Professional Documents
Culture Documents
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©1999 by Smithsonian Institution.
ISBN 0-9677554-0-9
Please cite as:
Manual of Leaf Architecture - morphological description and categorization of
dicotyledonous and net-veined monocotyledonous angiosperms by Leaf Architecture
Working Group. 65p.
We gratefully acknowledge funding from Michael Sternberg and Jan Hartford for the printing
of this manual.
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Names and addresses of the Leaf Architecture Working Group in alphabetical order:
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INTRODUCTION
Since the time of Linnaeus the identification and reconstruction of relationships between
plants have been based largely on features of the reproductive organs. Although flower and fruit
characters have proved very useful in both botany and paleobotany, there are situations in which
these organs are not available for study. For example, leaf compression and impression fossils
are the most common macroscopic remains of plants, but they are generally not attached to other
plant organs. Because of their abundance and dense stratigraphic occurrence, fossil leaves can
provide an enormous amount of information about the composition and diversity of past floras - if
they can be used to recognize species reliably and assign them to higher taxa. Tropical botanists
also find themselves confronted with the need to identify and classify plants using vegetative
characters because so many long-lived tropical plants flower infrequently and irregularly. In spite
of the success of Linnaeus’s sexual system and its descendants, there is a great need to be able to
identify and classify dispersed leaves. The overall purpose of this manual is to help you do that.
The problem of working with isolated leaves is a long-standing one in paleobotany. Lacking
both an accepted system of terms for describing leaf form, and a knowledge of the systematic
distribution of leaf features among living angiosperms, and in many cases faced with poorly
preserved fossils, most early workers focused on overall characters of leaf shape and size that
ultimately have not proven very useful in recognizing species or higher taxa. Names of living
genera were widely applied to fossils so that there are, for example, many taxonomically valid
fossil species of Ficus, Populus, and Aralia based on poorly preserved leaves with only vague
similarities to the living members of these genera. Late nineteenth and early twentieth century
angiosperm paleobotanists left a legacy of poorly defined taxa with botanically misleading names.
In the last half of the twentieth century two new approaches have helped rectify this problem.
One has been to study multiple organs, including leaves, thought to represent the same plant
species, either because they are preserved in attachment or because they occur together at many
localities. This approach allows traditional characters of flowers and fruits to be used in defining
extinct taxa and determining their relationships (e.g., Manchester 1986). Studying characters of
multiple organs of the same plant allows fossil taxa to be described more comprehensively and
systematic relationships to be established with greater certainty than can be gained from leaves
alone. However, there are many types of fossil leaves that have not been found attached to or
consistently associated with other organs. The second approach has been to identify systematically
informative leaf features (Hickey and Wolfe 1975, Wolfe 1989, Hickey and Taylor 1991) that
allow species to be recognized on the basis of dispersed leaves; these features may also permit the
fossil to be assigned to a family or higher taxonomic category. This approach has been used
principally in dicotyledonous angiosperms with complex vein systems. Among living dicots,
foliar characters may or may not offer conclusive evidence of the generic or higher-level affinities
of a plant, but generally they do allow even closely related species to be distinguished (e.g., Merrill
1978).
The main goal of this manual is to define and illustrate for the reader an unambiguous and
standard set of terms for describing leaf form and venation, particularly of dicots. This manual
also provides a template and set of instructions that show how descriptive information can be
entered into a standardized database of fossil and extant leaves. The Leaf Architecture Working
Group (LAWG) adopted and in some cases added or modified the definitions and terms found in
this manual and developed its format.
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The purpose of any terminology or method for quantifying leaf form is to allow objective
description of and comparisons among different types of leaves. Many sets of terms and methods
have been devised for describing leaves (e.g., Ettingshausen 1861; Melville 1937, 1976; Dale et al.
1971; Hickey 1973, 1977, 1979; Mouton 1966, 1967; Dickinson et al. 1987; Jensen 1990; Ray
1992). These will not be reviewed here. Terms for the description of leaf form and venation are
largely from the leaf architectural system of Hickey (1973, 1977, 1979). The terms and drawings
illustrating leaf cuticle features have been taken without modification from Dilcher (1974). These
terminologies have been adopted because they are in wide use among botanists and paleobotanists,
including the members of the LAWG. Although fully quantitative methods for describing leaf
shape exist and are presumably more objective than the qualitative and semi-quantitative terms
described here, they have several disadvantages. Quantification of leaf shape through, for ex-
ample, Fourier or landmark methods is still time consuming when compared with semi-quantita-
tive characterization. It is also difficult to apply these techniques to typically incomplete fossil
specimens. Further, we wished this system to be applicable across all types of dicot and net-
veined monocot leaves, thus eliminating methods that require recognition of homologous points or
vein patterns. Finally, we decided against a fully quantitative approach because many of the most
systematically valuable features of leaves are in the venation, and quantification of vein networks
is even more time-consuming than quantification of leaf shape.
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THE LEAF ARCHITECTURE WORKING GROUP (LAWG) DATABASE
This manual is a companion to a database developed by the LAWG and is intended to help
researchers describe fossil leaves in a consistent way and to make it easier to compare leaves de-
scribed by different researchers working on floras from different ages or regions. The database is a
FileMaker®Pro application with 57 fields to contain information about each given morphotype. A
blank database form is shown in Figure 5. The first section of the entry form has 13 fields for
recording basic information about the morphotype and the fossils on which it is based. This in-
cludes fields for the higher taxonomic category, the describer, the localities at which the
morphotype has been found, and specimen numbers. The next 43 fields in the form are for descrip-
tors of the morphotype. Each field corresponds to a character of the size or shape of the leaf blade,
the course of the venation, the form of the margin, and the leaf cuticle. Each field is provided with a
pull-down list of character states that the character might have. These character state lists are based
on our collective experience with living and fossil dicot leaves, but the lists are not exhaustive. The
data-entry format of the fields will permit you to enter character states not on the pull-down lists,
but clearly the comparability of descriptions by different workers, and therefore the usefulness of
searching this database, will be enhanced if the defined terms are used whenever possible. If you
would like additional characters or character states to be added to this manual and the database,
please contact Scott Wing at the address listed on the third page of this manual. The final field in
the form is formatted to hold digital images which should include both a low magnification image to
show leaf shape and a higher magnification image to show details of venation and/or marginal teeth.
The digital version of the FileMaker®Pro entry form is on the CD that contains this manual.
This manual is organized in the same order as the database form. Characters, the field names
from the database, are numbered and in gray boxes. Choices of character states from the pull-down
lists are in boldfaced type. Definitions of terms used in the manual are in italics. For an explana-
tion and illustration of any character or character state simply go to the section of the manual that
corresponds to its number (see p. 10). The figures in the manual are numbered to link them to the
character names, thus figures 14.1 – 14.4 illustrate character states of the character “leaf attach-
ment,” which is the 14th field in the database. The figures used in the introductory section, and in
the definitions of terms rather than characters, are numbered sequentially through the manual.
Wherever possible we have tried to illustrate character states with real specimens rather than
idealized drawings. Almost all of the specimens used in illustrations are from the United States
Geological Survey/National Museum of Natural History cleared leaf collection. Slide numbers of
the cleared leaves are available in a spreadsheet file, and the original black and white images of the
figured specimens are stored in .jpg format on the CD with the digital version of this manual. The
images were recorded with a digital camera.
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3. Begin to sort the leaves into groups based on shared leaf-architectural characteristics. As each
group is defined, select the best specimen (most complete and well preserved) of that group to
be the “holomorphotype.” Assign a unique morphotype number to the holomorphotype (e.g.,
HC1) and sequester the type where it is accessible for comparison. This specimen should be
assigned a museum specimen number and its status as the holomorphotype noted on the speci-
men tag. It is also useful to maintain a running list or spreadsheet that records the information
about the individual holomorphotypes. One major distinction between holotype specimens (the
formal name-bearing specimen in Linnean taxonomy) and holomorphotype specimens (the
informal number-bearing specimen in this system) is that holotypes are permanent whereas
holomorphytpes may be replaced with better specimens or sunk into other morphotypes.
Proceed to identify all of the remaining specimens that can be referred to the morphotype based
on the holomorphotype and label them accordingly. It is usually best to start with the best-
preserved and most abundant morphotypes and work toward the poorly preserved and less
common types. In practice, as work proceeds on a fossil flora, some of what were originally
recognized as sharply delineated morphotypes will be shown to belong to a continuum, while
others will remain as discrete entities.
4. The initial sorting of a collection is usually done on the basis of toothed versus entire margins,
primary and secondary vein patterns, and the presence and types of lobes. These characters are
usually stable within morphotypes. The least reliable characters are leaf size and shape. Once
the fossils are grouped into broad categories, it is much easier to separate them by higher-order
venation pattern and tooth type [see Hickey 1973, 1979 and Hickey and Wolfe, 1975]. To
highlight the characters that define your groups, it is helpful to sketch and/or photograph the
holomorphotype and note diagnostic features and the range of variation. It is useful to print
photos or scanned slides as full page images that can be mounted on the wall of your work area.
This allows increased familiarity with the various morphotypes. In one varition on this tech-
nique, Kirk Johnson makes two sets of holomorphotype images. The first set is mounted on the
wall in numerical order and the second set is placed in folders in the following categories:
pinnate toothed leaves; pinnate entire leaves; palmate toothed leaves; palmate entire leaves;
palmately lobed leaves; pinnately lobed leaves; fruits, seeds and cones; gymnosperm leaves;
ferns and fern allies. This allows a large number of images to be searched visually or by major
architectural group.
5. Describe the morphotype using the holomorphotype as the basic reference. Expand the circum-
scription, when necessary, using additional specimens that show clear overlap in their morpho-
logical characters with the holomorphotype specimen. Use the fossil-leaf database and this
manual as a guide in this process.
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REFERENCES CITED AND ADDITIONAL SOURCES
Dale, M. B., Groves, R. H., Hull, V. J., O’Callaghan, J. F. 1971. A new method for describing leaf shape.
New Phytologist 70:437-442.
Dickinson, T. A., Parker, W. H., Strauss, R. E. 1987. Another approach to leaf shape comparisons. Taxon
36:1-20.
Dilcher, David L. 1974 Approaches to the identification of angiosperm leaves. The Botanical Review
40(1).
Ettingshausen, C. 1861. Die Blatt-Skelete der Dicotyledonen. Vienna 1,21.
Hickey, Leo J. 1973. Classification of the architecture of dicotyledonous leaves. American Journal of
Botany 60:17-33.
Hickey, Leo J. 1974. A revised classification of the architecture of dicotyledonous leaves. Pp. 25-39 in
C.R. Metcalfe and L. Chalk, eds. Anatomy of the Dicotyledons, Volume I, Second Edition.
Clarendon Press, Oxford.
Hickey, Leo J., and Wolfe, Jack A. 1975. The bases of angiosperm phylogeny: vegetative morphology.
Annals of the Missouri Botanical Garden 62(3):538-589.
Hickey, Leo J. 1977. Stratigraphy and paleobotany of the Golden Valley Formation (Early Tertiary) of
western North Dakota. Geological Soceity of America Memoir 150.
Hickey, L. J., Taylor, D. W. 1991. The leaf architecture of Ticodendron and the application of foliar
characters in discerning its relationships. Annals of the Missouri Botanical Garden 78:105-130.
Jensen, R. J. 1990. Detecting shape variation in oak leaf morphology: a comparison of rotational-fit
methods. American Journal of Botany 77:1279-1293.
Johnson, Kirk R. 1989. A high resolution megafloral biostratigraphy spanning the Cretaceous-Tertiary
boundary in the northern Great Plains. Unpublished Ph.D. dissertation, Yale University.
Johnson, K. R. 1992. Leaf-fossil evidence for extensive floral extinction at the Cretaceous-Tertiary
boundary, North Dakota, USA. Cretaceous Research 13:91-117.
Little, John R., and Jones, C. Eugene 1980. A Dictionary of Botany.
Manchester, S. R. 1986. Vegetative and reproductive morphology of an extinct plane tree (Platanaceae)
from the Eocene of western North America. Botanical Gazette 147:200-226.
Melville, R. 1937. The accurate definition of leaf shapes by rectangular coordinates. Annals of Botany
1:673-679.
Melville, R. 1976. The terminology of leaf architecture. Taxon 25:549-561.
Merrill, E. K. 1978. Comparison of mature leaf architecture of three types in Sorbus L (Rosaceae).
Botanical Gazette 139:447-453.
Mouton, J. A. 1966. Sur la systematique foliaire en paleobotanique. Bulletin de la Société Botanique de
France 113:492-502.
Mouton, J. A. 1967. Architecture de la nervation foliaire. Congres national des sociétés savantes 92:165-
176.
Ray, T. S. 1992. Landmark eigenshape analysis: homologous contours: leaf shape in Syngonium
(Araceae). American Journal of Botany 79:69-76.
Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Clarendon Press, Oxford.
Spicer, R. A. 1986. Pectinal veins: a new concept in terminology for the description of dicotyledonous
leaf venation patterns. Botanical Journal of the Linnean Society 93:379-388.
Webb, L.J. 1955. A physiognomic classification of Australian rain forests. Journal of Ecology 47:551-
570.
Wilf, P. 1997. When are leaves good thermometers? A new case for leaf margin analysis. Paleobiology
23(3):373-390.
Wilf, P., Wing, S. L., Greenwood, D. R., Greenwood, C. L. 1998. Using fossil leaves as
paleoprecipitation indicators: an Eocene example. Geology 26(3):203-206.
Wolfe, J. A. 1978. A paleobotanical interpretation of Tertiary climates in the northern hemisphere.
American Scientist 66:694-703.
Wolfe, J. A. 1993. A Method of Obtaining Climatic Parameters from Leaf Assemblages. USGS Bulletin
2040.
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Basic Terminology
admedial - toward the midline of the lamina (Fig. 4).
apex - usually the upper ~25% of the lamina (see Character 24).
base - usually the lower ~25% of the lamina (see Character 23).
concave - curving toward the center of the lamina or tooth (Fig. 3). Fig. 1 Simple Leaf
convex - curving away from the center of the lamina or tooth (Fig. 3).
costal vein - primary and secondary veins that extend from the base of
the leaf or from a primary toward the leaf margin.
lamina (blade) - the expanded, flat part of a leaf or leaflet (Fig. 1).
midvein - medial primary, in pinnate leaves this is the only primary. Fig. 2
node - the place where a leaf is (or was) attached to the axis (stem) (Figs. 1, 2).
primary vein - the widest vein of the leaf and any others of like width and/or
course. Primaries usually originate at or just above the petiole.
Symbolized 1o (Fig. 1, see Section III).
Fig. 3
rachis - the prolongation of the petiole of a pinnately compound leaf
upon which leaflets are attached (Fig. 2).
secondary - the next narrower class of veins after the primary, originating
from the primary or primaries. Symbolized 2o (Fig. 1, see Section III).
tertiary vein - the next narrower class of veins after the secondaries, originating
from the secondaries or primaries. Symbolized 3o (see Section III).
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LIST OF FIELDS (CHARACTERS) and OPTIONS ON PULL-DOWN LISTS
(CHARACTER STATES)
1. MORPHOTYPE NAME – text field 33. 2º VEIN ANGLE – abruptly increasing toward base,more
2. MORPHOTYPE # – text field acute on one side, one pair acute basal secondaries,
3. MAJOR PLANT GROUP – DIC, MON, CON, CYC, smoothly decreasing toward base, smoothly increasing
PTE, SPE, LYC, BRY toward base, two pair acute basal secondaries, uniform
4. ORGAN TYPE – leaf, root, axis, reproductive, seed, fruit 34. INTER-2º VEINS – absent, strong, weak
5. MORPHOTYPER – text field 35. 3º VEIN CATEGORY – alternate percurrent,
6. TYPE LOCALITY # – text field dichotomizing, mixed opp/alt, opposite percurrent,
7. RECORD DATE – fills automatically random reticulate, regular polygonal reticulate
8. PLANT FAMILY – pull-down list is too long to 36. 3º VEIN COURSE – admedially ramified, convex,
reproduce here (>400 family names) exmedially ramified, sinuous, straight
9. CIC (COMPENDIUM INDEX CATEGORY) – see 37. 3º (VEIN) ANGLE TO 1º - acute, obtuse, perpendicular
categories at back of booklet 38. 3º VEIN ANGLE VARIABILITY – decreasing
10. LOCS. (OTHER LOCALITIES) - text field medially, inconsistent, increasing basally, increasing
11. TYPE SPECIMEN # – text field exmedially, uniform
12. MQI – 1, 2, 3, 4, 5 39. 4º VEIN CATEGORY – alternate percurrent,
13. DIAGNOSTIC FEATURES OF MORPHOTYPE - text dichotomizing, opposite percurrent, regular polygonal
field reticulate
14. LEAF ATTACHMENT - alternate, decussate, opposite, 40. 5º VEIN CATEGORY – dichotomizing, regular
whorled polygonal reticulate
15. LEAF ORGANIZATION – palmately compound, 41. AREOLATION – lacking, moderately developed,
pinnately compound, simple, ternate, bipinnate, paxillate, poorly developed, well-developed, 3 to 4
tripinnate sided, 5 or more sided
16. PETIOLE FEATURES – text field, but striations, 42. F.E.V.S – absent, unbranched, 1-branched, 2 or more
pulvinate and base swollen available on pull-down list branched
17. LAMINAR SIZE - leptophyll, nanophyll, microphyll, 43. HIGHEST ORDER - text field
notophyll, mesophyll, macrophyll, megaphyll 44. HIGHEST EXCURRENT - text field
18. LAMINAR SHAPE – elliptic, oblong, obovate, ovate, 45. MARGINAL ULTIMATE (VENATION) – fimbrial
special vein, incomplete loops, looped
19. LAMINAR SYMMETRY – asymmetrical, base 46. LEAF RANK - 1r, 2r, 3r, 4r
asymmetrical, symmetrical 47. # OF ORDERS (OF TEETH) - 1, 2, 3
20. LAMINAR L:W RATIO - text field 48. TEETH/CM - text field
21. BASE ANGLE – acute, obtuse, wide obtuse, circular 49. (TOOTH) SPACING - regular, irregular
22. APEX ANGLE – acute, obtuse, wide obtuse 50. (TOOTH) SHAPE – cv/cv, cv/st, cv/cc, cv/fl, cv/rt, st/cv,
23. BASE SHAPE – complex, concave, concavo-convex, st/st, st/cc, st/fl, st/rt, cc/cv, cc/st, cc/cc, cc/fl, cc/rt, fl/cv,
convex, cordate, cuneate, decurrent, hastate, lobate, fl/st, fl/cc, fl/fl, fl/rt, rt/cv, rt/st, rt/cc, rt/fl, rt/rt
rounded, sagittate, truncate 51. SINUS (SHAPE) – angular, rounded
24. POSITION OF PETIOLAR ATTACHMENT - 52. (TOOTH) APEX – foraminate, mucronate, non-specific
marginal, peltate-central, peltate-eccentric glandular, papillate, setaceous, simple, spherulate, spinose
25. APEX SHAPE – acuminate, complex, convex, 53. TOOTH VENATION - text field
emarginate, lobed, retuse, rounded, straight, truncate 54. LEAF TEXTURE – chartaceous w/ cuticle, chartaceous
26. MARGIN TYPE – crenate, dentate, entire, erose, w/o cuticle, coriaceous w/ cuticle, coriaceous w/o cuticle,
revolute, serrate membranaceous w/ cuticle, membranaceous w/o cuticle,
27. LOBATION – unlobed, bilobed, palmately lobed, not apparent
pinnately lobed. 55. STOMATA – actinocytic, amphianisocytic,
28. 1° VEIN CATEGORY – basal acrodromous, basal amphibrachyparacytic, amphibrachyparatetracytic,
actinodromous, campylodromous, flabellate, amphicyclocytic, amphidiacytic, amphiparacytic,
palinactinodromous, parallelodromous, pinnate, suprabasal amphiparatetracytic, amphipericytic, anisocytic,
acrodromous, suprabasal actinodromous anomocytic, anomotetracytic, axillocytic, brachyparacytic,
29. 2º VEIN CATEGORY – basal acrodromous, brachyparahexacytic, brachyparatetracytic, coaxillocytic,
brochidodromous, cladodromous, craspedodromous, copericytic, copolocytic, cyclocytic, desmocytic, diacytic,
eucamptodromous, festooned brochidodromous, festooned hemiparacytic, hexacytic, paracytic, parahexacytic,
semicraspedodromous, interior, intramarginal vein, paratetracytic, pericytic, polocytic, polycytic, staurocytic,
reticulodromous, semicraspedodromous, suprabasal tetracytic
acrodromous, weak brochidodromous 56. (CUTICULAR) FEATURES – hair bases, multicellular
30. AGROPHIC VEINS – compound, none, simple hairs, papillae, peltate hairs, simple hairs, stellate hairs,
31. # OF BASAL VEINS – enter a number striations, thickened areas, trichomes, unicellular hairs
32. 2º VEIN SPACING – decreasing toward base, increasing 57. PHOTO
toward base, irregular, uniform
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Fig. 5
GENERAL INFORMATION MORPHOTYPE NAME MORPHOTYPE #
MAJOR PLANT GROUP ORGAN TYPE MORPHOTYPER TYPE LOC. # RECORD DATE
PLANT FAMILY CIC L
O
TYPE SPEC. # MQI
C
S.
DIAGNOSTIC FEATURES OF MORPHOTYPE:
LEAF TEXTURE
C STOMATA
U
T
I
FEATURES
C
L
E
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Section I: Catalog information
The first section in the database contains basic information about the morphotype and where it
was found.
1. MORPHOTYPE NAME
Scientific binomial (valid or invalid) or nickname.
2. MORPHOTYPE #
Number assigned to the morphotype. This number consists of a two letter prefix, which is
usually an abbreviation for a stratigraphic unit or research area, followed by a number. (For
example: “FU37” would be the 37th morphotype designated for the Fort Union formation.)
5. MORPHOTYPER
The name of the person describing the morphotype.
6. TYPE LOC. #
Museum or personal locality number where the holomorphotype was found. Use identifying
initials for institution or collection.
7. RECORD DATE
The date the morphotype record was created.
8. PLANT FAMILY
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9. CIC
Compendium Index Categories. These categories are used in the North American Paleobotany
Compendium Index of Fossil Plants at Yale University to sort the major plant groups into mor-
phological groups. CICs exist for all dicot leaves and may be used as a device to sort a fossil
flora. (This database has room for up to 4 CIC entries per morphotype.) Look up this number by
using the key in Appendix A.
10. LOCS.
12. MQI
Extremely well preserved means that the fossil has at least fifth order veins.
Well preserved means that the fossil has at least fourth order veins.
Poorly preserved means that the fossil has less than fourth order veins.
Complete means that the fossil has an apex, base and greater than 1/2 the margin.
This field is used to state the characteristics of the morphotype that distinguish it from other
leaves at the same locality or in the same formation. This field is also useful for describing
features that don’t conform to the categories in the form.
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Section II: Leaf
Description of the shape, size and organization of the leaf.
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15. LEAF ORGANIZATION
petiolule
petiole
sessile petiolulate
Chorisia insignis (Bombacaceae)
Fig. 15.2a Fig. 15.2b
palmately compound - a leaf
with separate subunits (leaflets)
attached at the apex of a petiole.
Fig. 15.1 Fig. 15.3
simple - consisting of a ternate (trifoliate) - a compound
single lamina. leaf with three leaflets.
petiolule
rachis
rachis
rachis
petiolule
petiole petiole
rachilla
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16. PETIOLE FEATURES
Note distinctive features of the petiole (e.g., width, length, base swollen, base inflated, sessile or
other).
The following chart shows the ranges of areas for the different leaf classes (Webb 1955).
The two entries in the database field should contain the minimum and maximum size categories
observed for the morphotype.
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Fig. 6
3:1
Notophyll
2:1
1.5:1
1 cm
1:1
Microphyll
Mesophyll
Choose the size
of figure that
the leaf fits into
completely.
Nanophyll
Leptophyll
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18. LAMINAR SHAPE
The simplest way to describe the overall shape of the lamina is to locate the axis or, in some
cases, the zone of greatest width that lies perpendicular to the axis of greatest length (long axis):
- 18 -
19. LAMINAR SYMMETRY
Measure the length of the lamina (L - see Fig. 7) and divide this number by the width of the
lamina. Report the full range of ratios (e.g., 3:1 - 6:1).
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DEFINITIONS
Midvein length, lm = distance from proximal most to the distal most point of the midvein (Figs. 7a-7d).
Apical extension length, la = distance on a perpendicular from the distal most point of the midvein to the
distal most extension of leaf tissue (Figs. 7c, 7d). Can equal zero (Figs. 7a, 7b).
Basal extension length, lb = distance on a perpendicular from the proximal most point of the midvein to
the proximal most extension of leaf tissue (Figs. 7b, 7d). Can equal zero (Figs. 7a, 7c).
Leaf Length, L = lm + la + lb
Mucronate – apex terminating in a sharp point that is the continuation of the midvein. Character goes
in diagnostic features field if observed.
la la
L = lm
lm L lm L lm L
lb lb
Fig. 7a Fig. 7b Fig. 7c Fig. 7d
The vertex of the base angle lies in the center of the petiole at the point where the basal most
laminar tissue touches the petiole. Base angle is the angle from the vertex to the points where a
line perpendicular to the midvein at 0.25lm from the base intersects the margin (Fig. 21.1, 21.2).
In leaves with a basal extension (lb>0), the base angle should be measured from the same vertex
point to the basal most points of the leaf on each side (Fig. 21.3). The base angle is always mea-
sured on the apical side of the rays even in leaves where the angle is greater than 180°. Peltate
leaves are defined as having a circular angle.
0.25lm
0.25lm
vertex
Schumacheria castaneafolia (Dilleniaceae) Aristotelia racemosa (Elaeocarpaceae) Asarum europaeum (Aristolochiaceae)
Fig. 21.1 Fig. 21.2 Fig. 21.3
acute - base angle <90o. obtuse - base angle >90o. wide obtuse - a special case of
obtuse such that the base angle is
>180o.
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22. APEX ANGLE
Apex angle is the angle from the apical termination of the midvein to the pair of points where a
line perpendicular to the midvein and 0.75lm from the base intersects the margin (Fig. 22.1, 22.2).
In leaves with an odd number of lobes, measure the apex angle as in unlobed leaves (Fig. 22.4,
Fig. 22.5). In leaves with an apical extension (la>0) the apex angle should be measured using the
termination of the midvein as the vertex, and the apices of the lobes on either side (Fig. 22.3). The
apical angle is always measured on the basal side of the rays, even in leaves where the angle is
greater than 180°.
0.75lm 0.75lm
0.75lm 0.75lm
- 21 -
23. BASE SHAPE
These states apply to the basal 25% of the lamina (0 - 0.25L as in Fig. 7).
- 22 -
23. BASE SHAPE CONTINUED
petiole attachment
- 23 -
25. APEX SHAPE
These states apply to the apical 25% of the lamina (0.75L - 1L as in Fig. 7).
- 24 -
26. MARGIN TYPE
DEFINITIONS
TEETH are marginal projections with sinuses indented less than 1/4 of the distance to the
midvein or long axis of the leaf. Teeth can be either dentate, serrate or crenate.
Note: If there is a single tooth of any size, the leaf is considered to be toothed.
27. LOBATION
LOBES are marginal indentations that reach 1/4 or more of the distance to the midvein, measured
parallel to the axis of symmetry of the lobe.
Gouania longispicata (Rhamnaceae) Liriodendrites bradacii Dioscoreophyllum strigosum (Menispermaceae) Stenocarpus sinuatus (Proteaceae)
- 25 -
Section III: Vein Orders
The first step in describing the pattern of venation in a leaf is to recognize discrete categories
or orders of veins that have similar widths and courses. Most angiosperm leaves have between
four and seven orders of venation. The first step in describing venation is to recognize the first
three orders of veins. In general, the primary and secondary veins are the major structural veins of
the leaf, while the tertiary veins are the largest veins that fill the field of the leaf. The primary vein
or veins are somewhat analogous to the main trunk or trunks of a tree--they are the widest veins,
they usually taper along their length, and they generally run from at, or near, the base of the leaf to
the margin. Secondary veins are analogous to the major limbs of a tree. They are the next set in
width after the primary(s), they also usually taper along their course, and they ordinarily run from
either the base of the leaf or from a primary vein toward the margin. For tertiary and higher order
veins the analogy with the branching system of a tree breaks down. Tertiary veins are usually
considerably narrower than the secondary set and have courses that connect primary and second-
ary veins to one another in a similar fashion throughout the leaf. Tertiaries are usually the widest
veins that form a more or less organized “field” over the great majority of the leaf area. Generally
it is fairly easy to recognize the primaries and tertiaries, but sometimes the secondaries consist of
several subsets with different widths and courses. Nevertheless, all the subsets of veins between
the primaries and the tertiaries are considered to be secondaries.
After the three lowest vein orders have been demarcated, the observer can proceed to discrimi-
nating the higher orders of venation (4-7) present in the leaf. Each of these higher vein orders can
be highly variable among species and higher taxa in its degree of distinctness from both the next
higher, and the next lower vein order. Good diagnostic features for distinguishing higher vein
orders from one another are excurrent origin from their source veins and a distinctly narrower
gauge. If they arise dichotomously or appear to be of the same, or nearly the same, width as their
parent vein, they are of the same order as the source vein.
Obviously the simultaneous use of two criteria for the determination of vein order introduces a
degree of ambiguity into the process because some veins may have the width typical of one vein
order but the course typical of a different vein order. However, recognizing orders based solely on
their width or solely on their course leads to illogical situations where veins that appear to have
different functions and developmental origins are assigned to the same order. Assigning veins to
orders also has a somewhat arbitrary aspect because variation in width and course is not discrete -
for example, a vein may be intermediate in width between the primary vein and the secondary
veins. However, there do appear to be natural breaks in the variation in width and course, so that
most veins can be assigned to an order unambiguously. In our experience, vein orders can usually
be defined in a repeatable manner for a given leaf by different observers who follow a consistent
set of rules.
Leaves with veins that form a high number of discrete orders or that have regular courses, are
considered to be more organized or “higher rank” leaves. The concept of leaf rank is discussed
and illustrated in Character 46. Figures 8 and 9 demonstrate designation of vein orders for two
leaves.
- 26 -
Vein orders continued
General rule: All vein orders should be recognized in sequence from lowest to highest. The
sole and rare exception is that some leaves with extremely acrodromous primary veins may lack
secondaries (Fig. 28.6). To recognize the primary, secondary and tertiary veins, take the
following steps.
1. Find the widest vein(s) in the leaf; this is the primary vein. Most leaves have a single primary
vein and are called pinnate (if so, go to step 3). If more than one vein originates at or near the
base of the leaf, then proceed to step 2 to determine if the leaf has one or more primary veins.
2. After recognizing the widest single vein of the leaf as a primary (generally the midvein), other
primaries are recognized by being 74% or more of the width of the the widest primary (at the
point of origin of the widest primary). These veins are basal or nearly basal. If these veins
enter lateral lobes or run in strong arches towards the apex, they are generally easily
recognized as primaries. But if the lateral primaries curve toward the midline apically (Fig.
28.6) or branch toward the margin (Fig. 28.3), they may be hard to distinguish from
secondaries. In pinnatifid leaves, primaries may be difficult to distinguish from costal
secondary veins.
If there is more than one primary vein (based on vein width) other veins originating at the
base may be considered primaries if their course and function is similar to that of the
previously defined primaries, even if their width falls into the range of 25-75% of the widest
primary vein. The width of these may fall within the width range of the secondary or tertiary
veins. If these veins are narrower than 25% of the widest primary vein, they are not
considered primaries.
3. Find the widest veins that fill the field of the leaf; these are the tertiary veins (refer to
Character 35, 3° Vein Category). Proceed to step 4. (Watch out for rare exceptions such as
Clusiaceae where secondary veins fill the field of the leaves.)
4. Having recognized the limits of the primary and tertiary vein sets, identify the intermediate
set. These veins are secondary veins and may consist of costals (the rib forming veins that
originate on the primary and run to the margin), interior secondaries, intersecondaries, outer
secondaries, and intramarginal veins (refer to Character 29, 2° Vein Category). The
secondaries will fall within a smooth continuum of width and behavior. Proceed to step 5.
As noted above and illustrated in Figure 37.1, secondaries may be absent rarely.
5. Once you have recognized the first three orders of venation, proceed in sequence to determine
the higher orders venation using the criteria of vein width and course.
- 27 -
3o 4o
3o 2o 1o 5o
○
2o ○
○
○
○
○
○
1o 1o
○
2o
1o 3o 2o
○
o
2 ○
○
○
○
○
4o 1o 2o 4o 3o
○
○
○
○
○
○
1o ○
○
○
○
○
○
○
○
○
○
○
○
○
6o
○
○
5o
○
○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○
○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○ ○
○
- 28 -
28. 1° VEIN CATEGORY
- 29 -
29. 2° VEIN CATEGORY
- 30 -
29. 2° VEIN CATEGORY CONTINUED
- 31 -
30. AGROPHIC VEINS
Agrophic - a comb-like complex of veins comprised of a lateral 1° or 2° backbone with 2 or more
exmedial 2° veins that travel roughly parallel courses towards the margin. These veins may be
straight or looped. Agrophic veins are similar to pectinal veins defined by Spicer (1986).
- 32 -
32. 2° VEIN SPACING
- 33 -
33. 2° VEIN ANGLE
- 34 -
34. INTER-2° VEINS
Intersecondary veins have a width and course similar to the 2°s, but they are usually thinner than
the costal 2°s and do not reach the margin.
- 35 -
35. 3° VEIN CATEGORY
- 36 -
36. 3° VEIN COURSE
2° 1° 2° 1° 2° 1°
3° 3°
3°
- 37 -
37. 3° (VEIN) ANGLE TO 1°
1° 3° 2° 1° 2°
3°
3°
- 38 -
38. 3° VEIN ANGLE VARIABILITY
Fig. 38.1
- 39 -
39. 4° VEIN CATEGORY
Fourth and higher order venational characters should be scored in the portion of the leaf that is
roughly half way between the base and the apex unless the area is lacking.
- 40 -
40. 5° VEIN CATEGORY
- 41 -
41. AREOLATION
Areoles are the smallest areas of the leaf tissue surrounded by veins; taken together they form a
contiguous field over most of the area of the lamina. Any order of venation can form one or more
sides of an areole.
- 42 -
42. F. E. V. S
“FEVs” are the freely ending ultimate veins of the leaf. The two database fields should contain
the extreme states observed.
Fig. 42.1 Fig. 42.2 Fig. 42.3 Fig. 42.4 Fig. 42.5 Fig. 42.6
Highest vein order showing excurrent branching; that is, having true lateral branches rather
than those produced by forking of the vein.
- 43 -
45. MARGINAL ULTIMATE (VENATION)
Eucryphia glandulosa (Eucryphiaceae) Carissa bispinosa (Apocynaceae) Mollinedia floribunda (Monimiaceae) Picramnia krukovic (Simaroubaceae)
Fig. 45.1 Fig. 45.2 Fig. 45.3 Fig. 45.4
teeth no teeth teeth no teeth
incomplete loops - freely ending veinlets looped - marginal ultimate vein recurved
adjacent to the margin. to form loops.
leaf margin
leaf margin
fimbrial vein
fimbrial vein
- 44 -
46. LEAF RANK
Leaf rank is a semiquantitative description of the regularity of the leaf’s vein system, from an
arbitrary level of 1r for the lowest rank or level of organization to 4r for the highest. The rank
number corresponds to the highest order of veins that is well organized. The table on the next
page gives the characters that define the ranks.
1r 2r
~3X ~3X
2° 2°
1°
~7X ~7 X
~1X ~1X
3r 4r
~3X ~3X
2° 2°
~7X ~7X
~1X ~1X
- 45 -
Elements 1r 2r 3r 4r
intercostal area shapes vary shapes similar shapes similar shapes similar
Fig. 46.5
- 46 -
Section 4: Teeth
DEFINITIONS
(1°, 2° or 3°) If the teeth can be separated into different size groups, they are called compound.
1° tooth
1° tooth
1° tooth
2° tooth
2° tooth
3° tooth
The number of teeth/cm in the This refers to the interval between corresponding
middle 50% of the leaf. points on the teeth or crenations.
cm
4/cm 2
- 47 -
50. (TOOTH) SHAPE
Tooth shape is described in terms of the shape of the apical side and the basal side. The possible
combinations are shown in the chart below. In the database, the following abbreviations are used:
cv (convex) st (straight) cc (concave) fl (flexuous) rt (retroflexed)
basally convex and apically convex and
apically concave basally concave
The apical shape is listed first. For example, cc/fl would be concave on the apical side and
flexuous on the basal side of the tooth. Note that a given leaf can exhibit more than one tooth
shape.
APICAL SIDE
B
A
S
A
L
S
I
D
E
Fig. 50.1
- 48 -
51. SINUS (SHAPE)
- 49 -
52. (TOOTH) APEX
There are three major types of tooth apex: simple, spinose, and glandular. In living leaves and
some fossils, it may be possible to distinguish the following subsets of glandular: spherulate,
papillate, foraminate, mucronate, and setaceous. For situations in which a more specific identifica-
tion is not possible, use non-specific glandular.
- 50 -
53. TOOTH VENATION
This describes the venation that is associated with the tooth. The principal vein is the thickest
vein entering the tooth. Other veins in the tooth are accessory veins. Describe their characteristic
arrangement.
Section 5: Cuticle
Leaf texture is difficult to compare between localities. This is a relative scale for leaves
preserved in a similar rock type.
membranaceous w/cuticle - compression appears to be very thin compared with other leaf types
preserved in the same matrix; cuticle present.
membranaceous w/o cuticle - compression appears very thin compared with other leaf types
preserved in the same matrix.
chartaceous w/cuticle - compression appears moderately thin compared with other leaf types
preserved in the same matrix; cuticle present.
chartaceous w/o cuticle - compression appears moderately thin compared with other leaf types
preserved in the same matrix.
coriaceous w/ cuticle - compression appears thick compared with other leaf types preserved in
the same matrix; cuticle present.
coriaceous w/o cuticle - compression appears thick compared with other leaf types preserved in
the same matrix.
- 51 -
55. STOMATA
Figures 55.1 - 56.3 are reprinted with permission from Botanical Review, vol. 40, no. 1,
copyright 1974, The New York Botanical Garden, Approaches to the identification of
angiosperm leaf remains by David Dilcher.
Anatomy of
Stomata: T-piece at
stomatal pole
stomatal aperture
poral
epidermal
} walls of
guard-cells
walls of
outer stomatal ledge
radial
tangential
} subsidiary
cells
Fig. 55.1
- 52 -
anisocytic types - 3 cells, may be unequal in size, enclosing the guard cell.
amphianisocytic - double
anisocytic - single ring of cells enclosing the
ring of 3 cells (2 guard cells with the inner
larger, one smaller) ring consisting of 3 cells (2
enclosing the guard larger, one smaller); outer
cells. ring may be incomplete
consisting of 2-3 or 4 cells.
diacytic types - 2 cells enclsing the guard cells at right angles to the long axis of guard cells.
paracytic types - 1 or 2 cells adjacent to the guard cells with their long axis parallel to
the long axis of the guard cells.
- 53 -
paracytic types continued
brachyparatetracytic - 2 amphibrachyparatetracytic -
short cells lateral and parallel 2 short cells lateral and parallel
to the guard cells, 2 wide polar to the guard cells, 2 wide polar
cells. cells, all of which is sur-
rounded by a ring of small
cells.
- 54 -
hexacytic types - 4 cells adjacent to the guard cells with 2 additional (lateral or polar)cells which
can be distinguished from the epidermal cells.
Fig. 55.25
amphipericytic - one cell enclosing both
guard cells enclosed by a second single cell.
- 55 -
polocytic types - one cell nearly but not completely enclosing the two guard cells.
- 56 -
56. (CUTICULAR) FEATURES
hair bases
trichomes
unicellular hair
multicellular hairs
peltate hairs
simple hairs
stellate hairs
- 57 -
APPENDIX A
COMPENDIUM INDEX OF NORTH AMERICAN FOSSIL PLANTS
First Field
1— ANGIOSPERMS
Leaves with several orders of venation, cross-veins and vein anastamoses at
several orders.
- 58 -
125 Lamina pinnately veined, with pectinal vein, unlobed, elliptic or oblong, toothless
126 Lamina pinnately veined, with pectinal vein, unlobed, ovate, toothed
127 Lamina pinnately veined, with pectinal vein, unlobed, ovate, toothless
128 Lamina pinnately veined, with pectinal vein, unlobed, obovate
129 Lamina acrodromously veined, elliptic or oblong, toothless
130 Lamina acrodromously veined, elliptic or oblong, toothless
131 Lamina acrodromously veined, ovate, toothed
132 Lamina acrodromously veined, ovate, toothless
133 Lamina acrodromously veined, obovate
134 Lamina actino- or palinactinodromously veined, unlobed, elliptic, or oblong, toothed
135 Lamina actino- or palinactinodromously veined, unlobed, elliptic or oblong, toothless
136 Lamina actino- or palinactinodromously veined, unlobed, ovate, toothed
137 Lamina actino- or palinactinodromously veined, unlobed, ovate, toothless
138 Lamina actino- or palinactinodromously veined, unlobed, obovate
139 Lamina actino- or palinactinodromously veined, 2-lobed or lobes in multiples of 2
140 Lamina actino- or palinactinodromously veined, 3-lobed
141 Lamina actino- or palinactinodromously veined, 5 or more lobes
142 Lamina definitely palinactinodromously veined, 3 lobed
143 Lamina definitely palinactinodromously veined, 5 or more lobes
144 Lamina campylodromously veined
145 Lamina flabellately veined, very weakly pinnately or palmately veined or multistranded
midvein
146 Lamina flat and unlobed, veins parallelodromous, pinnately attached to a costa
147 Lamina flat and unlobed, veins parallelodromous from a zone at the blade base
148 Lamina plicate or breaking into narrow-segments, venation parallelodromous, leaf shape and
vein origin unknown
149 Lamina plicate and lobed, fan-shaped, venation palmate
150 Lamina plicate and lobed, feather-shaped, venation pinnate
- 59 -
Other Organs
170 Flowers occurring as single units
171 Flowers aggregated into catkins or aments
172 Flowers aggregated into heads or capitulas
180 Fruits, dry,indehiscent, seed-containing portion relatively small (generally <5mm) or, if
winged, the winged portion exceeding the size of the seed (achenes, caroyapsis, utricles,
cypselas, samaras, etc.)
181 Fruits, dry, indehiscent, large (>5mm) or, if winged, the winged portion smaller than the seed
bearing portion (acorns, balaustas, calybiums, nuts)
182 Fruits, dry, dehiscent capsules, follicules, or siliques
183 Fruits, dry dehiscent legumes or loments
184 Fruits, fleshy (berries, drupes, pomes, etc.)
185 Fruits, aggregate or multiple
186 Fruits, other - or of indeterminate characters
190 Wood or stems
2— GYMNOSPERMS
200 Pteridosperms (including Caytoniales)
210 Cycadophytes,leaves dissected, toothless, veins parallel except convergent at pinna apex and
base, mainly forked
211 Cycadophytes,leaves dissected, toothless, veins parallel except convergent at pinnule apex
and base, mainly unforked, pinnules <3cm long
212 Cycadophytes, leaves dissected, toothless veins parallel except convergent at pinnule apex
and base, mainly unforked, pinnules >3cm long
213 Cycadophytes, leaves dissected, toothless, veins pinnate or radiating throughout length of
pinnule
214 Cycadophytes, leaves dissected, pinnules toothed
215 Cycadophytes, leaves undissected, veins parallel, unforked
216 Cycadophytes, leaves undissected, veins parallel, forked
217 Cycadophyte leaves, habit indeterminable
218 Cycadophyte seeds, cones, and “flowers”
219 Cycadophyte stems and wood
220 Ginkgophytes, leaves fan-shaped, veins flabellate, includes the Neoggerathiales,
Czekanowskiales
230 Conifers, scaley foliage, leaves appressed to stem for more than 1/2 of their length
231 Conifers, short needles: average <3cm
232 Conifers, long needles: average >3cm
233 Conifers, leafy blades 1<3cm, 1/w ratio >10:1 or 1>3cm l/w ratio 1.5 or less
234 Conifers, cones
235 Conifers, cone scales
236 Conifers, seeds
237 Coniferophytes, wood
238 Conifers, characters uncertain
240 Gnetophytes
- 60 -
300 ALGAE
350 FUNGI
400 BRYOPHYTES
5— FERNS
500 Blades dissected, veins open, ultimate laminar division w/o midribs
501 Blades dissected, w/ midribs, veins unforked
502 Blades dissected, veins closed, ultimate laminar divisions w/ midribs, veins forked
503 Blades dissected, veins closed, ultimate laminar divisions w/o midribs
504 Blades dissected, veins closed, ultimate laminar divisions with midribs
505 Blades undissected
506 Venation obscure or uncertain
507 Specialized fertile pinnae, fertile part much exceeding sterile tissue in at least a part of the
leaf
508 Fern stems and rhizomes
509 Fragments too small to determine
600 SPHENOPSIDS
7— LYCOPSIDS
700 Lycopodium and Selaginella
710 Isoetales
- 61 -
- 62 -
- 63 -
- 64 -
- 65 -
- 66 -
- 67 -