The Terminology of Leaf Architecture

R. Melville

Taxon, Vol. 25, No. 5/6. (Nov., 1976), pp. 549-561.

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TAXON ~ ~ ( 5 1:6549-561.
) NOVEMBER 1976

THE TERMINOLOGY OF LEAF ARCHITECTURE

Summary
A comparative study of leaf architecture throughout the Angiosperms has been made
and six classes of venation patterns are recognised. The terms employed are defined and
arranged in a systematic order for convenience of reference. Some new terms have been
introduced where greater precision was needed and for little known venation patterns.

Introduction
I n the early sixties the Systematics Association set up a committee for de-
scriptive terminology, the objectives of which were to consider the morphological
terms in use in descriptive plant taxonomy and to make recommendations for
their standardisation. A start was made with the simple plane shapes of leaves
for which W. B. Turrill had already drawn u p a scheme. This was modified
slightly and submitted to botanists in many countries for criticism. An agreed
list of terms was then drawn u p with illustrations and published in Taxon in
1962. I t had been the intention of the committee to pass on to a consideration of
the terminology of more complex shapes and to characters of leaf margins,
indumentum, glands and other features. Unfortunately they were unable to find
volunteers able to devote the time necessary to survey the literature and produce
preliminary proposals for other character systems. The need to review and
standardise botanical terminology has since remained an outstanding problem.
Recently L. J. Hickey (1973, 1974) has made a valuable contribution in this
field in a paper on the architecture of dicotyledonous leaves. I n this he considers
the shapes of leaf bases and apices, the marginal teeth and the terminology of
the venation patterns of leaves. Mouton (1970) covered much the same field,
giving terms in French only and mingling marginal characters with purely venation
characters in his terminology in a manner which is apt to be confusing. Neither
treatment is exhaustive as Hickey deals only with Dicotyledons and Mouton
incompletely with the whole of the Angiosperms. Both authors approached the
subject from the point of view of palaeobotany, but go back in the fossil record
only to the Cretaceous. Both, also, have taken up the terminology of von
Ettingshausen (1861), another palaeobotanist. Von Ettingshausen appears to have
set out to devise a new series of terms without giving full consideration to terms
already in use. Unfortunately he based his terms on Greek roots which results
in an ungainly series of polysyllables which do not harmonise with the Latin
terminology used in taxonomic descriptions.
The purpose of this paper is to attempt a systematisation of the terminology of
leaf architecture, introducing a few additional terms, where necessary, based on
Latin roots. The majority of terms used in the description of leaves have been
defined and many illustrated by Stearn (1966) and it is not necessary to repeat
here those appertaining to leaf shapes, margins, apices and bases or indumentum.
With one exception, attention will be restricted to leaf venation patterns, a field
which has been much neglected in the past. I t is necessary, however, to consider
the evolution of megaphyllous leaves from the time of the first invasion of the
land by vascular plants in the Silurian, if a proper understanding of the leaves

Royal Botanic Gardens, Kew, Surrey, England.

Tab. I . I, Arbuscular fastigiate leaf - Stirlingia tenuifolia. 2, Simple flabellate - Cir-
caeaster agrestis. 3, Pedati-flabellate - Leucadendron argenteum. 4 , Pedati-flabellate -
Petal of Ploiarium alternifolium. 5-7, Rectipalmate leaves of Acer: 5, A . monspessulanum;
6, A. campestre; 7, A. palmaturn. 8, Pedate - Platanus occidentalis. 9-12, Convergate
(Curvipalmate) leaves of Dioscorea: 9-10,D. spicata; 1 1 , D . alata; 12, D . bulbifera. 13,
Palmati-pinnate - Thespesia populnea.

of modern Angiosperms is to be attained. The studies on which these proposals

are made have led me to examine cursorily all Angiosperm families and a
number of genera in some detail as well as to scan the fossil record, especially
that of the Glossopteridae, which play a key role in Angiosperm evolution.
Considerable simplification of our terminology can be attained if we pay
attention to the simple fact that veins may be straight or curved. Then it is
necessary to recognise six primary systems of venation patterns of which pal-
mate and pinnate are the most familiar. To this must be added some consider-
ation of evolutionary processes and the ontogeny and physiology of development
of leaves, but the latter aspects will be treated in another paper.

Primavy systems of venation patterns

The nomenclature of the primary systems takes into account the order in which
veins are formed during leaf development and to some extent the manner in which
they are formed. Thus, in a pinnate leaf the midrib is the only primary vein
and the lateral pinnately arranged costae are secondaries. The intercostal areas
are normally divided up by a ternary series of veins and the areolae so formed
may be further divided by a quaternary series and further by a quinary series.
The symbol IO, zO, 3 O , 4 O and 5' will be used to indicate this order. In palmate
leaves veins of each order are initiated simultaneously, or nearly so, but the
vein orders follow one another successively in time.

5 so T A X O N VOLUME 2
 In palmate leaves the number of primary veins varies from 3-y-7-9-11 ...
Probably these primaries are always initiated in r a p ~ dsuccession, but any attempt
to discriminate between them on this account would lead to confusion. Between
these primaries the intercostal areas may be divided up successively by secondary
and higher orders of veins in much the same manner as for the intercostal areas
between the secondaries of pinnate leaves. Somewhat different considerations
arise in connection with the remaining primary venation systems and these are
discussed separately. They are taken in the following order:
I Arbuscular (arbuscularis) 4 Pinnate (pinnatus)
2 Flabellate (flabellatus) 5 Collimate (collimatus)
3 Palmate (palmatus) 6 Conglutinate (conglutinatus)

I Arbusctllav (Arbuscularis)
This term is proposed for leaves that branch repeatedly by regular dichotomy
to give rise to a three dimensional bush-like structure consisting of linear
segments. Two classes can be recognised:
1.1 Arbuscular simple: Segments containing a single vascular strand. Found in Rhynia
spp., Actinoxylon banksii Matton and common in the Psilophytes. Not found in
Angiosperms.
These structures are strictlv branch svstems which functioned as leaves. Thev are
the proto-phyllomes from which megaphyllous leaves evolved.
1.2 Arbuscular, fastigiate. Segments containing a fascicle of dichotomous vascular
strands; e.g. Stirlingia tenuijolia (R. Br.) Endl. (Fig. I), Isopogon villosus Meissn.
(Proteaceae)

z Flabellate (Flabellatus)
Primary veins straight or only slightly curved, idverging frrom the base in a
fan-like manner.
2.1 Simple jlabellate: Vein anastomoses absent or rare. e.g. Ginkgo biloba L., Adiantum
capillus-veneris L. (leaflets), Circaeaster agrestis Maxim. (Fig. 2), Kingdonia uni-
flora Balf. f. & W. W. Sm.
2.2 Reticuli-jlabellate: Vein anastomoses frequent, usually forming a dichotomo-
reticulate mesh.
2.21 Cuneati-flabellate: Primary veins diverging from I to 3 veins at the base. e.g.
Leucadendron platyspermum R.Br., Myrothamnus jlabellijolius (Sond.) Welw.,
Euryops latijolius B. Nord., Stylidium striatum Lindl.
2.22 Pedati-jlabellate: Primary veins forming a broad band at the base, often uniting
into pedate basal branches attached to a single trace at a unilacunar node, as in
Aeonium. e.g. Aeonium arboreum Webb & Berth., Gangamopteris spp., Leucaden-
dron argenteum (L.) R.Br. (Fig. 3). Also in petals, e.g. Ploiariurn alternijolium
(Vahl) Melchior (Fig. 4), Calophyllum inophyllum L.
The exact mode of attachment in Gangamopteris is not known, but it is possible
that some species had pedate bases uniting into a single trace as in Aeonium.

3 Palmate (Palmatus)
There is no sharp break between flabellate and palmate venation systems and
comparative studies indicate that palmate leaves have evolved from flabellate
ancestors. When the primary veins are straight and z 0 veins are absent the
separation is sim le, but if z0 veins are present the dichotomo-reticulate venation
is the distinguisting character of the flabellate system. Leaves with curved
primary veins that might justifiably be called curvi-flabellate, are not readily
distinguishable by means of superficial characters if the venation pattern is
obscured by a thick cuticle or dense indumentum. For this reason all leaves with
curved primaries are treated here as palmate.
Palmate leaves with rectilinear and curvilinear primary veins can be recog-
Tab. 2 . 14, Rectipinnate - Carpinus betulus. ~ j C , ompound rectipinnate - Corylus
avellana. 1 6 , Curvipinnate - Cornus sanguinea. 17, Co-arcuate - Prunus svium. I 8,
Paxillate - Calophyllum inophyllum. 19, Curvi-paxillate - Calathea zebrina. zo, Lirate -
Adenanthe bicarpellata Z I , Collimate - Hordeum vulgare. 22, Sub-collimate - Olyra
latifolia.

nised. In the rectilinear series the I' veins are straight or nearly so and they
radiate from a node which may be basal o r supra-basal. The I' veins of the
curvilinear series diverge from a basal node and arch towards a second node
a t the leaf apex. In either series, starting with three I " veins additional veins can
be added and the leaf expanded laterally, the base passing from acute to truncate
to cordate and finishing with overlapping basal lobes or by becoming peitate. The
shapes resulting from this process form natural intergrading series, which can
readily be reduced to a mathematical formula (D'Arcy Thompson 1942). The
slight changes in growth rhythm underlying the process are often expressed in
different species of a genus and are illustrated here by a rectilinear series in
Acev (Figs. 7, 6 , 7) and a curvilinear series in Dioscovea (Figs. 9-12).
3.1 Rectipalmate (Recti-palmatus).
e.g. Acer monspessularium L. (Fig. I), A campestre L. (Fig. 6 ) , A paimatum
Thunb. (Fig. 7 ) . Synonym: Actinodromous.
3.2. Pedate (Pedatus)
Leaf shape palmatifid or palmatisect, with the upper lobes supplied by I " veins
but lower lobes on either side supplied not by I " veins, but by 2 " recti-pinnate
laterals of the lower primaries. e.g. Platanus occidentalis L. (Fig. S ) , Cucurbita
pep0 L. Synonyms: Palmactinodromous (Von Ettingshausen), pedalCe (Mouton).
3.3 Convergate (Convergatus) or Curvi-palmate (Curvi-palmatus).
Although the curvilinear primaries of this series can be correctly described as
curvipalmate, it is convenient to have a simple alternative and Convergate, meaning
"bowed together", which implies both the curvature of the veins and the presence
of basal and apical nodes is here proposed. e.g. the series in Dioscorea (Figs. 9-11).
D. spicata Roth with elliptic-acute convergate leaves with 3 or 5 I " veins.
D. alata L. with ovate truncate convergate leaves with 7 I " veins.
D. bulbijera L. with cordate convergate leaves with 9 I " veins.
Other examples:
Plantago major L.; Melaleuca leucadendron (L.) L. Synonyms: Acrodromous,
campylodromous.

TAXON VOLUME Z j
4 Pinnate (pinnatus)
With a single primary vein, the midrib, along which straight or arching 2'
veins are arranged at f regular intervals.
4.1 Rectipinnate 4.4 Lirate
4.2 Curvipinnate 4.s Palmati-pinnate
4.3 Paxillate 4.6 Meta-pinnate
4.1 Rectipinnate (rectipinnatus)
Secondary veins straight or nearly so.
4." Simple rectipinnate - z o veins terminating at marginal teeth.
e.g. Ulmus glabra Huds., Alnus glutinosa (L.) Gaertn., Carpinus betulus L. (Fig.
14), Quercus castaneifolia C. A. Mey. Synonyms: Craspedodromous Craspedo-
drome aristCe. (Mouton).
4.12 Compound rectipinnate - 2" veins branching near the margin to supply several
teeth. e.g. Corylus avellana L. (Fig. I S ) , Viburnum lantana L. Synonym: Cras-
pedodrome composCe (Mouton).
4.2 Curvipinnate (curvipinnatus)
Secondary veins curving gradually towards the margin and not supplying a
marginal tooth directly.
4.21 Simple curvipinnate - z o veins curving gradually towards the margin and often
forming marginal or submarginal veins (see below). e.g. Cornus sanguinea L.
(Fig. 16), Bridelia ferruginea Benth. (Fig. 37). Synonym: Camptodromous.
4.22 Co-arcuate (co-arcuatus). - 2' veins connected by arching loops some distance
from the margin forming a strong infra-marginal vein; e.g. Prunus avium L.
(Fig. 17), Ficus populifolius Vahl, Ligustrum ovalzfolium Hassk., Aucuba japonlca
Thunb. Synonym: Brochidodromous.
4.23 Multi-arcuate (multi-arcuatus) - z o veins forming a coarcuate infra-marginal vein
and breaking up into a series of small arching loops forming a zone between the
infra-marginal vein and the margin. e.g. Napoleonea leonensis Hutch. & Dalz.
Synonym: Brochidodrome arch (Mouton).
4.3 Paxillate (paxillatus)
Secondary veins numerous, closely ~ a r a l l e lto one another, generally making angles
of 60-90 degrees t o the midrib. Derived from paxillus, "a little stake or pale",
hence paxillate, made of little stakes or pales and forming a small fence or palings.
4.31 Recti-paxillate - 2" veins unbranched or with a few basal dichotomies, with
free ends at the margin or looped together at the margin to form a simple marginal
vein. N o specialised areolae along the midrib.
4.311 Vein endings free e.g. Taeniopteris spp., Glossopteris clarkei Feistm.
4.312 Veins terminating in a marginal vein, e.g. Oleandridium brackebuschianum Kurtz
(Kurtz 1921), Calophyllurn inophyllum L. (Fig. 18). These constitute the
taeniopteroid leaves of earlier papers (Melville 1969, 1971).
Synonym: Parallelodrome transverse (Mouton).
4.32 Dichotomo-paxillate - 2" veins dichotomous and anastornosing to form a simple
reticulum (dichotomo-reticulate see below) and with specialised types of areolae
associated with the midrib. (Fig. 76-83) e.g. Glossopteris communis Feistm.
(Fig. 23), Gyrinops walla Gaertn., Roucheria calophyllum Planch. (Fig. 24).
Angiosperm leaves of this class have hitherto been called glossopteroid in earlier
papers (Melville 1969, 1971).
4.33 Sub-paxillate - Derivatives from simple paxillate types, lacking specialised areolae
along the midrib, but with some 2" veins stronger than others and the intercostal
areas dichotomo-reticulate or occupied by pendent or dendroid 3" veins. Inter-
mediate between simple paxillate and more advanced reticulate venation. e.g.
Tyleria tremulea Mag. & Wurd., Schuurmansiella angustijolia (Hook. f.)
Hallier f., Qualea gestasiana St. Hil., Lorostemon bombyciflorus Ducke.
4.34 Reticuli-paxillate - Covers stages in the conversion of a simple paxillate venation
pattern into a coarcuate or similar pinnate type but with the 3" reticulum
imperfect and the paxillate veins more or less reorientated in relation to the
3' reticulum. This, at times, produces a fingerprint-like pattern. At present only
 known from Rubiaceae. e.g. Timonius avenis Val., T . affinis A. Gray (Fig. 29).

Curvi-paxillate - With numerous closely parallel 2" veins, arching t o nearly

straight, except near the margin where they curve more or less abruptly into

a marginal vein. e.g. Strelitzia reginae Banks, Musa spp., Heliconia spp., Calathea

zebrina Lindl. (Figs. 19, 36). Synonym: Parallelodrome transverse.

Lirate (liratus)

With numerous parallel forwardly directed (oblique) 2" veins making angles of

10-30" with the midrib. Derived from lira, a ridge between two furrows, as on a

ploughed field.

Palaeo-lirate with the midrib consisting of a single strand as in some fossils. e.g.

. I

lirate (or neo-lirate) with the midrib consisting of a few to many closely aggre-
gated strands, e.g. Hanguana malayana (Jack) Merrill, Dracaena spp., Cordyline
spp., In Adenanthe bicarpellata Maguire & Wurdack (Fig. 20) the veins
occasionally dichotomise.
Palmati-pinnate.
Intermediate between palmate and pinnate, with the distal part of the leaf

pinnate and a basal or suprabasal pair of pinnated major veins extending for

'I3- ' 1 3 of the length of the lamina. e.g. Corchorus olitorius L., Tilia cordata Mill.,

Apeiba tibourbou Aubl., Grewia spp., Thespesia populnea (L.) Soland. (Fig. 13).

Synonym: Actinodrome composCe (Mouton, in part).

Meta-pinnate (meta-pinnatus)

Composite veins formed by the approximate alignment of cross connections

between the longitudinally parallel veins of collimate or lirate leaves giving rise

to regular, irregular or broken veins arranged transversely t o the principal vein

system in a pinnate manner.

Normal meta-pinnate, when they may be ascending in the upper part of the leaf,

spreading in the middle and declining in the basal part. e.g. Hanguana malayana

(Jack) Merr. (Figs. 43,44,45).

Broken meta-pinnate (fracti-meta-pinnatus). With the veins interrupted and f

irregular e.g. Helmholtzia acorifolia F. Muell. (Fig. 42).

Gradate (gradatus). Broken meta-pinnate veins in which the cross veins are

separated and arranged like the steps of a stairway e.g. Pandanus adinobotrys Merr.

& Perry (Figs. 40, 41).

5 Collimate (Collimatus)
Leaves w i t h numerous longitudinally parallel p r i m a r y veins arising f r o m a
transverse meristem.

Tab. 3. 23-24, Dichotomo-paxillate veins: 23, Glossopteris communis; 24, Roucheria

calophyllum. 25, Dichotomo-reticulata venation - Lilium martagon. 26, Pendulous veins -
Qualea gestasiana. 27, Zigzag vein - Degeneria vitiensis.

5 54 TAXON VOLUME 25
Tab. 4. 28, Irregular polygonal reticulum - Roupala montana. 29, Reticuli-paxillate
venation - Timonius affinis. 30, Sinuous dichotomo-reticulate - Senecio scorzoneroides.

The term is derived from collimatus, directed straight forward. Compare the
parallel usage in physics for collimator, an instrument for collecting light and
projecting it as a parallel beam. Synonyms: parallelodromous, paralleledrome
longitudinale (Mouton).
f . 1 Transverse meristem at the base of the lamina. e.g. Endymion, Haemanthus.
5.2 Transverse meristems at the bases of the lamina and of the leaf sheath. e.g.
Gramineae, Hordeum vulgare L. (Fig. 21).
f.3 Transverse meristem at the base of the lamina, where the longitudinal veins
are attached t o pedate transverse basal veins in the adult leaf. e.g. Richea spp.
5.4 Transverse meristem at the leaf apex, growth continuing until arrested by drought.
Found in a few Proteaceae e.g. Hakea multilineata Meissn.
1.5 Sub-collimate may be applied t o leaves in which the longitudinal veins become
bowed by differential growth and the leaf shape changed t o lanceolate or elliptic.
Then distinguished from convergate by the larger number of longitudinal veins and
the presence of a secondary series of longitudinal parallel veins between the pairs
of the primary series. e.g. Olyra latifolia L. (Fig. 22), Zeugites mexicana Trin.

6 Conglutinate (Conglutinatus)
Simple leaves or leaflets evolved by the coalescence of leaflets of compound
pinnate, bipinnate or tripinnate leaves with fusion of the formerly separate
venation systems producing zones of interaction along the lines of the original
leaflet margins.
All stages in the condensation of a complex frond into a simple leaf occurred
in Gigantopteridaceae (See Asama, I 9 5 9).
6.1 Simple conglutinate. One stage of fusion between leaflets.
a. leaflets: e.g. Gigantonoclea lagrelii (Halle) Koidzumi, Thelypteris megaphylla

(Mett.) Iwatsuki (Fig. 35) and other Thelypteroid ferns, Touroulia guianensis

Aubl.

This is the unicoherent leaf segment stage of Asama.

6.2 Bi-conglutinate. T w o stages of fusion, simple conglutinate leaflets fused t o form

a complex leaf or leaflet. e.g. Bicoemplecopteris hallei Asama. Quiina guianensis
Aubl. Synonym: Bicoherent leaf segment stage of Asama. Plumose-reticulate of
Foster. (Foster 1950).
6.3 Tri-conglutinate. Three stages of fusion, biconglutinate leaflets fused t o form
a simple leaf. e.g. Gigantopteris nicotianifolia Schenk. The tri-coherent leaf stage
of Asama.
Tab. j. 31, Dendroid veins - Lorostemon bombyciflorus. 3 2 , Incomplete dichotomo-
reticulate - Heliophila linearis. 33, Pinnati-reticulate - Trochocarpa laurina. 34, Corrivate
veins - Astroloma tectum. 3 5 , Compound corrivate - Thelypteris megaphylla.

7 T h e Ternary and higher order reticulum

The finer reticulum of Angiosperm leaves can be analysed into three classes.
The first of these is a direct descendant of the dichotomous branch systems of
early land plants which by planation and webbing gave rise to the flat mega-
phyllous leaf with flabellate venation (Fig. 2). The next step was the development
of anastomoses between the dichotomous veins which gave rise to a simple
reticulum, here called dichotomo-reticulate. The other two classes are the product
of diffusion reactions which take place early in the ontogeny of the leaf (Mel-
ville 1971). These diffusion reactions are responsible for the spatial distribution
of the morphogens which determine the positions of procambial strands and
subsequently of the vascular tissue of the veins. With a uniform distribution of
morphogens over the meristematic tissue of the developing leaf a hexagonal
pattern is produced. I n the climax condition of this system a pattern of regular
hexagons should be produced, but interfering factors rarely allow the climax
to be achieved and the common result is a system of polygons in which pentagons
and hexagons predominate. For this reason, the second class is here called
polygonal. When the diffusion reaction is constrained between previously formed
veins, as for example between the approximately parallel secondary veins of a
pinnate leaf, a sine-wave-like distribution of morphogens may be produced
which gives rise to regularly spaced transverse veins between the secondaries.
The arrangement of veins is like the rungs of a ladder and hence this class is
called scalariform. Interfering factors often cause irregularities in the pattern
and the continuation of meristematic activity may separate the procambial
strands to the point where an additional vein may be intercalated into the system,
as in the example of Hosta ventricosa (Salisb.) Stearn ( H . coerulea auct.), studied
bv Prav, I ~,,,,
\
sqti.
Transitions between the three principal classes of reticulum are frequent.
There is a tendency for the dichotomo-reticulate system to break down partially
and become incomplete before one of the other systems takes over in an evolution-
ary sequence. The areolae may be invaded by free simple or dendroid veins before
a link up to form a closed reticulum is achieved. Scalariform 3' veins may be
arching or irregular in transitions between scalariform and polygonal systems or
there may be a gradual change from scalariform 3' veins to polygonal j 0 veins
in one leaf.
7.1 Dichotomo-reticulate (Dichotomo-reticulatus). Reticulum formed by vein dicho-
tomy and anastomosis usually giving rise to elongated areolae in which the vein
junctions are similar to those of Glossopteridae. See vein junctions of the glos-
sopterid syndrome (Melville 1969).
7.11 Normal dichotomo-reticulate. Areolae of the reticulum complete, veins not inter-
rupted. e.g. Glossopteris communis Feistm. (Fig. 23), Gangamopteris spp., Rou-

5 l6 T A X O N VOLUME I f
 cheria calophyllum Planch. (Fig. 24), Lilium martagon L. (Fig. r ~ ) Gyrinops
, walla
Gaertn.
Incomplete dichotomo-reticulate. Some of the areolae incomplete with some vein
ends free and failing to link up with neighbouring veins. This occurs frequently
in the transitional evolutionary stages between glossopteroid venation patterns
and more advanced Angiosperm types. I t often occurs in petals as a result of
reduction in size and complexity. e.g. Leaves: Heliophila linearis DC. (Fig. 32),
Mairea coriacea Bolus, Lepidium draba L. Petals: Ranunculus spp., Roucheria
calophyllum Planch.
Sinuous dichotomo-reticulate. Veins of the areolae sinuous. e.g. In leaf tip of
Senecio scorzoneroides Hook. f. (Fig. 30).
Polygonal (Polygono-reticulatus). Areolae formed by 3" or higher order veins
angular, commonly with 5-6 sides, but varying from 3-7.
Regular. Areolae of k uniform size and shape tending towards regular hexagons
(climax condition), but commonly pentagonal or hexagonal. e.g. Dryandra nivea
R.Br. (Fig. j8), Laurus nobilis L., Cinnarnonum zeylanicum Bl., Glossopteris
retifera Feistm.. Hemionitis arifolia
, \
(Burm.) Moore.
Irregular. Sides of the areolae of unequal length and areolae of unequal size.
Very common in many families. e.g. Roupala montana Aubl. (Fig. 28).
Polygono-scalariform. Intermediate between polygonal and scalariform types of
reticulum with polygonal passing into irregular scalariform often within a single
intercostal area. Frequent in many families. e.g. Eriobotrya japonica (Thunb.) Lindl.
Scalariform (Scalariformis). Intercostal areas bridged at regular intervals by
transverse veins either at right angles or with a regular orientation and having
the appearance of rungs on a ladder. Common in many families. The process
repeated into 4" and j o order veins produces a fine meshed rectangular or
quadrangular reticulum e.g. Apeiba tibourbou Aubl. (Fig. 39).
Regular scalariform. Bridging veins straight or nearly so and regularly spaced. e.g.
many Melastomaceae, in curvipinnate leaves of Bridelia ferruginea Benth. (Fig. 38),
in curvipaxillate leaves of Calathea zebrina Lindl., in Marantaceae and Musaceae
and in collimate leaves of Gramineae.
Irregular scalariform. Bridging veins arched, sinuous or angled passing into
polygono-scalariform (7.23), e.g. Hevea brasiliensis Muell. Arg., Corylus avellana
L. (Fig. 11).
Tesselate (tesselatus) Bridging veins in collimate leaves well spaced producing
brick-shaped areoleae, those between adjacent pairs of longitudinal veins alternating
and giving the appearance of brick wall or tesselate pavement, e.g. Zeugites
mexicana Trin. (Fig. 46), Arundinaria fastuosa (Mitf.) Makino.

T a b . 6. 36, Curvi-paxillate veins - Calathea zebrina. 37, Dendroid veins - Asarum

canadense. 38, Regular scalariform veins - Bridelia ferruginea. 39, Scalariform veins, 3 ° ,
4' & 5.' - Apeiba tibourbou. 40-41, Gradate veins - Pandanus adinobotrys. 42, Broken
meta-pinnate veins - Helmholtzia acorifolia. 43-45, Normal, meta-pinnate veins -
Hanguana malayana: 43, at tip, 44, at middle, 45, at base of lamina. 46, Tesselate
veins - Zeugites mexicana.
 Gradate Bridging veins between adjacent pairs of longitudinal veins arranged
like the steps in a stairway. (see 4.63).
Accessory types o j minor venation. A variety of accessory types of vein archi-
tecture occur in the intercostal areas of leaves in intermediate stages of evolution.
Some of the more frequent and significant types are enumerated here.
Dendroid (Dendroideus). Regularly or irregularly dichotomous veins occupying
an areolus and attached to the areolar veins at one point. e.g. Asarum canadense
L. (Fig. 37), Lorostemon bombycijlorus Ducke (Fig. 31), Melaleuca globijera R.Br.
Dendro-reticulate (Dendro-reticulatus). Dendroid intra-areolar veins in which
additional branches unite with the areolar veins or with one another. A stage
in the evolution of a polygonal reticulum in some families. e.g. Mareya spicata
Baill.
Free pinnate (Libero-pinnatus). Pinnately branched veins standing free in an
intercostal area and connected to the costae only at their base, occur in Proteaceae
(leaves and bracts) and in Epacridaceae. e.g. Petrophtle carduacea Meissn. (Fig. 59).
Pinnati-reticulate (Pinnati-reticulatus). Pinnatelv branched veins some of the
lateral branchlets of which anastomose with adjacent veins to produce an incom-
plete and irregular reticulum. e.g. Trochocarpa laurina R.Br. (Fig. 33).
Corrivate (Corrivatus). Veins from opposite sides of an intercostal area which run
together and anastomose to form a gamma-shaped vein. Derived from corrivo -
"running together like two streams of a river". e.g. Astvoloma tectum R.Br.
(Fig. 34).
Compound corrivate. A series of corrivate veins united by their apices. In
Thelypteroid ferns compound corrivate veins mark the line of union of adjacent
leaflets in conglutinate leaf segments (6.1). e.g. Thelypteris megaphylla (Mett.)
Iwatsuki (Fig. 35).
Zig-zag (Fractiflexus). Medianly placed intercostal veins running approximately
parallel with the costae are frequent in pinnate leaves. They commonly take a
zig-zag course and may be formed by the approximate alignment of certain
veins in a polygonal reticulum or in intergrades between polygonal and scalariform
patterns. e.g. Degeneria vitiensis Bailey & Smith. (Fig. 27). Synonym: composite
intersecondary (Hickey).
Pendulous. Branching veins lying free in an intercostal area or an areolus, attached
at their distal ends and appearing to be pendulous from a submarginal vein or
costal vein. e.g. Qualea gestasiana St. Hil. (Fig. 26).
Digammoid (Digammoideus). When a sinus extends into a dichotomous vein system
it is held up at a dichotomy, which is commonly attached by a short vein to a
second dichotomy. This combination has the appearance of two Y's (or gammas)
one inside the other. It is a feature of the initial disruption of an entire margin in
certain primitive leaf types. e.g. Petrophile carduacea Meissn. (Fig. fg), Stylidium
barleei F. Muell., Isopogon villosus Meissn.
Flabellate leaj apex. In some species of Glossopteris transitional between Ganga-
mopteris and Glossopteris the midrib of the dichotomo-paxillate leaf breaks up at
the apex and spreads out into a flabellate pattern like that of Gangamopteris. A
similar phenomenon occurs in some relatively primitive paxillate angiosperm leaves.
e.g. Glossopteris antarctica Plumstead, Lepinia solomonensis Hemsl. (Fig. 60).
8 Marginal V e i n s
Veins associated with the leaf edge produce a number of characteristic patterns.
I n primitive leaves the excurrent veins remain simple and straight and run up
directly to the leaf edge. With the diversification of the leaf reticulum, veins
parallel with the leaf edge have developed a t different levels so that marginal,
sub-marginal and infra-marginal types can be distinguished.
Recently Hickey (1973) has proposed the use of the term "fimbriate" for
simple linear marginal veins, but this conflicts with its normal usage in botanical
terminology. Although it appears that the original classical meaning of fimbria
was the hem of a garment, the hem was often decorated with a fringe. Latin
dictionaries of the 17th and 18th centuries, while referring to the classical usage,

5~~ TAXON VOLUME 2 5

translated fimbriatus as "fringed o r jagged". I t is in t h e latter sense t h a t t h e t e r m
has a l w a y s been used i n Botany, a s i n Dianthus fimbriatus Lam., C a r e x fimbriata
Schkuhz, Gaillardia fimbriata Michx, C y p e r u s fimbriatus Nees a n d Habenaria
fimbriata R.Br. T o a l t e r t h e well established application n o w could o n l y l e a d
t o confusion.
Marginal vein absent.
Excurrent veins simple, running direct to the leaf edge and remaining free from
one another. e.g. Glossopteris spp. (Fig. 47), Gangamopteris spp., sepal of Tyleria
floribunda Gleason (Fig. 48).
Excurrent veins branched or dendroid at the leaf edge. e.g. Beauprea spathulifolia
Brongn. & Gris (Fig. 49).
Marginal vein simple, situated close t o the leaf edge and without any other veins
extending beyond it.
Linear: with a single continuous vein "linking" the ends of all of the excurrent
veins at the margin. e.g. Calophyllum inophyllum L. (Fig. IS), Enkleia siamensis
(Kurz) Nevling (Fig. 54), Qualea gestasiana St. Hil., Asarum canadense L. (Fig. 57).
Incomplete: marginal vein broken, linking some of the excurrent veins but leaving
others free. e.g. Glossopteris angustifolia (Fig. ~ o ) ,Stylidium striatum Lindl..
(Fig. 51).
Arcuate: marginal vein formed of arching veins linking the ends of the excurrent
veins. e.g. Ranunculus parnassiifolius L. (Fig. yz), Qualea multiflora Mart. (Fig. 56).
Irregular: marginal vein irregular, formed by the random union of veins adjacent
to the leaf edge. e.g. Grindelia buphthalmoides DC. (Fig. 53).
Submarginal veins. Any vein running parallel with the leaf edge, but separated
from it by other veins must be classified as a submarginal vein. One, two or
occasionally more submarginal veins may exist together in a leaf. The submarginal
zone between two adjacent submarginal veins may have distinctive features.
Fimbriate submarginal. Submarginal vein simple, linear, with a fringe of simple

Tab. 7 . 47-48, Simple excurrent veins: 47 - Glossopteris damudica Feistm.; 48 - sepal of

Tyleria floribunda. 49, Branched excurrent veins - Beauprea spathulifolia. 50-51,
Incomplete marginal veins: 50 - Glossopteris angustifolia; 5 1 - Stylidium striatum. $2,
Arcuate marginal vein - Ranunculus parnassiifolius. 5 3 , Irregular marginal vein -
Grindelia buphtalmoides. 54, Linear marginal vein - Enkleia siamensis. $5, Fimbriate
submarginal vein - Tyleria floribunda. 56, Paxillate submarginal zone - Qualea mul-
tiflora. $7, Submarginal zone irregularly reticulate - Jacquinia pungens. 58, Regular
polygonal reticulum - Dryandra nivea. 59, Digammoid veins at base of a sinus and free
pinnate veins Petrophile carduacea. 60, Flabellate leaf apex - Lepinia solomonensis.
Tab. 8. 61-69, Vein junction types: 61 - gamma, 62 - lambda, 63 - eta, 64 - kappa,
65 - Psi, 66 - Psi-lambda, 67 - chi, 68 - Zeta, 69 - zeta series. 70-75, Free vein islands
(areolae): 70 - biconvex lenticular; 71 - planoconvex lenticular; 72-73 - hexagonal;
74-75 - hydroid. 76-83, Attached vein islands: 76 - deltoid; 77 - arching; 78 - angled;
79 - triangular; 80 - truncate triangular; 81-82 - curvi-triangular; 83 - prismatic.

veins projecting towards the margin. This occurs in some paxillate leaves. e.g.
Tyleria floribunda Gleason (Fig. 55).
8.4 Submarginal zone. A submarginal zone may be formed between a marginal and a
submarginal vein, or between adjacent submarginal veins. The following conditions
have been noted.
8.41 Marginal and submarginal veins simple, linear, intervenal zone irregularly reti-
culate. e.g. Jacquinia pungens A. Gray. Fig. 57).
8.42 Marginal veins arcuate, submarginal vein linear, intervenal zone paxillate. e.g.
Qualea multiflora Mart. (Fig. 56).
8.5 Inframarginal veins. Arching veins connecting the 2" veins at some distance from
the margin (10-zoolo of the leaf width) are common in curvipinnate leaves
producing the coarcuate condition (4.22). The infra-marginal vein so formed may
be regular or irregular.
8.51 Regular. Loops of the infra-marginal vein uniformly arching. e.g. Prunus avium
L. (Fig. 17).
8.12 Irregular. Infra-marginal vein angled or irregular. e.g. Persoonia elliptica R.Br.
9 V e i n junctions and vein islands
For mechanical reasons only a limited number of different types of vein
junction is possible in a dichotomo-reticulate system of venation. In the absence
of a midrib the number of vein island types (areolae) was also limited, but with
the development of a midrib a new diversification was possible and led to a
number of new types in the Glossopteridae and primitive Angiosperms. Some
of these are shared with other groups, such as the ferns, which also produce types
of vein island not observed in Angiosperms and not discussed here.
Vein junctions. In the dichotomo-reticulate system, vein junctions resemble certain

letters of the Greek alphabet which can conveniently be used to describe them.

Gamma junction. The result of a simple dichotomy of a vein. y (Fig. 61).

Lambda junction. The result of the simple anastomosis of two approximately

parallel veins. (Fig. 62).

Eta junction. Formed by a transverse vein between two approximately parallel

veins. H (Fig. 63).

Kappa junction. The result of a dichotomy and an anastomosis resembling K or its

(Fig. 64) mirror image.

Psi junction. Formed by 3 veins diverging from one point. Y (Fig. 65).

Psi-lambda junction A psi-junction combined with an anastomosis. (Fig. 66) or

its mirror image.

Chi junction. Formed by the union of veins at an oblique crossing X (Fig. 67).

Zeta junction. Formed by an oblique transverse vein between two approximately

parallel veins. (Fig. 68).

Zeta series. A series of zeta junctions side by side frequent in some species of

Glossopteris and occasional in dichotomo-reticulate Angiosperm leaves. (Fig. 69).

5 60 TAXON VOLUME 25
 Free vein islands (Areolae). The following types of vein islands occur free in the

reticulum, i.e. not associated with a midrib or major vein, in leaves of Glossop-

teridae and in primitive Angiosperm leaves with dichotomo-reticulate venation.

Lenticular - shaped like the cross-section of a lens, either biconvex or plano

convex. (Figs. 70, 71).

Hexagonal. Either regular hexagonal or elongate hexagonal. (Figs. 72, 73).

Hydroid. Vein islands tapering to the proximal end and enlarged towards the

distal end t o which 4-7 other veins are attached giving a Hydra-like appearance.

e.g. in Gangamopteris and Glossopteris leaves; leaves of Laccopetalum giganteum

(Wedd.) Ulbrich, Pringlea antiscorbutica R.Br. (Figs. 74, 75). Petals of Rosa

spp., Geranium spp.

Attached vein islands. Vein islands attached to the midrib or to a major vein. e.g.

Glossopteris spp. Gyrinops walla Gaertn., Roucheria calophyllum Planch., Timonius

affinis A. Gray, Lepinia solomonensis Hemsl.

Deltoid. Erect, or slightly oblique, triangular areolae attached to a major vein

by a smaller side (Fig. 76).

Arching. Elongate curvi-triangular areolae (Fig. 77).

Angled. Oblique triangular areolae with a vein inserted along one side which is

angled. (Fig. 7.8).

Triangular. Tr~angular areolae attached by a broad base. (Fig. 79).

Truncate triangular. Triangular areolae with the apex truncated. (Fig. 80).

Curvi-triangular. Triangular areolae with the free sides arched (Figs. 81, 82).

Prismatic. Usually oblique areolae with parallel sides capped by a lambda junction.

(Fig. 83).

Acknowledgments
This paper has been read by senior members of the staff of the Royal Botanic Gardens,
Kew, and by Dr. N . K. B. Robson of the British Museum (Natural History), to all of
whom I am indebted for constructive suggestions. I am especially grateful to Mr.
H . K. A. Shaw for discussing the correct latin form of the new terms.

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The Terminology of Leaf Architecture
R. Melville
Taxon, Vol. 25, No. 5/6. (Nov., 1976), pp. 549-561.
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References

Morphology and Venation of the Leaf in Quiina acutangula Ducke

Adriance S. Foster
American Journal of Botany, Vol. 37, No. 2. (Feb., 1950), pp. 159-171.
Stable URL:
http://links.jstor.org/sici?sici=0002-9122%28195002%2937%3A2%3C159%3AMAVOTL%3E2.0.CO%3B2-S

Classification of the Architecture of Dicotyledonous Leaves

Leo J. Hickey
American Journal of Botany, Vol. 60, No. 1. (Jan., 1973), pp. 17-33.
Stable URL:
http://links.jstor.org/sici?sici=0002-9122%28197301%2960%3A1%3C17%3ACOTAOD%3E2.0.CO%3B2-2

Foliar Venation of Angiosperms. IV. Histogenesis of the Venation of Hosta

Thomas R. Pray
American Journal of Botany, Vol. 42, No. 8. (Oct., 1955), pp. 698-706.
Stable URL:
http://links.jstor.org/sici?sici=0002-9122%28195510%2942%3A8%3C698%3AFVOAIH%3E2.0.CO%3B2-E

Systematics Association Committee for Descriptive Biological Terminology. II. Terminology of

Simple Symmetrical Plane Shapes (Chart 1)
Taxon, Vol. 11, No. 5. (Jun., 1962), pp. 145-156.
Stable URL:
http://links.jstor.org/sici?sici=0040-0262%28196206%2911%3A5%3C145%3ASACFDB%3E2.0.CO%3B2-G