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A modi f i ed t ermi nol ogy f or
angi osperm l eaf archi t ect ure
Mi ke Pol e
a

b
a
Depar t ment of Geol ogy, Uni ver si t y of Ot ago, P. O. Box 56,
Dunedi n, New Zeal and
b
Depar t ment of Pl ant Sci ence, Uni ver si t y of Tasmani a, GPO
Box 252C, Hobar t , 7001, Aust r al i a
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Jour nal of t he Royal Soci et y of New Zeal and, 21: 4, 297- 312
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Journal of the Royal Society of New Zealand.
Volume 21. Number 4, December 1991, pp 297-312
A modified terminology for angiosperm leaf architecture
Mike Pole*
Amodifiedterminologyfor describingleaf architectureis presented,whichincorporates
lamina development and gives greater emphasis to the patterns formed by groups of
venationelements than to their size. Leaf architectureis described usinga hierarchyof
venation "elements", which are grouped into "patterns", which are located in "zones"
and "segments" of the leaf lamina.
Keywords: macropaleobotany. leaf venation, leaf ontogeny, constructional morphology
INTRODUCTION
In the study of angiosperm leaf impressions within paleobotany it soon becomes apparent
that there is a need to develop a comprehensive terminology to describe leaf "architecture".
This term was defined by Hickey (1979: 25) as
"... the position and form of the elements constituting the outward expression of leaf
structure. These include venation pattern, marginal configuration, leaf shape, and gland
position. Architecture is the aspectof morphologywhichappliesto the spatialconfiguration
and coordinationof those elements making up part of a plant without regard to histology,
function, origin, or homology."
Leaf fossils are usually found as impressions only, with no anatomical detail, and the
outline of the leaf and the pattern of lines left by the venation on fossilisation is all one has to
work with. An understanding of venation, its formation and the taxonomic distribution of
different venation types and patterns, is then very important (a taxonomic survey of venation
patterns based on the modifications presented here is beyond the scope ofthis paper, but is in
progress and will be presented later). The addition of anatomical information, such as
cuticular structure and stomatal distribution, adds a new range of characters and requires
further terminology.
The terminology of leaf architecture must develop to a point where it can allow plants
known only from leaf impressions to be integrated into the accepted Linnean-style taxonomic
edifice. I propose a modified leaf architecture terminology which I suggest is an improvement
on all existing systems.
DEFICIENCIES OF EXISTING LEAF ARCHITECTURE TERMINOLOGY
Several different terminologies have been devised for describing leaf architecture, of
which the most comprehensive is that presented by Hickey (1973,1979) and Dilcher (1974).
Melville (1976) objected to this terminology because it is restricted to dicotyledons and also
because its basis is in Greek rather than Latin. He presented a new terminology as a
substitute. Melville's criticisms were answered by Hickey (1979) and his terminology is not
considered further here.
The terminology of Hickey (1973, 1979) and Dilcher (1974) has as its cornerstone the
recognition of vein orders, i.e.
* Department of Geology, Universityof Otago, P.O. Box 56, Dunedin, NewZealand.
Present address: Department of Plant Science, Universityof Tasmania, GPOBox 252C, Hobart 7001,
Australia.
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298 Journal of the Royal Society ofNew Zealand, Volume 21,1991
"... the recognition of vein orders is essential in describing leaf architecture ... "
and
"... The fundamental rule for the determination of the order of a vein is its relative size at
its point of origin" (Hickey, 1979: 32).
The patterns formed by the venation have secondary importance in Hickey's scheme, i.e.
.".. Subsidiary to the rule of size at the point of origin in the recognition of vein order is a
consideration of the behaviour of a vein in relation to veins of other orders and to marginal
features of the leaf blade" (Hickey 1979: 32).
I agree with Hickey on the importance of establishing vein orders, but believe that his use
of size at the point of origin to determine the orders introduces a serious problem into his
terminology. Hickey himself (1979) noted that important ambiguities may arise when applying
these rules, for instance in distinguishing a "lateral primary vein" from a "basal secondary
vein". If such ambiguity arises, it may not be possible to ordinate veins to at least the sixth
order, as Hickey does. Inan ideal terminology one needs to know that something referred to
as a "secondary" vein in one leaf is homologous to a "secondary" in another. In order to
reveal phylogenetic relationships, a terminology must incorporate knowledge of these
homologies.
Thickness of veins at the point of origin is surely a variable thing, depending on the extent
of growth. Troll (1935) and Arber (1950) both refer to the effects on venation of differing
growth rates over the lamina, even to the point where lateral veins may dominate over the
midrib (see below). It is the behaviour of the veins which expresses their identity, not their
thickness.
Spicer (1986) criticised Hickey's terminology, including the intergradation of pinnate and
palmate organisation and the difficulty of distinguishing between primary and secondary
veins. His solution places emphasis (p. 379) "... upon the pattern of vein courses and
branching hierarchy which provides unambiguous reference points 00'" I agree with his
comments, but disagree with the details of his solution. Spicer proposed the term "pectinal
veins" for lateral veins which
'''00 differ from the main secondaries in that they produce abmedial veins almost throughout
their length and a greater number of abmedial branches than any other laterals" (Spicer
1986: 382).
In his scheme only one pair of pectinals is recognised in anyone leaf, and this could be
used as a point of reference. 1 do not accept that Spicer's "pectinal" veins are significantly
different from other lateral veins, and believe that they result only from a greater amount of
progressive, or ramifying, growth (see below). Often there is a complete gradation apically in
the number of abmedial branches on a lateral, but sometimes there is not. To single out the
laterals with the most branches on them is similar to singling out those which are thickest as
primaries. Both phenomena result from a localised increase in development (the summary of
Arber's comments below is relevant to this discussion) furthermore, Spicer's terminology
does not deal with abmedial branches on his "superior secondaries".
Differential development of the lamina results in some end-member conditions which
Hickey and Dilcher have included along with venation patterns in their terminology. In leaf
architecture, "venation pattern" refers to the interrelationships between veins or between
veins and the margin. The amount of lamina development between veins, or the thickness of
the veins themselves, is a separate phenomenon and should not be incorporated in a
classification of venation pattern.
Careful consideration should be made before slotting features into a terminology. For
instance, study of many leaves, both fossil and extant, has revealed that the "loops" of
venation, usually referred to as "brochidodrornous" (e.g. by Hickey, 1979), are in fact almost
always composite structures, formed of several elements. Brochidodromous in the sense of a
loop formed by the joining of just two elements may be rare.
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Pole - Modified leaf architecture terminology 299
WAYS OF LOOKING AT VENATION
The underlying principle of my approach is to look at leaf venation as a coherent
configuration or set of patterns. Emphasis should be placed on describing the patterns formed
by groups of elements rather than the behaviour of individual elements. I take the stance that
features of venation, or other aspects of leaf architecture, can be labelled and have taxonomic
or phylogenetic importance. They are comparable with the various diagnostic features in
bones used in osteology or vertebrate zoology.
ONTOGENY OF VENATION
Looking at leaves from a developmental point of view may provide a conceptual tool for
ordering the patterns of leaf venation and allow the construction of a comprehensive
terminology. Once the patterns are understood, homologies may be postulated.
Leaf development in dicotyledons (Esau, 1965; Pray, 1955, 1963; Slade, 1957, 1959)
begins with the progressive formation of the midrib during apical growth, controlled by the
apical meristem. Laterals develop progressively away from the midrib during marginal
growth, controlled by the marginal meristems. Lateral veins continue to be formed until the
end of apical growth. In plants with well-developed areolation, higher-order veins develop
both simultaneously and progressively during intercalary growth, controlled by the intercalary
meristem. Vein endings within these areoles appear to differentiate progressively. In plants
with imperfect or poor areolation, all vein orders appear to differentiate progressively. From
the point of view of the terminology developed here, the division of venation into "progressive"
and "simultaneous" may be better expressed as "ramifying" and "non-ramifying". The term
ramifying applies to any vein which can be related to a hierarchical system of branching
within the lamina, while non-ramifying applies to those veins which cross-link ramifying
veins or other non-ramifying veins.
In several stimulating essays on plant form, Arber (1950) discussed leaf architecture. She
advanced the "partial-shoot theory of the leaf' which holds that the shoot is the basic unit "for
the fundamental interpretation of the plant body" and that the leaf itself is a partial shoot. This
contrasts with the concept of root, stem, and leaf as separate units. She states
"... The wholeplant maybe said to consist of a series of shoot generations, togetherwitha
series of root generations; every individual lateral branch is a repetition, modified in
varyingdegrees, of the originalprimaryshoot, whileevery lateral root similarlyrepeatsthe
primaryroot" (Arber, 1950: 77).
And with reference to the leaf
"... thepartial-shoot hasaninherent urgetowards thedevelopment of whole-shoot characters"
(Arber 1950: 78).
With particular relevance to the problem of "lateral primaries", "palmate venation", and
"acrodromous venation" sensu Hickey (1979), Arber writes of the
.".. tendencyfor themembersof eachgenerationtocompetewith, andattemptto dominate,
those of the previous generation" (Arber, 1950: 93).
This results in the phenomenon of "daughter-shoot dominating parent-shoot" and
"ascendancy oflaterals in shoot and leaf'. Leaf examples given include Bauhinia yunnanensis
Franch. and Passiflora capsularis L. var. acutiloba , bi-lobed leaves in which the terminal
pinna or lobe are reduced to points. The leaf of Cercis siliquastrum L. apparently simple, is
surmised to "consist of two lateral pinnae in a state of union, the median pinna being absent".
According to Arber,
"... Palmatevenationmaybe envisagedas a type inwhichthe mainveinis shortenedbythe
suppression of its earlier 'internodes'; the lateralsare thus more importantrelatively to the
midrib than in pinnate leaves, in which the midrib is elongated and dominates its own
branches" (Arber, 1950: 102).
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300 Journal of the Royal Society of New Zealand, Volume 21, 1991
...
-.
-.
Increasing dcvclopmcnt of the looping zone
Apical
lamina
dcvclopmcnt
-.
-+
Mid
lamina
development
-.
Basal
lamina
development
I
Development of externals
-.
Development of externals and counter-externals
Fig. 1 - Schematic leaves showing the effects of differential development of various portions of the
lamina.
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Pole - Modified leaf architecture terminology 301
Examples are given of Campanula rotundiflora L. and Gaya lyallii E.G. Baker, which
show a range of palmate to pinnate venation even within a single plant. According to
Meyerhoff (1952: 31), Troll (1935) also believed that palmate venation resulted from more
rapid development of the basal one or two pairs of secondary veins. Arber (1950) also
pointed out, using TWa tomentosa Moench as an example, the effects of repetition of shoot
generations and symmetry on leaf venation. Because the generations of veins remain
permanently attached to one another and within the constraints of a two dimensional lamina,
the production of successive generations of laterals (external laterals in the present terminology)
takes place along the side of the parent vein remote from the midrib, the abmedial side. Inthe
case of the basal laterals, this is the side on which growth can develop freely. Cordate bases
are the result of extended growth of vein generations rotating about the pivot of the midrib
base.
"Cinnamomum" venation, i.e., two strong basal acrodromous veins extending for about
two-thirds of the length of the lamina with a few closely-spaced laterals near the apex, may
be an example of Arber's "ascendancy of laterals". Alternatively, it may be that the bulk of
leaf expansion during intercalary growth is centered in the basal region of the leaf. This
phenomenon was discussed by Comer (1958) with respect to a brochidodromous-acrodromous
gradation in Ficus. Total reduction of the apical laterals would lead to acrodromous veins,
running all the way to the apex, as in various Myrtaceae and Melastomataceae. Elimination
of externals and of most growth outside the lateral veins by cessation of marginal growth
immediately after initiation of the lateral veins (Carr et al., 1986) would lead to an
"intramarginal" vein of the Eucalyptus type.
Carr et al. (1986) show that two fundamentally different venation patterns may co-exist
within the genus Eucalyptus, and sometimes even within a single species. Most juvenile
leaves of Eucalyptus are brochidodromous, their lateral veins linking to form a "paramarginal
vein", while adult leaves are acrodromous, their "intramarginal veins" ontogenetically distinct
and forming before the finer veins along the midrib. They describe ways in which the two
forms could be distinguished in fully developed leaves. The brochidodromous type is
distinguished by
"... the diminishing thickness of the lateral veins in approaching the leaf margin, as well as
the gaps often found in the pararnarginal vein, where it has failed to be completed by the
thickening of interconnecting minor veins" (Carr et al., 1986: 57).
The acrodromous type is distinguished by the intramarginal veins being
.".. of uniform thickness and as thick or thicker than the primary lateral veins."
They discuss the problem of what ordination to give the intramarginal veins, and conclude
that since they were
" ... independent of the midrib at the base of the lamina ... (they are therefore) ... undoubtedly
primary" (Carr et al., 1986: 59).
Despite this they recommend the use of the non-ordinated term "intramarginal" to describe
them. Non-ordinated terminology is attractive, but not very useful if used to circumvent
problems of homology. Inthe presently proposed terminology, a single midrib is recognised
and the two intramarginal veins are described as first order laterals. I prefer not to use the
term "intramarginal vein" as I feel it leads to semantic problems when there is a significant
amount of venation external to this vein. Instead, such veins will be termed "longitudinal"
(see below).
The terms "outer secondaries" as used by Romero and Dibbern (1985), and "secondary or
tertiary branches" sensu Hickey (1979), refer to craspedodromous venation, mostly when the
laterals terminate in a tooth. However, there seems to be a link between a toothed margin and
craspedodromous venation, and between an entire margin and brochidodromous venation
(van Steenis, 1953). Inthe leaves of several plants, normally dentate and with craspedodromous
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302 Journal of the Royal Society of New Zealand, Volume 21,1991
a
-+
....
Development of a single pair of longitudinal veins by expansion of the
basal interlateral segment
b
-+
-+
Development of several longitudinal veins by expansion of basal interlateral
and basal loop segments
Fig. 2 - Possible development of leaves with single (a), or multiple (b), longitudinal veins from an
original camptodromous condition.
secondaries and externals (eg Nothofagus ), occasionally there is no tooth for these veins to
enter. The result is that the external vein loops back to its "parent" lateral and/or the lateral
vein loops up to the next more apical lateral. This suggests that "outer secondary loops"
(Hickey, 1979) and the loops in "festooned brochidodrornous" leaves are homologous with
external veins. This condition is referred to here as "external veins looped".
The classic work of D' Arcy Thompson (1952) on growth and form has particular relevance
to leaf architecture. He proposed that
"... The rate of growth deserves to be studied as a necessary preliminary to the theoretical
study of form, and the organic form itself is found, mathematically speaking, to be a
function of time ... We might call the form of an organism an event in space-time, and not
merely a configuration in space" (D' Arcy Thompson, 1952: 283).
Figs 1 and 2 illustrate some of the changes in basic leaf architecture resulting from
different directions and amounts of laminar and marginal growth, either ontogenetically or
phylogenetically, and the effect these may have on the venation and form of a leaf. I stress
that these sequences are hypothetical. The terms used are explained below.
A REVISED VENAnON TERMINOLOGY
Unless redefined below, terminology follows Hickey (1973, 1979).
A simple set of working assumptions may be stated to distinguish growth phases in a leaf,
and to incorporate these distinctions in a terminology:
1. Progressive growth is indicated by decreasing thickness along a vein and/or by free
ends.
2. Simultaneous development is indicated by veins of equal width along their length
which terminated against a lower order vein at each end.
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Second Order
External Loop
Third Order
External Loop
Lateral
Int ernal Loop
Basiscoplc
Internal Loop
Firs t Order
External Loop
a
Pole - Modifi ed leaf architecture terminology 303
Firs t Order Lateral
Second Order Lateral
~ Third Order Latera l
Counter-external
\ c ( Joining Vein
;;:s:: Firs t Order External
e:f Interangular Vein
' ~ r - Second Order Exte rnal
Basic Venation Elements
Looping Zone It
IShaded!
Lateral Lone
(Un.hadtdl
Interloop Segment
Iut erlateral Segment
Joining Vein Segment
~ '-Y =r - Interangular Segment
---"
: Interexternal Segment
b Lamina Zones and Segments
Fig. 3 - Schemat ic leaves to illustr ate terminology of (a) basic venat ion clements and (b) laminar zones
and segments.
3. Thicker veins are ont ogenet ically older than thinner ones in the immediate
neighbourhood.
"Ramified" tertiar y veins sensu Hickey (1979) are considered to be a result of progressive
growth , and "percurrent " tertiar ies to be produced by simultaneous growth. There will no
doubt be subjectivity in their recognition. Veins often show some taper, but. for reasons of
their general location and behaviour , one might neverthel ess suspect that they developed
simult aneously. For instance, external veins or second or third order laterals in some leaves
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304 Journal of the Royal Society of New Zealand, Volume 21,1991
may appear to intergrade with non-ramifying veins. Nevertheless, relative vein size should
provide a simple way of estimating the pattern of growth over a leaf lamina. It should be
emphasised that this procedure is an estimation with the purpose of providing a conceptual
tool for describing leaf venation. The actual details of how venation develops must be studied
on growing plants.
HIERARCHY IN LEAF ARCHITECTURE
For ease of comprehension, this revised terminology of leaf venation is presented in a
hierarchical fashion.
A. Elements
The veins themselves are termed "elements" and, depending on their behaviour and size,
can be divided into ramifying or non-ramifying; first, second, third order elements; etc. Non-
ramifying elements are most conveniently discussed as being portions of patterns (Figs 3 and
7).
B. Patterns
The way in which venation elements interact with each other provides "patterns". Depending
on the order of the elements concerned, and their location, these patterns can be divided into
first, second, third order patterns, etc (Figs 6 and 7).
C. Zones and Segments
These both refer to areas of the lamina which are defined by the course of elements and
filled in with patterns. "Zone" is the most general category, and it may be divided into
"segments" (Fig. 3).
BASIC VENATION ELEMENTS (see Fig. 3)
Midrib
Only one primary vein exists per leaf, for which the term "midrib" is used. The midrib is
the central vein of a leaf, acting more or less as an axis of symmetry, and it is the main point
of reference for the following vein terminology. The midrib need not be the strongest vein in
the leaf.
Laterals
Any ramifying veins developed along the midrib, and when applicable, the largest pair at
the midrib base, are referred to as "laterals". The largest of these ("secondaries" sensu
Hickey, 1979) are here termed "first order laterals".
Higher order laterals, i.e. second or third order laterals (here termed "extra-laterals"), refer
to Hickey's "intersecondary veins". Higher order laterals can usually be recognised by their
behaviour as well as by their relative thickness. Often they do not extend across the full
length of the lateral segment, or else join the lateral loops with little effect on them, whereas
first order laterals join at a sinus, or kink. At other times they will be recognised because they
interrupt the normal style of joining veins in a percurrent pattern. Collectively, higher order
lateral veins are termed "extra-laterals".
Sometimes a vein which fits in with a pattern of "interangular veins" (see below) will
progressively thicken as it nears the midrib. It may be difficult to distinguish depending on
how pronounced is the thickening. The two categories may well intergrade.
Depending on the growth pattern along the midrib the spacing between first order laterals
may decrease apically, increase apically, decrease both apically and basally from mid-
lamina, or be relatively constant.
The pattern of higher-order laterals, which form subsequent to the first order laterals,
depends on how the lateral segment expands (Fig. 6c). New higher-order laterals may be
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Pole - Modified leaf architecture terminology 305
lateral Loop formed by
the junction of First
Order Lateral \YUh
Second Order External
1. =first Order External Loop
2. =Second Order External Loop
3. = Third Order External Loop
f. O. E. =First Order.External
S. O. E.. SKOnd Orde r
T. O. E.. Third Ordf'r
Fig. 4 - Annotated venation of the extant Muehlenbeckia australis.<Polygonaceae)showi ng interpretation
of the "loops" as complexes of several venation elements.
intercalated symmetrically as if there were a mirror line running Ihrough the lateral segment
at right angles to the midrib, or they may be initiated only in the basal, or axial, region of the
segment. The two patterns formed are termed "symmetrical" and "axial".
External Veins
All ramifying veins developin g from the basiscopi c side of lateral veins are termed "fi rst
order external veins". External veins normall y develop along the distal parts of the laterals, or
along the complete length of the most basal laterals. Often they will develop from the point
where the lateral connects with the midrib. Successive orders of externals may develop
rotationally from this point. On occasions mid-lamina first order laterals may develop an
external at their junction with the midrib . External veins include the "outer secondaries" of
Romero and Dibbern (1985) and the "pectinal veins" of Spicer ( 1986).
Any vein developing progressively from a first order external vein is a "second order
external vein" , and any vein developing progressively from a second order external vein is a
"third order external vein" (Fig. 4). Ramifying veins developing from the apical side of
lateral veins are called "co unter-external veins", the term used by Meyerhoff (1952) .
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306 Journal of the Royal Society ofNew Zealand, Volume 21,1991
Hyphodrornous I'arallclodromous Craspcdoclroruous M.ixcd
Fig. 5 - Schematic leaves illustrating four kinds of first order venation patterns.
Loops
Loops refer to closed circuits of ramifying venation (Figs. 3a, 4). As mentioned previously,
they are often composite structures formed by the linkage of several elements. The thickest
vein in the loop is the "base" from which the "origins", the next thickest veins, depart. The
opposite end of the loop from the base is the "arch".
a) Lateral loops
These are loops formed by first order lateral veins (the origins) growing to join an adjacent
first order lateral or an external branch of it. The midrib forms the base, and the arch may
include first order lateral, first order external, second order external, and possibly higher
order elements.
b) External loops (Figs. 3 and 4)
1. First-order external loops. These loops have first order externals as their origins and
are based on first order laterals. The arch is frequently a second order external.
2. Second-order external loops. These have second order externals as their origins and
are based on a first order external. The arch is frequently a third order external.
3. Third-order external loops. These have third order externals as their origins and are
based on a second order external.
c) 1nternalloops
These loops are found within the interlateral segments (cf, Fig. 3b) defined by the lateral
loops.
1. Lateral internal loops. These are similar to lateral loops but are formed of higher order
lateral veins, generally second order laterals.
2. Basiscopic internal loops. These are formed by the linking up of external veins,
exactly the same as external loops, but within the lateral loop. Included are second order
lateral veins which loop up to an external.
Loops may not show any particular distortion or may be elongated widthwise (perpendicular
to the midrib), or be elongated lengthwise (parallel to the midrib).
FIRST ORDER VENATION PATTERNS
The "traditional" patterns of ramifying venation are somewhat arbitrary with little taxonomic
significance, and intergradation is common as the distinctions may be due merely to varying
amounts of meristematic growth. The recognition of a single primary vein, or midrib, in the
present terminology requires the redefinition of some of the basic types of venation, as given
in Hickey (1979) where one or more primaries are recognised. These "basic" venation
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Pole - Modified leaf architecture terminology 307
a
l
Externodromous
' ~
~
Eucamptodromous
(lI nkr d by r nl. rgr d joini ng
veins)
I ~
l
Eucamptodromous
IUn krd by j o i n l n ~ . rin51
5
E
Brochidodromous
b
Joining veins from
devel. of N.E.
quadrant.
Cascade venation
from devel, of
S.W. quadrant.
Development of
all quadrants.
Development of
S. E. and S.W.
quadrants.
c
~
l
.....
.....
Development of
symmetrically
placed
extra-laterals
resulting in the
"symmetrical"
pattern
Development of
basally initiated
extra-laterals
resulting in the
"axial" pattern
Fig. 6 - (a) types of the camptodromous first order venation pattern, (b) development of four different
types of second order venation as a result of differential laminar growth (based in stippled areas) from
a basic "quartered" interl ateral segment (modified from Comer, (958), (c) symmetrical growth within
an interl ateral segmen t result ing in the "symmetrical" pattern of extra-laterals (top) and growth in the
axils of first order laterals result ing in the "axial" pattern of ext ra-laterals (bottom).
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308 Journal of the Royal Society of New Zealand, Volume 21, J991
patterns, formed by the midrib and the first order laterals, are the first order patterns of a leaf.
Seven first order venation patterns are recognised.
a) Hyphodromous venation
Used in the sense of Hickey (1979) where all veins but the midrib are absent, rudimentary,
or concealed within a coriaceous or fleshy mesophyll (Fig. 5).
b) Parallelodromous venation
Several veins originating beside each other at the leaf base and running parallel to the apex
where they converge, with no obvious decrease in thickness along their length, and no strong
connecting or joining veins (Fig. 5). [It should be noted that Hickey's illustration (1973, fig.
45) of parallelodromous venation is not consistent with the definition given in his text. In his
illustration the veins do not originate beside each other at the leaf base.]
c) Craspedodromous venation
All first order lateral veins terminate at the margin (Fig. 5). Similar to Hickey (1979).
d) Mixed venation
Some first order lateral veins terminate at the margin and some otherwise (Fig. 5).
e) Camptodromous venation
First-order lateral veins not terminating at the margin. Three basic conditions exist here
(Fig. 6a), but in practice there may be occasions where it is difficult or impossible to
distinguish between them.
1. Brochidodromous venation
This term is reserved for that condition where adjacent first order laterals directly join one
another.
2. Externodromous venation
This is a new term for the condition where a first order lateral joins only to an external
branch of an adjacent first order lateral (see also Fig. 4).
3. Eucamptodromous venation
This is as used by Di1cher(1974) and Hickey (1979) where first order laterals are upturned
and gradually diminish apically with no direct lateral-lateral contact. They are linked to
adjacent laterals by joining veins, and although the laterals diminish to the thickness of
joining veins, there is no break in the pattern of the joining veins. Sometimes a particular
joining vein may be thicker than the rest, forming a "pseudo-lateral loop". It is possible that
such veins are externals ontogenetically.
Hickey's "cladodromous" and "reticulodromous" are rejected. Hickey (1973: 24) applied
these terms respectively to laterals "freely ramified toward the margin", and laterals "losing
their identity toward the leaf margin by repeated branching into a vein reticulum". I believe
that closer observation would indicate an externodromous pattern.
SECOND ORDER VENATION PATTERNS
Second order venation pattern refers to the most basic pattern formed in the region
between first order lateral veins, externals, or loops (Figs 6 and 7). The word "pattern" as
used here refers to a group of veins showing some coherence in their order. Groups of
thinner/higher order veins may be found within an area defined by one pattern, but not be part
of that pattern. They could be "noise" or separate patterns of a higher order.
Astute observations on the development of the lateral segment venation of Ficus have
been published by Corner (1958). He noted that simplified secondary segment venation
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Pole - Modified leaf architecture terminology 309
-- Ramifying
Lateral Vein
Lattice
Percurrcnt
Joining Vein
Transverse
Second Order Patterns
Third Order Patterns
Fig. 7 - Diagrams to illustrate common second and third order venation patterns.
could be found in the unexpanded segments towards the apex of a leaf. In effect, an
ontogenetic sequence is preserved from the apex to the widest part of the lamina. Using this
as a guideline, Corner showed how several apparently different venation patterns in Ficus
were related by the initial pattern of subdivision of the segment. The divergent venation
patterns could be produced by differential expansion of different parts of the segment (Fig.
6b). The principle corresponds to that of the similarity of juvenile forms of related taxa
known in the field of zoology.
Percurrent venation
This refers to non-ramifying venation which runs directly between two veins. It may be
simple or it may branch, but the term percurrent does not apply to relatively continuous
networks of veins which run between two veins and are also interconnected with veins of
similar size. In a leaf showing percurrent venation, two kinds of percurrent veins (Fig. 3) may
be defined. These were named by Meyerhoff (1952), who grouped them as "tertiaries":
1. Joining veins
Non-ramifying veins which join two adjacent lateral veins (abbreviated to j.v.).
2. lnterangular veins
Non-ramifying veins which join lateral veins with the midrib (abbreviated to i.a.v.). The
relative development of these veins within a lateral segment depends on the amounts of
growth along the midrib between two laterals, and across the lamina between the midrib and
leaf margin or lateral loop.
Cascade venation
Corner (1958) recognised a special case where interangular veins are not directly percurrent,
but are staggered so that the end of a vein originating at the midrib terminates along a vein
originating on a lateral, and vice-versa. Growth producing this pattern is initiated in the angle
between midrib and laterals. It can also be recognised in loops.
Ramifying venation
The segments between the veins forming the first order patterns are occupied by a
ramifying vein network. A ramifying set is anchored to the first order pattern at some point
and then generally branches in a transverse, admedial, or exmedial direction (equivalent to
Hickey's (1979) ramified veins).
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310 Journal of the Royal Society of New Zealand, Volume 21,1991
THIRD ORDER VENATION PATTERNS
Third order refers to repeating patterns of venation which fill the segments defined by
veins of the second order pattern (Fig. 7), for instance a pattern which is consistently found
between two percurrent veins.
A large number of third order patterns may be recognised, but a comprehensive treatment
of them and their interelationships will require comprehensive study of recent plants and is
beyond the scope of this paper. The three most obvious are as follows:
Lattice venation
Found as segment "fill", often between percurrent veins, but also found between first
order laterals. It is formed by a zigzag vein connecting two rows of offset veins projecting
from the two veins defining the segment. In this sense it is similar to the construction of a
cascade vein, but there is no marked trend in the size of its constituent veins in any particular
direction, and it is not oriented towards the axil between two major veins.
Percurrent venation
Similar to a percurrent second order pattern, except that the veins run between adjacent
joining veins.
Transverse venation
The veins have a somewhat ramified appearance but are aligned parallel to enclosing
joining veins.
FIMBRIAL VEIN
Hickey (1973, 1979) and Dilcher (1974) proposed "fimbrial vein" for "higher vein orders
fused into a vein running just inside of the margin". I retain this term but suggest that it may
represent an independent entity rather than fusion of higher vein orders.
BASIC LAMINA ZONES AND SEGMENTS (see Fig. 3b)
1. Lateral zone
The area of lamina between the lateral loops and the midrib. In a craspedodromous leaf,
where lateral loops are absent, the whole lamina forms the lateral zone.
a) Interlateral segments
The area of lamina bounded by two lateral veins, the midrib, and either the margin or the
lateral loop.
b) Joining vein segments
The area of lamina bounded by two joining veins and two lateral veins.
c) Interangular (vein] segments
The area of lamina bounded by two interangular veins, the midrib, and a lateral vein.
d) Interexternal {vein] segments
The area of lamina bounded by at least two external veins and the margin.
2. Looping zone
This refers to the area of lamina between the lateral loops and the margin. It contains the
"external loops". It may be nearly absent on some leaves, or so developed in others that it
makes up almost half the width of the lamina.
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Pole - Modified leaf architecture terminology 311
Inter/oop segments
The area of lamina bounded on all sides by a loop.
LAMINA DEVELOPMENT
1. Basic development
This proceeds as a smooth gradation of development (expansion) over the lamina.
2. Actinodromous development
This is a condition of craspedodromous venation where the basal laterals are strongly
developed often with concurrent development on them of external veins and counter-external
veins, sometimes forming lobes in the lamina, and/or a cordate base (Fig. 1).
3. Acrodromous development
This refers to disproportional development of the basal, or suprabasal, interlateral segments.
It is normally a condition of camptodromous venation (Fig. 2).
SUMMARY
I consider that the currently-accepted leaf architecture terminology of Hickey (1973,
1979) and Dilcher (1974) needs modification, for several reasons, including: (1) ambiguous
ranking of low order veins; (2) confusion of vein relationships with lamina development; (3)
non-resolution of important elements of architecture, such as "loops"; and (4) low level
importance of patterns.
The terminology proposed here: (1) utilises the difference between veins resulting from
"ramifying" growth and "non-ramifying" growth; (2) recognises only one primary vein, the
midrib, per lamina; (3) recognises the difference between vein relationships and the effects of
differential lamina development (expansion); (4) recognises that venation "loops" of earlier
terminologies are often complex structures of several venation elements, and introduces a
new term, "externodromous", for this condition; (5) recognises the equivalence of "external"
veins ("outer secondaries" and "pectinal" veins of earlier terminologies) with the elements of
"loops"; (6) introduces a number of terms referring to areas of the lamina, for instance the
"looping zone", which are helpful in describing the difference between architectural patterns;
(7) recognises three distinct orders of venation patterns in an attempt to produce a terminology
which expresses more closely the ontogenetic development of the leaf; and (8) emphasises
the importance of patterns in recognising venation elements and growth directions in the
lamina.
ACKNOWLEDGEMENTS
I would like to thank my supervisor, J.D. Campbell, and also R.S. Hill, J.D. Lovis and
M.M. Dettmann, and two anonymous referees, who reviewed the manuscript at various
stages. This work was financed by a UGC scholarship and University of Otago Bridging
Finance.
REFERENCES
Arber, A., 1950. The natural philosophy of plant form. Cambridge University Press. Cambridge.
Carr, DJ., Carr, S.G.M. and Lenz, J.R., 1986. Leaf venation in Eucalyptus and other genera of
Myrtaceae: implications for systems of classification of venation. Australian Journal of Botany 34:
53-62.
Comer, E.J.H., 1958. Transference of function. Botanical Journal of the Linnean Society 56: 33-40.
Dilcher, D.L., 1974. Approaches to the identification of angiosperm leaf remains. The Botanical
Review 40: 1-85.
Esau, K., 1965. Plant Anatomy, second edition. John Wiley and Sons Inc., New York.
Hickey, L.J., 1973. Classification of the architecture of dicotyledonous leaves. American Journal of
Botany 60: 17-33.
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312 Journal of the Royal Society of New Zealand, Volume 21,1991
1979. A revised classification of the architecture of dicotyledonous leaves. In: Metcalfe, e.R.
and L. Chalk (eds.) Anatomy ofthe Dicotyledons. Vol. I. second edition, pp. 25-39. Clarendon Press,
Oxford.
Hickey, L.l. and Wolfe, l.A., 1975. The bases of Angiosperm phylogeny: vegetative morphology.
Annals of the Missouri Botanical Garden 62: 538-589.
Melville, R., 1976. The terminology of leaf architecture. Taxon 25: 549-561.
Meyerhoff, A.A., 1952. A study of leaf venation in the Betulaceae, with its application to paleobotany.
Unpublished PhD thesis, Stanford University.
Pray, T.R., 1955. Foliar venation of angiosperms, II. Histogenesis of the venation of Liriodendron.
American Journal of Botany 42: 18-27.
--- 1963. Origin of vein endings in angiosperm leaves. Phytomorphology 13: 60-81.
Romero, E.l. and Dibbern, M.e., 1985. A review of the species described as Fagus and Nothofagus by
Dusen. Palaeontographica Abt B 197: 123-138.
Slade, B.F., 1957. Leaf development in relation to venation as shown in Cercis siliquastrum L., Prunus
serrulata Lind!., and Acer pseudoplatanus L. New Phytologist 56: 28 -300.
--- 1959. The mode of origin of vein-endings in the leaf of Liriodendron tulipifera L. New
Phytologist 58: 299-305.
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.
Steenis, C.G.G.J. van, 1953. Results of the Archbold expeditions. Papuan Nothofagus. Journal of the
Arnold Arboretum 34: 301-373.
Thompson, D' Arcy, 1952. On growth andform, second edition. Cambridge University Press. Cambridge.
Troll, W., 1935. Vergleichende Morphologie der hoheren Pflanzen. Gebriider Borntaeger, Berlin.
Received 31 August 1990; accepted 10 May 1991
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