Manual of Leaf Architecture

Authors Beth Ellis Douglas C. Daly John D. Mitchell Kirk R. Johnson Leo J. Hickey Peter Wilf Scott L. Wing

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Manual of Leaf
Architecture

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Copyright © 2009 Cornell University.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
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Manual of Leaf
Architecture
Beth Ellis
Douglas C. Daly
Leo J. Hickey
Kirk R. Johnson
John D. Mitchell
Peter Wilf
Scott L. Wing

Published in Association with The New York Botanical Garden

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First published 2009 by Cornell University Press
First printing, Cornell Paperbacks, 2009

Printed in the United States of America

Library of Congress Cataloging-in-Publication Data

Manual of leaf architecture / Beth Ellis ... [et al.].
p. cm.
Includes bibliographical references and index.
ISBN 978-0-8014-7518-4 (pbk. : alk. paper)
1. Leaves--Morphology--Handbooks, manuals, etc. 2.
Leaves--Anatomy--Handbooks, manuals, etc. I. Ellis, Beth. II.
Title.

QK649.M326 2009
575.5’733--dc22

2008044216

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Introduction 1

Leaves
General Leaf Def initions 4

Leaf Characters 12

Veins
General Vein Def initions 44

Determining Vein Order 47

Vein Characters 57

Teeth
General Tooth Def initions 101

Tooth Characters 103

Appendix A: Outline of Characters and Character States 113

Appendix B: Examples of Fully Described Leaves 118

Appendix C: Vouchers 156

Appendix D: Instructions for Clearing Leaves 176

References 178

Index 188

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About the Authors
Beth Ellis is a Research Scientist at the Denver Museum of Nature & Science.

Douglas C. Daly is Director of the Institute of Systematic Botany
at The New York Botanical Garden.

Leo J. Hickey is Professor of Geology at Yale University and Curator of Paleobotany
at Yale Peabody Museum of Natural History.

Kirk R. Johnson is Vice President of Research and Collections and Chief Curator
at the Denver Museum of Nature & Science.

John D. Mitchell is a Research Fellow at The New York Botanical Garden.

Peter Wilf is Associate Professor of Geosciences at Pennsylvania State University.

Scott L. Wing is Research Scientist and Curator in the Department of Paleobiology
at the Smithsonian Institution.

Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Acknowledgments
KRJ and BE acknowledge support from the National Science Foundation under Grant no.
0345910. LJH acknowledges support from the National Science Foundation under Grant no.
0431258 and from Yale Peabody Museum. PW acknowledges support from the David and Lucile
Packard Foundation. Insightful reviews were provided by Robyn Burnham and Lawren Sack.
The authors thank the following individuals for their support in producing this document:
Bobbi Angell, Amanda Ash, Richard Barclay, Ellen Currano, Regan Dunn, Richard Ellis,
Carolina Gómez-Navarro, Katherine Kenyon Henderson, Fabiany Herrera, Rebecca Horwitt,
Carol Hutton, Ramesh Laungani, Stefan Little, Mandela Lyon, Dane Miller, Ian Miller,
Amy Morey,Daniel Peppe, Sandra Preston, Mary Ellen Roberts, Dana Royer,
and Caroline Strömberg.

Illustration credits
The following figures were drawn by Rebecca Horwitt: 1, 8, 10, 14, 22,
23, 24, 25, 26, 27, 28, 29, 30, 32, 33, 34, 35, 36, 37, 38, 39, 40, 42, 43,
44, 45, 46, 47, 48, 49, 50, 84, 85, 86, 87, 125, 126, and 128.

The following figures were drawn by Amanda Ash: 2, 17, 92, 244, 297, 298, 299, 300, and 301.

The following figures were drawn by Bobbi Angell: 13, 124, and 127.

The following figures were provided courtesy of The New York Botanical
Garden: 15, 16, 131, 153, 193, 200, 208, 217, 227, 228, 230, 233, 250, 251, 253,
254, 276, 292, 303, 304, Appendix 6, Appendix 16, and Appendix 17.

Figure 51 photograph courtesy of Dennis Stevenson.

Figure 81 photograph courtesy of Dana Royer.

Appendix 18 leaf photograph courtesy of the Denver Museum of Nature & Science.

Other images are included by courtesy of Smithsonian Institution and Yale Peabody Museum of
Natural History, being either photographed from the Smithsonian Institution collections that are
currently housed at both institutions, or reprinted from the
1999 edition of the Manual of Leaf Architecture.

Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Introduction

S ince the time of Linnaeus, compara-
tive analysis of reproductive characters
has been the principal morphological
technique for identifying and classifying an-
giosperms (e.g., Takhtajan, 1980; Cronquist,
tion, diversity, and paleoecology of past floras
(Chaney and Sanborn, 1933; MacGinitie,
1953; Burnham, 1994; Johnson and Ellis,
2002; Wang and Dilcher, 2006). Furthermore,
fossil leaf morphology is widely used to pro-
1981). The Linnaean system and its descen- duce estimates of paleoclimatic and paleoen-
dants have been very successful, but there are vironmental conditions (Bailey and Sinnott,
compelling reasons to increase the use of foliar 1915, 1916; Chaney and Sanborn, 1933; Wolfe,
characters in angiosperm identification and 1971, 1995; Utescher et al., 2000; Jacobs and
systematics. For example, living tropical plants Herendeen, 2004). Fossil identifications, in-
may flower and fruit infrequently, and repro- cluding those based on leaves, are also used
ductive organs may occur only high in the to estimate divergence times of clades (e.g.,
canopy when they are present, making foliar Richardson et al., 2000, 2001; Renner, 2004;
characters more practical for field identifica- Davis et al., 2005; Uhl et al., 2007).
tion (Gentry, 1993). Even when reproductive
organs are available, foliar features can provide Working with isolated fossil angiosperm leaves
information that enhances systematic analyses is a long-standing challenge in paleobotany.
(Levin, 1986; Keating and Randrianasolo, Late-nineteenth- and early-twentieth-century
1988; Högermann, 1990; Todzia and Keating, paleobotanists left a legacy of poorly defined
1991; Seetharam and Kotresha, 1998; Roth, taxa. Most early workers had neither an accept-
1999; González et al., 2004; Martínez-Millán ed lexicon for describing leaf form nor knowl-
and Cevallos-Ferriz, 2005; Wilde et al., 2005; edge of how leaf features are distributed among
Gutiérrez and Katinas, 2006; Doyle, 2007; living angiosperms (see discussions in Dilcher,
Manos et al., 2007). 1973; Hill, 1982, 1988). They focused mostly
on shape, size, and generalized vein charac-
One of the most critical uses of foliar char- ters that failed to discriminate species or even
acters is in interpreting the angiosperm fossil higher taxa accurately and routinely applied
record. Although fossil reproductive struc- names of living genera to fossils from unrelated
tures comprise an important source of data fossil genera based on poorly preserved leaves
(e.g., Friis and Skarby, 1982; Basinger and without diagnostic characters. Thus, modern
Dilcher, 1984; Herendeen et al., 1999; Crepet workers inherited a host of misidentified fossil
et al., 2004; Friis et al., 2006), compressions species incorrectly described as Ficus, Populus,
and impressions of leaves are the most com- Aralia, and other modern genera.
mon macroscopic angiosperm fossils. Because
of their abundance, fossil leaves provide a Two recently developed approaches address
great deal of information about the composi- some of these problems. One method is the

study of multiple organs, including leaves, architecture is a major goal, but not one that
thought to represent the same plant species ei- we undertake here.
ther because they are preserved in attachment
or because they co-occur at many fossil locali- The purpose of leaf terminology is to allow
ties (e.g., Dilcher and Crane, 1984; Crane and objective and reproducible description and
Stockey, 1985; Manchester, 1986; Boucher comparison. Descriptive systems have a long
et al., 2003; Manchester et al., 2004, 2006; history, which we acknowledge but do not
Zamaloa et al., 2006; Manchester and Hickey, attempt to review (e.g., von Ettingshausen,
2007). Traditional characters of flowers and 1861; Kerner, 1895; Lam, 1925; Melville,
fruits can thus be used along with leaves to 1937, 1976; Mouton, 1966, 1967; Hickey, 1973,
define extinct taxa and determine their rela- 1974, 1977, 1979; Dilcher, 1974; Dickinson et
tionships. However, most fossil leaf species are al., 1987; Pole, 1991). The system presented
found neither attached to, nor consistently as- here is a revision of Hickey’s leaf architectural
sociated with, other organs. terminology, which in turn is loosely based on
that of von Ettingshausen and has been used
The second approach identifies systematically extensively to characterize fossil floras.
informative characters of extant leaves that
allow taxonomic affinities to be recognized Fully quantitative methods for describing leaf
solely on the basis of isolated fossil leaves shape exist (e.g., Jensen, 1990; Ray, 1992;
(MacGinitie, 1953; Dilcher, 1974; Hickey and Meade and Parnell, 2003; Royer et al., 2005)
Wolfe, 1975; Wolfe and Wehr, 1987; Hickey and are presumably more objective than the
and Taylor, 1991; Meyer and Manchester, qualitative and semi-quantitative terms de-
1997; Candela et al., 1999; Meyer, 2003; scribed here, but leaf shape is well known
DeVore et al., 2004; Fuller and Hickey, 2005). to have a high degree of convergence among
This method, which has been used principally unrelated lineages (Doyle, 2007; Little et al.,
for angiosperms with net-venation, is our fo- 2007). Leaf venation holds many systematical-
cus here. ly valuable features, and although techniques
exist for quantifying the overall properties of
Foliar characters may or may not offer con- vein networks (Bohn et al., 2002; Couder et
clusive evidence of the generic or higher-level al. 2002), these have not yet been applied to
affinities of living angiosperms, but gener- problems in leaf identification or botanical
ally they allow closely related taxa to be dis- systematics. Thus, in spite of recent advances
tinguished from one another (e.g., Merrill, in quantifying leaf morphology, we think the
1978; Sajo and Rudall, 2002; Espinosa et al., descriptive system presented here will remain
2006). Doyle (2007) examined leaf architec- useful because it generates consistent results,
tural characteristics within the framework it can be applied to partial specimens such as
of molecular phylogenetic analysis to further incomplete fossil leaves and “sterile” extant
highlight some evolutionary trends across the plants, and it does not require intensive image
angiosperms. processing.

Cuticle is often preserved with leaf fossils. The potential contributions of leaf architec-
Analyzing the characteristics of leaf cuticle in ture to paleobotany, ecology and paleoecology,
combination with leaf architecture is a power- plant systematics, and conservation are only
ful tool for identifying species (Dilcher, 1963; beginning to be realized. The development
Dilcher and Crane, 1984; Upchurch, 1984; of interactive, image-driven keys emphasizing
Jacobs and Kabuye, 1989; Pole and MacPhail, leaf characters will lead to an expanded use
1996; Conran and Christophel, 1999; Kvac̆ek of this rich source of characters and charac-
and Manchester, 1999; Barnes et al., 2001; ter states for describing plant form and struc-
Carpenter et al., 2004), but synoptic guides ture. The purpose of this guide is to provide a
to cuticle classification are currently lacking. clearly defined and illustrated set of terms to
Integrating cuticular information with leaf support wider use of venation features. We look

to future work to elucidate not only the origin We intend this work to stimulate diverse re-
and directionality of the different vein orders search, including deeper studies of the taxo-
(Dimitriov and Zucker, 2006) and leaf shapes nomic distribution of character states; appli-
(Hay and Tsiantis, 2006), but also the function- cations of leaf characters in systematic studies
al ecological relationships among the character of a broader range of taxonomic groups; ap-
states described here (Roth et al., 1995; Roth plication of matrices of the characters in
and Mosbrugger, 1999; Sack and Frole, 2006, phylogenetic studies; and a greatly expanded
Sack et al., 2008; Taylor et al., 2008). Moreover, role for leaf architecture in dendrology, forest
investigators of parallel- and reticulate-veined management, and ultimately conservation.
monocot groups (Hickey and Peterson, 1978;
Wilde et al., 2005) will expand the system and The manual is organized into three major sec-
make it more broadly applicable. tions covering general leaf characters, vein
characters, and tooth characters. Each sec-
This work evolved from the 1999 Manual of tion contains a set of general definitions fol-
Leaf Architecture (Ash et al., 1999) printed and lowed by a hierarchical, illustrated list of the
distributed by the Leaf Architecture Working described characters and their character states.
Group. We have made numerous substantive Appendix A contains a summary outline of
changes and additions to the 1999 manual in character states; Appendix B shows examples
order to clarify existing terms, coin terms for of fully described leaves; Appendix C presents
previously unrecognized characters and char- voucher data for the leaf images; and Appendix
acter states, and reorder terms into a more D describes a method for clearing leaves.
logical and hierarchical progression. Some
of these changes reflect comments received
from users of the 1999 manual. We have also
significantly increased the number and qual-
ity of reference illustrations, using examples
selected from the more than twenty thou-
sand specimens in the National Cleared Leaf
Collections of the Smithsonian Institution
and other repositories. Following the scope
of coverage found in prior work (e.g., Hickey,
1979), we emphasize features of the leaf blade
and provide only superficial treatment of leaf
attachment, insertion, stipules, and so on. We
do not address monocots in detail because
many monocot groups have specialized fea-
tures that require separate study (Hickey and
Peterson, 1978; Wilde et al., 2005).

We identified and corrected inconsistent usage
of terminology by having two groups of four
observers each score a test set of seventy-five
cleared leaves of extant taxa using the schema
below. We analyzed the scores and refined the
definitions for character states that were in-
consistently applied. A second group of seven
researchers then scored the same leaves using
the updated definitions. This procedure was
followed iteratively with additional leaves to
improve the consistency of scores by different
observers.

T his section describes the shape, size, sur-
face, organization, and other general
features of leaves. Some suites of char-
acters are treated only briefly or are omitted
entirely because they have been well described
ogy, see Dilcher, 1974; and Wilkinson, 1979,
pp. 97–117. For more detailed treatment of
stipules, stipels, pseudostipules, and phyllo-
taxy (leaf arrangement), see Bell, 2008; and
Keller, 2004. For more detailed treatments of
by other researchers. For descriptions of mod- leaf domatia, see Wilkinson, 1979, pp. 132–
ern leaf surfaces including cuticular morphol- 140; and O’Dowd and Wilson, 1991.

General Orientation Terms

abaxial
Pertaining to the surface adaxial
of the leaf facing away
from the axis of the plant,
generally the underside
of the leaf (Fig. 1).

adaxial
Pertaining to the surface
of the leaf facing toward
the axis of the plant,
generally the upper
surface of the leaf (Fig. 1).

admedial abaxial
Toward the midvein
(Fig. 2).
apical
apical, distal distal Fig. 1
Toward the apex (tip)
of the leaf (Fig. 2).

basal, proximal
Toward the base
of the leaf (Fig. 2). admedial
exmedial
exmedial
Away from the
midvein (Fig. 2).

proximal
basal
Fig. 2

apex
The distal ~25% of the
lamina (Fig. 3). If the lamina
apex
has an apical extension
0.75lm (tissue distal to the point
where the primary vein
ends), the apex includes all
tissue distal to 0.75 lm, where
lm is the distance from the
proximal to the distal end
of the midvein. Note: See
Figure 17 for a description of
lamina length.

base
0.25lm The proximal ~25% of
the lamina (Fig. 3). If the
base lamina has a basal extension,
the base includes all tissue
proximal to 0.25 lm, where
lm is the distance from the
proximal to the distal end
Fig. 3 of the midvein. Note: See
Dipterocarpus verrucosus Figure 17 for a description
(Dipterocarpaceae) of lamina length.

concave
Curved inward relative to
convex the midvein (Fig. 4).

concave convex
Curved outward relative to
the midvein (Fig. 4).
Fig. 4
decurrent
Approaching an intersection
a in an asymptotic manner in
b the basal direction (Fig. 5).
Applies both to veins, as
shown in Figure 6, and to
laminar tissue, as shown in
Figure 7. Note that decurrent
secondary veins may simply
branch, as shown in Figure
5a, or may “steal” part of the
midvein, making the midvien
thinner above the secondary,
as shown in Figure 5b.

Fig. 5 Fig. 6 Fig. 7
Itea chinensis Berberis sieboldii
(Iteaceae) (Berberidaceae)

Parts of a Simple Leaf

lamina (blade)
The expanded, flattened midvein
portion of a leaf (Fig. 8).

leaf
The chief photosynthetic
organ of most vascular land
plants, usually a determinate
outgrowth of a primordium
produced laterally on an leaf
axis. Most leaves consist of
a petiole (stalk), a leaf base,
and a bifacial lamina (blade).
Leaves subtend axillary
buds and have a definite axillary bud
arrangement, or phyllotaxy,
in their insertion along the
axis (Fig. 8).

petiole petiole
The stalk that attaches a leaf
to the axis (Figs. 8, 10). lamina

insertion point
The place where the base of
the lamina joins the petiole Fig 8
(Fig. 9).

margin
The outer edge of the lamina
(Fig. 10).

margin
leaf

midvein

petiole
insertion
point

Fig. 9
Alangium chinense Fig. 10
(Cornaceae)

midvein
The medial primary vein.
midvein In pinnate leaves, it is the
lobes only primary vein (Figs. 8,
10, 11; see Section II, below,
sinus for further discussion of
primary veins).

lobe
A marginal projection with a
corresponding sinus incised
25% or more of the distance
from the projection’s apex
to the midvein, measured
parallel to the axis of
symmetry and along the
distal side of the projection
or the basal side of a
Fig. 11 terminal projection (Fig. 11).
Liquidambar styraciflua
(Altingiaceae) sinus
A marginal embayment,
incision, or indentation
between marginal
projections of any sort,
typically lobes (Fig. 11),
teeth (Fig. 12), or the base of
cordate leaves (Fig. 12).

leaf domatia
Cavities or hollow structures
on the laminar, stipular, or
tooth sinus Fig. 12 leaf base sinus petiolar surfaces of the leaf,
Acalypha pringlei inferred to be habitable by
(Euphorbiaceae) insects or mites (Fig. 13).

Fig. 13
Leaf domatia

Parts of a Compound Leaf

compound leaf
A leaf with two or more
noncontiguous areas of
laminar tissue (Fig. 14). leaflet
leaflet
A discrete, separate laminar
segment of a compound
leaf. Leaflets never subtend
axillary buds (Fig. 14).
midvein
rachis
The prolongation of the
petiole of a pinnately
compound leaf, to which
leaflets are attached
(Fig. 14). In cross-section
the rachis may be terete insertion point
(round), semiterete, angular,
canaliculate (having
longitudinal channels), or
winged (see Figs. 46–49 for leaf
the analogous characters
in petioles). A second-order
rachis is a rachilla (see rachis
Fig. 32).

petiolule
The stalk that attaches a
leaflet of a compound leaf to
its rachis (Fig. 14).

insertion point petiolule
The point where the leaf is
attached to the axis or where
a leaflet is attached to the axillary bud
petiole or petiolule (Fig. 14). petiole

Fig. 14

Stipels and Stipules
stipel
A stipule-like structure
located at the base of the
petiolule of some leaflets or
extrafloral nectaries. Stipels
may occur on the petiolule
or at the juncture of the
petiolule and rachis (Fig. 15).

leaflet stipel

stipule
On dicotyledonous plants,
Fig. 15 an outgrowth (scale, laminar
Andira mandshurica structure, or spine) usually
(Fabaceae) associated with the point
of insertion of a leaf on a
stem (Fig. 16). Stipules may
occur on or along part of
the base of the petiole but
are more often on the axis
near the petiole base, where
they can be intrapetiolar
(between petiole and stem),
leaf-opposed, lateral, or (for
opposite leaves) interpetiolar.
They are usually paired
but may be fused to form a
single sheathing or perfoliate
structure. Stipules are usually
deciduous, often leaving
behind a characteristic
scar. Domatia, tendrils, or
extrafloral nectaries may
occupy stipule positions.
Stipules may be difficult
to distinguish from
intrapetiolar pseudostipules, stipule-like
stipule paired outgrowths on the
petiole toward or rarely
at the base of pinnately
Fig. 16 compound leaves that are
Malus baccata morphologically distinct
(Rosaceae) from the leaflets.

Measurements

lamina length, L = lm + la + lb (Fig. 17).

apical extension length, la = Distance from the most
distal point of the midvein to the most distal extension of leaf
tissue, the latter projected to the trend of the midvein (Fig.
17c, d). In most leaves, la = 0 (Fig. 17a, b).

basal extension length, lb = Distance from the most
proximal point of the midvein to the most proximal extension
of leaf tissue, the latter projected to the trend of the midvein
(Fig. 17b, d; Fig. 18). In many leaves lb = 0 (Fig. 17a, c). When
lb is longer on one side of the leaf than the other, always use
the larger value when calculating lamina length (Fig. 18).

midvein length, lm = Distance from the proximal end of
the midvein to the distal end (Fig. 17).

width ratio, x/y = The ratio of the smaller to the larger of the
two distances measured perpendicularly from the midvein to the
margin on each side of the leaf at the position of maximum leaf
width (Fig. 19). On a lobed leaf, the width ratio is measured to
the outermost portion of the leaf (Fig. 20).

basal width ratio, similar to width ratio but measured only
in the widest portion of the base of the leaf (Fig. 21).

la la

L=l m
lm L lm L lm L

lb lb
lb
a b c d

Fig. 17

L b1

L b2
y x

Fig. 18 Fig. 19
Basal extension Width ratio
Tilia chingiana Discocledidion rufescens
(Malvaceae) (Euphorbiaceae)

y x
y x

Fig. 20 Fig. 21
Lobed leaf width ratio Basal width ratio
Croton lobatus Aleurites remyi
(Euphorbiaceae) (Euphorbiaceae)

1.1 Petiolate – A petiole attaches the leaf to the axis (Figs. 8, 10, 13, 22).

1.2 Sessile – Leaf attaches directly to the axis without a petiole (Fig. 23).

Fig. 22 Fig. 23
Leaf attachment petiolate, leaf Leaf attachment sessile
arrangement alternate (distichous)

2. Leaf Arrangement – The placement of adjacent leaves on the nodes of the axis (more
than one may apply). Note: For more detailed treatments of phyllotaxy, see Bell, 2008; or
Keller, 2004.

2.1 Alternate – Adjacent leaves occur above or below others on the axis with one
leaf per node (Fig. 22). The arrangement may be distichous (in one plane in two
ranks on opposite sides of the axis) or helical (in a spiral along the axis).

2.2 Subopposite – Adjacent leaves occur in pairs that are nearly but not strictly
opposite (Fig. 24). These pairs may be decussate (leaf pairs inserted at ~90 o to
those above and below), distichous (leaf pairs are aligned with those above and
below), or spirodecussate (successive leaf pairs inserted at angles >90 o to those
above and below).

2.3 Opposite – Leaves occur in opposed pairs that arise from the same node
along the axis. Leaf pairs may be decussate (Fig. 25), distichous (Fig. 26), or
spirodecussate.

2.4 Whorled – Three or more leaves are borne at each node (Fig. 27).

Fig. 24 Fig. 25
Leaf arrangement subopposite (distichous) Leaf arrangement opposite (decussate)

Fig. 26 Fig. 27
Leaf arrangement opposite (distichous) Leaf arrangement whorled

3. Leaf Organization

3.1 Simple – Leaf consists of a single lamina attached to a simple petiole (Fig. 28).
This is the most common case.

3.2 Compound – Leaf consists of two or more leaf lets (laminae not intercon-
nected by laminar tissue.) Note: Ternate, a term used for various types of orga-
nization of leaflets (and leaves) into threes, is not treated here.

3.2.1 Palmately compound – Leaf has more than two separate laminar sub-
units (leaflets) attached at the apex of a petiole (Fig. 29). The description
should include the number of leaflets.

3.2.2 Pinnately compound – Leaf has leaflets arranged along a rachis.

3.2.2.1 Once compound – With a single order of pinnate leaflets
(Fig. 30, 31).

3.2.2.2 Twice, or bipinnately compound – Dissected twice with
leaflets arranged along rachillae that are attached to the rachis
(Fig. 32).

3.2.2.3 Thrice, or tripinnately compound – Leaflets are attached
to secondary rachillae that are in turn attached to rachillae,
which are borne along the rachis (Fig. 33).

Fig. 28 Fig. 29
Leaf organization simple Leaf organization palmately compound

Fig. 30 Fig. 31
Leaf organization once-pinnately compound (odd) Leaf organization once-pinnately compound (even)
Hymenaea courbaril
(Fabaceae)

Fig. 32 Fig. 33
Leaf organization twice-pinnately compound Leaf organization thrice-pinnately compound

4. Leaflet Arrangement – These character states apply only to pinnately compound leaves.
Note that odd-pinnately compound (imparipinnate) leaves have a single terminal leaflet,
and even-pinnately compound (paripinnate) leaves do not. These terms are illustrated
for opposite leaflets but may apply to subopposite leaflets as well.

4.1 Alternate – Leaflets are arranged alternately on the rachis (Fig. 34).

4.2 Subopposite – Leaflets are in pairs that are nearly, but not strictly, opposite (Fig. 35).

4.3 Opposite – Leaflets are in pairs that arise on opposite sides of the rachis.

4.3.1 Odd-pinnately compound (Fig. 36).

4.3.2 Even-pinnately compound (Fig. 37).

4.4 Unknown – fossil only; not preserved (Fig. 38).

5. Leaflet Attachment – These character states apply only to compound leaves.

5.1 Petiolulate – Leaflet is attached to the rachis by means of a petiolule (stalk),
analogous to the petiole of a leaf (Figs. 34–38).

5.2 Sessile – Leaflet is attached directly to the rachis (Fig. 39).

Fig. 34 Fig. 35 Fig. 36
Leaflet arrangement alternate; Leaflet arrangement Leaflet arrangement
petiolulate subopposite; petiolulate opposite (odd)

Fig. 37 Fig. 38 Fig. 39
Leaflet arrangement Leaflet arrangement Leaflet attachment
opposite (even) unknown sessile

6. Petiole Features

6.1 Petiole base

6.1.1 Sheathing – Petiole expands to clasp
the stem (Fig. 40).

6.1.2 Pulvin(ul)ate – Having an abruptly
swollen portion near the node around
which the leaf(let) can flex (Fig. 41); may
occur with or without an abscission joint
(Fig. 42). On compound leaves, a pulvi-
nulus may occur at the proximal and/or
distal end of the petiolule and sometimes Fig. 40
only on the terminal leaflet (Fig. 43). Petiole base sheathing

terminal
leaflet

distal

proximal

Fig. 41 Fig. 42 Fig. 43
Petiole base pulvinate Petiolule base pulvinulate Position of pulvinulus
Antrocaryon amazonicum
(Anacardiaceae)

6.2 Glands (see also I.22 and III.53) – Swollen areas of secretory tissue, often paired.

6.2.1 Petiolar – Glands are borne along the petiole (Fig. 44).

6.2.2 Acropetiolar – Glands are borne at the distal end of the petiole, below
the base of the leaf (Fig. 45).

6.3 Petiole-cross section

6.3.1 Terete – Round (Fig. 46).

6.3.2 Semiterete – Semicircular (Fig. 47).

6.3.3 Canaliculate – Having a longitudinal channel or groove (Fig. 48).

6.3.4 Angular (Fig. 49).

6.3.5 Alate or Winged – With lateral ridges or flanked by laminar tissue
(Fig. 50).

6.4 Phyllodes – Petiole or rachis is expanded to make a lamina (Fig. 51).

Fig. 44 Fig. 45
Petiolar glands Acropetiolar glands

Fig. 46 Fig. 47 Fig. 48
Terete petiole cross-section Semi-terete petiole cross-section Canaliculate petiole cross-section

Fig. 49 Fig. 50
Angular petiole cross-section Alate petiole and rachis

Fig. 51
Phyllode
Acacia mangium
(Fabaceae-Mimosoideae)

7. Position of Lamina Attachment – The point from which the lamina is borne.

7.1 Marginal – Leaf is attached at its margin (Fig. 52).

7.2 Peltate central – Leaf is borne from a position near the center of the lamina
(Fig. 53).

7.3 Peltate excentric – Leaf is borne from a position within the boundaries of the
lamina but not near its center (Fig. 54).

Fig. 52 Fig. 53 Fig. 54
Marginal petiole attachment Peltate central Peltate excentric
x Mahoberberis neubertii Brasenia schreberi Macaranga bicolor
(Berberidaceae) (Cabombaceae) (Euphorbiaceae)

8. Laminar Size – The area of the leaf blade. When possible, the area should be measured
directly (e.g., digitally) or approximated by multiplying the length by the width by 0.75
(Cain and Castro, 1959). Alternatively, laminar size can be approximated by size classes
(Raunkiaer, 1934; Webb, 1959). Figure 55 shows outlines of the maximum sizes of five
of the smallest size classes; the leaf belongs in the smallest size class into which its area
fits completely. The template, which can be photocopied onto clear acetate and placed
over a leaf, is included for paleobotanists, who often work with incomplete fossil leaves
and must approximate leaf area.

Areas of leaf size classes (Webb, 1959):

8.1 Leptophyll <25 mm2
8.2 Nanophyll 25–225 mm2
8.3 Microphyll 225–2,025 mm2
8.4 Notophyll 2,025–4,500 mm2
8.5 Mesophyll 4,500–18,225 mm2
8.6 Macrophyll 18,225–164,025 mm2
8.7 Megaphyll >164,025 mm2

Notophyll
2:1

1 cm

1.5:1

Microphyll
1:1

Mesophyll

Nanophyll

Leptophyll

Fig. 55
Leaf size template

9. Laminar L:W Ratio – Ratio of laminar
length to maximum width perpendicular
to the axis of the midvein (Fig. 56).

Fig. 56
Trichosanthes formosana
(Curcurbitaceae)

10. Laminar Shape (in compound leaves, this applies to the shape of the leaflets) –
To determine the shape of the lamina, locate the midvein and determine the zone of
greatest width measured perpendicular to the midvein. In lobed leaves, draw a line from
the apex to the widest point on either side of the midvein and determine the shape by
finding the zone of greatest width based on this outline (Fig. 57). Historically, botanists
combined leaf shape with imprecisely defined L:W ratios to create additional character
states (e.g., von Ettingshausen, 1861). Some common historical terms are italicized be-
low but not illustrated.

10.1 Elliptic – The widest part of the leaf is in the middle one-fifth (Fig. 58). Note:
The terms orbiculate and oblate have been used to describe unlobed, elliptic leaves
that are very wide. We suggest using orbiculate for elliptic leaves with a L:W ratio
ranging from 1.2:1 to 1:1 and oblate for elliptic leaves with a L:W ratio <1:1.

10.2 Obovate – The widest part of the leaf is in the distal two-fifths (Fig. 59). We sug-
gest defining oblanceolate leaves as obovate leaves with a L:W ratio between 3:1
and 10:1.

10.3 Ovate – The widest part of the leaf is in the proximal two-fifths (Fig. 60). Note:
Lanceolate has been used to describe ovate leaves that are long and narrow. We
suggest defining lanceolate leaves as ovate leaves with a L:W ratio between 3:1
and 10:1.

10.4 Oblong – The opposite margins are roughly parallel for at least the middle one-
third of the leaf (Fig. 61).

10.5 Linear – The L:W ratio of a leaf is ≥10:1, regardless of the position of the
widest part of the leaf (Fig. 62).

10.6 Special – Outlines that do not fall readily into one of the shape classes above;
for example, the pitcher-shaped leaf apex of Nepenthes.

1
5

Fig. 57 Fig. 58
Measuring lobed leaves Elliptic leaf shape
Dioscoreophyllum strigosum Cheiloclinium anomalum
(Menispermaceae) (Celastraceae)

2
5

1
3

2
5

Fig. 59 Fig. 60 Fig. 61 Fig. 62
Obovate leaf shape Ovate leaf shape Oblong leaf shape Linear leaf shape
Alstonia congensis Parinari sp. Ficus citrifolia Xylomelum angustifolium
(Apocynaceae) (Chrysobalancaceae) (Moraceae) (Proteaceae)

11. Medial Symmetry – Determined by the width ratio in the middle of the leaf (see
Measurements, above).

11.1 Symmetrical – Width ratio (x/y) > 0.9 from 0.25L to 0.75L (Fig. 63).

11.2 Asymmetrical – Width ratio (x/y) < 0.9 from 0.25L to 0.75L (Fig. 64).

x y

Fig. 63 Fig. 64
Leaf medially symmetrical Leaf medially asymmetrical
Maytenus aquifolium Ramirezella pringlei
(Celastraceae) (Fabaceae)

12. Base Symmetry – Base symmetry and basal width asymmetry are determined by the
width ratio in the base of the leaf (see Measurements, above). Leaf bases can be asym-
metrical in insertion, extension, and width.

12.1 Base Symmetrical – Base lacks any of the asymmetries identified below
(Fig. 65).

12.2 Base Asymmetrical

12.2.1 Basal width asymmetrical – Basal width ratio (x/y) < 0.9 (Fig. 66).

12.2.2 Basal extension asymmetrical – Basal extension length on one side
is <0.75 of the other side (Lb1/Lb2 < 0.75) (Fig. 67).

12.2.3 Basal insertion asymmetrical – Insertion points of lamina base on
either side of the petiole are separated by >3 mm (Fig. 68).

x y x y

Fig. 65 Fig. 66
Basal width symmetrical Basal width asymmetrical
Aleurites remyi Lunania mexicana
(Euphorbiaceae) (Salicaceae)

L b1

L b2 >3mm

Fig. 67 Fig. 68
Basal extension asymmetrical Basal insertion asymmetrical
Tilia chingiana Fraxinus floribunda
(Malvaceae) (Oleaceae)

13. Lobation – A lobe is a marginal projection with a corresponding sinus incised 25% or
more of the distance from the projection apex to the midvein, measured parallel to the
axis of symmetry and along the apical side of the projection (or the basal side of a termi-
nal projection). A leaf is considered lobed even if it has only one marginal projection that
fits the definition. If the sinus described above is incised less than 25% of the width, the
projection is considered a tooth (see Section III).

13.1 Unlobed – The leaf has no lobes (Figs. 69, 70). Note that the leaf in Figure 69
is also called “entire” because it lacks lobes and teeth. The term entire is useful
because it describes the majority of angiosperm leaves. For further discussion
of entire, see I.14.

13.2 Lobed

13.2.1 Palmately lobed – Major veins of the lobes are primary veins that
arise from the base of the leaf (Fig. 71).

13.2.1.1 Palmatisect – Special case of palmately lobed in which the incision
goes almost to the petiole but without resulting in distinct leaflets
(Fig. 72). Palmatifid and palmatipartite are variously used terms for
leaves with incised palmate lobes that are not treated here.

13.2.2 Pinnately lobed – Major veins of the lobes are formed by costal sec-
ondaries (Fig. 73).

13.2.2.1 Pinnatisect – Special case of pinnately lobed in which the in-
cision goes almost to the midvein but without resulting in distinct
leaflets (Fig. 74). Pinnatifid and pinnatipartite are variously used terms
for leaves with pinnately-incised lobes that are not treated here.

13.2.3 Palmately and pinnately lobed – At least one lobe in a palmately
lobed leaf is pinnately lobed (Fig. 75).

13.2.4 Bilobed – Leaf has two lobes (Fig. 76).

Fig. 69 Fig. 70
Unlobed (entire) Unlobed (with teeth)
Parinari campestris Melanolepis multiglandulosa
(Chrysobalanaceae) (Euphorbiaceae)

Fig. 71 Fig. 72 Fig. 73
Palmately lobed Palmatisect Pinnately lobed
Adenia heterophylla Potentilla recta Stenocarpus sinuatus
(Passifloraceae) (Rosaceae) (Proteaceae)

Fig. 74 Fig. 75 Fig. 76
Pinnatisect Palmately and pinnately lobed Bilobed
Dryandra longifolia Cucurbita cylindrata Bauhinia madagascariensis
(Proteaceae) (Curcurbitaceae) (Fabaceae)

14. Margin Type – Features of the edge of the lamina. Section I.13 describes how to distinguish
lobes and teeth.

14.1 Untoothed – Margin has no teeth (Fig. 77). Note: The term entire describes a
leaf with no teeth and no lobes (Fig. 69). Leaf Margin Analysis and other physi-
ognomic methods of paleoclimate inference score lobed leaves without teeth in
the same category as entire leaves (Wolfe 1995), thus the category “entire” has
sometimes been inferred to include lobed, untoothed leaves. We prefer the word
untoothed for this category because it provides the clearest alternative to toothed and
does not conflict with the standard botanical meaning of entire, which excludes all
lobed leaves.

14.2 Toothed – Margin has vascularized projections (Figs. 78–80) separated by sinuses
that are incised less than 25% of the distance to the midvein or long axis of the leaf
as measured parallel to the axis of symmetry from the apical incision of the projec-
tion. Note that a leaf with a single tooth of any size is considered toothed. Also, both
lobes and teeth may be present on the same leaf (but see notes below).

14.2.1 Dentate – Majority of the teeth have axes of symmetry directed perpen-
dicular to the trend of the leaf margin (Fig. 78).

14.2.2 Serrate – Majority of the teeth have axes of symmetry directed at an
angle to the trend of the leaf margin (Fig. 79).

14.2.3 Crenate – Majority of the teeth are smoothly rounded, without a pointed
apex (Fig. 80). Note: Crenate margins are also either dentate or serrate.

Fig. 77 Fig. 78
Untoothed margin Dentate margin
Caraipa punctulata Casearia ilicifolia
(Clusiaceae) (Salicaceae)

Fig. 79 Fig. 80
Serrate margin Crenate and serrate margin
Betula lenta Viola brevistipulata
(Betulaceae) (Violaceae)

Notes: The difference between lobes and teeth is sometimes am-
biguous. Some leaves have geometrically similar projections that
could be scored as lobes or teeth using the 25% rule above. When
at least one definitive lobe is present, we suggest scoring such pro-
jections as lobes and not as teeth (Fig. 81) (Royer et al., 2005). Some
toothed leaves have projections at the apex that are incised more
than 25%. We suggest scoring these projections as teeth rather
than lobes (Fig. 82).

lobe

tooth

Fig. 81 Fig. 82
Lobe that looks like a tooth Tooth that looks like a lobe
Quercus alba Rubus mesogaeus
(Fagaceae) (Rosaceae)

15. Special Margin Features

15.1 Appearance of the edge of the leaf blade

15.1.1 Erose – Margin is minutely irregular, as if chewed (Fig. 83).

15.1.2 Sinuous – Margin forms a series of shallow and gentle curves that lack
principal veins. These projections are not considered teeth (see above or
Section III) (Fig. 84).

15.2 Appearance of the abaxial-adaxial plane of the leaf blade

15.2.1 Revolute – Margin is turned down or rolled (in the manner of a scroll)
in the abaxial direction (Fig. 85).

15.2.2 Involute – Margin is turned up or rolled in the adaxial direction (Fig. 86).

15.2.3 Undulate – Margin forms a series of smooth curves in the abaxial-
adaxial plane (in and out of the plane of the leaf ) (Fig. 87).

Fig. 83 Fig. 84
Erose margin Sinuous margin
Bridelia cathartica
(Phyllanthaceae)

Fig. 85
Revolute margin

Fig. 86 Fig. 87
Involute margin Undulate margin

16. Apex Angle
The vertex of the apex angle lies at the center of the midvein where it terminates at the
apex of the leaf. The apex angle is formed by the two rays that depart this vertex and
are tangent to the leaf margin without crossing over any part of the lamina (Figs. 88, 89).
The apex angle is always measured on the proximal side of the rays. If the leaf is toothed,
draw the lines along the edge of the margin, connecting the marginal tissue (Fig. 89). If
the midvein terminates between two lobes, the angle is formed as in unlobed leaves but
is greater than 180° (Fig. 90). If the midvein terminates at the apex of a lobe, the rays
need only be tangent to the margin of the terminal lobe and may pass over lateral lobes
(Fig. 91). Leaves with retuse apices (see 20.3) are considered to have an obtuse apex angle.
The following categories are useful for scoring apex angles:

16.1 Acute – Apex angle <90° (Fig. 88).

16.2 Obtuse – Apex angle between 90° and 180° (Fig. 89).

16.3 Reflex – Apex angle >180° (Fig. 90).

Fig. 88 Fig. 89
Acute apex angle Obtuse apex angle
Ostrya guatemalensis x Mahoberberis neubertii
(Betulaceae) (Berberidaceae)

Fig. 90
Reflex apex angle
Bauhinia madagascariensis
(Fabaceae)

Fig. 91
Acute apex angle on a lobed leaf
Dioscoreophyllum strigosum
(Menispermaceae)

17. Apex Shape – These states apply to the shape of the distal 25% of the lamina. On a
toothed leaf, a smoothed curve through the tips of the teeth determines the shape (Fig. 93).
For leaves with an apical extension (la > 0), follow the guidelines in Figure 92. If the apex is
retuse (see also 20.3), it can still be scored for the other shape features given below.

17.1 Straight – Margin between the apex and 0.75L has no significant curvature
(Fig. 93).

17.2 Convex – Margin between the apex and 0.75L curves away from the midvein
(Fig. 94).

17.2.1 Rounded – Subtype of convex in which the margin forms a smooth arc
across the apex (Fig. 95).

17.2.2 Truncate – Apex terminates abruptly as if cut, with margin perpen-
dicular to midvein or nearly so (Fig. 96).

17.3 Acuminate – Margin between the apex and 0.75L is convex proximally and
concave distally, or concave only. This category, especially when the distal portion
of the apex abruptly narrows, accommodates most apex types called “drip tips”
(Figs. 97, 98).

17.4 Emarginate – lm is 75–95% of lm + la (Fig. 99); see also retuse (20.3).

17.5 Lobed – lm is <75% of lm + la (Fig. 90).

Note: If the leaf has a different apex shape on either side, both shapes should be
recorded (Fig. 100).

100%

75%

0%
Retuse Emarginate Lobed
la < 5% L la =5–25% L la < 25% L

Fig. 92
Definitions of apex shapes for leaves that have an apical extension

Fig. 93 Fig. 94
Apex shape straight Apex shape convex
Aristotelia racemosa Saurauia calyptrata
(Elaeocarpaceae) (Actinidiaceae)

Fig. 95 Fig. 96 Fig. 97
Apex shape rounded Apex shape truncate Apex shape acuminate (with drip tip)
Ozoroa obovata Liriodendron chinense Neouvaria acuminatissima
(Anacardiaceae) (Magnoliaceae) (Annonaceae)

Fig. 98 Fig. 99 Fig. 100
Apex shape acuminate (without drip tip) Apex shape emarginate Apex shape acuminate on the left
Corylopsis veitchiana Lundia spruceana and straight on the right
(Hamamelidaceae) (Bignoniaceae) Tapura guianensis
(Dichapetalaceae)

18. Base Angle – The vertex of the base angle lies in the center of the midvein next
to the point where the basalmost laminar tissue joins the petiole (or joins the proxi-
mal margin in the case of sessile leaves). The base angle is formed by the two rays
that depart this vertex and are tangent to the leaf margin without crossing over any
part of the lamina. The base angle is independent of base shape (see Base Shape, I.19).

For consistency, the base angle is always measured on the distal side of the vertex, even
when the angle is greater than 180° (Fig. 103–104). The following categories are useful
for scoring base angles

18.1 Acute – Angle <90° (Fig. 101).

18.2 Obtuse – Angle >90° but <180° (Fig. 102).

18.3 Reflex – Special case of obtuse in which angle is >180° but <360° (Figs. 103, 104).

18.4 Circular – Special case of reflex in which angle is >360°. This includes leaves
in which the basal extension overlaps across the midline, as well as peltate leaves
(Fig. 105).

Fig. 101 Fig. 102
Acute base angle Obtuse base angle
Schumacheria castaneifolia Mauria heterophylla
(Dilleniaceae) (Anacardiaceae)

Fig. 103 Fig. 104
Reflex base angle Reflex base angle
Tilia chingiana Asarum europaeum
(Malvaceae) (Aristolochiaceae)

Fig. 105
Circular base angle
Cissampelos owariensis
(Menispermaceae)

19. Base Shape – These states apply to the shapes of the proximal 25% of the lamina. On
a toothed leaf, a smoothed curve through the tips of the teeth determines the shape.

19.1 If there is no basal extension (lb = 0), the following base types are
recognized

19.1.1 Straight (cuneate) – Margin between the base and 0.25L has no signifi-
cant curvature (Fig. 106).

19.1.2 Concave – Margin between the base and 0.25L curves toward the mid-
vein (Fig. 107).

19.1.3 Convex – Margin between the base and 0.25L curves away from the midvein
(Fig. 108).

19.1.3.1 Rounded – The margin forms a smooth arc across the base
(Fig. 109).

19.1.3.2 Truncate – The base terminates abruptly as if cut perpendicu-
lar to the midvein or nearly so (Fig. 110).

19.1.4 Concavo-convex – Margin between the base and 0.25L is concave
proximally and convex distally (Fig. 111).

19.1.5 Complex – Margin curvature has more than one inflection point (change
of curvature) between the base and 0.25L (Fig. 112).

19.1.6 Decurrent – Special case in which the laminar tissue extends along
the petiole at a gradually decreasing angle (Figs. 113, 114); can occur in
concave, concavo-convex, or complex bases.

19.2 If there is a basal extension (lb > 0), the following base types are
recognized

19.2.1 Cordate – Leaf base forms a single sinus with the petiole generally in-
serted at the deepest point of the sinus (Figs. 115, 116).

19.2.2 Lobate – Leaf base is lobed on both sides of the midvein. The lobes are
defined by a central sinus containing the petiole as in cordate leaves, and
by sinuses on their distal sides such that the nadirs of the distal sinuses
are within the base of the leaf (Figs. 117, 118). The following terms have
been used historically for some leaves that have two basal projections.
We consider them to be subtypes of lobate bases.

19.2.2.1 Sagittate – Leaf base has two narrow, usually pointed projec-
tions (technically these may not qualify as lobes because they are
not bounded by distal sinuses) with apices directed proximally at
an angle 125º or greater from the midvein (Fig. 119).

19.2.2.2 Hastate – Leaf base has two narrow lobes with apices directed
exmedially at 90º–125º from the midvein (Fig. 120).

19.2.2.3 Runcinate (not pictured) – A lobate lamina with two or more
pairs of downward-pointing (>110 o) angular lobes.

19.2.2.4 Auriculate (not pictured) – A lobate lamina having a pair of
rounded basal lobes that are oriented downward, with their
axes of symmetry at an angle >125o from the midvein of the
leaf. If the lateral sinuses that define the lobes extend more than
50% of the distance to the midvein, such laminar bases may be
referred to as panduriform.

Fig. 106 Fig. 107
Base shape cuneate Base shape concave
Carya leiodermis Sassafras albidum
( Juglandaceae) (Lauraceae)

Fig. 108 Fig. 109
Base shape convex Base shape rounded
Prunus mandshurica Carissa opaca
(Rosaceae) (Apocynaceae)

Fig. 110 Fig. 111
Base shape truncate Base shape concavo-convex
Populus dimorpha Diploclisia chinensis
(Salicaceae) (Menispermaceae)

Fig. 112 Fig. 113 Fig. 114
Base shape complex Base shape decurrent Base shape decurrent
Adelia triloba Alstonia plumosa Berberis sieboldii
(Euphorbiaceae) (Apocynaceae) (Berberidaceae)

Fig. 115 Fig. 116
Base shape cordate Base shape cordate
Phyllanthus poumensis Cercidiphyllum japonicum
(Phyllanthaceae) (Cercidiphyllaceae)

Fig. 117 Fig. 118
Base shape lobate Base shape lobate
Acer saccharinum Liquidambar styraciflua
(Sapindaceae) (Hamamelidaceae)

Fig. 119 Fig. 120
Base shape sagittate Base shape hastate
Sagittaria sp. Araujia angustifolia
(Alismataceae) (Apocynaceae)

20. Terminal Apex Features – The following characters describe the region where the
midvein terminates.

20.1 Mucronate (apiculate) – The midvein terminates in an opaque, peg-shaped,
nondeciduous extension of the midvein (Fig. 121).

20.2 Spinose – The midvein extends through the margin at the apex; the spine may
be short or long, but it is not always sharp (Fig. 122).

20.3 Retuse – The midvein terminates in a shallow sinus such that lm is 95–99% of lm + la
(Fig. 123).

Fig. 121
Terminal apex mucronate
Cocculus ferrandianus
(Menispermaceae)

Fig. 122 Fig. 123
Terminal apex spinose Terminal apex retuse
Bauhinia rubeleruziana Fitzalania heteropetala
(Fabaceae) (Annonaceae)

21. Surface Texture (see Stearn, 1983)

21.1 Smooth – Lacking indentations, projections, hairs, or other roughness.

21.2 Pitted – Having indentations.

21.3 Papillate – Having small projections originating from the laminar surface.

21.4 Rugose – Rough; for example, from vein relief.

21.5 Pubescent – Having hairs (see Theobald et al., 1979, or Hewson, 1988, for pu-
bescence categories).

22. Surficial Glands – Placement of secretory structures.

22.1 Laminar – Glands present on the surface (may be clustered) (Fig. 124).

22.2 Marginal – Glands present only near or on the blade margin (Fig. 125).

22.3 Apical – Glands present only near the blade apex (Fig. 126).

22.4 Basilaminar – Glands present only near the base of the blade (Fig. 127).

5 mm
Fig. 124
Surficial glands laminar

Fig. 125
Surficial glands marginal

5 mm

Fig. 126 Fig. 127
Surficial glands apical Surficial glands basilaminar

T his section describes the orders, fabric,
and course of leaf venation.

costal veins
General Vein
intercostal area
Def initions
costal
Literally, “pertaining
to the ribs.” Used here
for secondary veins that
originate directly from
primary veins and are
typically, together with the
primaries, the principal
structural supports for the
leaf blade (Fig. 128). Costal
secondaries are also called
major secondaries.

decurrent
Referring to a vein junction Fig. 128
at which one vein’s course
asymptotically converges Fig. 129
on another (usually of larger Decurrent secondary veins
gauge) (Fig. 129). Itea chinensis
(Iteaceae)
dichotomous
Branching into two veins
of equal gauge; commonly
both branches have a
thinner gauge than the vein
from which they branched.
Dichotomous vein systems
generally ramify freely
(Figs. 130, 131).

Fig. 130
Dichotomous branching

Fig. 131
Dichotomous tertiary veins
Astronium graveolens
(Anacardiaceae)

excurrent
Branching laterally without
significant deflection of the
main vein trunk. A vein
segment with only excurrent
branches is monopodial
(Figs. 132–135).

gauge
Width of a vein measured
perpendicular to its course
and in the plane of the
Fig. 132 lamina (bundle sheath
Excurrent secondaries included).

intercostal areas
Sections of the leaf lying
between adjacent major
(costal) secondaries
(Fig. 128).

Fig. 133 vein fabric
Excurrent branching The overall appearance of
Carpinus fargesii the network of tertiary and
(Betulaceae) higher order veins. Fabric
is characterized by the
gauge, orientation, spacing
and course of the higher
order veins with respect
to one another and the
whole leaf. In leaves with
distinct orders of veins
above tertiary it may be
useful to describe the fabric
of the tertiary, quaternary,
and even quinternary veins
separately.
Fig. 134
Monopodial primary monopodial
Having a single main vein
trunk whose lateral veins
do not deflect the course.
Most primary veins are
monopodial (Figs. 134, 135).

Fig. 135
Monopodial primary
Bixa orellana
(Bixaceae)

ramified
Branching into higher-order
veins without rejoining veins
of the same or lower orders
(Fig. 136).

sympodial
Type of branching in
which the main vein axis
is deflected at each branch
point (Figs. 137, 138).

vein course
The path of a vein.

Fig. 136
Ramified veins
Comocladia cuneata
(Anacardiaceae)

Fig. 137
Sympodial branching

Fig. 138
Sympodial branching of primary
Griselinia scandens
(Griseliniaceae)

T he first step in describing the pattern of
venation in a leaf is to recognize catego-
ries, or orders, of veins that have visu-
ally distinct gauges and courses. Most angio-
sperm leaves have between four and seven
it is fairly easy to recognize the primaries
and tertiaries, but the secondaries sometimes
consist of several subsets with different gauges
and courses. Nevertheless, all the subsets of
veins between the primaries and the tertiaries
orders of venation, which are conventionally are considered to be secondaries.
numbered sequentially, starting with 1º for
the primary vein or veins. After the three lowest vein orders have been
demarcated, the higher orders of venation (4º–
In general, the primary (1º) and secondary 7º) present in the leaf can be identified. Each
(2º) veins are the major structural veins of of these higher vein orders may be highly vari-
the leaf, and the tertiary (3º) veins are the able among species and higher taxa in its de-
largest-gauge veins that constitute the mesh, gree of distinctness from both the next higher
or “fabric,” of the vein system. The primary and next lower vein orders. This may be true
vein or veins are somewhat analogous to the even within a single leaf. Good diagnostic fea-
main trunk or trunks of a tree—they have tures for distinguishing higher orders of veins
the largest gauge, they usually taper along from one another are (1) excurrent origin from
their length, and they generally run from the their source veins and (2) a distinctly narrower
base or near the base of the leaf to its margin gauge. If they arise dichotomously or appear
at the apex. Secondary veins are analogous to have the same, or nearly the same, gauge
to the major limbs of a tree. They are the as their parent vein, they are considered the
next set in gauge after the primary or pri- same order as the source vein.
maries, they also usually taper along their
course, and they ordinarily run either from The simultaneous use of two criteria for de-
the base of the leaf or from a primary vein termining vein order introduces a degree of
toward the margin. The tree analogy breaks ambiguity into the process, because some
down for 3º and higher-order veins because veins may have the gauge typical of one vein
these veins maintain a similar gauge along order but the course typical of a different vein
their courses, and because they may form a order. On the other hand, recognizing orders
reticulum, or net. based solely on their gauge or solely on their
course leads to illogical situations in which
Tertiary veins usually have a narrower gauge veins that appear to have different functions
than the secondary set, have courses that often and developmental origins are assigned to the
connect 1º and 2º veins to each other through- same order. Assigning veins to orders also has
out the leaf, and are the veins of highest gauge a somewhat arbitrary aspect because varia-
that form a more or less organized “field” over tions in gauge and course are not mathemati-
the great majority of the leaf area. Generally, cally discrete (Bohn et al., 2002); for example,

a vein may be intermediate in gauge between veins may be difficult to distinguish in leaves
the 1º vein and the 2º veins. Natural breaks in of low rank (Fig. 139).
gauge usually occur at vein branching points,
however, so most veins can be assigned to an In our experience, different observers fol-
order unambiguously using visual cues. lowing a consistent set of rules can usually
define vein orders in a repeatable manner for
The regularity of vein systems varies widely, a given leaf (Figs. 140, 141). It is generally
but it can be described semi-quantitatively in good practice to discriminate vein orders at
terms of “leaf rank” (Hickey, 1977). Leaf rank the point where they are expressed at their
has practical significance for recognizing vein widest gauge, usually nearest to the center
orders because vein systems that are less well or base of the leaf. The following is a set of
organized (i.e., have lower rank) also tend to guidelines for recognizing vein orders; see
have less distinct vein orders. Even 2º and 3º also the definitions that follow.

Fig. 139
Delphinium cashmerianum
(Ranunculaceae)

3˚ 2˚

1˚ 2˚

1˚

1˚
3˚
4˚
5˚

Fig. 140
Acer argutum
(Sapindaceae)

1˚

2˚
1˚
3˚
2˚
4˚

1˚

3˚

4˚

5˚

2˚

Fig. 141
Fagus longipetiolata
(Fagaceae)

General rules
Most leaves have a continuous sequence of vein orders that are typically easy to recognize by
starting at the thickest (1º vein) and progressing to the finest. To recognize the 1º, 2º, and 3º veins,
take the following steps

1. Find the vein or veins of the largest gauge: the primary vein(s) (some leaves have more than one).
Most leaves have a single primary vein that gives rise to pinnately arranged secondaries or costal
veins (in this case, go to step 3). If more than one vein originates at or near the base of the leaf,
follow step 2 to determine if the leaf has more than one 1º vein.

2. After recognizing the single vein of greatest gauge as the primary (generally the midvein),
other primaries are recognized by being at least 75% of the gauge of the widest primary (at the
point of divergence from the widest primary). These veins are basal or nearly basal. If these
veins enter lateral lobes or run in strong arches toward the apex, they are generally easily rec-
ognized as primaries. But if the lateral primaries curve toward the midvein distally (Figs. 142,
143) or branch toward the margin (Fig. 144), it may be hard to designate them as primaries or
secondaries.

1˚ 1˚

2˚ 1˚

2˚

Fig. 142 Fig. 143 Fig. 144
Meriania speciosa Loreya arborescens Givotia rottleriformis
(Melastomataceae) (Melastomataceae) (Euphorbiaceae)

Note: If there is more than one 1º vein (based on vein gauge),
other veins originating at the base may be considered primaries
if their course is similar to that of the previously defined prima-
ries, even if their gauge falls into the range of 25–75% of the
widest 1º vein. If these veins are narrower than 25% of the wid-
est 1º vein, they are not considered primaries (Figs. 145, 146).

2˚ 1˚ 2˚

Fig.145
Pinnate venation
Tannodia swynnertonii
(Euphorbiaceae)

1˚ 1˚ 1˚ 1˚ 1˚ 2˚

2˚

Fig. 146
Palmate venation with five 1º veins
Tetracentron sinense
(Trochodendraceae)

3. Find the veins of greatest gauge that form the vein field mesh
or fabric of the leaf: the tertiary veins (Figs. 140, 141). Note that
in some instances 2º veins fill the field (Figs. 142, 147). Tertiary
veins are considered:

• epimedial if they intersect a 1º vein (Fig. 148).

• intercostal if they intersect a 2º vein but no primary (Fig. 148).

• exterior if they are exmedial to all 2º veins (Fig. 148).

intercostal epimedial

exterior

Fig. 147 Fig. 148
Field filled by secondaries rather than tertiaries Tertiary veins
Calophyllum calaba Sassafras albidum
(Clusiaceae) (Lauraceae)

4. Having recognized the limits of the 1º and major 2˚
3º vein sets, identify the set of veins that is in- or costal 2˚
termediate in gauge. These are the secondary
veins, and they may vary substantially in both
gauge and course. Typical types of secondary
veins include the following:

• major (or costal) secondaries,
the rib-forming veins that
originate on the primary and run
toward the margins (Fig. 149).

• minor secondaries, which
branch from lateral primaries
or major secondaries and run minor 2˚
toward the margins (Fig. 149).
Note: these are often the “tines”
of agrophic veins (see II.26).

• interior secondaries, which run
between primaries in palmately
veined leaves (Figs. 150, 151).
Fig. 149
• intersecondaries, which Major and minor 2º veins
have courses similar to major Parrotia jacquemontiana
secondaries but have a gauge (Hamamelidaceae)
intermediate between secondaries
and tertiaries and do not reach
the margin (Fig. 152).

• intramarginal secondaries,
which run parallel to the leaf
margin with laminar tissue
exmedial to them (Figs. 153).

• marginal secondaries, veins
of secondary gauge that run on interior 2˚
the margin of the leaf with no
exmedial laminar tissue (Fig. 154;
marginal veins of tertiary gauge
are called fimbrial veins, see II.30.3).

Fig. 150
Interior 2º veins

interior
secondary vein
intersecondary
vein

Fig. 151 Fig. 152
Interior secondaries Intersecondaries
Buxus glomerata Croton hircinus
(Buxaceae) (Euphorbiaceae)

intramarginal
vein

Fig. 153
Intramarginal secondary
Spondias globosa
(Anacardiaceae)

5. Once you have recognized the first three orders of venation,
proceed in sequence to determine the higher orders of venation
using the criteria of vein gauge and course. Each successive vein
order should have a distinctly narrower gauge, and the course
may differ as well.

6. In most leaves, the veins of the finest gauge are freely ending
veinlets (FEVs). FEVs enter, but do not cross, the smallest vein-
bounded regions of leaf tissue, the areoles. FEVs can be un-
branched, but they most often ramify within the areole. The
boundaries of most areoles are formed by the highest order of
excurrently branched veins.

marginal secondary vein

Fig. 154
Marginal secondary
Diploclisia kunstleri
(Menispermaceae)

23.1 Pinnate – Leaf or leaflet has a single 1º vein (Figs. 155–158).

23.2 Palmate – Leaf has three or more basal veins, of which at least two are primaries
(i.e., at least one has 75% of the gauge of the thickest vein, which is usually the
midvein, see Determining Vein Order and Type, above). It can be difficult to
distinguish pinnate from palmate primary frameworks near the 75% cutoff.

23.2.1 Actinodromous – Three or more 1º veins diverge radially from a single
point.

23.2.1.1 Basal – Primary veins radiate from the petiolar insertion point
(Figs. 159, 160).

23.2.1.2 Suprabasal – Primary veins radiate from a point distal to peti-
olar insertion (Fig. 161).

23.2.2 Palinactinodromous – Three or more primaries diverge in a series of
branches rather than from a single point (Figs. 162, 163).

23.2.3 Acrodromous – Three or more primaries originate from a point and
run in convergent arches toward the leaf apex.

23.2.3.1 Basal – Primary veins radiate from the petiolar insertion point
(Figs. 164, 165).

23.2.3.2 Suprabasal – Primary veins radiate from a point distal to peti-
olar insertion (Fig. 166).

23.2.4 Flabellate – Several to many equally fine basal veins diverge radially at
low angles to each other and branch distally (Fig. 167).

23.2.5 Parallelodromous (typically only in monocot leaves) – Multiple paral-
lel 1º veins originate collaterally at the leaf base and converge toward the
leaf apex (Fig. 168).

23.2.6 Campylodromous (typically only in monocot leaves) – Multiple
parallel 1º veins originate collaterally at or near the leaf base and run in
strongly recurved arches that converge toward the leaf apex (Fig. 169).

Fig. 155 Fig. 156
Pinnate Pinnate
Ostrya guatemalensis Carrierea calycina
(Betulaceae) (Salicaceae)

Fig. 157 Fig. 158
Pinnate Pinnate
Dalechampia cissifolia Croton hircinus
(Euphorbiaceae) (Euphorbiaceae)

Fig. 159 Fig. 160
Basal actinodromous Basal actinodromous
Dombeya elegans Tetrameles nudiflora
(Malvaceae) (Datiscaceae)

Fig. 161 Fig. 162 Fig. 163
Suprabasal actinodromous Palinactinodromous Palinactinodromous
Phoebe costaricana Platanus racemosa Trichosanthes formosana
(Lauraceae) (Platanaceae) (Curcurbitaceae)

Fig. 164 Fig. 165
Basal acrodromous Basal acrodromous
Paliurus ramosissimus Sarcorhachis naranjoana
(Rhamnaceae) (Piperaceae)

Fig. 166 Fig. 167
Suprabasal acrodromous Flabellate
Topobea watsonii Paranomus sceptrum
(Melastomataceae) (Proteaceae)

Fig. 168 Fig. 169
Parallelodromous Campylodromous
Potamogeton amplifolius Maianthemum dilatatum
(Potamogetonaceae) (Ruscaceae)

24. Naked Basal Veins

24.1 Absent (Figs. 165, 166).

24.2 Present – the exmedial side of one or both lateral primaries, or of basal second-
aries, forms part of the leaf margin at the base (Fig. 170).

Fig. 170
Naked basal primary veins
Trichosanthes formosana
(Curcurbitaceae)

25. Number of Basal Veins – Total number of 1º and 2º veins that originate in the base
of the leaf and have courses similar to the course(s) of the primary or primaries. The leaf
in figure 171 has six basal veins; the leaf in figure 172 has one basal vein.

Fig. 171
Six basal veins
Acer miyabei
(Sapindaceae)

Fig. 172
One basal vein
Sorbus japonica
(Rosaceae)

26. Agrophic Veins – A comblike complex of veins composed of a lateral 1° or 2° vein with
two or more excurrent minor 2° veins that originate on it and travel roughly parallel
courses toward the margin. The latter may be straight or looped and are only exterior
(not bilaterally paired along the vein of origin). Agrophic veins are similar to pectinal
veins as defined by Spicer (1986).

26.1 Absent (Figs. 172, 173)

26.2 Present

26.2.1 Simple – One or two agrophic veins. These may be paired (Fig. 174)
or appear on only one half of the leaf.

26.2.2 Compound – More than two agrophic veins (Figs. 175, 176).
These secondaries are
dichotomizing, so they
are not agrophic veins.

Fig. 173 Fig. 174
Agrophic veins absent Simple agrophic veins
Eucryphia glandulosa Alchornea tiliifolia
(Cunoniaceae) (Euphorbiaceae)

Fig. 175 Fig. 176
Compound agrophic veins Compound agrophic veins
Parrotia jacquemontiana Cissus caesia
(Hamamelidaceae) (Vitaceae)

27. Major Secondary Vein Framework – To describe 2º vein framework characters,
examine the courses of the 2º veins in the middle of the lamina. There is no obligate
relationship between secondary course and margin type: all major types of secondary
course occur in both entire-margined and toothed leaves. When secondaries branch di-
chotomously, the branches are also considered to be secondaries. This is important in dis-
tinguishing eucamptodromous from semicraspedodromous secondaries, for example.

27.1 Major secondaries (or their branches) reach the margin.

27.1.1 Craspedodromous – Secondaries terminate at the margin (Figs. 177,
178) or at the marginal vein. It is possible, although rare, to have both
craspedodromous secondaries and an entire margin (Fig. 179).

27.1.2 Semicraspedodromous (usually in toothed leaves) – Secondaries
branch near the margin; one of the branches terminates at the margin,
and the other joins the superjacent secondary (Figs. 180–182).

27.1.3 Festooned semicraspedodromous – Secondaries form more than
one set of loops, with branches from the most exmedial loops terminat-
ing at the margin (Figs. 183–185).

27.2 Major secondaries and their branches do not reach the margin and
lose gauge by attenuation.

27.2.1 Eucamptodromous – Secondaries connect to superjacent major second-
aries via tertiaries without forming marginal loops of secondary gauge
(Figs. 186–188). Three special cases are noted.

27.2.1.1 Basal eucamptodromous – All eucamptodromous secondar-
ies arise from the base of the leaf (<0.25L; Fig. 189). May be
difficult to distinguish from acrodromous primaries (II.23.2.3;
Figs. 164–166).

27.2.1.2 Hemieucamptodromous – All eucamptodromous secondar-
ies arise from the proximal half of the leaf (Fig. 190).

27.2.1.3 Eucamptodromous becoming brochidodromous dis-
tally – Proximal secondaries are eucamptodromous, but distal
secondaries form loops of secondary gauge (Fig. 191).

27.2.2 Reticulodromous – Secondaries branch into a reticulum of higher-
order veins (Fig. 192).

27.2.3 Cladodromous – Secondaries freely ramify exmedially (Fig. 193).

27.3 Major secondaries form loops of secondary gauge and do not reach
the margin.

27.3.1 Simple brochidodromous – Secondaries join in a series of prominent
arches or loops of secondary gauge. Junctions between secondaries are
excurrent and the smaller vein has >25% of the gauge of the larger vein
at the junction (Figs. 194–196).

27.3.2 Festooned brochidodromous – Secondaries branch into multiple
sets of loops of secondary gauge, often with accessory loops of higher
gauge (Figs. 197–199).

27.4 Mixed – Major secondary course varies within the leaf (Fig. 200).

Fig. 177
Major secondaries craspedodromous
Corylopsis glabrescens
(Hamamelidaceae)

2˚

Fig. 178 Fig. 179
Major secondaries craspedodromous Major secondaries craspedodromous
Desfontainea spinosa Cyclea merrillii
(Desfontaineaceae) (Menispermaceae)

Fig. 180
Major secondaries semicraspedodromous
Aphaerema spicata
(Salicaceae)

Fig. 181 Fig. 182
Semicraspedodromous Semicraspedodromous
Cercidiphyllum japonicum Casearia ilicifolia
(Cercidiphyllaceae) (Salicaceae)

Fig. 183
Major secondaries festooned semicraspedodromous
Laurelia novae-zelandiae
(Atherospermataceae)

Fig. 184 Fig. 185
Festooned semicraspedodromous Festooned semicraspedodromous
Tetracentron sinense Mahonia wilcoxii
(Trochodendraceae) (Berberidaceae)

Fig. 186
Major secondaries eucamptodromous
Tetracera rotundifolia
(Dilleniaceae)

Fig. 187 Fig. 188 Fig. 189
Eucamptodromous Eucamptodromous Basal eucamptodromous
Cornus officinalis Isoptera lissophylla Tococa aristata
(Cornaceae) (Dipterocarpaceae) (Melastomataceae)

brochidodromous

eucamptodromous
Fig. 190 Fig. 191
Hemieucamptodromous Eucamptodromous becoming brochidodromous distally
Cleistanthus oligophlebius Rhamnidium elaeocarpum
(Phyllanthaceae) (Rhamnaceae)

Fig. 192 Fig. 193
Reticulodromous Cladodromous
Eucryphia moorei Cotinus obovatus
(Cunoniaceae) (Anacardiaceae)

Fig. 194
Major secondaries brochidodromous
Baccaurea staudtii
(Phyllanthaceae)

Fig. 195 Fig. 196
Brochidodromous Brochidodromous
Santiria samarensis Aextoxicon punctatum
(Burseraceae) (Aextoxicaceae)

Fig. 197
Major secondaries festooned brochidodromous
Antigonon cinerascens
(Polygonaceae)

craspedodromons

brochidodromons

Fig. 198 Fig. 199 Fig. 200
Major secondaries festooned Major secondaries festooned Major secondaries mixed
brochidodromous brochidodromous Comocladia glabra
Capsicodendron pimenteira Tapura guianensis (Anacardiaceae)
(Canellaceae) (Dichapetalaceae)
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
72 Manual of Leaf Architecture

28. Interior Secondaries

28.1 Absent (Figs. 181, 197, 205).

28.2 Present – These secondaries cross between 1º veins or between 1º and perimar-
ginal 2º veins (see II.30) but do not reach the margin (Figs. 201–203). They are
typically arched or straight and are present in the central part of many palmately
lobed leaves, where they may have a course similar to adjacent 3º veins. Interior
secondaries may also occur in leaves with acrodromous 1º veins, intramarginal
2º veins (Fig. 203), or basally eucamptodromous secondaries.

interior 2˚

Fig. 201
Interior secondary
Filipendula occidentalis interior 2˚
(Rosaceae)

interior 2˚

Fig. 202 Fig. 203
Interior secondary Interior secondary
Triplochiton scleroxylon Scaphocalyx spathacea
(Malvaceae) (Achariaceae)

29. Minor Secondary Course

29.1 Craspedodromous – Terminating at the margin (Fig. 204).

29.2 Simple brochidodromous – Joined together in a series of prominent arches
or loops of secondary gauge. Junctions between secondaries are excurrent and
the smaller vein has >25% of the gauge of the larger (Fig. 205).

29.3 Semicraspedodromous – Minor secondaries branch near the margin. One
of the branches eventually terminates at the margin, and the other joins the
superjacent minor secondary (Fig. 206).

minor 2˚

Fig. 204 Fig. 205
Minor secondary course Minor secondary course
craspedodromous simple brochidodromous
Viburnum setigerum Bixa orellana
(Adoxaceae) (Bixaceae)

Fig. 206
Minor secondary course
semicraspedodromous
Philactis zinnioides
(Asteraceae)

30. Perimarginal Veins – When present, these veins closely parallel the leaf margin and
lose little gauge distally.

30.1 Marginal secondary – Vein of 2° gauge running on the leaf margin (Fig. 207).
There are no veins exmedial to a marginal secondary.

30.2 Intramarginal secondary – Vein of 2º gauge running near the margin with
laminar tissue exmedial to it (Figs. 208, 209). Intramarginal veins typically are
intersected by major secondaries.

30.3 Fimbrial vein – Vein of consistent 3° gauge running on the margin with no
laminar tissue exmedial to it (Fig. 210).

marginal 2˚ intramarginal 2˚ intramarginal 2˚

Fig. 207 Fig. 208 Fig. 209
Marginal secondary Intramarginal secondary Intramarginal secondary
Securidaca marginata Spondias bivenomarginalis Graffenrieda anomala
(Polygalaceae) (Anacardiaceae) (Melastomataceae)

fimbrial vein

Fig. 210
Fimbrial vein
Castanea sativa
(Fagaceae)

31. Major Secondary Spacing – Variation in the distance between adjacent secondaries,
measured at their intersections with the midvein.

31.1 Regular – Secondary spacing proportionally decreases distally and proximally
(Fig. 211).

31.2 Irregular – Secondary spacing varies over the lamina (Fig. 212).

31.3 Decreasing proximally – Secondary spacing decreases toward base (Fig.
213); may be regular or irregular.

31.4 Gradually increasing proximally – Secondary spacing increases gradually
toward base (Fig. 214).

31.5 Abruptly increasing proximally – Secondary spacing increases abruptly
toward base (Fig. 215).

Fig. 211 Fig. 212
Secondary spacing regular Secondary spacing irregular
Vitex limonifolia Kermadecia sinuata
(Lamiaceae) (Proteaceae)

Fig. 213
Secondary spacing decreasing proximally
Glochidion bracteatum
(Phyllanthaceae)

Fig. 214 Fig. 215
Secondary spacing gradually increasing proximally Secondary spacing abruptly increasing proximally
Populus jackii Apeiba macropetala
(Salicaceae) (Malvaceae)

32. Variation of Major Secondary Angle to Midvein – Each angle measured on the
distal side of the junction (the vertex) of the secondary with the midvein. One ray of the
angle follows the midvein distal to the junctions, and the other follows the secondary for
25% of its length. The major secondary angle should be evaluated proximal to 0.75 lm.

32.1 Uniform – Major 2° angle varies <10° from the base to 0.75 lm (Fig. 216).

32.2 Inconsistent – Major 2° angle varies >10° from the base to 0.75 lm (Fig. 217).

32.3 Smoothly increasing proximally (Fig. 218).

32.4 Smoothly decreasing proximally (Fig. 219).

32.5 Abruptly increasing proximally (Fig. 220).

32.6 One pair of acute basal secondaries (Figs. 215, 221).

Fig. 216 Fig. 217
Secondary angle uniform Secondary angle inconsistent
Tilia heterophylla Alchornea polyantha
(Malvaceae) (Euphorbiaceae)

Fig. 218 Fig. 219
Secondary angle smoothly increasing proximally Secondary angle smoothly decreasing proximally
Pseudolmedia laevis Popowia congensis
(Moraceae) (Annonaceae)

Fig. 220 Fig. 221
Secondary angle abruptly increasing proximally One pair of acute basal secondaries
Banisteriopsis laevifolia Microcos tomentosa
(Malpighiaceae) (Malvaceae)

33. Major Secondary Attachment to Midvein

33.1 Decurrent – Major secondaries meet the midvein asymptotically (Fig. 129, 222).

33.2 Proximal secondaries decurrent – Major secondaries near the lamina base
are decurrent on midvein, though distal secondaries are excurrent (Fig. 223).

33.3 Excurrent – Major secondaries join the midvein without deflecting it, midvein
monopodial (Fig. 224).

33.4 Deflected – Midvein is deflected at junctions with major secondaries and is thus
sympodial (Fig. 225).

Fig. 222 Fig. 223
Decurrent secondary attachment Proximal secondaries decurrent
Itea chinensis Crataegus brainerdii
(Iteaceae) (Rosaceae)

Fig. 224 Fig. 225
Excurrent secondary attachment Deflected secondary attachment
Tetracera podotricha Celtis cerasifera
(Dilleniaceae) (Cannabaceae)

34. Intersecondary Veins – Veins with courses similar to those of the major secondar-
ies, but generally shorter in exmedial extent and intermediate in gauge between major
secondaries and tertiaries (Fig. 226).

34.1 Intersecondary proximal course

34.1.1 Parallel to major secondaries (Fig. 227).

34.1.2 Perpendicular to midvein (Fig. 228).

34.2 Intersecondary length

34.2.1 Less than 50% of subjacent secondary (Fig. 229).

34.2.2 More than 50% of subjacent secondary (Fig. 230).

34.3 Intersecondary distal course

34.3.1 Reticulating or rami- intersecondary
fying – Branching and
losing a defined course
(Fig. 231).

34.3.2 Parallel to a major sec-
ondary (Fig. 232).

34.3.3 Perpendicular to a sub-
jacent major second-
ary (Fig. 233).

34.3.4 Basiflexed but not join-
ing the subjacent sec-
ondary at right angles
(Fig. 234).

34.4 Intersecondary frequency –
Average number of intersecond-
ary veins per intercostal area

34.4.1 Less than one per inter-
costal area (Fig. 235).

34.4.2 Usually one per inter-
costal area (Fig. 236).

34.4.3 More than one per in-
tercostal area (Fig. 237).
Fig. 226
Intersecondary veins
Couepia paraensis
(Chrysobalanaceae)

intersecondary

Fig. 227 Fig. 228
Proximal course of intersecondary Proximal course of intersecondary
parallel to major secondaries perpendicular to midvein
Protium subserratum Dacryodes negrensis
(Burseraceae) (Burseraceae)

intersecondary

Fig. 229 Fig. 230
Length of intersecondary <50% of subjacent secondary Length of intersecondary >50% of subjacent secondary
Protium opacum Santiria griffithii
(Burseraceae) (Burseraceae)

2˚

intersecondary

intersecondary
Fig. 231 Fig. 232
Distal course of intersecondary Distal course of intersecondary
reticulating or ramifying parallel to major secondary
Comocladia cuneata Ancistrocladus tectorius
(Anacardiaceae) (Ancistrocladaceae)

intersecondary intersecondary

Fig. 233 Fig. 234
Distal course of intersecondary perpendicular Distal course of intersecondary basiflexed
to subjacent major secondary Stemonoporus nitidus
Canarium ovatum (Dipterocarpaceae)
(Burseraceae)

Fig. 235 Fig. 236
Frequency of intersecondary veins Frequency of intersecondary veins
<1 per intercostal area ~1 per intercostal area
Guarea tuberculata Cedrela angustifolia
(Meliaceae) (Meliaceae)

Fig. 237
Frequency of intersecondary veins >1 per intercostal area
Ouratea aff. garcinioides
(Ochnaceae)

35. Intercostal Tertiary Vein Fabric – The three major categories are percurrent (35.1),
reticulate (35.2), and ramified (35.3).

35.1 Percurrent – Tertiaries cross between adjacent secondaries.

35.1.1 Course of percurrent tertiaries

35.1.1.1 Opposite – Majority of tertiaries cross between adjacent sec-
ondaries in parallel paths without branching (Figs. 238–241).

35.1.1.1.1 Straight – Passing across the intercostal area without
a noticeable change in course (Fig. 238).

35.1.1.1.2 Convex – Middle portion of the vein arches exmedi-
ally, without an inflection point (Fig. 239).

35.1.1.1.3 Sinuous – Changes direction of curvature (Fig. 240).

35.1.1.1.4 Forming a chevron – Most tertiary courses have a
markedly sharp bend (Fig. 241).

35.1.1.2 Alternate – Majority of tertiaries cross between secondaries
with regular offsets (abrupt angular discontinuities) near the
middle of the intercostal area (Fig. 242).

35.1.1.3 M ixed – Tertiaries have both opposite and alternate percur-
rent courses (Fig. 243).

35.1.2 Angle of percurrent tertiaries – Angle formed between the midvein
trend and the course of a percurrent 3º vein projected to the midvein
(Fig. 244).

35.1.2.1 Acute – Angle <90° (Fig. 245).

35.1.2.2 Obtuse – Angle >90° (Fig. 246).

35.1.2.3 Perpendicular – Angle ~90° (Fig. 247).

35.2 Reticulate – Veins anastomose with other tertiary veins or secondary veins to
form a net (Figs. 248, 249).

35.2.1 Irregular – Tertiaries anastomose at various angles to form irregular
polygons (Fig. 248) or non-polygonal nets.

35.2.2 Regular – Tertiaries anastomose with other tertiaries at regular angles
to generate a regular polygonal field (Fig. 249).

35.2.3 Composite admedial – Tertiaries connect to a trunk that ramifies
admedially toward the axil of the subjacent costal secondary (Fig. 250).

35.3 Ramified – Tertiaries branch without forming a tertiary reticulum.

35.3.1 Admedially ramified – Multiple tertiary veins branch toward the
primary or midvein (Fig. 251).

35.3.2 Exmedially ramified – Tertiary branching is oriented toward the leaf
margin (Fig. 252).

35.3.3 Transversly ramified – Opposed 3º veins from adjacent major sec-
ondaries ramify and join at a higher vein order (Fig. 253).

35.3.4 Transversly freely ramified – Tertiary veins originating on one
secondary vein branch toward adjacent secondary but do not join other
veins from the opposing secondary (Fig. 254).

Fig. 238 Fig. 239 Fig. 240 Fig. 241
Straight Convex Sinuous Forming a chevron
Melicytus fasciger unknown genus Aphelandra pulcherrima Vallea stipularis
(Violaceae) (Dipterocarpaceae) (Acanthaceae) (Elaeocarpaceae)

3˚
2˚
2˚
3˚

Fig. 242 Fig. 243
Alternate percurrent tertiary fabric Mixed percurrent tertiary fabric
Santiria samarensis Davilla rugosa
(Burseraceae) (Dilleniaceae)

1˚
3˚

acute
3˚

3˚ angle

obtuse

3˚

Fig. 244 Fig. 245
Measurement of tertiary angle with Acute tertiary angles
respect to the 1º vein Nectandra cuspidata
(Lauraceae)

3˚ 3˚

Fig. 246 Fig. 247
Obtuse tertiary angle Perpendicular tertiary angle
Sloanea eichleri Bhesa archboldiana
(Elaeocarpaceae) (Celastraceae)

1˚

3˚

Fig. 248 1˚ 2˚ 3˚
Irregular reticulate tertiary fabric
Piranhea trifoliata
(Picrodendraceae)

1˚

Fig. 249
Regular reticulate tertiary fabric
3˚ Afrostyrax kamerunensis
(Huaceae)

2˚

Fig. 250
Composite admedial
Sorindeia gilletii
(Anacardiaceae)

1˚ 2˚

1˚ 3˚

3˚

2˚

Fig. 251 Fig. 252
Simple admedially ramified tertiary fabric Exmedially ramified tertiary fabric
Protorhus nitida Ouratea thyrsoidea
(Anacardiaceae) (Ochnaceae)

1˚ 2˚ 3˚

Fig. 253 Fig. 254
Transversely ramified tertiary fabric Transversely freely ramified tertiary fabric
Comocladia glabra Rhus taitensis
(Anacardiaceae) (Anacardiaceae)

36. Intercostal Tertiary Vein Angle Variability – Applies only to leaves with percur-
rent tertiaries; see 35.1.2 for measuring the angle. A leaf may exhibit more than one
character state.

36.1 Inconsistent – Angles of the tertiaries vary randomly over the lamina (Fig. 255).

36.2 Consistent – Angles of the tertiaries do not vary over the surface of the lamina
by more than 10% (Fig. 256).

36.3 Increasing exmedially – Angles of the tertiaries become more obtuse away
from the midvein (Fig. 257).

36.3.1 B asally concentric – Special case of “increasing exmedially” such
that the tertiaries form a “spider web pattern” around the primary vein(s)
at the base of the leaf (Fig. 258).

36.4 Decreasing exmedially – Angles of the tertiaries become more acute away
from the midvein (Fig. 259).

36.5 Increasing proximally – Angles of the tertiaries become more obtuse toward
the base of the lamina (Fig. 260).

36.6 Decreasing proximally – Angles of the tertiaries become more acute toward
the base of the lamina (Fig. 261).

3˚ angle

Fig. 255 Fig. 256 Fig. 257
Inconsistent tertiary angle Consistent tertiary angle Tertiary angle increasing
Viburnum sempervirens Diospyros maritima exmedially
(Adoxaceae) (Ebenaceae) Eriolaena malvacea
(Malvaceae)

Fig. 258 Fig. 259
Basally concentric Tertiary angle decreasing exmedially
Macaranga bicolor Juglans boliviana
(Euphorbiaceae) ( Juglandaceae)

Fig. 260 Fig. 261
Tertiary angle increasing proximally Tertiary angle decreasing proximally
Odontadenia geminata Flacourtia rukam
(Apocynaceae) (Salicaceae)

37. Epimedial Tertiaries – Tertiaries that intersect a 1º vein.

37.1 Epimedial tertiary fabric

37.1.1 Percurrent – Epimedial veins cross between 1º and 2º veins.

37.1.1.1 Opposite percurrent – Majority of tertiaries cross between
primary and secondaries in parallel paths without branching
(Fig. 262).

37.1.1.2 Alternate percurrent – Majority of tertiaries cross between
primary and secondaries with regular offsets (abrupt angular
discontinuities) (Fig. 263).

37.1.1.3 Mixed – Approximately equal numbers of opposite and alter-
nate percurrent tertiaries (Fig. 264).

37.1.2 Ramified – Epimedial tertiaries branch toward the leaf margin (Fig. 265).

37.1.3 Reticulate – Epimedial tertiaries anastomose with other 3° veins to
form a net (Fig. 266).

37.1.4 Mixed – Epimedial tertiaries do not consistently exhibit one character-
istic (Fig. 267).

37.2 Course of percurrent epimedial tertiaries

37.2.1 Proximal/admedial course of the epimedial tertiaries – Course
of the epimedial tertiaries from their junction with the midvein to their
approximate midpoint. More than one character state may apply.

37.2.1.1 Parallel to the subjacent secondary (Fig. 268).

37.2.1.2 Parallel to the intercostal tertiaries (Fig. 269).

37.2.1.3 Perpendicular to the midvein (Fig. 270).

37.2.1.4 Parallel to the intersecondary (Fig. 271).

37.2.1.5 Obtuse to the midvein (Fig. 272).

37.2.1.6 Acute to the midvein (Fig. 273).

37.2.2 Distal/exmedial course of the epimedial tertiaries – Course of
the epimedial tertiaries from their approximate midpoint to their inter-
section with the adjacent secondary (if not ramifying or reticulating).
Note: More than one character state may apply.

37.2.2.1 Parallel to intercostal tertiary – Epimedial tertiaries match
pattern of adjacent intercostal tertiaries (Fig. 274).

37.2.2.2 Basiflexed – Course bends toward the base of the leaf and may
either join the secondaries or lose gauge (Fig. 274, 275).

37.2.2.3 Acroflexed – Course bends toward the apex of the leaf and
may either join the secondaries or lose gauge (Fig. 276).

Fig. 262 Fig. 263 Fig. 264
Opposite percurrent Alternate percurrent Mixed percurrent
epimedial tertiaries epimedial tertiaries epimedial tertiaries
Actinidia latifolia Alangium chinense Bixa orellana
(Actinidiaceae) (Cornaceae) (Bixaceae)

Fig. 265 Fig. 266 Fig. 267
Ramified epimedial tertiaries Reticulate epimedial tertiaries Mixed epimedial tertiaries
Ouratea thyrsoidea Mahonia wilcoxii Aphelandra pulcherrima
(Ochnaceae) (Berberidaceae) (Acanthaceae)

1˚ 3˚ 2˚ 1˚ 2˚ 3˚

Fig. 268 Fig. 269
Proximal course of the epimedial tertiaries Proximal course of the epimedial tertiaries
parallel to subjacent secondary parallel to intercostal tertiary
Capparis lundellii Callicoma serratifolia
(Capparaceae) (Cunoniaceae)

1˚ 1˚ 1˚

inter –
3˚
2˚
2˚
3˚

3˚

2˚

Fig. 270 Fig. 271 Fig. 272
Proximal course of the epimedial Proximal course of the Proximal course of the epimedial
tertiaries perpendicular to epimedial tertiaries parallel to tertiaries obtuse to the midvein
the midvein the intersecondary Sloanea eichleri
Lunania mexicana Celastrus racemosus (Elaeocarpaceae)
(Salicaceae) (Celastraceae)

3˚

Fig. 273 Fig. 274
Proximal course of the epimedial Distal course of the epimedial tertiaries
tertiaries acute to midvein parallel to intercostal tertiaries
Bixa orellana Theobroma microcarpa
(Bixaceae) (Malvaceae)

3˚

Fig. 275 Fig. 276
Distal course of the epimedial tertiaries basiflexed Distal course of the epimedial tertiaries acroflexed
Spiropetalum erythrosepalum Commiphora aprevalii
(Connaraceae) (Burseraceae)

38. Exterior Tertiary Course – Configuration of the third-order veins that lie exmedially
to the outermost secondaries but do not necessarily form the marginal ultimate veins.

38.1 Absent – Leaf does not have exterior tertiaries (Fig. 277).

38.2 Looped – Tertiaries form loops (Figs. 278, 279).

38.3 Terminating at the margin – Tertiaries terminate at the margin (Figs. 280, 281).

38.4 Variable – Pattern is not consistent (Fig. 282).

3˚ 3˚

Fig. 277 Fig. 278 Fig. 279
Exterior tertiaries absent Exterior tertiaries looped Exterior tertiaries looped
Hedyosmum costaricense Picramnia krukovii Mollinedia floribunda
(Chloranthaceae) (Picramniaceae) (Monimiaceae)

terminates
at margin

3˚

looped
3˚

Fig. 280 Fig. 281 Fig. 282
Exterior tertiaries Exterior tertiaries Exterior tertiaries variable
terminating at margin terminating at margin Gymnosporia senegalensis
Barringtonia reticulata Carissa bispinosa (Celastraceae)
(Lecythidaceae) (Apocynaceae)

39. Quaternary
Vein Fabric – Pattern formed by fourth-order vein courses. This and
other higher-order venation characters should be scored near the center of the blade.

39.1 Percurrent

39.1.1 Opposite – Most quaternary veins cross between adjacent tertiary veins
in parallel paths without branching (Fig. 283).

39.1.2 Alternate – Most quaternary veins cross between adjacent tertiaries
with an offset (an abrupt angular discontinuity) (Fig. 284).

39.1.3 Mixed percurrent – Quaternaries are alternate and opposite in equal
proportions (Fig. 285).

39.2 Reticulate – Quaternaries anastomose with other veins to form a net.

39.2.1 Regular – Angles formed by the vein intersections are regular (Fig. 286).

39.2.2 Irregular – Angles formed by the vein intersections are highly variable
(Fig. 287).

39.3 Freely ramifying – Quaternaries branch freely and are the finest vein-order
the leaf exhibits (Fig. 288).
2˚ 3˚ 4˚ 1˚ 2˚ 3˚ 4˚ 1˚ 3˚ 4˚ 2˚

Fig. 283 Fig. 284 Fig. 285
Opposite percurrent quaternaries Alternate percurrent quaternaries Mixed percurrent quaternary
Shorea congestiflora Theobroma microcarpa Alangium chinense
(Dipterocarpaceae) (Malvaceae) (Cornaceae)

1˚ 3˚ 4˚ 2˚ 1˚ 2˚ 3˚ 4˚ 1˚ 3˚ 2˚ 4˚

Fig. 286 Fig. 287 Fig. 288
Regular reticulate quaternaries Irregular reticulate quaternaries Freely ramifying quaternary
Afrostyrax kamerunensis Diospyros pellucida Comocladia cuneata
(Huaceae) (Ebenaceae) (Anacardiaceae)

40. Quinternary Vein Fabric – Pattern formed by 5° vein courses, when present. This and
other higher-order venation characters should be scored near the center of the blade.

40.1 Reticulate – Quinternaries anastomose with other veins to form polygons.

40.1.1 Regular – Angles formed by vein intersections are regular (Fig. 289).

40.1.2 Irregular – Angles formed by vein intersections are highly variable (Fig. 290).

40.2 Freely ramifying – Quinternaries branch freely and are the finest vein-order
the leaf exhibits (Fig. 291).

2˚ 3˚ 4˚ 5˚ 2˚ 3˚ 4˚ 5˚

Fig. 289 Fig. 290
Regular reticulate quinternaries Irregular reticulate quinternaries
Pseudolmedia laevis Diospyros hispida
(Moraceae) (Ebenaceae)

3˚ 2˚ 4˚

5˚

Fig. 291
Freely ramifying quinternaries
Stemonoporus nitidus
(Dipterocarpaceae)

41. Areolation – Areoles are the smallest areas of the leaf tissue that are completely sur-
rounded by veins; taken together they form a contiguous field of polygons over most of
the area of the lamina. Any order of venation can form one or more sides of an areole.

41.1 Lacking – Venation ramifies into the intercostal area without producing closed
meshes (Fig. 292).

41.2 Present

41.2.1 Poor development – Areoles many-sided (often >7) and of highly ir-
regular size and shape (Fig. 293).

41.2.2 Moderate development – Areoles of irregular shape, more or less
variable in size, generally with fewer sides than in poorly developed
areolation (Fig. 294).

41.2.3 Good development – Areoles of relatively consistent size and shape
and generally with 3–6 sides (Fig. 295).

41.2.4 Paxillate – Areoles occurring in distinct oriented fields (Fig. 296; defini-
tion is more general than in Hickey, 1979.)

Fig. 292 Fig. 293 Fig. 294
Areolation lacking Areole development poor Areole development moderate
Rhus taitensis Chloranthus glaber Clusiella pendula
(Anacardiaceae) (Chloranthaceae) (Clusiaceae)

Fig. 295 Fig. 296
Areole development good Areole development paxillate
Piranhea trifoliata Afrostyrax kamerunensis
(Picrodendraceae) (Huaceae)

42. Freely Ending Veinlets (FEVs) – Highest-order veins that freely ramify.

42.1 FEV branching

42.1.1 FEVs absent (Fig. 297).

42.1.2 Mostly unbranched – FEVs present but unbranched, may be straight
or curved (Fig. 298).

42.1.3 Mostly with one branch (Fig. 299).

42.1.4 Mostly with two or more branches

42.1.4.1 Branching equal (dichotomous) (Fig. 300).

42.1.4.2 Branching unequal (dendritic) (Fig. 301).

42.2 FEV terminals

42.2.1 Simple (Fig. 302).

42.2.2 Tracheoid idioblasts – FEV endings are club-shaped and consist of
tracheal cells with spiral wall thickenings (Foster, 1956; called dilated tra-
cheal cells in Tucker, 1964) (Fig. 303).

42.2.3 Highly branched sclereids – FEVs branch densely (10+) out of the
plane of the veins; the finer branches often stain differently because they
are sclereids, not tracheids (Fig. 304).

Fig. 297 Fig. 298 Fig. 299 Fig. 300 Fig. 301
FEVs absent FEVs unbranched FEVs one branched FEVs dichotomous FEVs dendritic
branching branching

Fig. 302 Fig. 303 Fig. 304
Simple FEV terminals Tracheoid idioblasts Highly branched sclereids
Melicytus fasciger Bursera inaguensis Tetragastris panamensis
(Violaceae) (Burseraceae) (Burseraceae)

43. Marginal Ultimate Venation – Configuration of the highest-order veins at the mar-
gin (see also II.29 on perimarginal veins)

43.1 Absent – Ultimate veins join perimarginal veins (Fig. 305).

43.2 Incomplete – Marginal ultimate veins recurve to form incomplete loops (Fig. 306).

43.3 Spiked – Marginal ultimate veins form outward-pointing spikes (Fig. 307).

43.4 Looped – Marginal ultimate vein recurved to form loops (Figs. 308, 309).

Fig. 305 Fig. 306 Fig. 307
Marginal ultimate venation absent Marginal ultimate veins Marginal ultimate veins
Pycnocoma littoralis incomplete form spikes
(Euphorbiaceae) (line drawing) (line drawing)

Fig. 308 Fig. 309
Marginal ultimate venation looped Marginal ultimate venation looped
Mollinedia floribunda Picramnia krukovii
(Monimiaceae) (Picramniaceae)

L eaf teeth contain a great number of system-
atically informative characters (Hickey
and Wolfe, 1975; Hickey and Taylor,
1991; Doyle, 2007) and are extremely useful
for circumscribing fossil leaf taxa. Their preva-
Generally, a tooth can be recognized by its
projection from the leaf margin (see I.13
and I.14) and its associated vasculature.
Recognizing the boundaries of a tooth along
the leaf margin can be difficult when sinuses
lence in fossil floras provides reliable proxy data are absent or teeth are widely separated.
about pre-Quaternary terrestrial paleotemper- Some lab-tested, reproducible rules for de-
atures (Wolfe, 1971, 1995; Wilf, 1997; Utescher fining tooth boundaries when high precision
et al., 2000). Tooth size and shape appear to is necessary are found in Royer et al. (2005).
be useful variables for increasing precision in Hickey and Taylor (1991) used tissue-level fea-
paleoclimate estimates and for paleoecological tures to define admedial and conjunctal veins.
interpretation of fossil floras (Royer et al., 2005;
Royer and Wilf, 2006).

Def initions

distal flank
The portion of the margin between the tooth’s sinus
apex and the nadir of the superjacent sinus
(Fig. 310). tooth apex
proximal flank
The portion of the margin between the tooth’s
apex and the sinus on the proximal side. The
proximal sinus is recognized as the point where
the curve of the tooth departs from the curve of distal flank
the leaf margin, and may or may not coincide
with the nadir of the subjacent sinus (Fig. 310).

sinus
A marginal embayment, incision, or
indentation between marginal projections of
any sort, typically lobes (Fig. 11), teeth (Figs.
11, 310) or the base of cordate leaves (Fig. 12) proximal flank
tooth apex
The point of sharpest change in direction
along the tooth margin, commonly but Fig. 310
not always occurring at the most distal or The parts of a tooth
exmedial point on the tooth (Fig. 310).

principal vein
The vein of widest gauge that enters the
tooth (Fig. 311).

admedial vein
The first branch from the principal admedial vein
vein below the tooth apex that is of
the same order or one order finer
than the principal, and has >60%
of its vascular tissue at its junction principal vein
with the principal directed
admedially or toward the mid-line
of the leaf (Fig. 311).
accessory vein,
accessory veins conjunctal vein
All the veins between the tooth
apex and the admedial vein that
either branch from or merge with
the principal vein. Typically the
accessory veins of larger gauge
have consistent courses in relation
to the principal vein, admedial
vein, and other tooth features, and
such accessory veins commonly
are conjunctal veins as defined
below (Fig. 311). Fig. 311
Aporusa frutescens
conjunctal veins (Phyllanthaceae)
Accessory veins that converge
on or merge with the principal
vein, contribute vascular tissue to
the tooth apex, and have > 60% gland
of their vascular tissue directed
toward the tooth apex at their
point of convergence or fusion
with the principal vein. They
may occur singly or in pairs that
arise opposite or alternate to one
another (Fig. 311).

gland
A discrete area of specialized
cells that secrete by-products
of plant metabolism. In fossils
and cleared or dried leaves, the
glands typically appear darker
than the surrounding tissue. In
addition to occurring on the
lamina and petiole, they may oc-
cur in or be attached to the apex
of the tooth (Fig. 312).
Fig. 312
Gland
Gouania velutina
(Rhamnaceae)

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III. Tooth Characters
44. Tooth Spacing – Distance between the corresponding points on adjacent teeth

44.1 Regular – Minimum intertooth distance is >60% of the maximum intertooth
distance (Fig. 313).

44.2 Irregular – Minimum intertooth distance is <60% of the maximum intertooth
distance (Fig. 314).

Fig. 313 Fig. 314
Regular tooth spacing Irregular tooth spacing
Dichroa philippinensis Campylostemon mucronatum
(Hydrangeaceae) (Celastraceae)

45. Number of Orders of Teeth – Number of discrete sizes of teeth. Sometimes, second-
and third-order teeth occur in a regular series between first-order teeth.

45.1 One – All teeth are the same size or vary in size continuously (Fig. 315).

45.2 Two – Teeth are of two distinct sizes (Fig. 316).

45.3 Three – Teeth are of three distinct sizes (Fig. 317).

1st order
1st order

2nd order

Fig. 315 Fig. 316
One order of teeth Two orders of teeth
Leea macropus Aristotelia racemosa
(Vitaceae) (Elaeocarpaceae)

2nd order

3rd order

1st order

Fig. 317
Three orders of teeth
Crataegus brainerdii
(Rosaceae)

46. Number of Teeth per Centimeter – Measured in the middle 50% of the leaf; that is,
between 0.25 and 0.75 L (Fig. 318).

47. Sinus Shape

47.1 Angular (Fig. 319).

47.2 Rounded (Fig. 320).

2X
Fig. 318
Three teeth per cm
Dichroa philippinensis
(Hydrangeaceae)

Fig. 319 Fig. 320
Angular sinus Rounded sinus
Celtis cerasifera Phylloclinium paradoxum
(Cannabaceae) (Salicaceae)

48. Tooth Shape – Described in terms of the distal and proximal flank curvatures relative
to the midline of the tooth. The following states and abbreviations are used: convex (cv),
straight (st), concave (cc), flexuous (fl; tooth flank is apically concave and basally convex),
and retroflexed (rt; tooth flank is basally concave and apically convex). The distal flank
shape is given first: for example, cc/fl indicates that the tooth is concave on the distal
flank and flexuous on the proximal flank. The 25 possible combinations are shown in
Figure 321 below. Note that a given leaf often exhibits more than one tooth shape.

Distal f lank
CV ST CC FL RT

ST
Proximal f lank

Fig. 321
Chart of possible tooth shapes. Always list the distal flank first.

49. Principal Vein

49.1 Present (Figs. 322, 323, 324).

49.2 Absent – Generally occurs in teeth that are supplied
by two or more veins of equal gauge (Fig. 325). principal vein

conjunctal vein

principal vein

Fig. 322 Fig. 323
Principal vein present Principal vein present
Carpinus laxiflora Chloranthus serratus
(Betulaceae) (Chloranthaceae)
scale bar = 1 mm scale bar = 1 mm

admedial vein
principal vein

Fig. 324 Fig. 325
Principal vein present Principal vein absent
Martynia annua Lopesia lopezoides
(Martyniaceae) (Onagraceae)
scale bar = 1 mm scale bar = 100 µm

50. Principal Vein Termination

50.1 Submarginal (Fig. 326).

50.2 Marginal

50.2.1 At the apex of tooth (Fig. 327).

50.2.2 On the distal flank (Fig. 328).

50.2.3 At the nadir of superjacent sinus (Fig. 329).

50.2.4 On the proximal flank (Fig. 330).

principal
vein

Fig. 326 Fig. 327 Fig. 328
Principal vein termination Principal vein terminates Principal vein terminates on
submarginal at the tooth apex the distal flank
Fuchsia decidua Acer negundo Cupania vernalis
(Onagraceae) (Sapindaceae) (Sapindaceae)
scale bar = 100 μm scale bar = 1 mm scale bar = 1 mm

principal
vein
principal
vein
Fig. 329 Fig. 330
Principal vein terminates at Principal vein terminates
nadir of the superjacent sinus on the proximal flank
Elaeodendron glaucum Quercus alba × velutina
(Celastraceae) (Fagaceae)
scale bar = 1 mm scale bar = 5 mm
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Tooth Characters 109

51. Course of Major Accessory Vein(s)

51.1 Convex relative to principal vein (Fig. 331).

51.1.1 Looped – With multiple looping connections to principal vein (Fig. 332).

51.2 Straight or concave to principal vein (Figs. 333, 334).

51.3 Running from sinus to principal vein (Fig. 335).

accessory vein
accessory vein

Fig. 331 Fig. 332
Accessory veins convex Accessory veins looped
Melicytus fasciger Platanus orientalis
(Violaceae) (Platanaceae)
scale bar = 100 µm scale bar = 1 mm
accessory vein

accessory vein

accessory vein,
conjunctal vein

Fig. 333 Fig. 334 Fig. 335
Accessory vein straight Accessory vein concave Accessory vein running from sinus
Diphylleia grayi Vitis cavaleriei Vitis cavaleriei
(Berberidaceae) (Vitaceae) (Vitacea)
scale bar = 1 mm scale bar = 100 µm scale bar = 1 mm
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110 Manual of Leaf Architecture

52. Special Features of the Tooth Apex

52.1 Simple – No tissue or structure is present within or on the tooth apex (Fig. 336).

52.2 Specific tissue or structure present within the tooth apex

52.2.1 Foraminate – Having a bulb- or funnel-shaped cavity at the tooth apex
that opens to the outside (Fig. 337).

52.2.2 Tylate – Having clear tissue at the termination of the principal vein (Fig. 338).

52.2.3 Cassidate – Having opaque tissue at the termination of the principal
vein (Fig. 339).

52.3 Specific tissue or structure on the tooth apex

52.3.1 Spinose – Principal vein extends beyond the leaf margin; extension
may be short or long, usually sharp (Fig. 340).

52.3.2 Mucronate – An opaque, vascularized, peg-shaped, non-deciduous pro-
jection is present at the apex (Fig. 341).

52.3.3 Setaceous – An opaque, peg-shaped, deciduous projection is present at
the apex (Fig. 342).

52.3.4 Papillate – A clear, flame-shaped projection is present at the apex (Fig. 343).

52.3.5 Spherulate – A clear, spherical projection is present at the apex (Fig. 344).

52.4 Nonspecific – In fossils, it is often not possible to distinguish the type of gland
or structures at the tooth apex. This character state can be used for the descrip-
tion of fossil teeth with a visible concentration of material in or on the tooth apex
not assignable to the categories above (Fig. 345).

Fig. 336 Fig. 337
No special features Foraminate tooth apex
Melochia lupulina Circaea erubescens
(Malvaceae) (Onagraceae)
scale bar = 1 mm scale bar = 100 µm

Fig. 338 Fig. 339
Tylate tooth apex Cassidate tooth apex
Homalium racemosum Tetracentron sinense
(Salicaceae) (Trochodendraceae)
scale bar = 1 mm scale bar = 1 mm

Fig. 340 Fig. 341
Spinose tooth apex Mucronate tooth apex
Ilex dipyrena Trimeria alnifolia
(Aquifoliaceae) (Salicaceae)
scale bar = 1 mm scale bar = 1 mm

Fig. 342 Fig. 343
Setaceous tooth apex Papillate tooth apex
Thea sinensis Schumacheria castaneifolia
(Theaceae) (Dilleniaceae)
scale bar = 1 mm scale bar = 10 mm

Fig. 344 Fig. 345
Spherulate tooth apex Nonspecific tooth apex (fossil)
Idesia polycarpa Cercidiphyllum genetrix
(Salicaceae) (Cercidiphyllaceae)
scale bar = 1 mm scale bar = 5 mm

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please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Appendix A. Outline of
Characters and Character States
I. Leaf Characters 7. Position of Lamina Attachment
7.1 Marginal
1. Leaf Attachment 7.2 Peltate central
1.1 Petiolate 7.3 Peltate excentric
1.2 Sessile
8. Laminar Size
2. Leaf Arrangement 8.1 Leptophyll
2.1 Alternate 8.2 Nanophyll
2.2 Subopposite 8.3 Microphyll
2.3 Opposite 8.4 Notophyll
2.4 Whorled 8.5 Mesophyll
8.6 Macrophyll
3. Leaf Organization 8.7 Megaphyll
3.1 Simple
3.2 Compound 9. Laminar L:W Ratio
3.2.1 Palmately compound
3.2.2 Pinnately compound 10. Laminar Shape
3.2.2.1 Once 10.1 Elliptic
3.2.2.2 Twice 10.2 Obovate
3.2.2.3 Thrice 10.3 Ovate
10.4 Oblong
4. Leaflet Arrangement 10.5 Linear
4.1 Alternate 10.6 Special
4.2 Subopposite
4.3 Opposite 11. Medial Symmetry
4.3.1 Odd-pinnately compound 11.1 Symmetrical
4.3.2 Even-pinnately compound 11.2 Asymmetrical
4.4 Unknown
12. Base Symmetry
5. Leaflet Attachment 12.1 Symmetrical
5.1 Petiolulate 12.2 Asymmetrical
5.2 Sessile 12.2.1 Basal width asymmetrical
12.2.2 Basal extension asymmetrical
6. Petiol(ul)e Features 12.2.3 Basal insertion asymmetrical
6.1 Petiol(ul)e base
6.1.1 Sheathing 13. Lobation
6.1.2 Pulvin(ul)ate 13.1 Unlobed
6.2 Glands 13.2 Lobed
6.2.1 Petiolar 13.2.1 Palmately lobed
6.2.2 Acropetiolar 13.2.1.1 Palmatisect
6.3 Petiole cross-section 13.2.2 Pinnately lobed
6.3.1 Terete 13.2.2.1 Pinnatisect
6.3.2 Semi-terete 13.2.3 Palmately and pinnately lobed
6.3.3 Canaliculate 13.2.4 Bilobed
6.3.4 Triangular
6.3.5 Alate
6.4 Phyllodes
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
114 Manual of Leaf Architecture

14. Margin Type 20. Terminal Apex Features
14.1 Untoothed 20.1 Mucronate
14.2 Toothed 20.2 Spinose
14.2.1 Dentate 20.3 Retuse
14.2.2 Serrate
14.2.3 Crenate 21. Surface Texture
21.1 Smooth
15. Special Margin Features 21.2 Pitted
15.1 Appearance of the edge of the blade 21.3 Papillate
15.1.1 Erose 21.4 Rugose
15.1.2 Sinuous 21.5 Pubescent
15.2 Appearance of the plane of the blade
15.2.1 Revolute 22. Surficial Glands
15.2.2 Involute 22.1 Laminar
15.2.3 Undulate 22.2 Marginal
22.3 Apical
16. Apex Angle 22.4 Basal laminar
16.1 Acute
16.2 Obtuse II. Vein Characters
16.3 Reflex
23. Primary Vein Framework
17. Apex Shape 23.1 Pinnate
17.1 Straight 23.2 Palmate
17.2 Convex 23.2.1 Actinodromous
17.2.1 Rounded 23.2.1.1 Basal
17.2.2 Truncate 23.2.1.2 Suprabasal
17.3 Acuminate 23.2.2 Palinactinodromous
17.4 Emarginate 23.2.3 Acrodromous
17.5 Lobed 23.2.3.1 Basal
23.2.3.2 Suprabasal
18. Base Angle 23.2.4 Flabellate
18.1 Acute 23.2.5 Parallelodromous
18.2 Obtuse 23.2.6 Campylodromous
18.3 Reflex
18.4 Circular 24. Naked Basal Veins
24.1 Absent
19. Base Shape 24.2 Present
19.1 lb = 0
19.1.1 Straight (cuneate) 25. Number of Basal Veins
19.1.2 Concave
19.1.3 Convex 26. Agrophic Veins
19.1.3.1 Rounded 26.1 Absent
19.1.3.2 Truncate 26.2 Present
19.1.4 Concavo-convex 26.2.1 Simple
19.1.5 Complex 26.2.2 Compound
19.1.6 Decurrent
19.2 lb > 0 or lb ~ 0 27. Major 2° Vein Framework
19.2.1 Cordate 27.1 Major secondaries reach margin
19.2.2 Lobate 27.1.1 Craspedodromous
19.2.2.1 Sagittate 27.1.2 Semicraspedodromous
19.2.2.2 Hastate 27.1.3 Festooned semicraspedodromous
19.2.2.3 Runcinate
19.2.2.4 Auriculate

27.2 Major secondaries do not reach margin 34. Intersecondary Veins
and lose gauge by attenuation 34.1 Intersecondary proximal course
27.2.1 Eucamptodromous 34.1.1 Parallel to major secondaries
27.2.1.1 Basal eucamptodromous 34.1.2 Perpendicular to midvein
27.2.1.2 Hemieucamtodromous 34.2 Intersecondary length
27.2.1.3 Eucamptodromous 34.2.1 Less than 50% of subjacent
becoming brochidodromous secondary
distally 34.2.2 More than 50% of subjacent
27.2.2 Reticulodromous secondary
27.2.3 Cladodromous 34.3 Intersecondary distal course
27.3 Major secondaries form loops of 2° 34.3.1 Reticulating or ramifying
gauge and do not reach margin. 34.3.2 Parallel to major secondary
27.3.1 Simple brochidodromous 34.3.3 Perpendicular to subjacent major
27.3.2 Festooned brochidodromous secondary
27.4 Mixed 34.3.4 Basiflexed, not joining subjacent
secondary at right angle
28. Interior Secondaries 34.4 Intersecondary frequency
28.1 Absent 34.4.1 Less than 1 per intercostal area
28.2 Present 34.4.2 Usually 1 per intercostal area
34.4.3 More than 1 per intercostal area
29. Minor Secondary Course
29.1 Craspedodromous 35. Intercostal Tertiary Vein Fabric
29.2 Simple brochidodromous 35.1 Percurrent
29.3 Semicraspedodromous 35.1.1 Course of percurrent tertiaries
35.1.1.1 Opposite
30. Perimarginal Veins 35.1.1.1.1 Straight
30.1 Marginal secondary 35.1.1.1.2 Convex
30.2 Intramarginal secondary 35.1.1.1.3 Sinuous
30.3 Fimbrial vein 35.1.1.1.4 Chevroned
35.1.1.2 Alternate
31. Major Secondary Spacing 35.1.1.3 Mixed opposite-alternate
31.1 Regular 35.1.2 Angle of percurrent tertiaries
31.2 Irregular 35.1.2.1 Acute
31.3 Decreasing proximally 35.1.2.2 Obtuse
31.4 Gradually increasing proximally 35.1.2.3 Perpendicular
31.5 Abruptly increasing proximally 35.2 Reticulate
35.2.1 Irregular
32. Variation of Major Secondary 35.2.2 Regular
Angle to Midvein 35.2.3 Composite admedial
32.1 Uniform 35.3 Ramified
32.2 Inconsistent 35.3.1 Admedially ramified
32.3 Smoothly increasing proximally 35.3.2 Exmedially ramified
32.4 Smoothly decreasing proximally 35.3.3 Transverse ramified
32.5 Abruptly increasing proximally 35.3.4 Transverse freely ramified
32.6 One pair acute basal secondaries

33. Major Secondary Attachment to Midvein
33.1 Decurrent
33.2 Proximal secondaries decurrent
33.3 Excurrent
33.4 Deflected

36. Intercostal Tertiary Vein Angle Variability 40. Quinternary Vein Fabric
36.1 Inconsistent 40.1 Reticulate
36.2 Consistent 40.1.1 Regular
36.3 Increasing exmedially 40.1.2 Irregular
36.3.1 Basally concentric 40.2 Freely ramifying
36.4 Decreasing exmedially
36.5 Increasing proximally 41. Areolation
36.6 Decreasing proximally 41.1 Lacking
41.2 Present
37. Epimedial Tertiaries 41.2.1 Poor development
37.1 Epimedial tertiary fabric 41.2.2 Moderate development
37.1.1 Percurrent 41.2.3 Good development
37.1.1.1 Opposite 41.2.4 Paxillate
37.1.1.2 Alternate
37.1.1.3 Mixed 42. Freely Ending Veinlets (FEVs)
37.1.2 Ramified 42.1 FEV branching
37.1.3 Reticulate 42.1.1 FEVs absent
37.1.4 Mixed 42.1.2 Mostly unbranched
37.2 Course of percurrent epimedial 42.1.3 Mostly 1-branched
tertiaries 42.1.4 Mostly 2- or more branched
37.2.1 Admedial course 42.1.4.1 Branching equal
37.2.1.1 Parallel to subjacent (dichotomous)
secondary 42.1.4.2 Branching unequal
37.2.1.2 Parallel to intercostal (dendritic)
tertiaries 42.2 FEV terminals
37.2.1.3 Perpendicular to midvein 42.2.1 Simple
37.2.1.4 Parallel to intersecondary 42.2.2 Tracheoid idioblasts
37.2.1.5 Obtuse to midvein 42.2.3 Highly branched sclereids
37.2.1.6 Acute to midvein
37.2.2 Exmedial course 43. Marginal Ultimate Venation
37.2.2.1 Parallel to intercostal 43.1 Absent
tertiary 43.2 Incomplete loops
37.2.2.2 Basiflexed 43.3 Spiked
37.2.2.3 Acroflexed 43.4 Looped

38. Exterior Tertiary Course III. Tooth Characters
38.1 Absent
38.2 Looped 44. Tooth Spacing
38.3 Terminating at margin 44.1 Regular
38.4 Variable 44.2 Irregular

39. Quaternary Vein Fabric 45. Number of Orders of Teeth
39.1 Percurrent 45.1 One
39.1.1 Opposite 45.2 Two
39.1.2 Alternate 45.3 Three
39.1.3 Mixed percurrent
39.2 Reticulate 46. Number of Teeth/cm
39.2.1 Regular
39.2.2 Irregular 47. Sinus Shape
39.3 Freely ramifying 47.1 Angular
47.2 Rounded

48. Tooth Shape (cv, st, cc, fl, rt)
(distal flank listed first)

49. Principal Vein
49.1 Present
49.2 Absent

50. Principal Vein Termination
50.1 Submarginal
50.2 Marginal
50.2.1 At apex of tooth
50.2.2 On distal flank
50.2.3 At nadir of superjacent sinus
50.2.4 On proximal flank

51. Course of Ancillary Veins Relative to
Principal Vein
51.1 Convex
51.1.1 Looped
51.2 Straight or concave
51.3 Running from sinus

52. Special Features of the Tooth Apex
52.1 None
52.2 Within tooth apex
52.2.1 Foraminate
52.2.2 Tylate
52.2.3 Cassidate
52.3 On tooth apex
52.3.1 Spinose
52.3.2 Mucronate
52.3.3 Setaceous
52.3.4 Papillate
52.3.5 Spherulate
52.4 Nonspecific

T he eighteen examples in this appendix
are keyed to the numeric codes for each
character state described in the text.
The easiest way to score leaves or review the
scores that we assigned to these examples is to
is typed into the “score” column, the descrip-
tion field is populated automatically. (Note:
The shaded boxes are skipped and the actual
values for these character states are typed into
the description field.) Once the worksheet is
photocopy Appendix A and use it as a guide completed, the user can insert a verbal descrip-
to all of the possible character states. These tion of the leaf in the lower right-hand corner
numeric codes can also be used to quickly and or upload the data into a database. The blank
fully describe a leaf’s characteristics in a com- template on the facing page can be used to
puter database. quickly capture the numeric codes. Two boxes
are provided for a character state when a range
The examples are scored in a Microsoft® of choices is needed. Because the extant leaf
Excel® 2007 worksheet, shown on the facing images in Appendix B do not illustrate the leaf
page. Worksheets can be downloaded from attachment and organization characters, the
http://www.paleobotanyproject.org/. When first six characters were scored by reviewing
the number of the appropriate character state herbarium sheets.

Worksheets can be downloaded from http://www.paleobotanyproject.org/.

To improve data entry, we use the
following generic codes:

0 = Absent – The character is not present
in this leaf. For example, Tilia mandshurica
(Appendix B, example 1) does not have inter-
secondary veins, so this character is absent.

88 = Not visible – This character is not pre-
served and so cannot be scored.

99 = Not applicable (n/a) – The character
does not apply to this leaf. For example, tooth
type would score as n/a for a leaf that has a
smooth margin.

Excel Leaf Scoring Template

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1˚ Primary Vein Framework
Leaf Arrangement Naked Basal Veins
Leaf Organization Number of Basal Veins
Leaflet Arrangement Agrophic Veins
Leaflet Attachment 2˚ Major 2o Vein Framework
Petiole Features Interior Secondaries
Minor Secondary Course
Features of the Blade Perimarginal Veins
Position of Blade Attachment Major Secondary Spacing
Laminar Size Variation of Secondary Angle
Laminar L:W Ratio Major Secondary Attachment
Laminar Shape Inter-2˚ Proximal Course
Medial Symmetry Length
Base Symmetry Distal Course
Base Symmetry Vein Frequency
Lobation 3˚ Intercostal 3º Vein Fabric
Margin Type Angle of Percurrent Tertiaries
Special Margin Features Vein Angle Variability
Apex Angle Epimedial Tertiaries
Apex Shape Admedial Course
Base Angle Exmedial Course
Base Shape Exterior Tertiary Course
Base Shape 4˚ Quaternary Vein Fabric
Terminal Apex Features 5˚ Quinternary Vein Fabric
Surface Texture Areolation
Surficial Glands FEV branching
FEV termination
Marginal Ultimate Venation

III. Teeth Score Description Text Description:
Tooth Spacing
Number of Orders of Teeth
Teeth / cm
Sinus Shape
Tooth Shapes
Tooth Shapes
Tooth Shapes
Tooth Shapes
Principal Vein
Principal Vein Termination
Course of Accessory Vein
Features of the Tooth Apex

Example 1. Malvaceae - Tilia baccata var. mandshurica

Malvaceae - Tilia mandshurica

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.2.1 .1 basal actinodromous
Leaf Arrangement 2.1 alternate Naked Basal Veins 0 absent
Leaf Organization 3.1 simple Number of Basal Veins 8
Leaflet Arrangement 99 n/a Agrophic Veins 26.2 compound
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.1.2 semicraspedodromous
Petiole Features 88 not visible Interior Secondaries 0 absent
Minor Secondary Course 29.1 craspedodromous
Features of the Blade Perimarginal Veins 30 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.4 gradually increasing proximally
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.2:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 0 absent
Base Symmetry 12.2.1 extension asymmetry Distal Course 0 absent
Base Symmetry 12.2.1 extension asymmetry Vein Frequency 0 absent
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.1.2 convex opposite percurrent
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 35.1.2.2 obtuse to midvein
Special Margin Features 0 absent Vein Angle Variability 36.3.1 basally concentric
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.6 acute
Base Angle 18.2 obtuse Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.4 variable
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 39.1.2 alternate percurrent
Terminal Apex Features 88 not visible 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.3 good development
Surficial Glands 0 absent FEV branching 88 not visible
FEV termination 88 not visible
Marginal Ultimate Venation 88 not visible

III. Teeth Score Description Text Description:
Tooth Spacing 44.1 regular Leaf attachment petiolate. Marginal blade attachment. Laminar size
Number of Orders of Teeth 45.1 one mesophyll with L:W ratio of 1.2:1. Laminar shape elliptic to ovate,
symmetrical with basal extension asymmetry. Margin unlobed with ser-
Teeth / cm 2 rate teeth. Apex angle acute with acuminate shape. Base angle obtuse
Sinus Shape 47.2 rounded with cordate shape. Primary veins basal actinodromous with eight basal
veins. Compound agrophic veins present. Major secondary framework
Tooth Shapes cc/st semicraspedodromous, minor secondaries craspedodromous, major
Tooth Shapes cc/cc secondary spacing gradually increasing proximally with uniform angle
and excurrent attachment. Intersecondaries absent. Intercostal tertiary
Tooth Shapes cc/fl fabric opposite percurrent with convex course, obtuse angle to midvein,
Tooth Shapes with basally concentric tertiaries. Epimedial tertiaries opposite percur-
rent with acute admedial course, and exmedial course parallel to inter-
Principal Vein 49.1 present costal tertiary. Exterior tertiary course variable. Quaternary vein fabric
Principal Vein Termination 50.2.1 at apex of tooth alternate percurrent. Quinternary vein fabric regular reticulate. Areola-
tion shows good development but FEVs are not visible. Tooth spacing
Course of Accessory Vein 99 n/a regular, with a single order of teeth. Sinus shape rounded with tooth
Features of the Tooth Apex 52.1 none shapes: concave/straight, concave/concave, and concave/flexuous.
Principal vein terminates at apex of tooth. Accessory veins absent.

Example 2. Dilleniaceae - Davilla rugosa

Dilleniaceae - Davilla rugosa

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 0 absent
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.3.1 simple brochidodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.8:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.1 symmetrical Distal Course 99 n/a
Base Symmetry 12.1 symmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 sinuous opposite percurrent
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 35.1.2.2 obtuse to midvein
Special Margin Features not visible Vein Angle Variability 36.5 increasing exmedially
Apex Angle 16.2 obtuse Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.2.1 rounded Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.2 obtuse Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.1.3.1 rounded Exterior Tertiary Course 38.2 looped
Base Shape 19.1.3.1 rounded 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.1.2 irregular reticulate
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.3 looped

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal. Laminar size notophyll, L:W ratio 1.8:1,
Number of Orders of Teeth 99 n/a laminar shape elliptic with medial symmetry and basal symmetry. Mar-
gin is entire with obtuse apex angle, rounded apex, obtuse base angle,
Teeth / cm 99 n/a and rounded base shape. Primary venation pinnate with no naked basal
Sinus Shape 99 n/a veins, one basal vein, and no agrophic veins. Major secondaries simple
brochidodromous with regular spacing, uniform angle and excurrent
Tooth Shapes 99 n/a attachment to midvein. Interior secondaries absent, minor secondar-
Tooth Shapes 99 n/a ies absent, intersecondaries absent, fimbrial vein present. Intercostal
tertiary veins mixed percurrent with obtuse angle that increases exme-
Tooth Shapes 99 n/a dially. Epimedial tertiaries opposite percurrent with proximal course
Tooth Shapes 99 n/a perpendicular to the midvein and distal course parallel to the intercos-
tal tertiaries. Exterior tertiaries looped. Quaternary vein fabric irregu-
Principal Vein 99 n/a lar reticulate. Quinternary vein fabric irregular reticulate. Areolation
Principal Vein Termination 99 n/a shows moderate development. Freely ending veinlets have two or more
dendritic branches, and marginal ultimate venation is looped.
Course of Accessory Vein 99 n/a
Features of the Tooth Apex 99 n/a

Example 3. Dipterocarpaceae - Stemonoporus nitidus

Dipterocarpaceae - Stemonoporus nitidus

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23..1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 99 n/a 2˚ Major 2o Vein Framework 27.3.1 simple brochidodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 3:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.1 elliptic Inter- 2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.2.2 >50%
Base Symmetry 12.1 symmetrical Distal Course 34.3.4 basiflexed
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.2 ~1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 mixed percurrent
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features not visible Vein Angle Variability 36.2 consistent
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.2 obtuse Exmedial Course 37.2.2.2 basiflexed
Base Shape 19.1.3.1 rounded Exterior Tertiary Course 38.2 looped
Base Shape 19.1.3.1 rounded 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.2 freely ramifying
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Leaf attachment petiolate. Blade attachment marginal, laminar size no-
Number of Orders of Teeth 99 n/a tophyll, L:W ratio 3:1, laminar shape elliptic with medial symmetry and
basal symmetry. Margin entire with acute apex angle, acuminate apex,
Teeth / cm 99 n/a obtuse base angle, and rounded base shape. Primary venation pinnate
Sinus Shape 99 n/a with no naked basal veins, one basal vein, and no agrophic veins. Major
secondaries simple brochidodromous with regular spacing, uniform angle,
Tooth Shapes 99 n/a and excurrent attachment to midvein. Interior secondaries absent, minor
Tooth Shapes secondaries absent, and fimbrial vein present. Intersecondaries span more
99 n/a
than 50% of the length of the subjacent secondary, occur at slightly more
Tooth Shapes 99 n/a than one per intercostal area, proximal course is parallel to major second-
aries, and distal course is basiflexed and parallel to intercostal tertiaries.
Tooth Shapes 99 n/a
Intercostal tertiary veins mixed percurrent with obtuse angle that remains
Principal Vein 99 n/a consistently. Epimedial tertiaries opposite percurrent with proximal course
perpendicular to the midvein and distal course basiflexed. Exterior tertia-
Principal Vein Termination 99 n/a ries looped. Quaternary vein fabric irregular reticulate. Quinternary vein
Course of Accessory Vein 99 n/a fabric freely ramifying. Areolation shows moderate development. Freely
ending veinlets mostly two branched and marginal ultimate venation joins
Features of the Tooth Apex 99 n/a fimbrial vein.

Example 4. Fabaceae - Bauhinia madagascariensis

Fabaceae - Bauhinia madagascariensis

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.2.1.1 basal actinodromous
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 5
Leaflet Arrangement 99 n/a Agrophic Veins 26.2.1 simple
Leaflet Attachment 99 n/a 2˚ Major 2o Vein Framework 27.3.2 festooned brochidodromous
Petiole Features 6.1.2 pulvinate Interior Secondaries 28.2 present
Minor Secondary Course 29.2 simple brochidodromous
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.5 abruptly increasing proximally
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 0.85:1 Major Secondary Attachment 33.4 deflected
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.2 asymmetrical Length 34.2.2 >50% of subjacent secondary
Base Symmetry 12.1.1 basal width asymmetrical Distal Course 34.3.1 reticulating
Base Symmetry 12.1.1 basal width asymmetrical Vein Frequency 34.4.1 <1 per intercostal area
Lobation 13.2.4 bilobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 mixed percurrent
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 88 not visible Vein Angle Variability 36.5 increasing proximally
Apex Angle 16.3 reflex Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.5 lobed Admedial Course 37.2.1.6 acute to midvein
Base Angle 18.3 reflex Exmedial Course 37.2.2.2 basiflexed
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.2 looped
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 39.2 regular reticulate
Terminal Apex Features 20.2 spinose 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.3 good development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly unbranched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Leaf attachment petiolate. Blade attachment marginal, laminar
Number of Orders of Teeth 99 n/a size mesophyll, L:W ratio 0.85:1, laminar shape ovate with medi-
al asymmetry and basal width asymmetry. Margin is bilobed and
Teeth / cm 99 n/a untoothed with reflex apex angle, lobed apex shape, spinose apex,
Sinus Shape 99 n/a reflex base angle, and cordate base shape. Primary venation bas-
al actinodromous with no naked basal veins, five basal veins, and
Tooth Shapes 99 n/a simple agrophic veins. Major secondaries simple brochidodromous
Tooth Shapes 99 n/a with spacing that abruptly increases proximally, uniform angle, and
decurrent attachment to midvein. Interior secondaries present, mi-
Tooth Shapes 99 n/a nor secondaries simple brochidodromous, and fimbrial vein present.
Tooth Shapes 99 n/a Intersecondaries span more than 50% of the length of the subja-
cent secondary, occur at less than one per intercostal area, proximal
Principal Vein 99 n/a course is parallel to major secondaries, and distal course is reticu-
Principal Vein Termination 99 n/a lating or basiflexed. Intercostal tertiary veins mixed percurrent to
irregular reticulate. Epimedial tertiaries opposite percurrent with
Course of Accessory Vein 99 n/a proximal course acute to the midvein and distal course basiflexed.
Features of the Tooth Apex 99 n/a Exterior tertiaries looped. Quaternary vein fabric regular reticulate.
Quinternary vein fabric irregular reticulate. Areolation shows good
development. Freely ending veinlets mostly unbranched, and mar-
ginal ultimate venation is absent.

Example 5. Trochodendraceae - Tetracentron sinense

Trochodendraceae - Tetracentron sinense

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.2.1.1 basal actinodromous
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 7
Leaflet Arrangement 99 n/a Agrophic Veins 26.2.2 compound
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.1.3 festooned semicraspedodro-
mous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 29.3 semicraspedodromous
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.4 gradually increasing proximally
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.3:1 Major Secondary Attachment 33.4 deflected
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.2.1 basal width asymmetrical Distal Course 99 n/a
Base Symmetry 12.2.1 basal width asymmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.2 regular reticulate
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 0 absent Vein Angle Variability 99 n/a
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.1 straight Admedial Course 99 n/a
Base Angle 18.2 obtuse Exmedial Course 99 n/a
Base Angle 18.2 obtuse Exterior Tertiary Course 38.4 variable
Base Shape 19.1.3.2 truncate 4˚ Quaternary Vein Fabric 39.2.1 regular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 0 absent
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic

FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.3 looped

III. Teeth Score Description Text Description:
Tooth Spacing 44.1 regular Leaf attachment petiolate. Blade attachment marginal, laminar size
Number of Orders of Teeth 45.1 one notophyll, L:W ratio 1.3:1, laminar shape ovate with medial symme-
try and basal width asymmetry. Margin is unlobed and serrate with
Teeth / cm 4 acute apex angle, straight apex shape, obtuse base angle, and truncate
Sinus Shape 47.1 angular base shape. Primary venation is basal actinodromous with no naked
basal veins, seven basal veins, and compound agrophic veins. Major
Tooth Shapes cv/cv secondaries festooned semicraspedodromous with spacing that gradu-
Tooth Shapes ally increases proximally, uniform angle, and deflected attachment to
midvein. Interior secondaries absent, minor secondaries semicraspe-
Tooth Shapes dodromous, and perimarginal veins absent. Intersecondaries absent.
Tooth Shapes Intercostal tertiary veins regular reticulate. Epimedial tertiaries re-
ticulate. Exterior tertiaries variable. Quaternary vein fabric regular
Principal Vein 49.1 present reticulate. Areolation moderately developed. Freely ending veinlets
Principal Vein Termination 50.2.1 at apex of tooth mostly two or more branched with simple termination. Marginal ulti-
mate venation looped. Tooth spacing regular with one order of teeth
Course of Accessory Vein 51.1 convex and 4 teeth/cm. Sinus shape angular and tooth shape convex/convex.
Features of the Tooth Apex 52.2.3 cassidate Principal vein present and terminating at tooth apex. Accessory vein
course convex. Tooth apex cassidate.

Example 6. Anacardiaceae - Buchanania arborescens

Anacardiaceae - Buchanania arborescens

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.2.3 cladodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 n/a
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.3 decreasing proximally
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.3 smoothly increasing proximally
Laminar L:W Ratio 3:1 Major Secondary Attachment 33.1 decurrent
Laminar Shape 10.2 obovate Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.1 symmetrical Distal Course 99 n/a
Base Symmetry 12.1 symmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.3 composite admedial
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 88 not visible Vein Angle Variability 99 n/a
Apex Angle 18.1 acute Epimedial Tertiaries 37.1.2 ramified
Apex Shape 17.3 acuminate Admedial Course 37.2.1.1 parallel to subjacent secondary
Base Angle 18.1 acute Exmedial Course ramified
Base Shape 19.1.2 concave Exterior Tertiary Course 38.4 variable
Base Shape 19.1.2 concave 4˚ Quaternary Vein Fabric 39.3 freely ramifying
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 0 absent
Surface Texture 88 not visible Areolation 41.2.1 poorly developed
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size mesophyll, L:W ratio 3:1,
Number of Orders of Teeth 99 n/a laminar shape obovate with medial symmetry and basal symmetry.
Margin entire with acute apex angle, acuminate apex, acute base
Teeth / cm 99 n/a angle, and concave base shape. Primary venation is pinnate with no
Sinus Shape 99 n/a naked basal veins, one basal vein, and no agrophic veins. Major sec-
ondaries cladodromous with spacing that decreases proximally, angle
Tooth Shapes 99 n/a that smoothly increases proximally, and decurrent attachment to mid-
Tooth Shapes 99 n/a vein. Interior secondaries absent, minor secondaries absent, and fim-
brial vein present. Intersecondaries absent. Intercostal tertiary veins
Tooth Shapes 99 n/a composite admedial. Epimedial tertiaries ramified with admedial
Tooth Shapes 99 n/a course parallel to subjacent secondary and exmedial course ramified.
Exterior tertiaries variable. Quaternary vein fabric freely ramifying.
Principal Vein 99 n/a Areolation poorly developed. Freely ending veinlets have two or more
Principal Vein Termination 99 n/a dendritic branches.
Course of Accessory Vein 99 n/a
Features of the Tooth Apex 99 n/a

Example 7. Elaeocarpaceae - Aristotelia racemosa

Elaeocarpaceae - Aristotelia racemosa

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.3 opposite Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 5
Leaflet Arrangement 99 n/a Agrophic Veins 26.2.2 compound
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.1.3 festooned semicraspedodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 29.3 semicraspedodromous
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.4 gradually increasing proximally
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.4:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.1 symmetrical Distal Course 99 n/a
Base Symmetry 12.1 symmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 mixed percurrent
Margin Type 14.2.1 dentate Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 88 not visible Vein Angle Variability 36.5 increasing proximally
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.3 mixed percurrent
Apex Shape 17.1 straight Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.2 obtuse Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.1.3.2 truncate Exterior Tertiary Course 38.3 terminating at the margin
Base Shape 19.1.3.2 truncate 4˚ Quaternary Vein Fabric 39.2.1 regular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.3 good development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly unbranched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.4 looped

III. Teeth Score Description Text Description:
Tooth Spacing 44.1 regular Leaf attachment petiolate. Blade attachment marginal, laminar size
Number of Orders of Teeth 45.2 two mesophyll, L:W ratio 1.4:1, laminar shape elliptic to ovate with medial
symmetry and basal symmetry. Margin is unlobed and dentate with
Teeth / cm 2 acute apex angle, straight apex shape, obtuse base angle, and truncate
Sinus Shape 47.2 rounded base shape. Primary venation is pinnate with no naked basal veins,
five basal veins, and compound agrophic veins. Major secondaries fes-
Tooth Shapes st/st tooned semicraspedodromous with spacing that gradually increases
Tooth Shapes cc/cc proximally, with uniform angle and excurrent attachment to midvein.
Interior secondaries absent, minor secondaries semicraspedodromous,
Tooth Shapes and perimarginal veins absent. Intersecondaries absent. Intercostal
Tooth Shapes tertiary veins mixed percurrent with obtuse angle to midvein and
proximally increasing vein angle. Epimedial tertiaries mixed percur-
Principal Vein 49.1 present rent with proximal course perpendicular to the midvein and distal
Principal Vein Termination 50.2.1 at apex of tooth course parallel to intercostal tertiary. Exterior tertiaries terminate at
the margin. Quaternary and quinternary vein fabric regular reticu-
Course of Accessory Vein 51.2 straight or concave late. Areolation shows good development. Tooth spacing regular with
Features of the Tooth Apex 52.1 simple two orders of teeth and 2 teeth/cm. Sinus shape rounded and tooth
shape straight/straight to concave/concave. Principal vein present and
terminating at tooth apex. Accessory vein course straight or concave.
Tooth apex simple.

Example 8. Malvaceae - Bombacopsis rupicola

Malvaceae - Bombacopsis rupicola

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.2.1 palmately compound Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 5.1 petiolulate 2˚ Major 2 Vein Framework
o
27.3.2 festooned brochidodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 30.1 marginal secondary
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.3 decreasing proximally
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.2 inconsistent
Laminar L:W Ratio 2.2:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.2 obovate Inter-2˚ Proximal Course 34.1.3 perpendicular to midvein
Medial Symmetry 11.1 symmetrical Length 34.2.2 >50% of subjacent secondary
Base Symmetry 12.1 symmetrical Distal Course 34.3.1 reticulating
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.2 ~1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.1 irregular reticulate
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 88 not visible Vein Angle Variability 99 n/a
Apex Angle 16.3 reflex Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.2 convex Admedial Course 99 n/a
Base Angle 18.1 acute Exmedial Course 99 n/a
Base Shape 19.1.1 straight Exterior Tertiary Course 38.2 looped
Base Shape 19.1.1 straight 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 20.3 retuse 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.3 good development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly 1 branch
FEV termination 42.2.1 simple
Marginal Ultimate Venation 0 n/a

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size notophyll to mesophyll, L:
Number of Orders of Teeth 99 n/a W ratio 2.2:1, laminar shape obovate to elliptic with medial symme-
try and basal symmetry. Margin entire with reflex apex angle, convex
Teeth / cm 99 n/a apex shape and retuse apex, acute base angle, and straight base shape.
Sinus Shape 99 n/a Primary venation pinnate with no naked basal veins, one basal vein,
and no agrophic veins. Major secondaries festooned brochidodromous
Tooth Shapes 99 n/a with spacing that decreases proximally, inconsistent secondary angle,
Tooth Shapes 99 n/a and excurrent attachment to midvein. Minor secondaries absent and
marginal secondary present. Intersecondaries span more than 50% of
Tooth Shapes 99 n/a the length of the subjacent secondary, occur at roughly one per inter-
Tooth Shapes 99 n/a costal area, proximal course is perpendicular to midvein and distal
course is reticulating. Intercostal tertiary veins irregular reticulate.
Principal Vein 99 n/a Epimedial tertiaries reticulate. Exterior tertiaries looped. Quaternary
Principal Vein Termination 99 n/a vein fabric irregular reticulate. Quinternary vein fabric regular re-
ticulate. Areolation shows good development.
Course of Accessory Vein 99 n/a
Features of the Tooth Apex 99 n/a

Example 9. Gesneriaceae - Rhynchoglossum azureum

Gesneriaceae - Rhynchoglossum azureum

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.2.1 eucamptodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.3 decreasing proximally
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.3 smoothly increasing proximally
Laminar L:W Ratio 1.85:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.2 asymmetrical Length 99 n/a
Base Symmetry 12.2.1 basal width asymmetrical Distal Course 99 n/a
Base Symmetry 12.2.3 basal insertion asymmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.1.4 opposite percurrent
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features n/a Vein Angle Variability 36.4 decreasing exmedially
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.6 acute to midvein
Base Angle 18.2 obtuse Exmedial Course 37.2.2.2 basiflexed
Base Shape 19.1.2 concave Exterior Tertiary Course 38.2 looped
Base Shape 19.1.3 convex 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 n/a 5˚ Quinternary Vein Fabric 40.1.2 irregular reticulate
Surface Texture 21.5 pubescent Areolation 41.2.1 poor development
Surficial Glands 22.2 marginal FEV branching 42.1.3 mostly 1 branch
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.4 looped

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size notophyll, L:W ratio
Number of Orders of Teeth 99 n/a 1.85:1, laminar shape elliptic with medial asymmetry and basal
width and basal insertion asymmetry. Margin is entire with acute
Teeth / cm 99 n/a apex angle, acuminate apex shape, obtuse base angle, and concave
Sinus Shape 99 n/a to rounded base shape. Surface texture is pubescent with surface
glands. Primary venation is pinnate with one basal vein, and no
Tooth Shapes 99 n/a agrophic veins. Major secondaries eucamptodromous with spac-
Tooth Shapes 99 n/a ing that decreases exmedially, mostly attaching to the midvein
excurrently but with some apical deflection. Minor secondaries
Tooth Shapes 99 n/a and intersecondaries absent. Intercostal 3º veins form chevrons
Tooth Shapes 99 n/a with vein angles that decrease exmedially. Epimedial tertiaries
opposite percurrent with proximal course acute to midvein and
Principal Vein 99 n/a distal course basiflexed. Exterior tertiaries looped. Quaternary
Principal Vein Termination 99 n/a vein fabric irregular reticulate. Quinternary vein fabric irregular
reticulate. Areolation shows poor development. FEV’s are mostly
Course of Accessory Vein 99 n/a one branched with simple terminals. Marginal ulitmate venation
Features of the Tooth Apex 99 n/a looped.

Example 10. Nothofagaceae - Nothofagus procera

Nothofagaceae - Nothofagus procera

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 3
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 99 n/a 2˚ Major 2 Vein Framework
o
27.1.2 semicraspedodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 2.4:1 Major Secondary Attachment 33.2 basally decurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.1 symmetrical Distal Course 99 n/a
Base Symmetry 12.1 symmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.2 alternate percurrent
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 15.1.2 sinuous Vein Angle Variability 36.2 consistent
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.2 convex Admedial Course 99 n/a
Base Angle 18.2 obtuse Exmedial Course 99 n/a
Base Shape 19.1.3 convex Exterior Tertiary Course 38.3 terminating at the margin
Base Shape 19.1.3 convex 4˚ Quaternary Vein Fabric 39.2.1 regular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.3 good development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly unbranched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.4 looped

III. Teeth Score Description Text Description:
Tooth Spacing 44.2 irregular Blade attachment marginal, laminar size microphyll to notophyll,
Number of Orders of Teeth 45.2 two L:W ratio 2.4:1, laminar shape elliptic with medial symmetry
and basal symmetry. Margin unlobed, sinuous and serrate, with
Teeth / cm 6 acute apex angle, convex apex shape, obtuse base angle, and con-
Sinus Shape 47.2 rounded vex base shape. Primary venation pinnate with no naked basal
veins, three basal veins, and no agrophic veins. Major secondaries
Tooth Shapes st/st semicraspedodromous with regular spacing, uniform angle, and
Tooth Shapes cv/cv basally decurrent attachment to midvein. Interior secondaries
absent, minor secondaries absent, and perimarginal vein absent.
Tooth Shapes
Intersecondaries absent. Intercostal tertiary veins alternate per-
Tooth Shapes current with obtuse angle to midvein and consistent vein angle.
Epimedial tertiaries reticulate. Exterior tertiaries terminate at the
Principal Vein 49.1 present
margin. Quaternary and quinternary vein fabric regular reticu-
Principal Vein Termination 50.2.1 at apex of tooth late. Areolation shows good development. Tooth spacing irregular
with two orders of teeth and 6 teeth/cm. Sinus shape rounded and
Course of Accessory Vein 51.2 straight
tooth shape straight/straight to convex/convex. Principal vein ter-
Features of the Tooth Apex 52.1 simple minating at tooth apex.

Example 11. Sapindaceae - Acer franchetii

Sapindaceae - Acer franchetii

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.2.1.1 basal actinodromous
Leaf Arrangement 2.3 opposite Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 6
Leaflet Arrangement 99 n/a Agrophic Veins 26.2.2 compound
Leaflet Attachment 99 n/a 2˚ Major 2o Vein Framework 27.1.1 craspedodromous
Petiole Features 88 not visible Interior Secondaries 28.2 present
Minor Secondary Course 29.1 craspedodromous
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.5 abruptly increasing proximally
Laminar Size 8.6 macrophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.1:1 Major Secondary Attachment 33.4 deflected
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.3.2 parallel to major secondary
Base Symmetry 12.1 symmetrical Distal Course 34.2.1 <50% of subjacent secondary
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.3 >1 per intercostal area
Lobation 13.2.1 palmately lobed 3˚ Intercostal 3º Vein Fabric 35.1.1.2 alternate percurrent
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 0 absent Vein Angle Variability 36.2 consistent
Apex Angle 16.2 obtuse Epimedial Tertiaries 37.1.2 ramified
Apex Shape 17.2 convex Admedial Course 37.2.1.6 acute to midvein
Base Angle 18.3 reflex Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.2 looped
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 39.2.1 regular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.1.1 regular reticulate
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly unbranched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.3 spiked

III. Teeth Score Description Text Description:
Tooth Spacing 44.2 irregular Blade attachment marginal, laminar size microphyll to macro-
Number of Orders of Teeth 45.1 one phyll, laminar L:W ratio 1.1:1, laminar shape elliptic, blade me-
dially symmetrical, base symmetrical, palmately lobed, margin
Teeth / cm 3 serrate. Apex angle obtuse, apex shape convex, base angle reflex,
Sinus Shape 47.1 angular base shape cordate. Primary vein basal actinodromous, naked
basal veins absent, six basal veins, agrophic veins compound, ma-
Tooth Shapes st/st jor 2 ̊ veins craspedodromous, minor secondary course craspedo-
Tooth Shapes cv/cv dromous, interior secondaries present, major secondary spacing
abruptly increasing proximally, secondary angle uniform, major
Tooth Shapes secondary attachment deflected. Intersecondary length <50%
Tooth Shapes of subjacent secondary, distal course parallel to subjacent major
secondary, vein frequency >1 per intercostal area, intercostal ter-
Principal Vein 49.1 present tiary vein fabric opposite percurrent. Epimedial tertiaries rami-
Principal Vein Termination 50.2.1 at apex of tooth fied, admedial course acute to midvein, exmedial course parallel
to intercostal tertiary. Exterior tertiary course looped and occa-
Course of Accessory Vein 51.3 running from sinus sionally terminating at the margin. Quaternary vein fabric regu-
Features of the Tooth Apex 52.1 simple lar reticulate; quinternary vein fabric regular reticulate; areola-
tion development moderate. Tooth spacing irregular, one order
of teeth, 3 teeth/cm, sinus shape angular, tooth shapes st/st and
cv/cv. Principal vein terminates at apex of tooth, accessory veins
run from sinus.
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142 Manual of Leaf Architecture

Example 12. Malpighiaceae - Tetrapterys macrocarpa

Malpighiaceae - Tetrapterys macrocarpa

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.3 opposite Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 3
Leaflet Arrangement 0 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 0 n/a 2˚ Major 2o Vein Framework 27.3.2 festooned brochidodromous
Petiole Features 6.2.1 glands petiolar Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.3 decreasing proximally
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.4 smoothly decreasing
proximally
Laminar L:W Ratio 1.5:1 Major Secondary Attachment 33.1 decurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.2.1 <50% of subjacent secondary
Base Symmetry 12.1 symmetrical Distal Course 34.3.1 reticulating
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.1 <1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 mixed percurrent
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 0 absent Vein Angle Variability 36.1 inconsistent
Apex Angle 16.2 obtuse Epimedial Tertiaries 37.1.1.1 mixed percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.3 reflex Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.2 looped
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.2.2 irregular reticulate
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic
FEV termination 42.2.3 highly branched sclereids
Marginal Ultimate Venation 43.4 looped

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size notophyll, L:W ratio
Number of Orders of Teeth 99 n/a 1.5:1, laminar shape elliptic with medial symmetry and basal
symmetry. Margin entire with obtuse apex angle, acuminate apex
Teeth / cm 99 n/a shape, reflex base angle, and cordate base shape. Primary vena-
Sinus Shape 99 n/a tion pinnate with no naked basal veins, three basal veins, and no
agrophic veins. Major secondaries festooned brochidodromous
Tooth Shapes 99 n/a with spacing that decreases proximally, uniform secondary angle,
Tooth Shapes 99 n/a and decurrent attachment to midvein. Minor secondaries absent,
interior secondaries absent. Intersecondaries span less than 50%
Tooth Shapes 99 n/a of the length of the subjacent secondary, occur at less than one per
Tooth Shapes 99 n/a intercostal area, proximal course is parallel to major secondary
and distal course is reticulating. Intercostal tertiary veins mixed
Principal Vein 99 n/a percurrent with obtuse angle to midvein and inconsistent vein an-
Principal Vein Termination 99 n/a gle variability. Epimedial tertiaries mixed percurrent with proxi-
mal course perpendicular to the midvein and distal course paral-
Course of Accessory Vein 99 n/a lel to intercostal tertiaries. Exterior tertiaries looped. Quaternary
Features of the Tooth Apex 99 n/a vein fabric irregular reticulate. Quinternary vein fabric irregular
reticulate. Areolation shows moderate development. Freely ending
veinlets are two or more branched with highly branched sclereids.
Marginal ultimate venation forms incomplete loops.

Example 13. Cunoniaceae - Eucryphia glutinosa

Cunoniaceae - Eucryphia glutinosa

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 n/a 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.3 opposite Naked Basal Veins 24.1 absent
Leaf Organization 3.2.2.1 pinnately compounded Number of Basal Veins 1
Leaflet Arrangement 4.3.2 opposite-even Agrophic Veins 26.1 absent
Leaflet Attachment 5.1 petiolulate 2˚ Major 2 Vein Framework
o
27.1.2 semicraspedodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.2 irregular
smoothly decreasing
Laminar Size 8.4 notophyll Variation of Secondary Angle 32.4 proximally
Laminar L:W Ratio 1.8:1 Major Secondary Attachment 33.1 decurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.2.2 >50% of subjacent secondary
Base Symmetry 12.1 symmetrical Distal Course 34.3.2 parallel to subjacent major 2˚
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.1 <1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.1 irregular reticulate
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 0 absent Vein Angle Variability 99 n/a
Apex Angle 16.2 obtuse Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.2 convex Admedial Course 99 n/a
Base Angle 18.1 acute Exmedial Course 99 n/a
Base Shape 19.1.1 straight Exterior Tertiary Course 38.2 looped
Base Shape 19.1.1 straight 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.2.2 irregular reticulate
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.2 mostly unbranched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.2 incomplete loops

III. Teeth Score Description Text Description:
Tooth Spacing 44.1 regular
Blade attachment marginal, laminar size notophyll, laminar
Number of Orders of Teeth 45.1 one L:W ratio 1.8:1, laminar shape elliptic, blade medially symmetri-
cal, base medially symmetrical, margin unlobed with serrate teeth.
Teeth / cm 3
Apex angle obtuse, apex shape convex, base angle acute, base
Sinus Shape 47.1 angular shape straight. Primary venation pinnate with one basal vein and
no agrophic veins. Secondary veins semicraspedodromous with
Tooth Shapes cv/cv no interior secondaries, minor secondaries or perimarginal veins.
Tooth Shapes st/cv Major secondary spacing smoothly decreasing proximally, major
secondary attachment decurrent. Intersecondary proximal course
Tooth Shapes parallel to major secondaries, length >50% of subjacent secondary,
Tooth Shapes distal course parallel to subjacent secondary and frequency less
than one per intercostal area. Tertiary vein fabric irregular reticu-
Principal Vein 49.1 present late with reticulate epimedial tertiaries and looped exterior tertia-
Principal Vein Termination 50.1 submarginal ries. Quaternary vein fabric irregular reticulate. Quinternary vein
fabric irregular reticulate. Areolation moderately developed and
Course of Accessory Vein 51.1.1 looped FEVs mostly branched with simple terminals. Marginal ultimate
venation forms incomplete loops. Tooth spacing is regular with
Features of the Tooth Apex 52.1 simple
one order of teeth and three teeth per cm. Sinus shape is angular,
tooth shapes are convex/convex to straight/convex. Principal vein
is present with submarginal termination, looped accessory veins
and
Licensed under Creative Commons license CC BY-NC 4.0. To reprint a simple
this work intooth apex.
whole or in part for commercial purposes,
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146 Manual of Leaf Architecture

Example 14. Chrysobalanaceae - Licania michauxii

Chrysobalanaceae - Licania michauxii

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 1
Leaflet Arrangement 99 n/a Agrophic Veins 26.1 absent
Leaflet Attachment 99 n/a 2˚ Major 2o Vein Framework 27.3.1 simple brochidodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 30.3 fimbrial vein
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.2 irregular
Laminar Size 8.4 microphyll Variation of Secondary Angle 32.2 inconsistent
Laminar L:W Ratio 5:1 Major Secondary Attachment 33.1 decurrent
Laminar Shape 10.2 obovate Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.1 symmetrical Length 99 n/a
Base Symmetry 12.1 symmetrical Distal Course 99 n/a
Base Symmetry 12.1 symmetrical Vein Frequency 99 n/a
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.1 irregular reticulate
Margin Type 14.1 untoothed Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 15.1.1 erose Vein Angle Variability 99 n/a
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.2 convex Admedial Course 99 n/a
Base Angle 18.1 acute Exmedial Course 99 n/a
Base Shape 19.1.1 straight Exterior Tertiary Course 38.3 terminates at the margin
Base Shape decurrent 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.3 freely ramifying
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.3 mostly 1 branched
FEV termination 42.2.1 simple
Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size microphyll to notophyll,
Number of Orders of Teeth 99 n/a L:W ratio 5:1, laminar shape obovate with medial symmetry and
basal symmetry. Margin is entire and erose with acute apex angle,
Teeth / cm 99 n/a convex apex shape, acute base angle, and straight to decurrent base
Sinus Shape 99 n/a shape. Primary venation is pinnate with no naked basal veins, one
basal vein, and no agrophic veins. Major secondaries simple bro-
Tooth Shapes 99 n/a chidodromous with irregular spacing, inconsistent secondary angle,
Tooth Shapes 99 n/a and decurrent attachment to midvein. Minor secondaries, interior
secondaries, and intersecondaries absent. Fimbrial vein present.
Tooth Shapes 99 n/a
Intercostal tertiary veins irregular reticulate. Epimedial tertiaries
Tooth Shapes 99 n/a reticulate. Exterior tertiaries terminate at the margin. Quaternary
vein fabric irregular reticulate. Quinternary vein fabric freely rami-
Principal Vein 99 n/a
fying. Areolation shows moderate development. Freely ending vein-
Principal Vein Termination 99 n/a lets are mostly one-branched with simple termination. Marginal
ultimate venation is absent.
Course of Accessory Vein 99 n/a
Features of the Tooth Apex 99 n/a

Example 15. Moraceae - Morus microphylla

Moraceae - Morus microphylla

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.2.1.1 basal actinodromous
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.1 simple Number of Basal Veins 5
Leaflet Arrangement 99 n/a Agrophic Veins 26.2.1 simple
Leaflet Attachment 99 n/a 2˚ Major 2o Vein Framework 27.1.2 semicraspedodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 29.3 semicraspedodromous
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.2 irregular
smoothly decreasing
Laminar Size 8.3 microphyll Variation of Secondary Angle 32.4 proximally
Laminar L:W Ratio 1.3:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.2 asymmetrical Length 34.2.1 <50% of subjacent secondary
Base Symmetry 12.2.1 basal width asymmetrical Distal Course 34.3.1 reticulating
Base Symmetry 12.2.1 basal width asymmetrical Vein Frequency 34.4.1 <1 perintercoital
Lobation 13.2.1 palmately lobed 3˚ Intercostal 3º Vein Fabric 35.1.1.3 mixed percurrent
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 88 not visible Vein Angle Variability 36.4 decreasing exmedially
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.3 reflex Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.3 terminating at the margin
Base Shape 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.1.2 irregular reticulate
Surface Texture 21.5 pubescent Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 42.1.1 FEVs absent
FEV termination 99 n/a
Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 44.2 irregular Leaf attachment petiolate. Blade attachment marginal, laminar size
Number of Orders of Teeth 45.2 two microphyll to mesophyll, laminar L:W ratio 1.3:1, laminar shape
ovate, blade medially asymmetrical, basal width asymmetrical,
Teeth / cm 2 palmately lobed, margin serrate. Apex angle acute, apex shape
Sinus Shape 47.1 angular acuminate to convex, base angle reflex, base shape cordate. Sur-
face texture pubescent. Primary vein basal actinodromous, naked
Tooth Shapes cc/cc basal veins absent, five basal veins, simple agrophic veins, major
Tooth Shapes 2º veins semicraspedodromous, interior secondaries absent, minor
secondary course semicraspedodromous, major secondary spacing
Tooth Shapes
irregular, secondary angle smoothly decreasing proximally, major
Tooth Shapes secondary attachment excurrent. Intersecondary length <50% of
subjacent secondary, proximal course parallel to subjacent major
Principal Vein 49.1 present
secondary, distal course reticulating, vein frequency <1 per inter-
Principal Vein Termination 51.2 marginal costal area. Intercostal tertiary vein fabric mixed percurrent with
obtuse angle that decreases exmedially. Epimedial tertiaries oppo-
Course of Accessory Vein 51.1 convex
site percurrent, proximal course perpendicular to midvein, distal
Features of the Tooth Apex 53.1 none course parallel to intercostal tertiary. Exterior tertiary course ter-
minating at the margin. Quaternary and quintemary vein fabric
irregular reticulate, areolation development moderate. Tooth spac-
ing irregular, two orders of teeth, 2 teeth/cm, sinus shape rounded,
tooth shape concave/concave. Principal vein present with termina-
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this
tion workofintooth,
at apex wholeaccessory
or in part forconvex.
vein commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
150 Manual of Leaf Architecture

Example 16. Anacardiaceae - Comocladia dodonaea

Anacardiaceae - Comocladia dodonaea

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
Leaf Organization 3.2.2.1 once pinnately compound Number of Basal Veins 3
Leaflet Arrangement 4.3.1 opposite-odd Agrophic Veins 0 absent
Leaflet Attachment 5.1 petiolulate 2˚ Major 2o Vein Framework 27.1.1 craspedodromous
Petiole Features 88 not visible Interior Secondaries 0 absent
Minor Secondary Course 0 absent
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.3 microphyll Variation of Secondary Angle 32.3 smoothly increasing proximally
Laminar L:W Ratio 1:1 Major Secondary Attachment 33.1 decurrent
Laminar Shape 10.3 ovate Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.2.2 >50% of subjacent secondary
parallel to subjacent major
Base Symmetry 12.1 symmetrical Distal Course 34.3.2 secondary
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.2 ~1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.3.4 transverse freely ramified
Margin Type 14.2 toothed Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 15.1.2 sinuous Vein Angle Variability 36.4 decreasing exmedially
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.2 ramified
Apex Shape 17.3 acuminate Admedial Course 99 n/a
Base Angle 18.3 reflex Exmedial Course 99 n/a
Base Shape 19.2.1 cordate Exterior Tertiary Course 0 absent
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 0 absent
Terminal Apex Features 88 not visible 5˚ Quinternary Vein Fabric 0 absent
Surface Texture 88 not visible Areolation 41.2.1 poor development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic
FEV termination 42.2.2 tracheoid idioblasts
Marginal Ultimate Venation 43.2 incomplete loops

III. Teeth Score Description Text Description:
Tooth Spacing 44.1 regular Blade attachment marginal, laminar size microphyll, laminar L:W
Number of Orders of Teeth 45.1 one ratio 1:1, laminar shape ovate, blade medially symmetrical, base sym-
metrical, unlobed, margin toothed and sinuous. Apex angle acute,
Teeth / cm 1 apex shape acuminate, base angle reflex, base shape cordate. Primary
Sinus Shape 47.2 rounded vein pinnate, naked basal veins absent, three basal veins, no agrophic
veins. Major 2º veins craspedodromous, interior secondaries absent,
Tooth Shapes minor secondaries absent, major secondary spacing regular, secondary
Tooth Shapes angle smoothly increasing proximally, major secondary attachment de-
current. Intersecondary length >50% of subjacent secondary, proximal
Tooth Shapes
course parallel to major secondary, distal course parallel to subjacent
Tooth Shapes major secondary, vein frequency ~1 per intercostal area. Intercostal ter-
tiary vein fabric transverse freely ramified. Epimedial tertiaries rami-
Principal Vein 49.1 present
fied. Areolation development poor. Freely ending veinlets have two or
Principal Vein Termination 51.2.1 at apex of tooth more dendritic branches with tracheoid idioblasts. Marginal ultimate
venation loops incompletely. Tooth spacing regular, one order of teeth,
Course of Accessory Vein 0 absent
1 tooth/cm, sinus shape rounded. Principal vein present with termina-
Features of the Tooth Apex 53.3.1 spinose tion at apex of tooth, accessory vein absent. Tooth apex spinose.

Example 17. Anacardiaceae - Sorindeia gilletii

Anacardiaceae - Sorindeia gilletii

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 2.1 alternate Naked Basal Veins 24.1 absent
once pinnately com-
Leaf Organization 3.2.1.1 Number of Basal Veins 1
pound
Leaflet Arrangement 4.1 alternate Agrophic Veins 0 absent
Leaflet Attachment 5.1 petiolulate 2˚ Major 2o Vein Framework 27.2.1 eucamptodromous
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course n/a
Features of the Blade Perimarginal Veins 30.1 marginal secondary
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.5 mesophyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 3.3:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 0 absent
Medial Symmetry 11.2 asymmetrical Length 99 n/a
Base Symmetry 12.2.1 basal width asymmetrical Distal Course 99 n/a
Base Symmetry 12.2.3 basal insertion asymmetry Vein Frequency 99 n/a
Lobation 12.2.3 unlobed 3˚ Intercostal 3º Vein Fabric 35.2.3 composite admedial
Margin Type 13.1 untoothed Angle of Percurrent Tertiaries 99 n/a
Special Margin Features 0 absent Vein Angle Variability 99 n/a
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.3 reticulate
Apex Shape 17.3 acuminate Admedial Course n/a
Base Angle 18.1 acute Exmedial Course n/a
Base Shape 19.1.1 straight Exterior Tertiary Course 38.2 looped
Base Shape 19.1.1 straight 4˚ Quaternary Vein Fabric 39.2.2 irregular reticulate
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 40.2 freely ramifying
Surface Texture 88 not visible Areolation 41.2.1 poor development
Surficial Glands 88 not visible FEV branching 42.1.4.2 2 or more, dendritic

FEV termination 42.2.2 tracheoid idioblasts

Marginal Ultimate Venation 43.1 absent

III. Teeth Score Description Text Description:
Tooth Spacing 99 n/a Blade attachment marginal, laminar size mesophyll, L:W ratio
Number of Orders of Teeth 99 n/a 3.3:1, laminar shape elliptic with medial asymmetry and basal
width and insertion asymmetry. Margin is entire with acute apex
Teeth / cm 99 n/a angle, acuminate apex, acute base angle, and straight base shape.
Sinus Shape 99 n/a Primary venation is pinnate with no naked basal veins, one basal
vein, and no agrophic veins. Major secondaries eucamptodromous
Tooth Shapes 99 n/a with regular spacing, uniform angle, and excurrent attachment to
Tooth Shapes 99 n/a midvein. Interior secondaries absent, minor secondaries absent,
and marginal secondary present. Intersecondaries absent. Inter-
Tooth Shapes 99 n/a
costal tertiary veins composite admedial. Epimedial tertiaries
Tooth Shapes 99 n/a ramified. Exterior tertiaries reticulate. Quaternary vein fabric
freely ramifying. Quinternary vein fabric freely ramifying. Areo-
Principal Vein 99 n/a
lation moderately developed. Freely ending veinlets have two or
Principal Vein Termination 99 n/a more dendritic branches with highly branched sclereids. Marginal
ultimate venation absent.
Course of Accessory Vein 99 n/a
Features of the Tooth Apex 99 n/a

Example 18. Proteales - Leepierceia preartocarpoides

Proteales fossil - Leepierceia preartocarpoides

I. Leaf Characters Score Description II. Venation Score Description
Leaf Attachment 1.1 petiolate 1˚ Primary Vein Framework 23.1 pinnate
Leaf Arrangement 88 not visible Naked Basal Veins 24.1 absent
Leaf Organization 88 not visible Number of Basal Veins 7
Leaflet Arrangement 88 not visible Agrophic Veins 26.2.2 compound
Leaflet Attachment 88 not visible 2˚ Major 2o Vein Framework 27.4 mixed
Petiole Features 88 not visible Interior Secondaries 28.1 absent
Minor Secondary Course 29.2 simple brochidodromous
Features of the Blade Perimarginal Veins 0 absent
Position of Blade Attachment 7.1 marginal Major Secondary Spacing 31.1 regular
Laminar Size 8.3 microphyll Variation of Secondary Angle 32.1 uniform
Laminar L:W Ratio 1.75:1 Major Secondary Attachment 33.3 excurrent
Laminar Shape 10.1 elliptic Inter-2˚ Proximal Course 34.1.1 parallel to major secondaries
Medial Symmetry 11.1 symmetrical Length 34.2.1 <50% of subjacent secondary
parallel to subjacent major
Base Symmetry 12.1 symmetrical Distal Course 34.3.2 secondary
Base Symmetry 12.1 symmetrical Vein Frequency 34.4.1 <1 per intercostal area
Lobation 13.1 unlobed 3˚ Intercostal 3º Vein Fabric 35.1.1.1 opposite percurrent
Margin Type 14.2.2 serrate Angle of Percurrent Tertiaries 35.1.2.2 obtuse
Special Margin Features 0 absent Vein Angle Variability 36.3.1 basally concentric
Apex Angle 16.1 acute Epimedial Tertiaries 37.1.1.1 opposite percurrent
Apex Shape 17.3 acuminate Admedial Course 37.2.1.3 perpendicular to midvein
Base Angle 18.3 reflex Exmedial Course 37.2.2.1 parallel to intercostal tertiary
Base Shape 19.2.1 cordate Exterior Tertiary Course 38.2 looped
Base Shape 19.2.1 cordate 4˚ Quaternary Vein Fabric 39.1.3 mixed percurrent
Terminal Apex Features 0 absent 5˚ Quinternary Vein Fabric 88 not visible
Surface Texture 88 not visible Areolation 41.2.2 moderate development
Surficial Glands 88 not visible FEV branching 88 not visible
FEV termination 88 not visible
Marginal Ultimate Venation 88 not visible

III. Teeth Score Description Text Description:
Tooth Spacing 44.2 irregular Leaf attachment petiolate. Blade attachment marginal, laminar
Number of Orders of Teeth 45.1 one size microphyll, laminar L:W ratio 1.75:1, laminar shape elliptic,
blade medially symmetrical, base symmetrical, unlobed, margin
Teeth / cm 0.2 serrate. Apex angle acute, apex shape acuminate, base angle reflex,
Sinus Shape 47.2 rounded base shape cordate. Primary vein pinnate, naked basal veins absent,
seven basal veins, compound agrophic veins. Major 2º veins simple
Tooth Shapes st/st brochidodromous, interior secondaries absent, minor secondaries
Tooth Shapes cv/cv brochidodromous, major secondary spacing regular, secondary an-
gle uniform, major secondary attachment excurrent. Intersecond-
Tooth Shapes cc/cv
ary length <50% of subjacent secondary, proximal course parallel
Tooth Shapes to major secondary, distal course parallel to subjacent major sec-
ondary, vein frequency <1 per intercostal area. Intercostal tertiary
Principal Vein 49.1 present
vein fabric opposite percurrent with obtuse vein angle that is basally
Principal Vein Termination 51.2.1 at apex of tooth concentric. Epimedial tertiaries opposite percurrent with proximal
course perpendicular to midvein and distal course parallel to in-
Course of Accessory Vein 51.1.1 looped
tercostal tertiary. Exterior tertiary course looped. Quaternary vein
Features of the Tooth Apex 53.1 none fabric mixed percurrent, areolation development moderate. Tooth
spacing irregular, one order of teeth, 0.2 teeth/ cm, sinus shape
rounded, tooth shape straight/straight to concave/convex. Princi-
pal vein present with termination at apex of tooth.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Appendix C. Vouchers

M ost of the images are from the
National Cleared Leaf Collection,
Department of Paleobiology, National
Museum of Natural History, Smithsonian
Institution. Slides prefixed with NCLC-W
DMNH for the Denver Museum of Nature &
Science collection.

We used the Angiosperm Phylogeny Group
(A PG) website for fa m i ly a l ig nments
are housed at the Smithsonian; slides pre- (http://www.mobot.org/MOBOT/research/
fixed with NCLC-H are currently on long- APweb/). When nomenclature was in doubt
term loan at the Yale Peabody Museum. we used the International Plant Names Index
Additional abbreviations include NYBG for (IPNI) (http://www.ipni.org/), and to a lesser
the New York Botanical Garden, USNM extent, TROPICOS (http://mobot.mobot.
for the Smithsonian (fossil) collection and org/W3T/Search/vast.html).

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Dipterocarpus verrucosus A. D. E. Elmer 21650 (Brunei)
3 Dipterocarpaceae
Foxw. ex v. Slooten NCLC-W 1655

Peng 12615 (China)
6 Iteaceae Itea chinensis Hook. & Arn.
NCLC-H 3199

7 Berberidaceae Berberis sieboldii Miq. RWC ( Japan) NCLC-W 450

K. King 1926 (Kiangsu, China)
9 Cornaceae Alangium chinense (Lour.) Harms
NCLC-W 1225

Ruth 264 (Tennessee, USA)
11 Altingiaceae Liquidambar styraciflua L.
NCLC-H 815

T. H. Kearney 8000 (Pima
12 Euphorbiaceae Acalphya pringlei S. Watson
Co., Arizona) NCLC-H 6185

D. Daly (Madre de Dios,
15 Fabaceae Andira sp.
Peru) unvouchered

Malus mandshurica New York Botanical Garden
16 Rosaceae
(Maxim.) Kom. ex Juz. living collection (USA)

Chung 434 (Kiangsi, China)
18 Malvaceae Tilia chingiana Hu & Cheng
NCLC-W 8629

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Discocleidion rufescens W. Y. Chun 5021
19 Euphorbiaceae
(Fr.) Pax & K. Hoffm. (Hupeh, China) NCLC-W 3022
L. Krapovickas & Cristóbal
20 Euphorbiaceae Croton lobatus L. 12728 (Chaco, Argentina)
NCLC-W 11584
O. Degener 27455 (Hawaii,
21 Euphorbiaceae Aleurites remyi Sherff
USA) NCLC-H 709

Fabaceae- Cult. UCSC 259 (Puerto
31 Hymenaea courbaril L.
Caesalpinioideae Rico) NCLC-W 4284

Fabaceae- D. W. Stevenson s.n. (Vinh
51 Acacia mangium Willd.
Mimosoideae Phuc Province, Vietnam)

x Mahoberberis neubertii s.n. (North Dakota,
52 Berberidaceae
C. K. Schneid. USA) NCLC-H 1175

Deming s.n. (Connecticut,
53 Cabombaceae Brasenia schreberi J. F. Gmel.
USA) NCLC-H 6693

E. D. Merrill 1533 (Luzon,
54 Euphorbiaceae Macaranga bicolor Müll. Arg.
Philippines) NCLC-W 854

A. Henry 1952 (Taiwan)
56 Cucurbitaceae Trichosanthes formosana Hayata
NCLC-H 2050

J. Lebrun 2926 (Angodia,
57 Menispermaceae Dioscoreophyllum strigosum Engl.
Congo) NCLC-W 7814

B. A. Krukoff 6652 (Amazonas,
58 Celastraceae Cheiloclinium anomalum Miers
Brazil) NCLC-W 8251

W. T. S. Brown 2355
59 Apocynaceae Alstonia congensis Engl.
(Ghana) NCLC-W 5077

(without collector)
60 Chrysobalanaceae Parinari sp.
NCLC-W 12331

H. S. Irwin et al. (9/10/1960)
61 Moraceae Ficus citrifolia Mill.
(Amapá, Brazil) NCLC-W 10841

Xylomelum angustifolium C. C. Fauntleroy 2/17 (New South
62 Proteaceae
Kipp. ex Meissn. Wales, Australia) NCLC-W 6921

Y. Mexia 5241 (Minas Gerais,
63 Celastraceae Maytenus aquifolium Mart.
Brazil) NCLC-W 13582

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

C. G. Pringle 13822 (Guerrero,
64 Fabaceae-Faboideae Ramirezella pringlei Rose
Mexico) NCLC-W 14813

O. Degener 27455 (Hawaii,
65 Euphorbiaceae Aleurites remyi Sherff
USA) NCLC-H 709

J. A. Steyermark 47912
66 Salicaceae Lunania mexicana Brandeg.
(Guatemala) NCLC-H 1838

Chung 434 (Kiangsi,
67 Malvaceae Tilia chingiana Hu & Cheng
China) NCLC-W 8629

E. H. Wilson 2786 (Hupeh,
68 Oleaceae Fraxinus floribunda Wallich
China) NCLC-W 8963

G. T. Prance s.n. (Brazil)
69 Chrysobalanaceae Parinari campestris Aubl.
NCLC-H 4003

Melanolepis multiglandulosa E. D. Merrill 489 (Blanco,
70 Euphorbiaceae
Rchb. & Zoll. Philippines) NCLC-W 871

E. D. Merrill 5958 (Philippines)
71 Passifloraceae Adenia heterophylla (Blume) Koord.
NCLC-H 1935

Davidson 4197 (Iowa,
72 Rosaceae Potentilla recta Jacq.
USA) NCLC-H 3897

J. Wolfe, 1974 (cult. Missouri,
73 Proteaceae Stenocarpus sinuatus Endl.
USA, U815) NCLC-W 10238

Harvey Herb. (cult. Paris)
74 Proteaceae Dryandra longifolia R. Br.
NCLC-W 6334

I. L. Wiggins s.n. (Mexico)
75 Cucurbitaceae Cucurbita cylindrata L. H. Bailey
NCLC-H 2051

Fabaceae- Brion (1843) (Madagascar)
76 Bauhinia madagascariensis Desv.
Caesalpinioideae NCLC-W 5733

A. Ducke 35410, (Brazil)
77 Clusiaceae Caraipa punctulata Ducke
NCLC-H 1832

78 Salicaceae Casearia ilicifolia Vent. Miller 276 (Haiti) NCLC-H 1061

J. U. McClammer s.n. (Virginia,
79 Betulaceae Betula lenta L.
USA) NCLC-H 5415

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

H. Koidzumi 836 (Japan)
80 Violaceae Viola brevistipulata W. Becker
NCLC-H 2108

E. Kowalski and D. Dilcher
81 Fagaceae Quercus alba L.
126/132 (Millbrook, NY)

J. F. C. Rock 8636 (Yunnan,
82 Rosaceae Rubus mesogaeus Focke ex Diels
China) NCLC-W 12100

C. E. Tanner 3541 (Tanzania)
83 Phyllanthaceae Bridelia cathartica Bertol.f.
NCLC-W 11529

Le Sueur 1305 (Chihuahua,
88 Betulaceae Ostrya guatemalensis Rose
Mexico) NCLC-W 6773

x Mahoberberis neubertii (without collector) (North
89 Berberidaceae
C. K. Schneid. Dakota, USA) NCLC-H 1175

Fabaceae- Brion (1843) (Madagascar)
90 Bauhinia madagascariensis Desv.
Caesalpinioideae NCLC-W 5733

J. Lebrun 2926 (Angodia,
91 Menispermaceae Dioscoreophyllum strigosum Engl.
Congo) NCLC-W 7814

Anderson 260 (South Island,
93 Elaeocarpaceae Aristotelia racemosa Hook. f.
New Zealand) NCLC-W 9487

L. J. Brass 10908 (Papua New
94 Actinidiaceae Saurauia calyptrata Lauterb.
Guinea) NCLC-W 8944

Ozoroa obovata (Oliv.) A. Moura 43 (Mozambique)
95 Anacardiaceae
R. Fern. & A. Fern. NCLC-W 10067

Liriodendron chinense Chaney s.n. (Kiangsu,
96 Magnoliaceae
(Hemsl.) Sarg. China) NCLC-W 1553a

Neouvaria acuminatissima A. D. E. Elmer 21112 (Tawao,
97 Annonaceae
(Miq.) Airy-Shaw Philippines) NCLC-W 7851

JAW (7/6/64) (cult. Royal Botanic
98 Hamamelidaceae Corylopsis veitchiana Bean
Gardens, Kew) NCLC-W 1126

J. Steinbach 7333 (Santa Cruz,
99 Bignoniaceae Lundia spruceana Bur.
Bolivia) NCLC-W 218

Wachenheim (6/23/21) (French
100 Dichapetalaceae Tapura guianensis Aubl.
Guiana) NCLC-W 8070

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

S. Sohmer s.n. (Sri Lanka)
101 Dilleniaceae Schumacheria castaneifolia Vahl
NCLC-H 6793

Rimbach 38 (Ecuador)
102 Anacardiaceae Mauria heterophylla Kunth
NCLC-W4218

Chung 434 (Kiangsi,
103 Malvaceae Tilia chingiana Hu & Cheng
China) NCLC-W 8629

104 Aristolochiaceae Asarum europaeum L. Hawes s.n. (Poland) NCLC-H 6692

Cissampelos owariensis Beauv. ex Gilbert 2045 (Congo)
105 Menispermaceae
DC. (= C. pareira L.) NCLC-W 4498

W. Wolf (Alabama, USA)
106 Juglandaceae Carya leiodermis Sarg.
NCLC-W 8484

W. B. Marshall s.n. (New
107 Lauraceae Sassafras albidum (Nutt.) Nees
Jersey, USA) NCLC-W 6281

Prunus mandshurica E. H. Wilson 8775 (Korea)
108 Rosaceae
(Maxim.) Koehne NCLC-W 8775

U. Singh 136 (India)
109 Apocynaceae Carissa opaca Stapf. ex Haines
NCLC-W 13732

M. E. Jones (2-3-27) (Sinaloa,
110 Salicaceae Populus dimorpha Brandeg.
Mexico) NCLC-W 1262

Metcalf 2296 (Fukier)
111 Menispermaceae Diploclisia chinensis Merr.
NCLC-W 242

Steyermark 17489 (Panama)
112 Euphorbiaceae Adelia triloba Hemsl.
NCLC-W 2928

O. Degener 14673 (Fiji)
113 Apocynaceae Alstonia plumosa Labill.
NCLC-W 13703

R. W. Chaney (Japan)
114 Berberidaceae Berberis sieboldii Miq.
NCLC-W 450

H. McKee 4620 (New
115 Phyllanthaceae Phyllanthus poumensis Guillaumin
Caledonia) NCLC-W 11758

Cercidiphyllum japonicum
116 Cercidiphyllaceae (without collector) NCLC-W 9085
Sieb. & Zucc.

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Knowlton s.n. (Maine,
117 Sapindaceae Acer saccharinum L.
USA) NCLC-H 6861

H. Meyer (cult. Strybing Arb.
118 Hamamelidaceae Liquidambar styraciflua L.
66-125) NCLC-W 11912

119 Alismataceae Sagittaria sp. (without collector) NCLC-W 797

Palacios-Cuezzo 2233 (Corrientes,
120 Apocynaceae Araujia angustifolia Steud.
Argentina) NCLC-W 10244

Kruckeberg 97 (Oahu, Hawaii,
121 Menispermaceae Cocculus ferrandianus Gaudich.
USA) NCLC-W 10432

122 Fabaceae Bauhinia rubeleruziana J. D. Smith (without collector) NCLC-W 30221

F. von Mueller (Port Dennison,
123 Annonaceae Fitzalania heteropetala F. Muell.
Australia) NCLC-W 14543

131 Anacardiaceae Astronium graveolens Jacq. B. Wallnöfer 9567 (Peru) NY

Li 13081 (Anhui, China)
133 Betulaceae Carpinus fargesii C. K. Schneid.
NCLC-H 6455

J. Cuatrecasas 7403
135 Bixaceae Bixa orellana L.
(Colombia) NCLC-H 6255
R. A. & E. S. Howard 8249
Comocladia cuneata Britton
136 Anacardiaceae (Dominican Republic)
(syn.: C. acuminata)
NCLC-W 8197
E. Werdermann 923 (Coquimbo,
138 Griseliniaceae Griselinia scandens Taub.
Chile) NCLC-W 6513

(without collector) (Calcutta,
139 Ranunculaceae Delphinium cashmerianum Royle
India) NCLC-H 1477
E. H. Wilson 1914 (Botanic
Sapindaceae
140 Acer argutum Maxim. Garden Sapporo, Japan)
(ex-Aceraceae)
NCLC-W 8578
C. T. Hwa 36 (Szechuan,
141 Fagaceae Fagus longipetiolata Seemen
China) NCLC-W 1412

N. Espinal 3533 (Colombia)
142 Melastomataceae Meriania speciosa (Bonpl.) Naudin
NCLC-W 9286

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

R. E. Schultes & Cabrera
Loreya arborescens (Aubl.)
143 Melastomataceae 19755 (Amazonas, Colombia)
DC. (syn.: L. acutifolia)
NCLC-W 9280
Givotia rottleriformis R. G. Cooray 69100203R (Sri
144 Euphorbiaceae
Griff. ex Wight Lanka) NCLC-W 9046

J. McGregor M4/48
145 Euphorbiaceae Tannodia swynnertonii Prain
(Zimbabwe) NCLC-W 4631

(without collector) (China)
146 Trochodendraceae Tetracentron sinense Oliv.
NCLC-H 184

Hahm 150 (Martinique)
147 Clusiaceae Calophyllum calaba L.
NCLC-W 4372

R. K. Godfrey & Tryon 1443 (S
148 Lauraceae Sassafras albidum (Nutt.) Nees
Carolina, USA) NCLC-H 6280

JAW (7/6/64) (cult. Royal Botanic
149 Hamamelidaceae Parrotia jacquemontiana Decne.
Gardens, Kew) NCLC-W 1128

Buxus glomerata (Griseb.) A. H. Liogier 11086 (Dominican
151 Buxaceae
Müll. Arg. Republic) NCLC-H 6247

H. H. Pittier 5025 (Panama)
152 Euphorbiaceae Croton hircinus Vent.
NCLC-H 6223

Spondias globosa D. C. Daly et al. 7836
153 Anacardiaceae
J. D. Mitch. & Daly (Acre, Brazil) NY

(without collector) Nat. Col., B. Sci.
154 Menispermaceae Diploclisia kunstleri (King) Diels
2175 (Sarawak) NCLC-W 8815

P. Grant (May 22, 1963) (Nayarit,
155 Betulaceae Ostrya guatemalensis Rose
Mexico) NCLC-W 14869

E. H. Wilson 3227 (W. China
156 Salicaceae Carrierea calycina Franch.
[4000 ft.]) NCLC-W 7957

Dalechampia cissifolia D. M. Porter et al. 4887
157 Euphorbiaceae
Poepp. & Endl. (Panama) NCLC-W 11597

H. H. Pittier 5025 (Panama)
158 Euphorbiaceae Croton hircinus Vent.
NCLC-H 6223
J. Wolfe (6/26/64) (cult.
159 Malvaceae Dombeya elegans Cordem. Royal Botanic Gardens,
Kew) NCLC-W 1170

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

M. Jayasuriya 1336 (Sri
160 Datiscaceae Tetrameles nudiflora R. Br.
Lanka) NCLC-H 4947

M. E. Derdson 583 (Panama)
161 Lauraceae Phoebe costaricana Mez & Pittier
NCLC-W 5550

J. Wolfe (Berkeley, Calif.,
162 Platanaceae Platanus racemosa Nutt.
USA) NCLC-W 500

A. Henry 1952 (Taiwan)
163 Cucurbitaceae Trichosanthes formosana Hayata
NCLC-H 2050

Tsang 27855 (Kwangsin,
164 Rhamnaceae Paliurus ramosissimus Poir.
China) NCLC-W 1796

R. Lent 1586 (Costa Rica)
165 Piperaceae Sarcorhachis naranjoana Trel.
NCLC-W 12667

E. Contreras 6168 (Guatemala)
166 Melastomataceae Topobea watsonii Cogn.
NCLC-W 7585

N. S. Pillans 10899 (South
167 Proteaceae Paranomus sceptrum Kuntze
Africa) NCLC-W 5246

Pick s.n. (Oregon, USA)
168 Potamogetonaceae Potamogeton amplifolius Tuckerm.
NCLC-H 6777

Maianthemum dilatatum (Wood.) L. Roush (7/6/1919) (Washington
169 Ruscaceae
A. Nelson & J. F. Macbr. State, USA) NCLC-W 17896

A. Henry 1952 (Taiwan)
170 Cucurbitaceae Trichosanthes formosana Hayata
NCLC-H 2050

171 Sapindaceae Acer miyabei Maxim (without collector) NCLC-W 9072

Shiota 6315 (Mino, Japan)
172 Rosaceae Sorbus japonica (Decne.) Hedlund
NCLC-W 8671

Aravena (Linares, Chile)
173 Cunoniaceae Eucryphia glandulosa Reiche
NCLC -W 2468

A. Henry 12131C (China)
174 Euphorbiaceae Alchornea tiliifolia Müll. Arg.
NCLC-H 406
J. Wolfe (7/6/1964) (cult.
175 Hamamelidaceae Parrotia jacquemontiana Decne. Royal Botanic Gardens,
Kew) NCLC-W 1128

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Melville & Hooker 461 (Sierra
176 Vitaceae Cissus caesia Afzel.
Leone) NCLC-W 4948

Corylopsis glabrescens Walker 7663 (Pennsylvania,
177 Hamamelidaceae
Franch. & Sav. USA) NCLC-H 821

J. Cuatrecasas 11814
178 Desfontaineaceae Desfontainea spinosa Ruiz & Pav.
(Colombia) NCLC-H 4085

J. Clemens 16713 (Luzon,
179 Menispermaceae Cyclea merrillii Diels
Philippines) NCLC-W 4036

P. K. H. Dusén (11/25/14)
180 Salicaceae Aphaerema spicata Miers
(Paraná, Brazil) NCLC-W 1570

Cercidiphyllum japonicum R. W. Chaney s.n. (Japan)
181 Cercidiphyllaceae
Sieb. & Zucc. NCLC-W 26

182 Salicaceae Casearia ilicifolia Vent. Miller 276 (Haiti) NCLC-H 1061

A. K. Meebold s.n. (New
183 Atherospermataceae Laurelia novae-zelandiae A. Cunn.
Zealand) NCLC-H 6724

W. P. Fang 6705 (Szechuan,
184 Trochodendraceae Tetracentron sinense Oliv.
China) NCLC-W 6550

Mahonia wilcoxii R. S. Ferris 9991 (Arizona,
185 Berberidaceae
(Kearney) Rehder USA) NCLC-W 15043

Idrobo & Schultes 1320
186 Dilleniaceae Tetracera rotundifolia Sm.
(Colombia) NCLC-H 831

Li 13101 (Anhui, China)
187 Cornaceae Cornus officinalis Sieb. & Zucc.
NCLC-H 6496

de Silva 53 (Sri Lanka)
188 Dipterocarpaceae Isoptera lissophylla Liv.
NCLC-W 1662

H. A. Gleason 625 (Guyana)
189 Melastomataceae Tococa aristata Benth.
NCLC-W 9296

A. D. E. Elmer 21651
190 Phyllanthaceae Cleistanthus oligophlebius Merr.
(Brunei) NCLC-W 11559

191 Rhamnaceae Rhamnidium elaeocarpum Reiss. Pereira s.n. (Brazil) NCLC-H 4811

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

R. Schodde 3496 (New South
192 Cunoniaceae Eucryphia moorei F. Muell.
Wales, Australia) NCLC-W 2470

193 Anacardiaceae Cotinus obovatus Raf. W. Hess et al. 7511 (USA) NY

G. A. Zenker 568 (Cameroon)
194 Phyllanthaceae Baccaurea staudtii Pax
NCLC-H 11493

(without collector) (Camaris,
195 Burseraceae Santiria samarensis Merr.
Philippines) NCLC-H 208
(without collector) B. Sparre
196 Aextoxicaceae Aextoxicon punctatum Ruiz & Pav. & Constance 10742 (Osorno,
Chile) NCLC-W 2932
Antigonon cinerascens A. Ventura 4342 (Veracruz,
197 Polygonaceae
M. Martens & Galeotti Mexico) NCLC-W 14958

P. K. H. Dusén 1033a (Paraná,
198 Canellaceae Capsicodendron pimenteira Hoehne
Brazil) NCLC-H 238

Wachenheim s.n., 6/23/21
199 Dichapetalaceae Tapura guianensis Aubl.
(French Guiana) NCLC-W 8070

Comocladia glabra (Schult.) A. H. Liogier et al. 32748
200 Anacardiaceae
Spreng. (Puerto Rico) NY

L. F. Henderson (7/11/1882)
201 Rosaceae Filipendula occidentalis Howell
(Oregon, USA) NCLC-W 10707

A. J. M. Leeuwenberg 2877
202 Malvaceae Triplochiton scleroxylon K. Schum.
(Ivory Coast) NCLC-W 3656

(without collector) (Malaysia)
203 Achariaceae Scaphocalyx spathacea Ridl.
NCLC-H 953

Li 13015 (Anhui, China)
204 Adoxaceae Viburnum setigerum Hance
NCLC-H 6461

J. Cuatrecasas 7403
205 Bixaceae Bixa orellana L.
(Colombia) NCLC-H 6255

King 3446 (Chiapas, Mexico)
206 Asteraceae Philactis zinnioides Schrad.
NCLC-W 15130

G. T. Prance et al. s.n.
207 Polygalaceae Securidaca marginata Benth.
(Brazil) NCLC-H 2679

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Spondias bivenomarginalis
208 Anacardiaceae Liu Xingqi 27277 (China) MO
K. M. Feng & P. Y. Mao

J. Cuatrecasas 16820 (Valle,
209 Melastomataceae Graffenrieda anomala Triana
Colombia) NCLC-W 9273

(without collector) (Washington,
210 Fagaceae Castanea sativa Mill.
DC, USA) NCLC-H 1441

R. M. King 5488 (Kanchanaburi,
211 Lamiaceae Vitex limonifolia Wall.
Thailand NCLC-W 6656

Kermadecia sinuata M. Mackee 12877 (New
212 Proteaceae
Brongn. & Gris Caledonia) NCLC-W 6599

A. C. Smith 7366 (Fiji)
213 Phyllanthaceae Glochidion bracteatum Gillespie
NCLC-W 11666
Mairie-Victorin (6/30/33)
214 Salicaceae Populus jackii Sarg. (Montreal, Canada)
NCLC-W 1265
A. Ducke 18080 (Rio de Janeiro,
215 Malvaceae Apeiba macropetala
Brazil) NCLC-H 5343

J. Bright 9369 (Pennsylvania,
216 Malvaceae Tilia heterophylla Vent.
USA) NCLC-W 7734

Alchornea polyantha F. C. Lehmann (Cauca,
217 Euphorbiaceae
Pax & K. Hoffm. Colombia) USNH 1856534

B. A. Krukoff 10256 (La Paz,
218 Moraceae Pseudolmedia laevis Ruiz & Pav.
Bolivia) NCLC-W 10906

219 Annonaceae Popowia congensis Engl. & Diels Louis 724 (Congo) NCLC-W 5442

Banisteriopsis laevifolia Y. Mexia 5666 (Minas Gerais
220 Malpighiaceae
(A. Juss.) B. Gates Brazil) NCLC-W 6553

R. S. Toroes 1913 (Sumatra)
221 Malvaceae Microcos tomentosa Sm.
NCLC-W 11503

222 Iteaceae Itea chinensis Hook. & Arn. Peng 12615 (China) NCLC-H 3199

W. L. C. Muenscher &
223 Rosaceae Crataegus brainerdii Sarg. Lindsey 3373 (New York,
USA) NCLC-W 11964

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

H. J. R. Vanderyst 25190 (Sanga,
224 Dilleniaceae Tetracera podotricha Gilg.
Congo) NCLC-W 7841

G. Forrest 24471 (E Tibet/SW
225 Cannabaceae Celtis cerasifera C. K. Schneid.
China) NCLC-W 9000
J. J. Wurdack & Monachino
Couepia paraensis
226 Chrysobalanaceae 39893 (Bolívar, Venezuela)
(Mart. & Zucc.) Benth.
NCLC-W 4142
J. J. Pipoly & Gharbarran
227 Burseraceae Protium subserratum (Engl.) Engl.
10170 (Guyana) NY

Dacryodes negrensis G. T. Prance et al. 16147
228 Burseraceae
Daly & M. C. Martínez (Amazonas, Brazil) NY

B. A. Krukoff 4816 (Amazonas,
229 Burseraceae Protium opacum Swart
Brazil) NCLC-W 13245

230 Burseraceae Santiria griffithii Engl. Anta 56 (Indonesia) NY

R. A. & E. S. Howard 8249
Comocladia cuneata Britton
231 Anacardiaceae (Dominican Republic)
(syn.: C. acuminata Britton)
NCLC-W 8197 NY
H. Fung 20372 (Wen-Ch’ang,
232 Ancistrocladaceae Ancistrocladus tectorius Merrill
China) NCLC-H 5747

233 Burseraceae Canarium ovatum Engl. Molina 24514 (Philippines) NY

P. S. Ashton 2003 (Sri
234 Dipterocarpaceae Stemonoporus nitidus Thw.
Lanka) NCLC-H 4665

D. Vincent (Brazil)
235 Meliaceae Guarea tuberculata Vell.
NCLC-W 15406

Cedrela angustifolia Cooper & Slater (Panama)
236 Meliaceae
Moc. & Sessé ex DC. NCLC-H 640

T. Lasser 58 (Venezuela)
237 Ochnaceae Ouratea aff. O. garcinioides Ule
NCLC-H 5701

H. U. Stauffer 5827 (Fiji)
238 Violaceae Melicytus fasciger Gillespie
NCLC-W 3246

Note: genus and
239 Dipterocarpaceae P. S. Ashton (s.n.) NCLC-H 4552
species unknown

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

J. Cuatrecasas & Castañeda
240 Acanthaceae Aphelandra pulcherrima Kunth 25012 (Magdalena,
Colombia) NCLC-H 1207
E. L. Little 6129 (Pichincha,
241 Elaeocarpaceae Vallea stipularis L.f.
Ecuador) NCLC-H 5479

G. H. J. Wood 1791 (Brunei)
242 Burseraceae Santiria samarensis Merr.
NCLC-W 1733
E. G. Holt & Gehringer
243 Dilleniaceae Davilla rugosa Poir. 413 (Amazonas, Venezuela)
NCLC-H 845
Nectandra cuspidata T. G. Tutin 465 (Papua New
245 Lauraceae
Nees & Mart. ex Nees Guinea) NCLC-H 731

L. O. Williams 13211
246 Elaeocarpaceae Sloanea eichleri K. Schum.
(Venezuela) NCLC-H 5369

Bhesa archboldiana L. J. Brass 28105 (Papua New
247 Celastraceae
(Merr. & L. M. Perry) Ding Hou Guinea) NCLC-H 4421

B. A. Krukoff 5924 (Pará,
248 Picrodendraceae Piranhea trifoliata Baill.
Brazil) NY, NCLC-W 4626

Afrostyrax kamerunensis G. A. Zenker 365 (Cameroon)
249 Huaceae
Perkins & Gilg. NCLC-W 3257

J. M. & B. Reitsma 1420
250 Anacardiaceae Sorindeia gilletii De Wild.
(Gabon) NYBG

L. J. Dorr et al. 4617
251 Anacardiaceae Protorhus nitida Engl.
(Madagascar) NYBG

L. O. Williams 15365
252 Ochnaceae Ouratea thyrsoidea Engl.
(Venezuela) NCLC-H 5721

Comocladia glabra A. H. Liogier et al. 32748
253 Anacardiaceae
(Schult.) Spreng. (Puerto Rico) NYBG

254 Anacardiaceae Rhus (Melanococca) taitensis Guill. T. G. Yuncker 9332 (Tahiti) NYBG

S. K. Lau 3991 (Kiangsi,
255 Adoxaceae Viburnum sempervirens K. Koch
China) NCLC-H 1365

E. D. Merrill 9340 (Philippines)
256 Ebenaceae Diospyros maritima Blume
NCLC-W 13192

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Eriolaena malvacea A. Henry 12506 B (Yunnan,
257 Malvaceae
(H. Lév.) Hand.-Mazz China) NCLC-W 8045

E. D. Merrill 1533 (Luzon,
258 Euphorbiaceae Macaranga bicolor Müll. Arg.
Philippines) NCLC-W 854

Knowles & Bent s.n. (Metraro,
259 Juglandaceae Juglans boliviana Dode
Peru) NCLC-W 956b

Y. Mexia 6023 (Pará,
260 Apocynaceae Odontadenia geminata Müll. Arg.
Brazil) NCLC-W 9178

Taj 638 (Hainan, China)
261 Salicaceae Flacourtia rukam Zoll. & Mor.
NCLC-W 1577b

A. Petelot 8649 (Vietnam)
262 Actinidiaceae Actinidia latifolia Merr.
NCLC-W 8942

K. Ling (8/5/1926) (Kiangsu,
263 Cornaceae Alangium chinense (Lour.) Harms
China) NCLC-W 1225

C. F. Baker 2000 (Nicaragua)
264 Bixaceae Bixa orellana L.
NCLC-W 3234

L. O. Williams 15365
265 Ochnaceae Ouratea thyrsoidea Engl.
(Venezuela) NCLC-H 5721

R. S. Ferris 9991 (Arizona,
266 Berberidaceae Mahonia wilcoxii Rehder
USA) NCLC-W 15043

J. Cuatrecasas s.n. (Colombia)
267 Acanthaceae Aphelandra pulcherrima Kunth
NCLC-H 1297

D. E. Breedlove 42274 (Chiapas,
268 Capparaceae Capparis lundellii Standl.
Mexico) NCLC-W 15061b

R. Jaramillo & Dugand 4062
269 Dilleniaceae Dillenia indica Blanco
(Colombia) NCLC-H 918

C. A. Purpus 7381 (Chiapas,
270 Salicaceae Lunania mexicana Brandeg.
Mexico) NCLC-W 2693

C. G. Pringle (Mexico)
271 Celastraceae Celastrus racemosus Hayata
NCLC-H 4387

L. O. Williams 13211
272 Elaeocarpaceae Sloanea eichleri K. Schum.
(Venezuela) NCLC-H 5369

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

C. F. Baker (Nicaragua)
273 Bixaceae Bixa orellana L.
NCLC-W 3234

B. A. Krukoff 1644 (Mato
274 Malvaceae Theobroma microcarpa Mart.
Grosso, Brazil) NCLC-W 3654

Spiropetalum erythrosepalum G. A. Zenker 584 (Cameroon)
275 Connaraceae
Gilg. ex Schellen. NCLC-W 4198

P. Phillipson 1814
276 Burseraceae Commiphora aprevalii Guillaumin
(Madagascar) NYBG

Hedyosmum costaricense J. Luteyn & Stone 696 (Alajuela,
277 Chloranthaceae
C. E. Wood ex Burger Costa Rica) NCLC-H 6347B

B. A. Krukoff 5679 (Acre,
278 Picramniaceae Picramnia krukovii A. C. Sm.
Brazil) NCLC-W 13207

Y. Mexia 5098 (Minas Gerais,
279 Monimiaceae Mollinedia floribunda Tul.
Brazil) NCLC-W 10597

Escritor 21512 (Mindanao,
280 Lecythidaceae Barringtonia reticulata Miq.
Philippines) NCLC-W 12636

L. J. Brass 16182 (Malawi)
281 Apocynaceae Carissa bispinosa Desf.
NCLC-W 5044

Imperial Forest Institute 456
282 Celastraceae Gymnosporia senegalensis Loes.
(Tanzania) NCLC-H 4441

Shorea congestiflora P. S. Ashton 2022 (Sri
283 Dipterocarpaceae
(Thw.) P. S. Ashton Lanka) NCLC-H 4636

B. A. Krukoff 6203 (Amazonas,
284 Malvaceae Theobroma microcarpa Mart.
Brazil) NCLC-H 5641

K. Ling (8/5/1926) (Kiangsu,
285 Cornaceae Alangium chinense (Lour.) Harms
China) NCLC-W 1225a

Afrostyrax kamerunensis G. A. Zenker 365 (Cameroon)
286 Huaceae
Perkins & Gilg. NCLC-W 3257

A. D. Elmer s.n. (Luzon,
287 Ebenaceae Diospyros pellucida Hiern
Philippines) NCLC-H 5100
R. A. & E. S. Howard 8249
Comocladia cuneata Britton
288 Anacardiaceae (Dominican Republic)
(syn.: C. acuminata Britton)
NCLC-W 8197

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

Pseudolmedia laevis B. A. Krukoff 10256
289 Moraceae
(Ruiz & Pav.) J. F. Macbr. (La Paz, Bolivia) NCLC-W 10906

H. S. Irwin 32089 (Brazil)
290 Ebenaceae Diospyros hispida A. DC.
NCLC-H 5022

P. S. Ashton 2003 (Sri Lanka)
291 Dipterocarpaceae Stemonoporus nitidus Thw.
NCLC-H 4665

292 Anacardiaceae Rhus (Melanococca) taitensis Guill. T. G. Yuncker 9332 (Tahiti) NYBG

Chloranthus glaber Tsang 21487 (Kwangtung, China)
293 Chloranthaceae
(Thunb.) Makino NCLC-W 2329

Killip 34966 (Colombia)
294 Clusiaceae Clusiella pendula Cuatrec.
UCH967992 NCLC-W 2648

B. A. Krukoff 5924 (Pará, Brazil)
295 Picrodendraceae Piranhea trifoliata Baill.
NY, NCLC-W 4626

Afrostyrax kamerunensis G. A. Zenker 365 (Cameroon)
296 Huaceae
Perkins & Gilg. NCLC-W 3257

302 Violaceae Melicytus fasciger Gillespie Stauffer 5827 (Fiji) NCLC-W 3246

G. V. Nash & N. Taylor 1205
303 Burseraceae Bursera inaguensis Britton
(Bahamas) NYBG

Tetragastris panamensis S. A. Mori et al. 14969
304 Burseraceae
(Engl.) Kuntze (French Guiana) NYBG

A. V. Bogdan VB 622 (Kenya)
305 Euphorbiaceae Pycnocoma littoralis Pax
NCLC-W 3141

Y. Mexia 5098 (Minas Gerais,
308 Monimiaceae Mollinedia floribunda Tul.
Brazil) NCLC-W 10597

B. A. Krukoff 5679 (Acre, Brazil)
309 Picramniaceae Picramnia krukovii A. C. Sm.
NCLC-W 13207

Ramos 1364 (Brunei) NCLC-W
310 Phyllanthaceae Aporusa frutescens Blume
11487

H. Rombouts 662 (Brazil)
312 Rhamnaceae Gouania velutina Reiss.
NCLC-H 5324

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

A. D. E. Elmer 16177 (Luzon,
313 Hydrangeaceae Dichroa philippinensis Schltr.
Philippines) NCLC-W 2161

Campylostemon mucronatum A. J. M. Leeuwenberg 4118 (Ivory
314 Celastraceae
(Exell) J. B. Hall Coast) NCLC-W 6867

Leea macropus JAW (6/26/64) (cult. Royal Botanic
315 Vitaceae
Lauterb. & K. Schum. Gardens, Kew) NCLC-W 1151

L. Hickey s.n. (New Zealand)
316 Elaeocarpaceae Aristotelia racemosa Hook.f.
NCLC-H 6479

Muenscher & Lindsey 3373 (New
317 Rosaceae Crataegus brainerdi Sarg.
York, USA) NCLC-W 11964

A. D. E. Elmer 16177 (Luzon,
318 Hydrangeaceae Dichroa philippinensis Schltr.
Philippines) NCLC-W 2161

G. Forrest 24471 (E Tibet/SW
319 Cannabaceae Celtis cerasifera C. K. Schneid.
China) NCLC-W 9000

Achten 560 (Luebo, Congo)
320 Salicaceae Phylloclinium paradoxum Baill.
NCLC-W 7830

Carpinus laxiflora C. Y. Chiao 14466 (Chekiang,
322 Betulaceae
(Siebold & Zucc.) Blume China) NCLC-H 6212

Chloranthus serratus P. H. Dorsett & W. J. Morse 503
323 Chloranthaceae
Roem. & Schult. (Fujiyama, Japan) NCLC-H 658

Martynia annua I. S. Brandegee s.n. (Sinaloa,
324 Martyniaceae
L. ex Rehm. Mexico) NCLC-H 1706
Lopezia lopezoides (Hook. &
McVaugh 14350 (Jalisco, Mexico)
325 Onagraceae Arn.) Plitmann, P. H. Raven &
NCLC-H 1909)
Breedlove
D. E. Breedlove 15821 (Guerrero,
326 Onagraceae Fuchsia decidua Standl.
Mexico) NCLC-H 3852

C. L. Porter 3887 (Colorado, USA)
327 Sapindaceae Acer negundo L.
NCLC-W 14573

S. Venturi 5206 (Jujuy, Argentina)
328 Sapindaceae Cupania vernalis Cambess.
NCLC-H 2091

S. Ripley 67 (Sri Lanka)
329 Celastraceae Elaeodendron glaucum Pers.
NCLC-H 4425

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

without collector
330 Fagaceae Quercus alba × velutina
NCLC-W 1079

331 Violaceae Melicytus Fasciger Gillespie Stauffer 5827 (Fiji) NCLC-W 3246

E.W. Nelson 540 (Mexico)
332 Platanaceae Platanus oaxacana Standley
NCLC-H 3743

(without collector) (Shinano, Japan)
333 Berberidaceae Diphylleia grayi F. Schmidt
NCLC-H 1168B

(without collector) Maire 7462
334 Vitaceae Vitis cavaleriei H. Lév & Vaniot
(Yunnan, China) NCLC-W 289

Lei 541 (Hainan, China)
335 Iteaceae Itea macrophylla Wall.
NCLC-H 3250

D. R. Harris 11955 (Virgin Islands,
336 Malvaceae Melochia lupulina Sw.
USA) NCLC-H 5555

P. Raven s.n. (Japan)
337 Onagraceae Circaea erubescens Franch. & Sav.
NCLC-H 2154

E. L. Ekman 7984 (Pinar del Río,
338 Salicaceae Homalium racemosum Jacq.
Cuba) NCLC-H 1019

(without collector) (China)
339 Trochodendraceae Tetracentron sinense Oliv.
NCLC-H 184

G. Forrest 20680 (Yunnan, China)
340 Aquifoliaceae Ilex dipyrena Wall.
NCLC-H 4342

Rogers 18117 (Transvaal, South
341 Salicaceae Trimeria alnifolia Harv.
Africa) NCLC-H 1016

(without collector) (England, cult.)
342 Theaceae Hartia sinensis Dunn.
NCLC-H 5

S. Sohmer & Waas (Sri Lanka)
343 Dilleniaceae Schumacheria castaneifolia Vahl
NCLC-H 6793

(without collector) (Japan)
344 Salicaceae Idesia polycarpa Maxim.
NCLC-H 1005

Cercidiphyllum genetrix L. Hickey (Golden Valley Fm.)
345 Cercidiphyllaceae
(Newberry) Hickey USNM 43234

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

C. Y. Chiao 2721 (Shantung,
APP 1 Malvaceae Tilia mandshurica Rupr.
China) NCLC-H 5406
E. G. Holt & Gehriger s.n.
APP2 Dilleniaceae Davilla rugosa Poir. (Amazonas, Venezuela)
NCLC-H 845
P. S. Ashton 2003 (Sri Lanka)
APP 3 Dipterocarpaceae Stemonoporus nitidus Thw.
NCLC-H 4665

Fabaceae- Brion 1843 (Madagascar)
APP 4 Bauhinia madagascariensis Desv.
Caesalpinioideae NCLC-W 5733

W. P. Fang 6705 (Szechuan,China)
APP 5 Trochodendraceae Tetracentron sinense Oliv.
NCLC-W 6550

Buchanania arborescens Reynoso et al. s.n. (PPI 1403)
APP 6 Anacardiaceae
(Blume) Blume (Philippines) NY

L. Hickey s.n. (New Zealand)
APP 7 Elaeocarpaceae Aristotelia racemosa Hook.f.
NCLC-H 6479

L. Williams 11630 (Venezuela)
APP 8 Malvaceae Bombacopsis rupicola Robyns
NCLC-H 5493

Rhynchoglossum azureum D. E. Breedlove 1154 (Chiapas,
APP 9 Gesneriaceae
(Schltdl.) B. L. Burtt. Mexico) NCLC-H 1714

P. Moreau 62822 (Argentina)
APP 10 Nothofagaceae Nothofagus procera Oerst.
NCLC-H 1760

Fang 3924 (Szechuan)
APP 11 Sapindaceae Acer franchetii Pax
NCLC-W 7628

Tetrapterys macrocarpa C. O. Erlanson 405 (Panama)
APP 12 Malpighiaceae
I. M. Johnst. NCLC-H 2479

Eucryphia glutinosa Aravena (Linares, Chile)
APP 13 Cunoniaceae
(Poepp. & Endl.) Baill. NCLC-W 2468

Biltmore, 19496 (Florida, USA)
APP 14 Chrysobalanaceae Licania michauxii Prance
NCLC-H 4026

Wiggins 7033 (Sonora, Mexico)
APP 15 Moraceae Morus microphylla Buckley
NCLC-W 14883B

T. Zanoni et al. 30780
APP 16 Anacardiaceae Comocladia dodonaea (L.) Urban
(Dominican Republic) NY

Collector and field number
Fig. Family Genus and species
(where collected) slide no.

J. M. & B. Reitsma 3112
APP 17 Anacardiaceae Sorindeia gilletii De Wild
(Gabon) NY

Leepierceia preartocarpoides Johnson 571 (Hell Creek Fm.)
APP 18 Proteales
(Brown) Johnson DMNH 6359

L eaf clearing is the process of removing
all pigment and then staining a leaf so
that its vein architecture is clearly visible.
This procedure can be used on leaves removed
(with permission) from herbarium sheets or on
Acid fuchsin is a particularly successful stain,
although safranin dye can also be useful.
Staining with acid fuchsin involves washing
the leaves in 50% ethanol, staining them in
1% acid fuchsin for 3–8 minutes, and then
live material. Many methods are used for clear- putting the leaves through a dehydration se-
ing leaves; here we briefly describe one method. ries in 50%, 95%, and 100% ethanol. The
The following sources contain additional infor- first two dehydration steps destain the leaves
mation on leaf clearing techniques: Foster (1953), because the water-soluble dye diffuses out of
de Strittmatter (1973), Hickey (1973), Shobe and the leaf into the ethanol; the third step stops
Lersten (1967), Pane (1969), and Bohn et al. the process once there is proper contrast be-
(2002). Note that this process must be performed tween leaf lamina and stained veins. The
in a well-ventilated area because some of the specimens can then be rinsed in clove oil, then
chemicals are harmful to humans. xylene (a toxic solvent), and finally stored tem-
porarily in a solution of 1:1 xylene:HemoDe®.
Leaves are placed in glass containers, cov- Proceeding directly from dehydration to stor-
ered by a piece of fiberglass mesh to facilitate age in HemoDe® also gives good results, but
changing solutions, and submerged in 1–5% the leaves will eventually lose some pigment.
NaOH, the strength depending on the thick-
ness of the material. The NaOH solution is For photography, the leaves are floated in a
changed every 1–2 days during the clearing glass dish placed on the backlit platform of a
process, which generally takes 2–10 days. dissecting microscope with a digital camera
The clearing process is finished by a wash in attachment. The acid fuchsin dye fades over
commercial Clorox® (typically 5–30 seconds) time, and limited restaining may be necessary
followed by a final wash in water to stop the in order to attain the necessary contrast for im-
bleaching process. Clorox removes any re- aging. Leaves are then permanently mounted
maining pigment from the leaves in prepara- on glass slides using standard anatomical tech-
tion for staining. This step requires caution niques. The leaves used in this publication were
because the leaves are typically fragile from permanently mounted and then photographed
the NaOH treatment and may disintegrate if using a light table or converted enlarger con-
bleached for too long. denser as a source of transillumination.

Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
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Index
A compound leaf – 8
abaxial – 4 concave – 5
accessory vein – 102, 109 concave base – 38, 39
acrodromous primary – 57, 60 concavo-convex base – 38, 39
acropetiolar – 18 conjunctal vein – 102, 107
actinodromous primary – 57, 59 convex – 5
acuminate apex – 34–35 convex apex – 34–35
acute apex angle – 32, 33 convex base – 38, 39
acute base angle – 36 cordate – 38, 40
adaxial – 4 costal – 44
admedial – 4 costal secondaries – 54
admedial vein – 102, 107 craspedodromous – 64–65
agrophic veins – 63 crenate – 28
alate petiole – 18, 19 cuneate – 38, 39
apex – 5
apex angle – 32 D
apex shape – 34–35 decurrent – 5, 44, 79
apical – 4 decurrent base – 38, 40
apical extension – 10, 38 deflected – 79
apical extension length – 10 dentate – 28
areolation – 98 dichotomous – 44
auriculate – 39 distal – 4
distal flank – 101
B
basal – 4 E
basal extension – 10, 38 elliptic – 22, 23
basal extension asymmetry – 24, 25 emarginate apex – 34–35
basal extension length – 10, 11 epimedial tertiaries – 53, 91–94
basal insertion asymmetry – 24, 25 erose – 30
basal veins – 62 eucamptodromous – 64, 68
basal width asymmetry – 24, 25 excurrent – 45, 79
basal width ratio – 10, 11 exmedial – 5
base – 5 exterior tertiaries – 53, 95
base angle – 36–37
base shape – 38–41 F
base symmetry – 24, 25 fabric – 45, 47
bilobed – 26, 27 festooned brochidodromous – 65, 71
brochidodromous – 65, 70 festooned semicraspedodromous – 64, 67
FEV’s – 56, 99
C fimbrial vein – 74,
campylodromous – 57, 61 flabellate – 57, 60
canaliculate – 18, 19 foraminate – 110
cassidate – 110–111 freely ending veinlets – 56, 99
circular base angle – 36–37
cladodromous – 64, 69
complex base shape – 38, 40
compound agrophics – 63

G marginal secondary – 74
gauge – 45 marginal ultimate vein – 100
glands – 18 medial asymmetry – 24
glands, surface – 43 medial symmetry – 24
midvein – 7
H midvein length – 10
hastate – 38, 41 minor secondaries – 54, 73
hemieucamptodromous – 64, 69 monopodial – 45
mucronate – 42
I mucronate tooth apex – 110–111
insertion point – 6, 8
intercostal – 45 N
intercostal tertiaries – 53 naked basal primary veins – 61
interior secondaries – 54–55, 72 naked base – 61
intersecondaries – 54–55, 80–83
intramarginal vein – 54–55, 74 O
involute – 30, 31 oblong – 22, 23
obovate – 22, 23
L obtuse apex angle – 32
lamina – 6 obtuse base angle – 36
laminar shape – 22 orders of teeth – 104
laminar size – 20, 21 ovate – 22, 23
leaf arrangement alternate – 12, 13
leaf arrangement opposite – 12, 13 P
leaf arrangement subopposite – 12, 13 palinactinodromous – 57, 59
leaf arrangement whorled – 12, 13 palmate primary – 57–61
leaf domatia – 7 palmately lobed – 26, 27
leaf organization compound – 14, 15 palmatisect – 26, 27
leaf organization palmately panduriform – 39
compound – 14, 15 papillate surface – 42
leaf organization pinnately papillate tooth apex – 110, 112
compound – 14, 15 parallelodromous – 57, 61
leaf organization simple – 14 peltate central – 20
leaf rank – 48 peltate excentric – 20
leaflet – 8 percurrent – 84–86, 89–90
leaflet arrangement – 16 perimarginal – 74
leaflet attachment – 16 petiolate – 12
length:width ratio – 22 petiole – 6
linear shape – 22, 23 petiole base pulvinate – 17
lobate – 38, 41 petiole base pulvinulate – 17
lobation – 26, 27 petiole base sheathing – 17
lobe – 7 petiolulate – 16
lobed apex – 34–35 petiolule – 8
lobe-like teeth – 29 phyllodes – 18,19
low rank – 48 pinnate primary – 57, 58
pinnately lobed – 26, 27
M pinnatisect – 26, 27
major secondaries – 54 pitted surface – 42
major secondary spacing – 75–76 primary veins – 45–52, 57–61
margin – 7 principal vein – 102, 107–108
marginal secondaries – 54, 56, 64–71 proximal – 4

proximal flank – 101 toothed margin – 28
pubescent surface – 42 tooth-like lobes – 29
pulvinate – 17 truncate apex – 34–35
pulvinulate – 17 truncate base – 38-39
tylate – 110–111
Q
quaternary veins – 96 U
quinternary veins – 97 undulate – 30, 31
unlobed – 26
R untoothed margin – 28
rachis – 8
ramified – 46, 85, 88 V
reflex apex angle – 32, 33 vein course – 46
reflex base angle – 36–37 vein fabric – 45
reticulate – 84, 87 veins – 47–100
reticulodromous – 64, 69
retuse – 34, 42 W
revolute – 30, 31 width ratio – 10, 11
rounded apex – 34–35
rounded base – 38, 39
rugose surface – 42
runcinate – 39

S
sagittate – 38, 41
semicraspedodromous – 64, 66
semiterite – 18, 19
serrate – 28
sessile – 12, 16, 17
setaceous tooth apex – 110, 112
simple agrophics – 63
sinuous – 30
sinus – 7, 101, 105
special shape – 22, 23
spherulate tooth apex – 110, 112
spinose – 42
spinose tooth apex – 110–111
stipel – 9
stipule – 9
straight apex – 34–35
straight base – 38, 39
surface texture – 42
surficial glands – 43
sympodial – 46

T
teeth – 103–112
terete – 18,19
tooth apex – 101, 110
tooth shape – 106
tooth spacing – 103

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please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.
Licensed under Creative Commons license CC BY-NC 4.0. To reprint this work in whole or in part for commercial purposes,
please contact Cornell University Press: www.cornellpress.cornell.edu. Copyright © 2009 Cornell University.