A bract borne on a secondary axis, as on the pedicel or even on the
petiole; a bracteole.
Branchlet. The ultimate division of a branch; the current year's Bl'owth; a twig.
Bristle. A stiff hair or seta; like a hog's brisUe.
Bristly. Bearing stiff strong hairs or bristles.
Britton, Nathaniel L.. pp. 112-113.
Buck.wheat family. described, pp. 128-129.
Bud, types, pp. 39-40.
Bulb, p. 38.
Bulb-tunic. A usually membranous or fibrous coat envelopina a bulb; types of.
p. 38.
Bulbel, p. 38.
Bulbil, p. 38.
Bulbl." p. 38.
Bullate, p. 53.
Bu,,;cule. The pouchIike expansion of tho atigm&, as that of some orchida and in
which the caudicIe of the pollinium is iDscrtecL
Butten:up family, d=ribed, p. 130.

Cactaceae. described, pp. 136-137.

Caducous. Falling off early, p. 56.
Caesa1piniojdeae. described. p. 134.
CaesioW'. p. 56.
Cala/hi/arm. Cup-shaped.
Ca}clUole. Having or produced into a spur.
CaUd/orm. CalYXMlike.
Callosity. A thickened raised area, often paler in color; the presence of a callus.
Calyculate. Calyx-like; bearing a part resembling a calyx; more commonly, pro-.
vided with bracts against or beneath the calyx and resentbling a supplee
mentary calyx.
Calyculu.r. A simulated calyx, composed of bracts or bractlets.
Calyptra. A bood or lid; particularly that of a moss cap.iDle; the lid in the fruit
of eucalyptus, or the form of the calyx as in some papaveraccous flowers.
Calyx, p. 63.
Cal,x Tub!". The: tube of a pmosepalous calyx. sn p. 18; sometimes employed
for hypanthium. which see.
Campanulaceae, described, p. 144.
Campanulate, p. 64.
Campylotropous. p. 75 and Fig. 38M.
CaMlicu/tUe. With a channel or groove.
152 GWSSARY AND INDE,x
Cancellate. Resembling latticework.
Candolle, see de Candolle.
Canescent. Gray-pubescent; hoary or becoming so.
Cap. A convex removable covering, as of a capsule; in the grape, the coberin,
petals falling off as a cap.
Capillary. Hairlike; very slender.
Capitalization of species names, p. 95.
Capitate. Headed; in heads; formed like a head.
Capitulum. A dense inflorescence composed of an aggrea:ation of usually aeaeile
or near-sessile flowers, p. 59.
Caprifoliaceae, described. p. 143.
Capsular. Pertaining to or formed like a capsule.
Capsule, pp. 80, 81-
Carinal. Keeled, especially when the keel contains other floral parts.
CarinQJe. Keeled; provided with a projecting central longitudinal line or ridge.
often on the dorsal or adaxial surface.
Carpel. One of the foliar units of a compound ovary or pistil; a foliar, usually
ovule-bearing unit of a simple ovary. two or more combined by connation
in the origin or development of a compound ovary; a female- or mega-
sporophyll of an angiosperm flower. See Pistil. For discussion, see
Lawrence. Taxonomy of Vascular Plants. pp. 72-75.
Carpdlate. Possessing or composed of carpels.
Carpophore, pp. 81, 138.
Carrot family, described, pp. 137-138.
Cartilaginous. Tough and hard. but bony; Vistly.
Caruncle. A caUus-like appendage at or about the hilum of a seed, often paler
in color than the seed coat.
Caryophyllaceae. described, p. 130.
Caryopsis. p. 122.
Castaneous. Of chestnut color; dark brown.
Catkin. A scaly-bracted, usually flexuous. spikelike "inflorescence" of cymules;
prominent in the willows. birches, and oaks. See p. 60.
COJUl4Ie, p. 44 (Fig. 1St).
C"Jldu. Stem. p. 36.
Caudicle. The stalk of an or<:hid pollinium, usually thread- or strap-shaped. Not
to be confused with the stalk of the pollinium in some Asclepiadaceae,
which is termed a Translator Arm.
Caulescent. Having an evident stem above ground. See p. 36.
Coull"", p. 49.
Centralium. A central lengthwise cavity, as found in the seeds of some palms.
Centrifugal. Developing from the center outwards.
Centripetal. Developing from the outside toward the center.
Centrum. The center of a solid.
Cernuous. Nodding; drooping.
Cesalpinaceous Corolla, p. 63.
Cespitose. p. 36.
ChaO. A small thin, dry, membranous scale or bract; in particular, the bracts
in the Bower heads of the Composites.
Chalaza. The basal part of an ovule where it is attathed to the funiculus (Fig. 38).
Chapman, A. W., pp. 114-115.
CluJ1'Iaceous. Of papery or tissue..llke texture and not UlUally &reen in color.
Chenopodiaceu, described, p. 129.
cwss,my .4ND INDEX ISS
Choripeta/oUl. Having separate and distinct peWs; polypetalous.
Cilia. Hairs arising from the marsin; eyelash-like.
CilI"'., p. 47.
CiUolate. Diminutive of -ciliate; minutely ciliate.
Cincin.ruu, p. 59.
CiMr~us. Ash-colored.
Circlnat., p. 30.
Circum.rcissile. p. 81.
Ci"MIU. pp. 31,44.
Citation of authors' names, pp. 93-94.
C/adophyll, p. 38.
Clambering, p. 31.
Ckuping. Partly or wholly surrounding the stem.
Class, a unit of classification. pp. 21, 22.
CltJ.uI/ication, defined, p. 2; phylogenetic IY'tem.s of, pp. 6-18; unlta of. pp. 21-22.
CIa,"'e. Club..haped, like a baseball haL See p. 53, Fig. 22f.
Clave/late. Diminutive of clavate.
Claw. The long narrow, often petiole-like, base of petals and sepals of some flowers.
Clayton, John, p. 103.
Cleft, p. 48.
Cki.Jtogamous Flowers. Small, closed, self-fertilized flowers, as in some violets
and many other plants; they are mostly on. near, or under the ground.
Climbing, p. 37.
Clinandrium. The pouch or "'anther-bed" in monandrous orchids; that part of the
gynandrium of ao orcliid ~ow.,. in which the po1lin.ia are concOJlled.
C/o"., pp. 22, 24.
Coaetaneow. Flowering as the leaves expand.
CoaJucence. Cohesion; the incomplete fusion or sticking together of like parts
(as petals to petals); sometimes loosely used synonymously with connation,
which is the complete fusion of like parts.
Cocou.r (plural Cocci). A berry; in particular, one of the parts of a lobed or
deeply divided fruit when each part is on....eeded, sometimes these parts
may be leathery or even dry.
Cochleate. Spiral, 'ike a snail shell.
Code of nomenclature, p. 89.
Coherent. Like parts in close contact but Dot fused.
Collecting techniques, pp. 84-87.
Collinson, Peter, p. 104.
Colonial. Forming colonies, usually by underground parts.
Col"",ella. The central axis (carpophore) supporting one of the halves (meri-
carps) of the fruit of Umbelliferae, sometim.. restricted to the basal un-
branched portion.
Column, p. 125.
Coma. A tuft of hairs, as on Asclepias seeds; • leafy crown or head, .. in some
palm trees.
Commissural Face. p. 138.
Comose. Provided with a coma, or tuft of bairs, or resemblina such.
Comparium. p. 102.
Compositae, described, pp. 144-145,
Compound. Of two or more parts or components; leal. pp. 40-41; ovary or ptrtil,
pp.71-74.
Compre","'. Flattened laterally. See Obcomp~.
154 CWSSARY AJIID INDEX.
Concave. Shallowly hoIlowed; saucer-like.
Concolorous. Uniformly colored.
Conduplicate, p. 83.
Cone. A dense and usually elongated assemblase of fertile spolOphylls on a central
axis, -each often subtended by a bract.
Confluent. Merging or blending together.
Congeneric. Belonging to the same genus.
Coniferales, characteristics of, pp. 33-35.
ComUJte. United or joined; particularly when two or morc like parts are united,
as petals fused to one another by their margins in a gamopetalous corolla.
Connate-per/oUale, p. 46.
Connective. The filament or tissue connecting the two cells or thecae of an anther,
more obvious when the two thecae are separated by this intervening tissue.
Connivent. Coming together or converging but not fused; coherent; the parts
often arching (treated by some authors as synonymous with adherent).
Conoidal. Nearly conical.
Conservation of names, p. 91.
Contiguous, p. 30.
Contorted. Twisted; convolute in aestivation.
Convex. Rounded outward.
Convolute. Said of floral envelopes in the bud (as parts of corolla or calyx) when
one edge overlaps the next part (petal, sepal or lobe) while the other edge
or margin is overlapped by a preceding part; rolled, the margins over-
lapping. See Fig. 30c.
Cordate, p. 46.
Cordiform. Shaped like a heart.
Coriaceous. Of leathery texture.
Corm, p. 38.
Cormel, p. 38.
Corolla. p. 63; types of. pp. 63-65.
Corona, of Amaryllidaceae, p. as; of Asclepiadaceae, p. 140.
Corylaceae, p. 127.
Corymb. p. 61.
Costa. A rib, the midvein of a simple leaf; less commonly the rachis of a pin-
nately compound leaf.
Costapalmate. Said of a palmate palm leaf whose petiole continue.! through the
blade_, forming a distinct midrib.• as in the palmetto.
Costate. Ribbed; having one or more, usually raised, longitudinal ribs or nerves.
Cotyledon. The seed leaf; the primary leaf or leaves in the embryo; in some
plants the cotyledon always remains within the seed coat, while in others
(as in the bean) it emerges on germination. For cotyledon positions, see
p. 83 and Fig. 38.
Cotyloid. Concave or cup-shaped. as the receptacle of Calycanthus.
Coulter. J. M .• p. 111.
Coville, F., p. 117.
Crassulaceae, descnoed, pp. 131-132.
Crateriform. Shaped like a saucer or cup.
Creeper. A trailing shoot that takes root mostly throughout its length; sometimes
applied to a tight-clinging vine.
Cremocarp. A dry, dehiscent. two-seeded fruit of the Umbellifer family; a schizo..
carp.
Crenate. p. 47.
GWSS.4RY .4ND INDEX ISS
Crenulate. Diminutive of crenate; finely waved in a vertical plane.
Crested. With a. ridlc that is irregular. elevated and often toothed. or appearinJ
as if so.
Crispole. Curled, aD extreme form of undulate.
Crisp-hairy. Provided with hairs that arc curled and somewhat kinky. .
Cristate. Bearing a crestlike append_Ie.
Crosier. p. 30.
Crown. A corona; that part of the stem situated at the surface of the ground; a
series of appendages in the throat of a cowlJa.
Crucif.r••, described, p. 131; figured, p. 132.
Cruciform Corolla. Cross-.shaped. the petal or corolla-lobes usually four. See p. 63.
Cryptogam. A plant reproducing by spores instead of by seeds, as ferns, JIIossa,
algae, fungi. S.. pp. 27-31.
Cucullate. Hooded or hood-shaped,
Cu<:urbitaceae, deacribed, pp. 143-144.
Culm. Stem, particularlY that of a gr....
Cultigen. Plant or group known only in cultivation; presumed to have on,inated
under domesticatioD; the opposite of incti.cn.
Cultiva,., A variety or race that has originated and peraisted under cultivatioa,
not neceuarily referable to a botanical species.
Cu_e,p.45.
Cuneiform, p. 43.
CUpreaa...... p. 35.
Cupule. Cuplike structure at the base of aome fruits (u in aoJIIO palma) formed
by the dry and enlarged Iloral envelopes; aIoo the cup of an acorn.
Cuspidole, p. 45.
Cyathlum. A type of inIloresceru:e characteriatic of EuPhorbiD; the um.en,.!
Ilowe" condensed and conaeated within a bractule envelope from which
they emerge at anthesis.
Cycadales, characteristics of, p. n.
Cyclic. Arranged in whorls; the oppoaite of spiralled.
Cymbiform. Boat·ahaped.
Cyme, p. 57.
Cym",e. Cyme-like iD1Iorescence.
Cymuie, p. 59.
Cypcmceae, deacribed, p. 122.
Cyprela. An achene bcarinl an .dnate cal}X, IS in 8OJIIO Compoaitae.
Cy.rolilh. A .Ione-like concretion of calciUJII cIepoait in the epidermis of IIOIDO
plants, as in the nettles.
Cytogenetics, p. 101.

Darlington, William, p. 106.

Darwin, CharI.., p. 9.
de Candollo, A. P., p. s.
Deciduous. Falling at the end of one .cason of growth or life, .. the lea... of
Don-evergreen trees.
Decli1l4te. Bent downward or forward, the tips often rccurved.
Dtcompound. More than once compound (FiB. lSf, g).
Decumbent. Reclining or lying on the ground, but the end ascending (Fig. 106).
Decurrenl. Extending down and adnate to the stem, as the leaf base of Verbascum.
DtclUsale. Opposite leaves in four rows, up and down the stem, altefllating in pain
with each pair at right ansi.. to the one above and below (Fig. 19d).
156 GLOSSARY ..4ND INDEX
De/lexed. Reflexcd.
Dehiscence. The method or process of opening; for that of anthers, see p. 68,
and of fruits, see pp. 80-81.
de lussieu. classification system, 8.
Deliquescent. Having the primary axis or stem much-branched above. as in an elm
tree; the opposite of Excurrent; said also of perianth parts that quickly
s.often and deteriorate in a few minutes to hours after picking, in Eichornip
or Trade.rcanlia.
Deltoid. Triangular; delta-like, derived from the Greek tetter, delta (Fig. 14p).
Dentate. With sharp, spreading, rather 'COarse indentations or teeth that arc pcr~
pendicular to the margin (Fig. 17g).
Denticle. A minute tooth.
Denticulate. Minutely or finely dentate (Fig. 17h).
Depressed. More or less .flattened endwise or from above; pressed down.
Determinate Inflorescence. p. 57.
DextTorse. Clockwise; toward the right hand.
Diadelphous, p. 68, Fig. 32.
Diaiypetaious. Polypetalousj a corolla of separate and distinct petals.
Diandrous. Having two stamens. as in Veronica or Salvia.
Dichasjum~ p. 57.
Dichotomous. Forked, in ODe or morc pairs, see p. 36; a dichotomous key iI one
with C()uplets of opposing statements, see p. 8S.
Dicotyledoneae, p. 127.
Diclesium. An acbene enclosed within a free but persistent envelope.
Die/inous. Unisexual; requiring two flowers (or plants) to represent both. sexes.
Didymous. In pairs, twins.
Didynamous. Wjth four stamens. in two pairs and oD:c pair of a different lenath
(or height) than the other (Fig. 32.).
DiDuse. Loosely branching or sprcading; of open growth.
Digitate. Handlike; the members arising from one point, as the leaftets of a horse-
chestnut leaf. See Palmate.
Dimorphic. Occurring in two forms. as in ferns (see p. 31) or baving juvenile and
adult foliage types as in Eucalyptus or Hedera.
Dl.a«iaus. Having $l1HIli.aate and pistillate .floweIS' on diJIereat plants; a term
properly applied to a taxonomic unit and bot to flowers (see p. 61).
DiplOid. A plant having two sets of chromosomes in its somatic nuclei; the oppoooi
site of HaplOid,' Ii plant may have a diploid number of chromosomes in its
vegetative cells and a haploid Dumber in its male and female nuclei,
(gametes).
Dip/ostemonous. Having the stamms in two whorls, those of the outer whorl
alternate with the petals, and those of the inncr whorl opposite the petals.
Dipterous. Two~winged.
Di.IC. A more or less fleshy or elevated development of the receptac1e, or of the
coalesced ntctaries or staminodcs, about the pistil; the receptacle in the
head of the Composite Bower; a Battened extremity, as on tendrUs of
Virginia creeper. The term is an anglicizatjon of the Latin discus, and some~
times is spelled disk, and often so when used with reference to the Com~
positae.
Disc Floret. The tubul... llower of som, members of til, Compositae (Fis. 69d).
Disciform. Circular and like a disc.
Discoid. Resembling a disc: used to designate a head of members of the Com-
posit•• when only disc lIowen alie present (the opposite of Radiate).
GLOSSARY AND INDEX IS7
DUcr~t~. Separate, DOt coherent or adherent. See Distinct.
Dilk, see Disc.
Dinected. Divided into many slender segments.
Dissepiment. A partition, asiD an ovary or fruit (oot commonly used).
DissichoJU, p. 411.
Distinct. Separate; not canute nor coherent with other parts of the same series;
e.g.• the petals of a polypetalous Bower are distinct. When petals are BOp-
aratc from sepals and not adnatc to them, one is saw' to be free from the
other. Compare Pree..
Dlu,,",l, p. 56.
Divaricate. Spreading very far apart; widely divergent.
Divergent. Spreading very broadly, but less so than whea -divaricate.
Divided. Separated to very near the base (or to the midrib).
Division, unit of classification, pp. 21, 22.
Dogbane family, described, pp. 139-140.
DokJbri/orm. Shaped like the head of an axe or hatchet.
Dorsal. Back; adaxial; relating to the back. or outer surfa.ce of a part of aD or~
as the' lower side of a leaf. or the part of an organ away from the axis
supporting it; the opposite ventral and abaxial. See p. 71.
l)tmi/ixed. Attached by the back; otten, hut oot necessarily versatile, .. the
anthers in Lilium (Fig. HDb, Do).
Dordventral. Flattened and provided with a definite dorsal and ventraJ surface;
laminate; e.g., most foliac:eou., leaves are dorsiventral (or have doniventral
surfaces) whereas the fruit of an OTlUlgt does Dot have a dorsiventral.ur·
face.
Dollble...serrole. With coarse serrations bearing minute teeth OD their mar&iDs.
Downy. Covered with very short and weak soft hairs.
Drake, Francis, p. 103.
DrllpauolU. Resembling a drupe bur morphologicaJly not identical with one.
Drupe, p. 82 and Fig. 41 .. b.
/)",pelt/f. One drupe of a fruit composed of an aggregation of miniature drupes,
\s in the raspberry, the ,o"aned aeed withlD a drupelet belDg a pyron•.
Dryio, specim.... p. 87.
Duct. A tube or canal which carries latex, resin" or oil.
Durand, Elias, p. 108.
D_ion, to...... of, pp. 36, 37.
E. or ex-. A Latin prefix often meanlDg without, as oeiliate (withnut cilia) or
exstipular (..tipular) without stipul...
Ebracttate. Without bracts.
Eohinate, p. 52.
Ecilitu~.
Without eilia.
Ecospecies, p. 102.
Ecotype, p. 102.
Effective pUblication. p. 92.
Elater. A ribbon·like band whicb, in Equisl!lUm, is bysroscopic. somewhat clavate.
.and aids in dispersal of the spore! to each of whicb four elatets are
attached.
Elliptic, p. 43.
Elongate. Lengthened; stretched out.
Emarginate, p. 45.
Embryo. The rudimontary plant within a steel, usually developing from a zraote.
158 GLOSSARY AlVD INDEX
Embryot~ga. A disclike callosity on the seed coat in species of Commelinaceae
FJagelJariaceae, and MaYBeaceac. '
EMtion. An epidermal outgrowth. often one cell thick.
Endtuch. Said of a type of xylem whose development is toward the centcr of
the axis (centripetal); the opposite of exareh.
Endemic. Native or confined naturally to a particular and usually restricted area
or region; biologically a relic of once wide distribution.
Endocarp. The inner layer of the pericarp or fruit wall.
Endosperm. The starch~ and oil-containina: tissue of many seeds; often referred
to as the albumen.
Engelmann, George, p. 1 U.
Engler, Adolf, pp. lG-12.
Ensiform. Sword-shaped.
Entire. With a continuous margin, not in any way indented. See p. 46.
Entomophilous. p. 69.
Envelopes, floral. The two series composing the perianth; the corolla and the'
calyx.
EpbedraIes, p. 32.
Ephemeral, p. 56.
Epi-. A Greek prefix, signifying on or upon.
Epibiotic. A species which is a survival of a lost flora; a near-endemic.
Epigynous. Borne on or arising from the ovary; said of floral parts when the
ovary is inferior and the flower not perigynous. The term is not applicable
to the ovary itself.
Epip~talow. Borne on or arising from the corolla or petah.
Epiphyte. A plant growing on another or on some other elevated support.
Equi'anJ. Overlapping in two ranks, as the leaves of Irt. (Fia. 19j); astride, ..
one leaf astride another.
Ericaceae, descnbed, pp. 138-139.
Erose. Said of a margin when appearing eroded or gnawed. or of a jaagednesa too
small to be fringed and too irregular to be toothed.
E,tipulate. Without stipules.
Euphorbiaceae, described, p. 135.
Eusporangiate. Said of fern sporangia that have developed from a group of cells
that first divided periclinally to produce inner (sporogenous-forming) and
outer (sterile, sporangium-forming) layers of cells. Se~ LeptosporanBiate.
Evening-primrose family, described, p. 137.
Even-pinnau. A pinnately compound leaf lacking a terminal leaflet, with the total
leaflets of an even number, pp. 40, 41.
Evergreen. Remaining green in its dormant season; sometimes applied to plants
that are green throughout the year; properly applied to plants and not to
leaves, but due to the persistence of leaves.
Evolution and pbylogeny, pp. 98-IOI;-and units of classification, pp. 19-25.
Ex-. Prefix meaning without, or lacking.
EXIJ3peraie. Rough, with the papillae bearing hard, minutely projecting points.
Excu"ent. Extending beyond the margin or tip, as a midrib developing into a
mucro or awn; or, descriptive of the habit of a plant with a continuous
unbranched axis, as Picea or A bies. The opposite of deliquescent.
Exfoliate. To peel off in shreds, thin layers, or plates, as bark from a tree trunk.
Exine. p. 69.
Exocarp. The outer layer of the pericarp or fruit wall.
E;tplanat•• Flattened, spread out.
CWSS.4Rr AND INDEX 159
E:uerted. Projecting beyond. aa stamens from the mouth of a corolla. The oppo-
site of included.
Eutlpult!te. Without stipules.
Extrone. Lootina or facing outward; laid of anther dehiscence and best deter~
mined by making a cross section of an undehisccd anther. See p. 68.
Eye. The marked center of a flower. a bud on a tuber; a sinsle bud.-cuttinJ.,

F ......... desoribed, p. 128.

F.leal•. Sickle..baped.
FalU. Outer whorl or aeries of perianth parts ·of an iridaceous ftower. often
broader than those of the inner aeries (the standards) and, in some ITiI.
droop in, or flexuous.
Family, a unit of classification, pp. 21, 22-23; characterization of. pp. 119-145.
FarinoceoUl. Containing starch or starchlike materials; sometimes appliod to a
surface covered with a mealy coatina. as leaves of some Primula spp.
FOTUw6e. Covered with a meal~like powder. Su p. 56.
Ftuci4tt. Much flattened: an abnormal widening and flattening of the stem.
Fardcle. A condensed or close cluster, as of lowen. or of most pine leaves (Pia,
19f).
FIUCIc.Iot., p. 49.
FlUtigi4te. With branches erect and more or leas appressed. s.e p. 36.
F"""aI. Pertainin. to the throat, as of a coron•. F.uc.1 App.ndages are smaU
often toothlike appendages or enationi arising from a corolla at its mouth,
as in some BoraSe8 or members of CaryophyJlaceae.
Fal1eolale. Honeycombed. See Alveolate, .Foveolate.
Fttlther-veined. With the ribs an extending from the central midveiD toward the
margin and remainin, .imple or scarcely branched.
'_st,al" p. 55.
Pemsld, M. t., p. 1.12,
PeI'DI, characteristics of, pp. 27-31.
FemtgifWw. Colored rust-red.
Fertil•. Said of ponen·bearing stamens and seed-bearinS fruits.
Fertilir.aJion. The union of two gametes (as the male nucleus from a germinatinl
pollen grain, and the e,a-nucleus of an ovule) to form a zygote (which, in
seed plants, develop. to fonn the embryo within • seed); not to be confused
with pollination.
Fetid. Having. disasreeable odor.
FihrilltJ. Diminutive of fiber.
-fjd. Combining fonn denoting cleft; cut about halfway to the middle.
Figwort family, described, pp. 142-143.
Filament. Thread; particularly the .taIlt of • stamen (terminated by the anther).
S.e p. 65.
Filamentous. Composed of threadlike strands.
Filicinae, pp. 27, 29.
Filiform, p. 42.
Fimbriate. Fringed.
Fimbrillate. Provided wiC! a minute fringe.
FistultJ. Tubular; hollow and cylindrical.
Flahellate. Fanshaped, as the leaf of Oinks<> or of Palmelto palm; broadly wecIse-
shaped.
Flaccid. Limp, Hoppy; wilted.
Flag.llI/arm. Like a whiplash or a diminutive of IL
160 cwss,ml' AND INDEX
FlexuolU. Having: a morc or less zigzaa or wavy form; withy.
Floccos•• Covered with tufts of soft woolly hairs thai usually rub oft readily.
Floral-cup. p. 78.
Floret. An individual ftower, usually m\fiute. especially of &rasses and c{)mpositel;
any assemblaae of small flowers that compose a compact type of in-
ftorescence.
Florlc."". The !lowering and fruiting stem, especially of a bramble (Rubus), usu·
ally the second year's aro'Wth or development.
Floriferous. Flower-bearing.
Flower. An axis bearing onc or morc pistils, ODC or more stamens, or both; when
only the lormer, it is a pistillate /lower, when only the lattcr a Itam;na/e
flower, when both sex elements are present it is a perltel flower (i.e., bi.
sexual or hermaphroditic). When the pcdcc( Bower is surrounded by a
perianth composed of two floral envelopes (the inner envelope the coro~
the outer the calyx), it is a complete /lower. See also pp. 61-78.
Foliaceous. Leaflike; said particularly of sepals, calyx-lobes, and bracts that in
size. texture, and color resemble small or larae leaves.
Pollicetum. An aggregation of follitles, distinct or basally connate, representina
the product of an apocarpous multipistillate l)'Doecium.
Follicle, pp. 8()...$1.
Forma, a unit of clasailk:atioa. pp. 22, 24.
FOl'eolole. p. 55.
Fraser, 1., p. 104.
Free. Not adnate or adherent to other orgaQS; as petals free from the stamens Ot
calyx; or, the vein1eta (as in some ferns) not united. but are free. Some-
times, and especially in older literature, the IOrm is used in the sense 01
distinct.
Free-cenlral Plocentation. p. 74.
Frond. The leaf of a fero; sometim.. used in the sense of foliage, especially of
palms and cycads. See p. 30.
FrontlMe. Bearing fronds; leafy.
Fruit. The ripened ovary (pistil) with the adDale parts, the seed·bearina organ;
it is applied also to the aced·bearing organ 01 gymnosperms, although not
the product of a ripened ovary or pistil; in sOo..e works it is improperly
applied to the fertile portions of • fero-leaf. For fruit types. .ee pp.
7S-82.
Fl'Ulico... Shrubby or shrublike in the ..use 01 being woody.
FugaceoiU. p. 56.
Full'olU. Tawny in color.
Funiculus, p. 7S.
FUMelform. With the tube gradually widening upward and passiq insensibly into
Ibe limb, and in many kinds of Convolvulus: infundJbuliform (Fig. 28e).
Furcat•. Forked.
Furrowed. With longitudinal channels or grooves.
FuscolU. Grayish brown.
Fusiform. Spindle-shaped; narrowed both ways from a swollen mi6-region.

Old•. A helme4 as one sepal in an aconite ftower, or the upper lip of some
bilabiate corollas.
GtJ!eate. Provided with a galea; hooded.
 GWSSA.RY A.ND INDEX 161
Gamete. A minute protoplasmic unit cootaininl a single nucleus (whose chromo-
somes are usually of the haploid number) which on union with another, and
usually unlike, gamete forms the zygote .
. Gametophyre. The gcneration that bears the sex organs; in ferns, a thallus-like
small or minute body bearing archegonia (the egg-containing organs) and
antheridia (the sperm-producing organs); in the angiosperms reduced to
the three-nucleate pollen-tube and the eight-nucleate (or ~lled) embryo sac
imd: contents. See p. 27.
GamopetalolU, p. 63.
Gamophyllous. With leaveo, or foliar units, connate by their edgea.
GamosepaloU8. p. 63.
Geminate. Equal, in pairs.
Gemmtl. An asexual propagule sometimes appearinJ as, but not homologous with,
a bud.
Geniculate. Bent like a knee. See p. 36.
Ge1lOtype. A type (piant or population) determined by sonetical characters. See
Phenotype.
Genus, a unit of classification, pp. 22, 23; nomenclatural considerations of, p. 91.
Gibbous. p. 65.
Oinkgoales, characteristics of, p. 33.
Glnbrate (glabrescent). Nearly glabrous, or becoming glabrous with maturity or age.
Glnbrous. Not hairy; often incorrectly used in the sense of Imooth.
Gladtal<. Sword-sbaped, but may be straight or curved.
Gland. A secreting part or prominence or appendage, but often used in the sense of
a glandlike body.
Glandular. Having or bearing glands or secreting organs.
Glandulor·Pubescenl. With aIands and hairs intermixed, or hairs terminated by
pinhead·like glands.
Glandula,...Punctate. See Punctate.
Glanduliferous. Having or bearing glands.
GlGlICescent. Slightly glaucous.
Glnucous. Covered with a bloom or whitish substance that rubs off.
Glochid. A minute barbed spine or brisUe, often in tufts, as in some cacti (PiJ..
22d).
Glochidiate. Barbed at the tip.
Glomerate. In dense compact cluster or clusten.
Glumoceous. Provided with or resembling glumes.
Glume, p. 122.
Glutinous. Sticky, or covered with a sticky exudate.
Onetales, p. 32.
Ooodale, O. L., p. 111.
Ooosefoot family, described, p. 129.
Oourd family, described, pp. 143-144.
Gramineae, described, pp. 121-122.
Gronulose. Covered with very small grains, or appea.rinJ as if 10.
Orass family, described, pp. 121-122.
Orass Spikelet, p. 121.
Oray, Asa, p. 110.
Oreene, E. L., p. 116.
Oreenman, 1. M., pp. 115-116.
Gregarious. Growing in large colonies.
lliZ CLOss,mr AND INDB~

Oronovius, 1. p. p. 103.
Oymnospenns, characteristlco of, pp. 31-35.
GylUlltdrilUlJ, p. 125.
GylUlllllroU8. With tho stamono _ to or borDe OD tho pfstl1.
Gy_lUldrolU. With staminate and pistillate flowers in tho . - apit..1Il<. In-
lIoreaconco and with the pistillal. hoooath tho staminate.
Gynoblu•• An ODiaraemonl of tho receptacl. (axis) hoariD.I tho pistil.
Gy~ium. The fcmaJe element of a lower; a collective term employed for tho
several pistils of a single 1lower when referred to u a unit; when. only one
piati1 is present, pistil and lY1100Cium OR aynon)'mOUl. See pp. 69-78.
GYlIOphore. 'lbe oVlUJ'-hearinJ axis extended above tho re,lion of poriaDth attach-
menL
Gy_.mlum. 'lbe column (lYD8Ddrium) of an orchld IIower, formed by od.llm
of otam... and the styl. and atilJll8.

Habit. 'lbe ...,oral a~ of a planL ,

Habitat. 'lb. typo of localily and immediate onviro_ in whlch a plaut _ .
Haft. The narrow constricted part of an orJllUl or part.
Halb",J..haped. Sa Hastate, p. 46.
Half-lnferior Ovary, p. 76.
HatnIlI•• Hooted al tho tip.
Hastate, p. 46.
Hastula. T.rminal part of petiole on upper surface Ilf leaf blade of • palmate-
leaVed palm, sometim.. called the 1iau1. (but not homololOus with tho
IiauIo of tho grass leaf),
HfllUtoria. The absorbing or.... (often rootIIl<.) of some parasitic plants and
usually having the anatomy of a st.m rather than of a fOOL
Head. A abort donao in1IoroscoDCO of ....il. (or n.arly ....il.) flow.... .Ituated
on an axis that is much compressed vertically and sometimes flattened or
sauccr~haped. Set! p. 59.
H..th fanti1y, described, pp. 138-139.
Helicoid Cyme, p. 51.
Heml-. In Greek compounds, .ilPlifl'ing half.
HerbacwlU, p. 37.
Herbarium, pp. 86-87.
Hermaphrodite. With stamens and pistil in the same flower. a" p. 61.
H,aperidium. p. sa.
Hnero-. In Greek compounds, meanin8 two or more I0I1l.
Heterocarpous. Producing more than one kind of fruiL
Heterogamous. Bearing two or more types of flowers in one cluster, as in Com",...
itae.
HoterosparolU. A state of producing two kinds or sizes of spor.., as In S.loginello.
See p. 27.
Heterostyly. A .Iste of IIow.n of a pven planl or taxon poss...ing styl"'of dif-
ferent length ratios to the stamens or corolla, or other parta, as in Prlmula.
Hexa~. In Greek compounds, meaning in numbers of six.
Hexamerous. Its parts in sixes.
Hibernaculum (pl. hibernacula). A winter bud or an under8l'ound stem; a dormant.
resting bud.
Hickory fanti1y, described, p. 127_
Hlernal. Relaling to winter.
Hilum, pp. 75, 82.
CLOSSARY .mD INDEX 168
Hip~ The fruiting structure of a rose (Rosa). whose fleshy portion ill composed of
receptacle and hypanthiwn, and the achen.. included within.
Hlppocreplform. Ho....hoe-shaped.
Hirsute, pp. 51, 54.
Hirsutulous. SIiahtly or approachin, hirsute.
Hirtellous. Minutely hirsute.
Hispid, pp. n, 54.
HlsplduloUJI, pp. 52, 54.
Hoary. Orayish white and very fine p u _ ,
Holotype, p. 91.
HcmogamoUl. A head or cluster with flowers alI of one kind. with stijDlas becom~
ina receptive at the time pollen is shed in the same dower.
Homo/alow. Of similar origin, but differing in form or function, as the perianth
parts of a lower being homologous with the leaves.
Homonym. Two or more names of a plant (or epithets) with identical spelling,
but applied to two taxa. See auo p. 95.
Homosporous. The state of producing spores of ODe kind, as opposed to heterospo-
row; see alro p. 27.
HODOysuckle family, described, p. 143.
Hooker, 1. D., p. 9.
Howe, M. A., p. 113-114.
Humlfus•. Spreadina over the surface of the JI'OUDd.
Husk. An outer, usually looae, coveriq of aoma fruits, as In Phylallr.
Hutcl>inson, J. pp. 14-16.
Hy.line, p. 56.
Hybrid. A cross between two taxa; usually a cross between two species of 8
genua or genera.
Hybridization, defined, p. 19.
Hygroscopic. A changing of form or position through changes in moisture content.
Hyponthium, p. 78 and Fig. 39.
Hypocotyl. The axis or stem of a seedling below the cotyledoDS. S.e p. 82.
Hypocrattriform. SaIver-shaped, see salverform.
Hypogynium. A perianth-like structure subtendina and usually CDVe10plnJ the
ovary in some tnembcn of the Cyperaceae.
HyponnolU. Inserted below the gynoedum and free from it.
H.,stera1JlhoU4~ p. 56.

lcosandroUl. Wi.th twenty or mote stamens.

Identification, defined. p. 1; procedures of, pp. 85-86.
Iderttifyin. techniques, pp. 84-87.
megitima!e names, explained, p. 90; when rejected, pp. 94-95.
Imbricate, p. 49.
Immerged. Completely submerged. in water or plant tissues.
Imparipinnate. Odd-pinnate, with a terminal leaflet.
Imperfect Flower. A flower lactin, either male or female sex orsans. See p. 61.
Incarnate. Flesh colored.
Incised, p. 47.
Included. Not protruded beyond tho enveloping structure, as stamens may be
included within the corolla-tube; not exserted.
Incomplete Flower. A lIower lac:kins one of the perianth whnrls, as calyx or
corolla, or both. S•• Importee:! Flower.
164 cwss,my AND lIVJJEX
IncubolU. With the tip of one leaf overlapping the base of the leaf above, but Dot
to the extent of being imbricate.
Incumbent. Leaning or resting on; cotyledons are incumbent when the back of Doe
rests against the radicle; stamens are incumbent' when dehiscing toward the
IIorai axis (introrse).
Incurved. Gradually curvina inward.
lnilehbcent. Remaining closed; not splitting open at maturity along regular linea
of fusion.
Indetermillllle Inflorescence. p. 57.
lndigen. A native; as contrasted with a cuitigen (growing under cultivation). a
naturalized plant (thoroughly established but coming from a foreign
region), an adventive (somewhat naturalized but unable to compete in its
new environment).
lruJigenoUl. Native to the country or region.
Indumetnum, p. 49.
[ndupIicate. With the edges turned inward, and the external surfaces then coherent
or connate.
Indurate. Hard. as spin.. may become indurated (hardened) with maturity.
Jndusiate. Provided with an indusium.
Indusium, p. 30.
Induvime. Clothed with old and withered pans.
lnermis. Unarmed; not prickly; spineless.
Inferior Ovary. p. 76; origin of, pp. 7~78.
[nflesed. Turned abruptly inward.
Inflorescence, pp. 56-61; terms of, p. 57.
Infra-. Latin prefix, meaning below.
Infropetiolar Bud. described, p. 40.
Infraspecific Units of classification, pp. 24. 91.
Infundibular. p. 64.
Innate. Borne at the apex of the supportin, structure; an innate anther is one
attached at its base to the filament tip (i.... basi1ixed).
Innovation. A lateral branch or offshoot from the stem, usually restricted to
new growth.
InsecliverolU. Said of plants that capture insects.
Inserted. Growing out from, seemingly borne on.
Integument. A covering or envelop~ that which envelopes; specifically, the one
or two envelopes that covet (and are. a part of) the ovule; see p. 7S.
lme,.... Latin pretix, meaning between.
International Code of Botanical Nomenclature, p. 89.
Inlernode. That part of an axis between two nodes; see p. 36,
Intra-. Latin prefix, J;Ileaning within.
In/roTle, p. 68.
Invalid names, p. 90.
Involucel. A secondary involucre, as the umbellet of the Umbelliferae.
Involucre, p. 59.
Involute. Rolled inwards from the edges, the dorsal surface outermost.
Iridaceae. described. p. 12S.
Iris family. described. p. 12S.
l"egui.tu Flower, p. 62.
Isoetaceae, p. 28.
Isomerous. The same number in successive whorls.
Isotype. A specimen of the same collection as the holotype.
GWSSARf AND INDEX 165
Jepson, W. L., pp. 116-117.
lolnted. Separable into pieces at one or more points.
Ju&1andaceu. described. p. 127.
l",um. Latin, meaning a pair, as of leallets.

Kahn, p. p. 104.
Karyotype. The nature of the chromosomal complex characteristic of the indi-
vidual or taxon.
Keel. A ProjectinB lonaitudinal ridae, like the keel of a boat; carinate.
Keel Petal, p. 63.
Kermulne. Carmine red.
Key, and use of, pp. 85-86.
Key, a samara·type of dry indehiscent winaed fruit, as in Ash (Fraxinus).

Labellum. The usually enlaraed petal of an orchid flower. See p. 125.

Labels, data for, p. 87.
Labiatae, described, p. 142.
Lacerate, p. 47.
Lociniate, p. 47.
Loctescent. Producina milky juice, as does the Milkweed.
LacticiJerolD. Latex-containing; yielding a milky juice.
Lacuna. A chamber, hole, or gap; an air space within a tiasuc.
Lacustrine. Belonging to or occurring in lakes.
LaevigaJe. Smooth, as if polished.
Lamarckism, p. 20.
umella. A flat plate, sometimes a partial or complete partition.
Lamellate Placentation, p. 74.
Laminate. Expanded and flat, as the blade of a leaf.
LanoJe, p. 51.
Laneeolale, p. 42.
Lanuglnow, Cottony or woolly.
Lanulose. Woolly. sometimes spelled /anast.
LappaceoWl. Reaemblina a Burdack. burt; retrotaely ecb.ina1e.
Lateral. Belonging to the side.
weT/tous. Brick red in color.
Lax. Loose in texture or arransement; the opposite of crowded.
Leaf apex, terms of, pp. 44-45; bases, terms of, pp. 4.5-46; forms, pp. 42-44;
margins. terms of, pp. 46-48; position, terms of. pp. 48-49; structure. pp.
40-41; surface, terms of, pp. 54-56; venation, pp. 41-42.
Leaflet, p. 40.
Lectotype, pp. 91-92.
Leaitimate DaJDe, p. 90.
Legume. pp. 80, 81.
Leauminosae, described, p. 134.
Lemma, p. 122.
Lenticd. Lens..sbaped cork.y dots, or pits on young bark.
Lenticular. Lens-shaped, usually biconvexly so.
Lepidote. Pro¥ided with small scurfy scales.
Liane (Uana). A woody tropical vine that climbs or clambers Over other plants.
See p. 36.
Ligneous. Woody.
166 GLOSSARY AND INDEX
Ligulate Floret. A flower of the Compositac family whose corolla is strap-shaped.
See p. 65.
Ligule. The membranous append'ge at the top of leaf sheaths of moat arassea. Se,
Fig. 44c.
Ligulijorm. Strap-shaped.
Uliaceae, described, p. 123.
Lily family, described, p. 123. ,
Limb. The e:<panded portion of a petal or leaf, especially of • pmopetalous coroI1a.
Lindley. J" p. 8.
Linear, p. 42.
Lineate. p. SS.
Lineawt•• Bearing fine 1ineIike Dlorklup.
LingulaJe. Tongue-shaped.
Linnaeus. C., pp. 7-8.
Lip. The priocipal lobel of a bi!abiate corolla or eal)'L
Littoral. Belonging to the shore.
Livid. Pale gray. Iead-colored.
Lob., p. 47.
Lobulate. Cut into ama11 lobel.
Locule, p. 69.
Loculkidal. p. 81.
Lodicule. One of the small lCilelike prcc:un outside the Jtamens of a Il'ass
Dower, believed to repRotut the vestiJia1 perianth remaina.
Loment, pp. 80, 81.
Lorate. p. 42.
Lotoid..e, p. 134.
LUlUlIe. Cresccnt-shaped.
Lupuli.... Resembling hops.
Lurid. Dingy. dirty.
Lusus. An abnormal variation Or sport, a mutant form.
Luteolus. Yellowish, pale ye11o",.
Lycopsid. A taxon of the LycoPodium alliance, a club-moss or relative of it.
Lycopsida, p. 27.
Lyrate. Pinnately lobed, witb the terminal lobe tbe iarsest and laterala bec:omlq
progressively smaller toWard the base.

Macrosporangium. The sporc~ that produces macrosporea.

Macrospore. See Megaspore, p. 1.7.
Maculate. Blotched or spotted.
Madder family, described, p. 143.
Mailing specimens for identification, p. 87.
Mallow family, described, p. 13S.
MaJpighiaceous Hairs. Hairs that arc straight but whicb are attacbed at the middle,
often found on members of the Malpighiaceae, but also on plants of other
families.
Malvaceac, described, p. 135.
Mamillate. Bearing teat-shaped protuberances. ,
Manubrium. The constricted stalkuke basal portion Qf boat-shaped spathes in some
palms.
Marcescent, p. 51).
Marginal Placentation, p. 71.
Marmorate. Marbled.
GWSSA.RY A.ND INDEX 167
MarshaU, Humphrey, p. 104.
Marshall. Moses. p. 104.
Maxon, W. R .• pp. 117-118.
Megasporangium, p. 21.
Megaspore. p. 27.
Megasporophyll, p. 6!t
Membl'anolU~ p. 56.
Mer;carp. p. 81.
Meristem. An embryoDic u..ue of ceIIJ capable of dividinS aod produeinl DeW
crowth•
M.rrill. E. D .• p. 108.
M<socarp. Th. middl. I.y.r of • peri<:arp (ov.ry or fruit wall).
Mesophyte. Plants of medium moisture and Jight requirements. iDtermed.iate ...
tween those of very dry and very wet environments.
Mich.ux. Andre. p. 106; Mich.ux. F. A .• pp. 106-107.
Micropyle. pp. 75. 82.
Mlcrosporangium, p. 27.
Microspore, p. 27.
Mlcrosporophyll. p. 33.
Midrib. The middle or main nb of Ileal.
Milkweed family. described. p. 140.
Mimosoid.... p. 134.
Miniote. Dull vermilion, the color of red lead.
Mint family. described. p. 142.
MitTI/orm. Mitre-shapedj like a peaked cap.
Modified stema, types of. pp. 37-39.
MOlUJdelphora. p. 67.
Moniliform. p. 52 and Fig. 53,.
Monocorpic. pp. 37. 56.
Monocephalic. Bearing one head, often ODe headlikoe infiorescence.
Monocharium. A one-branched cyme, usually resuJting from reduction of laterals,
sometimes appearing raceme·lik~ as in lily-of-the--valley (Conl'allaria).
Monocltwmydeow. A perianth of a single envelope or whorl, i.e. of calyx or
corolla.
Monocolpate. One-furrowed, as the pollen arains of many monocots. See p. 69.
Monocotyledon•••• pp. 12G-126.
MOIIOecious, p. 61.
Monogynous. A flower boarinl a sinale pistil.
M01JOpetolou.r. Th. petals fused into a slnal. unit (obsolete).
MOllophy/elicism, p. 20.
Monotypic. A Benus of one species.
Morrison. Robert, p. 103.
Motile. Said of sperm when provided cilia, lIasell.. or other locomotion acces-
sories.
Mucok.J. p. S6.
Mucro. p. 45.
Mucronate. p. 4S.
Mucronulale. Diminuitive of mucronate. See Fia. IS h.
Muhlenberg. G. H. F .• p. 106.
Multicarpellate. A pistil of many carpels; sometimes used for a lYDoecium of many
pistils .
•1Iu/licipita/. Many headed.
 168 GLOSSARY ..4l'ID INDEX
Multifid. Cleft into many ......en.. of lobes.
Multiple Fruit, p. %G.
M.uTica/e. p. 55.
Muri/orm. Resembling courses of bricks in a walL
Mustard family, descnbed, p. 131.
MUlOIion, pp. 19-20.
MUlicous. Blunt, lacking a point.

NaJced Flo", ... One lacking a periaDth; a nsked bud it OlIO W:kin, bud _
Names, capitalization of, p. 95; conservation of, p. 91; when iIIeal!imate, pp. 94-95;
when rejected, pp. 94-95.
NQpi/onn. Turnip-shaped.
Nascent. In the act of being formed.
National Herbarium, pp. 117-118.
Natural Selection, p. 19.
Naturalized. Introduced from • foreign coun\Jy and ealabliabed natntally In dIO
new environment.
Navicular. Boat-shaped, as the glumcs of most grasscs.
NectariferolU. Having onc or more nectaries.
Nectary. A nectar-secreting organ. See p.- 66.
Nemorose. Inhabiting grovcs.
Neotype, p. 92.
Nerve. A vein. &Ii of a leaf-blade, perianth part, or carpe!.
Netted. Said of veins when branched to form a. network..
Neutral Flower. A "flower" having neither stamens Dor pistDa; sal...
New York Botanical Garden. p. 112.
New York &chootof taxonomy. p. 112.
Nightshade family, descn"bed, p. 142.
Nigrescenl. Becoming black, or nearly so.
Nltid. Shining.
Niveow. Snow white.
Nocturnal. Opening at night. See p. 56.
Nade, p. 36. •
Nodule. A small knob or knot.
Nomen conscrvandum, p. SIt.
Nomenclatore, p. 88; of cultivated plan... p. 97; de8necI, p. 2-
Nuce//w, p. 75.
Nuciform. Nut-shaped.
Nude FIo",er, p. 61.
NUl, p. 81.
Nutlet, p. 81.
Nuttall, T., pp. 107-108.
Nyctl-. Latio prefix denotiog night.

Ob-. A Latio prefix, signifying over, against, inverted.

otconical. Inversely tonical. attached at the apex of a cone-sbaped structure.
ObcordOle, p. 45.
ObdipJOS1emonous. Having the stamens in two whorls, those of the outer whorl
opposite the petals, and those of the inner whorl alternate with the petals.
Ob/anceolate, p. 43.
Ob/Ole. A sphere that has become Ilattened at the ends, .. in a tangerine.
Oblique, p. 46.
 GWSS..4RY AliD INDBlC 169
Oblo"" p. 43.
Obovate. p. 43.
Obovoid. A solid having the form of aD iIlvuted ell·
Ob.oIellCelll. Vestigial, nearly _ I .
Obsolete. Not evident, extinct.
Obtuse, pp. 45, 46.
Ochrea, p. 12~.
OchroleucolU. Yellowish white to buff colored.
OCTtolae. Small secondary leafooSbeaths, as in Eriogollum.
Odd-pi1lll4J•• Pinnately compoUnd with an odd number of leaftets. S •• p. 40.
Oligo--. A Greek. prefix, sisnifyinl few, as oligospermows (few seeded).
OlivaceolU. Olive-green.
Onaaraceae. described, p. 137.
Opaque. As applied to a surface- one that is dull and Dot ahinina.
Operculote. Provided with 11i~ or opel'CulPDL
Dperculum. p. '81.
Opposite, p. 49.
Orbicular, p. 44.
Order. discussion of. pp. 21. 22-23.
Orpin. family. descriptinn of. pp. 131-132-
Orrho.-. Greek prefix for .lrai.....
Orthotropous. p. 75.
Osseous. Of a bony texture.
Ovary. origin of, pp. 69-74; position, 76-7&.
Ovate, p. 42.
Ovulol•• BoarinB ovules, OVuUlVOlU • synonym.
Ovule, pp. 75-76.

PoIot,. A projection or IatruditJI fold that partially doses the throat of • cotoll•.
S.e p. 65.
Palea. p. 1Z2.
Paleaceoll4. Provided with chUf, or chaJIy in texture•
. Paleobotany. Fossil. botany, the .tudy of plants as they occurred in the geologic
past.
Palmate Venation. p. 42.
Palma/ely Compound. p. 40.
Palmalilid. p. 48.
Paludose. Inhabiting marshes.
Pandurale, p. 44.
Panicle, p. 60.
Pannose. Covered with a f.lt of woolly haiti.
Papaveraceac, described, p. 131.
PapillDniJuous. p. 63.
Papllionatac, see Lotoideae, p. 134.
Papillal•• p. 55.
Pappus, p. 63 and Fig. 42.
PtUallel Venation, p. 42.
Parasite. Botanically, a plant livinS on ud deriving nourish.ment from another
plllD!.
Paridal Placentation, p. 73.
Parlplnnate. Pinnate with an even Dumber of leallets.
Parted, p. 48.
 170 GLOSSARY AJIID INDEX
Parth~nogeM3is. The development, in pIanta, of a sporophyte from a female aameto
without fertilization or aYD.amy.
Patelliform. Rather thickly dise-shaped, lik.. the Im....,ap (paterla).
Patem. Open, spreading.
Pea family, described, p. 134.
PecliMte. Divided like the teeth of a comb.
Pedale. Like a bird.. foot, sometimes treated as a synonym of palmate.
Pedicel. p. S7. I
,Peduncle. p. S7•
. Peg. A stalk, applied to Ibe ,tall: of an ovary or fruit wben DOt composed of
ovarian (carpellary) tissue, as in the peanut (ANlGh!.).
Pellucid. Clear, transparent or nearly so.
. Peltate, p. 46.
Pendulou.r. Somewhat droopina.
I'enicillate. Tipped wilb a tuft of fine bain, as Ibe IIiJmas of IOIIlO gr......
Penninerved. Pinnately veined. See p. 42.
Pema-. A Gr..t prefix, ,isnifyinl five.
Pentamerous. In unita of five or multiples of five.
Pepo. p. 80.
Perennate. Lastina the wbole year Ihrouah; ..If·reDtwin. by lateral shoots from
the base.
Perennial, p. 37.
Perfect Flower, p. 61.
Perlollate, p. 46.
PergamlaceolU. Parchmental.ike.
PerianJh. pp. 6~3.
Perigynium. p. 122.
Perigone. A synonym of perlantb.
Perigynous. Borne or arising from around the ovary and DOt beneath it, as when
pcrianth segments and stamens scem.inaly arise from the edge of a cup-
shaped hypanthium; such cases are said to exhibit perigyoy.
Periphery. The outer wall or margin; the inner face or side of the ovary wall, as
opposed to Ibe faces of ita septa.
Persistent. Remaining attached; not faUiog all.
Personate. Said of a two-lipped corolla, Ibe Ihrost of which is closed by a palate,
as in toa~ See p. 65.
Perfuse. Perforated with a hole or slit.
PeruloJe. Scale--bearinS, as most buds.
Petal, p. 63.
Pdalody. Th. metamorpbosis of stam_ into pew., etc.
Petaloid. Petal-like; in color and shape reoemblinl a petal.
Petiole, P. 40.
Petlolule. p. 40.
Phanerogam. A seed plant or spermatophyte, as opposed to a cryptogam.
Phenotype. A plant or population determined by ita appearance, as opposed to
a senotypc, which is determined by genetic characters.
Phloem. A complex tissue of the vascular anatomy containing sieve tubes, phloem
parenchyma, and other elements such as fibers, stone cells, companion
cells, etc.; the inner bark of a woody plant; the outer two tissuea of a
vucular ,trand (th. xylem the inner on.).
Phy/Jary, p. 60.
GWSSARf AND INDEX, 171
Phyl/OCllld•• A branch, more or lea lIattened, fuDcIionln.I u • leaf, as in ChrisImu
caduI.
Phylllldium. LealIike petiole with no bWlo. as in lOme aA:8CW and tome other
planll.
Phyllolaxy. Tho arran.......,t of J••vea or floral porta OD an axis.
Phylopnetio classilioatioD&, discussed, pp. 100-101.
Phylogeny. functions of. pp. 98-101; .ystems of. p. 9.
Phylum. On. 01 the primary divisioos of the plant or anima1 I<in,dom whose
compoDenll are presumed to have had • common anceolry. S•• pp. 21-22.
PUo... p. 51.
Pink Iamily. described, p. 130.
Pinna. p. 31.
PhuuU~pp.4O.41.4~
PhuuUifiij. p. 48.
PhuuU16ecl. Cut dowD to the midrib in • pinnate IuhIon.
Plnnule. p. 3 I.
PUciform. Pea.peeI.
PUtII. pp. 69-70.
Pl.stillate. Femalej haviD, one or more pistils and no functional stamens.
Pinll/ode. A rudim.ntery or veatiaial pistil pr....,t in tom. staminate flowers.
Pllh. Th. 10ft IpODJY central eylincler of parencbyma tissue in most woody 8DJio-
oporm _ .
P/ocento. p. 70.
Ploc.ntDlion 'Typu. pp. 7G-74.
Plant claasification, pp. 2, 6-18.
Plant idonti1leation, procedures. p. 16.
PIIcDl•• Folded, as in a fan, or .pproaclllnJ this condition.
Plu""",e. pp. 52. 53.
Plumule. Of & yaunl plant, the fint ahoot abo.. the cotyledons.
Pod. A doblsc:ent dry fruiL
Poll.,. 0",1118. pp. 68-69.
Pollen &Ie. The microspor8DJium. ooDteiDiDl the po1lon; in moat 8DJiooporma oacb
anther is oomposed 01 four poneD ..... two in oacb lobo or half of the
antherr tho tissues separatin, them disintearatina: prior to antheais and the
resullinl anther _minaJr lWcKeUed or biloculate.
Pollination. Th. transfer of poU.., from the dehiacin, anther to the receptive
stiJma; the act of poUiD.1iDJ.
Polllnium. p. 69; in orchids. p. 126; in Aaclepiadaceae. p. 140.
Polyandry. Th. produ<linn of an indefinite Dumber 01 stamens in an androecium.
Polycarpic. p. 56.
Polygamodlo<clous. Said of a .peei.. th.t ... functionally dioeclous but bas • lew
flowers of the opposite aex or a few bisexual dowers on all plants at flower-
in, time.
Polygamous. Bearing unisexual and bisexual Bowers on the same plant.
POlyJOD.C.... described, pp. 128-129.
Polyp,'alous. p. 6J.
Polymorphic. Represented by two or more forma. as is a species 01 several closely
related infraspecific taxa.
Pol)lphyl"i3. A aituation represenlin,. polyphyletic orisin (... p. 20); that is. the
result of evolution alona two or more lines, havin, different origins.
Polyphyletic, p. 20.
17% Gwss.m.r AJIID INDEX
Polyploid. Having a chromosome complement of JIlOII) than two _ of tho mono-
ploid number.
Polypodiaceae, p. 120.
PolysepaJous. Having a calyx of separate sepals.
Pome. p. 80.
Poppy family, dcscnbed, p. 131.
POTickiol dehiscence. pp. 68, 81~
Preservation of plant specimens, pp. 86-87.
Pressing plants, p. 86.
Prickle. A small weak spinelike body on the bart or epidermis.
Primitive vs. advanced characters, pp. 99-101.
Primordium. A group of undifferentiated meristematic cells capablo of differentiat-
iag into various kinds of organs or tissues.
Primulaceae, described, p. 139.
Priority of names, p. 91.
Procumbent. p. 36.
Prostrale. A general term for lying flat on the around.
PrOlamirous (ProJerandrolR), p. 69.
ProtantheroU8. p. S6.
Protogynous. A flower whose stigma is receptive to pollen before pollen fa abed
from anth~l's of tho same flower. See p. 69.
Pruinose, p. 56.
Pseudoterminal Bud. p. 39.
Pseudobulb. The thickened or bulbiform stents of certain orchids, beIDa BOUd and
borne above the ground.
Psilopsida, p. 27.
Pteropsida. pp. IS, 17.
Puberulenl, pp. SO, 54.
PUbeTJJlous, p. SO.
Pubescent, p. SO.
Pull., A., p. 16.
Pulverulent, p. 56.
Pulvinate. Cushion-shaped.
Pulviniform. Havins the shape of a pulvinus; pad- or cushion_pod.
Pulvinus. A minute gland or swollen petiole {or petiolule) base responsive to
vibrations, beat, or some other stimuli, as in the leaves of Sensitive plant
(Mimosa).
Punctate, p. SS.
Pungent. Ending in a stiff' sharp point or tip; also, acrid to the taste.
Pursh, F~ederick, p. 107.
PustuUJr. Blistery, often minutely so.
Pyrene, p. 82.
Pyriform. Pear..hapecl.
Pyx;', p. 81.

Roceme. p. 60.
Racemose. Having flowers in raceme-like inflorescences that may or may not be
true racemes.
RachiUa. A secondary or diminutive axis, or rachis; in partleular, in the arasses
and sedges, the fIoret-bewg axis-
Rachis, p. 31.
.GWSSARY AND INDEX 173
Radiate. Standin, on and spreadina from a common center; also, having ray
lowers, as in the Compositae.
Radical. Said of leaves that are basal or in rosettesj lU'isinl from the root or ita
crown.
Radicle. The embryonic root of a aerminatin. oeed. See p. 82.
Ratlnesque, C. S., pp. 108-109.
Rami/orm. Branchina.
Rank. A vertical row; leaves that are two-ranked are in two vertical rows.
Ranunculacoae, described, pp. 130, 131.
Raphe. That portion of a fw:lic:ulus of an ovule that is adnate to the integument.
usually represented by a ridse, present in most anatropoWi ovules, diag-
nostic in Sa"acenia. See 4lso p. 82.
Rophid." p. 122.
Rattan, Y., p. 116.
Ray, 101m, p. 6.
Ray. Outer modified IIoret ot some compoaites, with an extended or strap-shaped
part 10 the corolla; aIao a branch (pedicel or peduncle) of an umbel or
umbel-like inIIoreacenco.
R«,ptacl,~ p. 60.
Re"'p/ocuwr Theory, p. 78.
Reell_e. Bent down or fallin, back from the perpendicular.
Reflex.d. Abruptly bent downward or backward, usually a 180' change of direc-
tion, or nearly 80.
Regular Flower. A aymmetrical flower with the parts in each series or set 80
arranged as 10 be divisible vel1ically inlo equal halves by two or more
planes.
Reniform. Kidney-shaped. See p. 44.
Repand. Weakly sinuate.
Repelll, p. 36.
Replurn. Partition between two loculi of cruciferous fruits. Se, Cruel/era,.
Resinous. Containing or producing resin, said of bud scales when coated with
a sticky exudate of resin (as in some .A.uculus spp.).
Resupinate. Twisted 180°, as the ovary (and tlower) of most orchids; upside down.
Reticulille. Netted. See p. 54.
Rericulote VeMtion, pp. 29, 31.
Retrorse. Bent or turned baU.ward or downward.
Retuse, p. 45.
Revolute. Rolled backward; with marain rolled toward lower side.
Rhitoid. A rootlike structure in function and general appearance, but not so in
anatomy.
Rhitomatous. Producing or possessing rhizomes.
Rhizome, p. 37.
Rhizophore. A leafless stem that produces roots, as in Setag/nella.
Rhombate, Rhomboidlli. Shaped like a rhomboid. See p. 44.
Rib. Any promint;D.t nerve or vein in a leaf or similar orpn.
llobinson, Benjamin L., p. 111.
Rodgers, A. D. lII, botanical biographies by, p. 118.
ROTlduwle, p. 56.
Rosaceae, described, p. 133.
ROItellmn. A small beak.
Rostrate. Havina a beak or beaklike projection.
174 GLOSSARY AND INDEX.
R""uldt•. In rooetIeI.
ROlate Corolla, p. 64.
Rotund. Orbicular, ineliaiD, to be oblona.
Rubiceae, described, p. 143.
Rudiment",.,. lmperfoctly developed, v~.
Ru_e,p. '4.
Rumina... Mottled in appearance; a tiau. of daB UIcI lIal!t _ dl irrquIar
outline.
Runel""'e, p. 43.
Ru1l1lllr. A _ tralliua obooI, rootin& al the DOdeo.
llydber.. Per Axel, p. 113.

Saccat•• Ba.-obaped, pouchy.

S.,Itt"'e, p. 46.
Solie...ae, <iescn'bed, p. 117.
Sa(verfomt) 9 .. 64,
Salviniac:eao, pp. 28, 110.
S.mara, p. 81.
Saprophyte. A plant (usually lKkinI chlorophyll) llvina OIl dead oqat>Ic ma_.
Sarment""e. ProduciniloDa lIoxuous 1'1JJIIIm or atolou.
Saxifr_ described, p. 132.
Scabrescent, p. 52.
ScabroUl, p. S2.
k.le. Any smaIl, usually dry, _reooed IeoI or b,""" often 0DIy VCIIlPaL
&:adent. Climbin..
Scape. pp. 57, 61.
Scapo.re. pp. 36, 6t.
ScariolU, p. '6.
Scbizaeaceae. p. 120.
Schiwcarp, p. 81.
Schmaltz, C. RaftDeoque, pp. 10&-109.
Scorp;oid Cyme, pp. 57. 59.
Scrophulariaceae, desen'bed, pp. 142.-'143.
Seillate. Lite a small shield.
S<cund. O .....ided, or aeeminaly 00.
Sed.. family, dcscn'bed, p. 122.
Seed. The ripened ovule; the essential part ia the embryo, usually contained within
one or two integuments. See pp. 82-83.
Segment. One of the diviaiooa of a leaf, peta1, sepal, or bract thaI ia divided but
not compound.
Sepal, p. 63.
Sepaloid. Said of an involucral bract or pets! thaI resembles a .epal.
Septate. Partitioned; divided by partitiona.
Septicidal. Dehiscence along or into the partition (seplum), not openinI direcdy
into the loeul•• Se. aLro p. 81.
Septum (pl. sepIa). A partition or cross·wall.
Serleeous. Silky. See pp. 51, 54.
Se""'e, p. 47.
Se"ulate, p. 47.
Sessile. Not stalked.
Seta. A briade.
Setaceous. Bearins bristlea.
GWSSARf AND INDEX 175
S~tose. pp. 52, 54.
Setultne, p. 52.
Shaw, Henry, p. 1U.
Sheath. Any Ion. mo~ or less tubular atructure. surrounding an orlan or put.
Shrub. A woody plant that remalns low and produces shoots or trunks from the
base; Dot treelike nor with .. smale bole; a descriptive term not subject
to precise circumscription.
Sigmoid. S-shaped.
SilkeolU. Containins minute particles of silica.
Silkle. Th. short fruit of certain Cruciferae, usually not more than twice as lana
as broad.
SilIqu•• The lona .Ieader fruit of certain Crucifer...
Sln"'e, p. 46.
S11IIII. The apace or recess between two lobes or divisions of a leaf or othc:r
expanded organ.
Small, 1. K., p. 114.
SoOOIlI"o"" p. 36.
Solanace8e, described, p. 142.
SOrl14 (pl. sori), p. 30.
Spadix, pp. 60, 122.
Spathoeeow. ROIemblina ur furnisbed with a apathe.
Spathe, pp. 60, 123.
SpoJh~Yalves. One or more herbaceous or acarious bracts that lubtend an inftorea-
cenee or fIowcr, and generally envelop the lubtcnded writ when in bud
Spatulale (Spathulale), p. 43.
Speoiea, pp. 22, 23-24.
Speoimcna, preparation of, pp. 8~7.
Spe,.m. A male gamete or reproductive cell.
Sph.nop,ld. Aoy of a &roup of l£qu/setum- or honetalI-lik. plants.
Sphenopsida. p. 27.
Spkate. Spit.lik•.
Spicule. A diminutive or aecondary apike.
Spike, p. 60.
Spikelet, p. 121.
Spine. A strona. sharp-polnted woody body mOlt1y arisinl from the wood of a
stem.
Spinucent. Tetminated by a spine, or bearin, apinelike teeth.
Spinulose. With small spinea over the surface.
SportJ1Jgium, p. 28.
Spo,., p. 21.
Sporocarp. A body or receptacl. containInB IPOl'RD8ia (&I in MarS/l.a).
Sporophyll. A rpore-bearin& leaf ( .. in ferns); a 1.a1lik. organ bearin& "'Pro-
ductive parts or organs.
Sporophyte, p. 21.
Spurge family, described, p. 135.
Squamate. With small scalelike leaves or bracts; scaly.
Squamellijorm, p. 52.
Squamose. Covered with small scales, more c:oanely so than when lepidote.
Stalk. The stem of any orpn, as the petiole, peduncle, pedicel, filament, ltipe.
Stalked Bud. On. whore outer scaI.. are attacbed above the b... of the bud
axis. See also p. 39.
StfllMn, pp. 6.5-69.
 C£OSS.4llF .lND INDB~

StlRniMt•• Furnished with stam..... but 110 functional platiI; male.

SttRninod., pp. 66, 68.
SttuUlard. The upper and broad, mote or 1... erect petal of • popillonaceoua tIower
(se. p. 63); the narrow, usually ezect, unit of the .biller ",riea of the
oerianth in an lru flower as opposod to the often dJoopinl falls. s.. p. 125.
Stellat., pp. 52, 53.
St.",.., pp. 35-39.
s.t6ile. Lacl<inJ any functional "'" or.....
Stigma, pp. 69, 74-75.
Stipe, p. 31.
Stipel. The lllipule subtendq a leallet.
Stiplde. A basal appeodage of a petiole; the thtee parts of a complete leaf ate
blade, petiole, aod stipules (usually two in number). S•• p. 40.
Stolon. A shoot that bends to the JOOund and takes root; mote commonly, a
borizontal stem at or below the surface of the JOOund that Ii- rise 10
• new plant at its tip; st%nijeTOIl3. See p. 36.
Sto_e. The pate in a leaf (usually on the lower side) formed by the c:oncavlly
of two facin, .ausage-like guard cells.
$10M. A large pytene; ..one fruit, a drupe or drupelet, as in Rosaceae.
Strrunineou.r. Straw colored.
Striat., p. 55.
Strict. Straight and upright. little if at all branched.
Strlgose, pp. $1, 54.
Strob& (mobilu.r), p. 33.
Styli. Within the style or stigma; pertainlng to the style.
Style, pp. 74, 75.
Styfopodium. A disclike enlargement at the base of the style, as in IIOIIle umbelllfen.
See p. 81.
Sub-. As a prefix usually .iJDifying someWhat. .lightly, or tathet.
Sub.""., p. 56.
Subinferlor Ovary, p. 76.
Subpetioll. Under the petiole b .... aod usually enveloped by it.
Subspecies, p. 24.
Subulate, p. 44.
Succulent. Juicy; fteshy; soft and thickened in texture.
SuDrutescent, p. 37.
Sukat., p. 55.
Superior Ovary, p. 76.
Superposed BUM, p. 40.
Supine. Lying ftat and with face upward.
Surface, terms of, pp. 54-56.
Suture. A line or mark of splitting open; a groove markinl a natural division
or union.
~conium. The "fruit" of a fig, actually an inverted and essentially closed receptacle
containing many achenes.
Sym-. &e under syn-.
Symmetrical. Said of an actinomorphic ftower that has the same number of parts
in each series or circle, as five petals, five stamens, etc.
Symph,m. The connation of like parts from time of meristematic development,
as petals in the pmopetalous type' of corolla. .
SympodiDl Inflorescence. A detennin~t~ in1lorescence that simulates BD: indetc:~.
nate inftorescence. as a SCOtplOld cyme (compare latter also With helICOid
cyme).
 CWSSARY AND INDEX 177
Syn-. A prefix from the Greek, meaning with. together, like. Syn- becomes sym-
before b, m, and p. and then has tho same meaning.
Synandrium. An androecium coherent by the anthers, as in some aroids; when
anthers are connate they are termed syngenesious.
Synantherous, p. 56.
Syncarpous. Having carpels united; applied to an ovary of t~'o or more carpels;
sometimes used when separate pistils within ODe flower are partially
united. See Apocarpous.
Syngenesious, p, 67.
Synonym. An equivaJent superseded name; a second name fOT a given taxon.
SYllpetaious. Gamopetalous; the petals marginally connate, at least basally.
Synscpa/ous. Gamosepalous; the sepals marginally connate. at least basally.

Tailed. Said of anthers having caudal (tail-like) appendages.

Taxaceae. p. 33.
Taxon (pI. taxa). A general term applied to any taxonomic element, popplation,
or group, irrespective of its classification level. See p. 13.
Taxonomy, defined. p. 1; in North America. pp. 103-118; opportunities in, p. 4;
significance of~ p. 3.
Tendril. A rotating or twisting threadlike process or extension by which a plant
grasps an obj~t and clings to it for support; morphologically it may be
stem or leaf or leaflet.
Tepai, p. 62.
Terete. Circular in transverse section.
Ternate. In threes.
Terrestrial. On the ground; a land plant, as opposed to aquatics. epiphytes. Of
saprophytes.
Testa. Outer coat of a seed (deve1oped from an integument).
TetTad. A group of fourj in angiosperms, a four-celled pollen-mother cell. See
p.69.
T etradynamou8, p. 67.
Tetramerous. Four·merous.
Tetrando"ow. With four stamens.
Texture, terms of, p. 56.
Thalamus. The receptacle of a flower; lbalamiftorae. a taXon whose floral parts
are hypogynous, distinct, and separate from one another on the receptacle.
ThnJJ= A flat J..rus. orgaA
Theca. The pollen sac of an anther. See p. 66.
Theophrastus. p. ?
Thorn. Same as spine.
Throat. In llowers. the opening or mouth of the tube of a gamopetalous corolla;
the place where the limb becomes differentiated from the tube.
Thyrse. p. 60.
Tippo, 0., pp. 16-18.
Tomentose. pp. SO, 54.
l'oJ'.'lenlu/o,u. p . .sO.
Torrey, John, p. J09.
Torulose. Twisted or knobby; irregularly swollen at cloee intervals.
Torus. Receptacle.
Tourne(ort, J. P., p. 7.
ransverse Dehiscence, p. 68.
i ree. A woody plant that produces one main tnmt or bole. and a more or leu
distinct and elevated head.
178 GLOSSARY AND INDEX
Trelease, Wm., p. 11S.
Tribe,p.22.
TrichOl]tt. A hair or bristle. See p. 49.
Trico/pate. Three·grooved. See p. 69.
T,JjoJiale. Three-leaved. as in Trillium. See p. 41.
'rri/oliolate. Having a leaf (or leaves) of three leaftets, as most clovers. See p ..i1.
Trigonal. Three-angled, without regard for the acuteness or bluntness of the anales.
Triquetrous. Three·angJl':d. and sharply so, the sides often som~what concave, as in
the stems of some sedges.
Trlternate. Three times tcrnate, as in some Meadow~rues or CtJlumbmes.
Trivial Name. The specific na.me, as opposed to the aencrlc name.
Truncale. p. 46.
Tuber. p. 38.
Tubular Corolla. p. 64.
Tumid. Swolle.. somewhat inlIated, as the portion 01 the trunk 01 the Belly palm.
Tunic. p. 38,
Turbinate. Inversely conical; top-shaped.
TUTgid. Swollen from fullness, but Dot usually distended.
Turion. A young sucker or shoot, as an emerginl stem of asparagus.
Twfg. A young woody stem; more precisely the current season'. growth of. branch.
Twining, p. 37.
Type Method, p. 91.
Typical element of • species, explained, pp. 95-96.

Umbel, p. 59.
Umbellate. With umbels, pertaining to umbels.
Umbellet. The secondary umbel of a compound umbellate inftoresceru::e. See p. $9.
Umbelliferae, descn1>ed, pp. 137-138.
Umbo. A small conical projection from the surface; a boss.
Umbracu[iform. Umbrella-shaped.
Uncinale, pp. 52, 53.
Underground Sterru. illustrated. ". 38.
U ndukue. p. 46.
Unguiculate. Narrowed into a claw or petiole-like base.
Unigeneric. Said of a family composed of a sinale seous, as Ginkgoaceae.
Unijugau. Said of a comt"'und leaf composed of one pair of leaflets.
Unilocular. Containing & singJe chamber or "cell."
Units of claasification, discussed" pp. 21-24.
Urceolate, p. 64.
Utricle. p. 81.

Vaginate. Sheathed.
Valid Name, p. 90; valid publication, p. 92.
V (J/1ecu/ar. Pertaining to the grooves between abe rld,es. as in buita of man),
Umbelliferae.
Valvate. Opening by valves; meeting by the edges without ovedappm,.
J'a/ve. A separabJe part of II pod; the units or piccl.3 into wbicb a capsule splits
or sep irates on dehiscing; a flaplik.e cover or lid as in some stamens.
Jlariety (varirtas), pp. 22, 24, 95-96.
Yascular. Pertaining to the presence ot vessels in the conducting tissues of the
stele.
Vascular cryptogams, pp. 27-31.
GWSS.4RY AND INDEX. 179
Vasey. George, p. 117.
Vad/ot'm. Of elongated funnel shape.
Ve/ulinolU, PI'. 50, 54.
Venation, types of, pp. 41-42.
Ventral, p. 71 (footnote); ventral placentation, p. 71.
Ventricose, p. 65.
Verbenactac, described, p. 141.
Verrucose. Having a wartlike or nodular surface.
Versatile. p. 66.
Verticel. A whorl; in the Mint family. a false wborl of flowers that ~e in cymulcs.
Verlicillute. Arranged in whorls, or seemingly but falsely so. See p. S9.
Vesicle. A small bladdery sac or cavity filled with gas or fluid.
Yestiture Types, pp. 5~54.
Vexillum. The broad upper petal (standard) of a papilionaceous torolla.
Villous, pp. 51, 54.
Violaceae, descnDcd, pp. 135-136.
VirgaU. Wandlikc; Jon& straight, md slender.
Viscid. Sticky. or with appreciable viscosity.
Voluble. Twinina.

Walnut family, described, p. 127.

Walter, Thomas, p. 104.
Watson, Sereno, pp. 110-111.
WeJwitschiales, p. 32.
Wettstein. R. von, p. IS.
Whorl. Three or more leaves or flowers at a node. See p. 49.
Wing Petal. Two of the lateral petals c.t a papilionaeeous flower. See p. 134.
Woolly, pp. 51, 54.

Zygomorphic, p. 62.

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