Chapter 9 Plant Morphology

Plant Morphology
From: Simpson, M. G. 2006, 2010. Chapter 9.
Plant Systematics.
Elsevier-Academic Press
2 Systematic Evidence and Descriptive Terminology

shoot
apex shoot apical
meristem vein

axillary bud
internode bud primordium
node

lateral
branch shoot tip

lateral branch

root tip

shoot

root hairs

lateral roots
root
root apical
meristem

root cap

Figure 9.1 General plant structure, showing primary root and primary shoot. Note that all parts of both root and shoot are derived
from cell divisions of the root or shoot apical meristem.
Chapter 9 Plant Morphology 3

parasite
primary host
root adventitious
roots
lateral
root

A B C D
taproot fibrous prop haustorial

E F G H
Figure 9.2 Root Types. A. Tap root. B. Fibrous root system. C. Prop roots. D. Haustorial roots. E. Storage roots; Raphanus sati-
vus, radish. F. Buttress roots; Ficus rubiginosa, fig. G-H. Pneumatophores; Avicennia germinans, black mangrove.
Stem/Shoot Type
protective
scale leaves

succulent
storage leaves
ground
stem level
l.s.
cactus
spines
areole bulb caudex caudiciform stems

cladode protective
scale leaves

flower(s) scale leaves

or leaf scars
stem
leaf
l.s.
cladode
bract
leaf subtending
(reduced) leaf

cladodes corm culm fascicle / short shoot rhizome

leaf scar

scape stolon thorn tiller tendril tuber

Figure 9.3 Stem Types. (l.s. = longitudinal section)
4 Systematic Evidence and Descriptive Terminology

inflorescence

prostrate

acaulescent caulescent cespitose

repent

new
growth

old
growth

arborescent suffrutescent decumbent

Figure 9.4 Stem Habit.

Branching Pattern

axis derived
from multiple sympodial
apical meristems unit
sympodial units
sympodial

teminal apical meristem abortive

bud divides equally terminal meristem
lateral lateral
bud branch

lateral lateral branch

bud sympodial
unit

monopodial dichotomous sympodial

axis derived from axis derived from
single apical meristem multiple apical meristems
Figure 9.5 Stem Branching Patterns.
Chapter 9 Plant Morphology 5

Twig Parts Bud Types / Parts

terminal (apical) bud

leaf
outer, primordia
protective
bud scales
apical
stipule scar lateral (axillary) bud meristem
node

lenticels
bud longitudinal section collateral superposed
internode
abortive apical
meristem

leaf scar
vascular bundle scars
petiole
encircling base
stipule scar leaf scar
terminal bud
scale scars
pseudoterminal naked infrapetiolar
Figure 9.6 Twigs parts and Bud types. (l.s. = longitudinal section)

inflorescence/
flower axis

bractlets petiolar
scale spine bud
bract

stipular
leaf spines
glochidia spines
(glochids) tendril (leaf) unifacial leaf
phyllode spine
Figure 9.7 Leaf structural types.
6 Systematic Evidence and Descriptive Terminology

vestigial
rachillae
phyllode

phyllode leaf
tendril
A B C
E petiole

stipular
spines leaf
spines
petiole
D spine F

G H I J

bract
scale
flower
calyx
epicalyx

leaf
K L M N

Figure 9.8 Leaf Structural Types. A-B. Phyllode, Acacia longifolia. A. Mature. B. Young, with vestigial, caducous rachillae, repre-
sentative of ancestral condition. C. Tendril, Lathyrus vestitus. D-F. Spines. D. Stipular spines, Euphorbia sp. E. Petiolar spines, Fouquieria
splendens. Note mature leaf (above), dehiscence of blade and upper tissue of petiole, leaving petiolar spine (below). F. Leaf spines, cactus
areole. G-K. Leaf modifications of carnivorous plants. G-H. Pitcher leaves. G. Nepenthes sp. H. Sarracenia purpurea. I. Tentacular leaves of
Drosera capensis. Note glandular trichomes (arrow). J. Trap leaf, Dionaea muscipula. K. Showy flower bracts, Bougainvillea. L. Epicalyx,
Lavatera bicolor. M. Bud scale, Liquidambar styraciflua. N. Unifacial leaf, Juncus phaeocephalus.
Chapter 9 Plant Morphology 7
Leaf Parts / Type
leaflet petiolule

midrib rachis
rachis

blade

imparipinnate
secondary
veins midvein
bud
petiole petiolule
stipels
petiole rachilla petiole
stipules
stipules
simple pinnately compound paripinnate bipinnately compound

rachis
petiolule petiolule petiolule
leaflet petiole

petiolule
rachillae
petiole petiole petiolule petiole

palmate-ternate pinnate-ternate biternately compound palmately compound

ternately compound (biternate)
(trifoliolate)

petiolule petiole

geminate

petiolule costa
rachilla petiole
petiole
rachilla petiole petiole

unifoliolate bigeminate geminate-pinnate costa-palmate

Figure 9.9 Leaf Types/Parts.

8 Systematic Evidence and Descriptive Terminology
Leaf Attachment

node adnation
petiole sheath

connate-
petiolate sessile sheathing decurrent amplexicaul perfoliate perfoliate

Figure 9.10 Leaf Attachment.

2
vein

mid-vein
(1) 1 veins

uninervous
pinnate-netted palmate-netted ternate-netted
ultimate veinlets reticulate (netted)
VENATION

2 vein
transverse
vein 1 vein
dichotomous parallel penni-parallel palmate-parallel
ultimate veinlets transverse
Figure 9.11 Leaf Venation, generalized terminology.
Chapter 9 Plant Morphology 9

2 vein 2 vein 2 vein 2 vein

1 vein 1 vein 1 vein 1 vein

simple semi- eucampto- reticulo- hyphodromous parallelo-
craspedodromous craspedodromous brochidodromous dromous cladodromous dromous dromous
craspedodromous camptodromous
pinnate venation

2 vein
1 veins 1 veins 1 veins
basal suprabasal basal suprabasal marginal reticulate flabellate
marginal reticulate
perfect imperfect
actinodromous

1 veins 1 veins

1 veins basal suprabasal basal suprabasal

1 veins
perfect imperfect
palinactinodromous campylodromous acrodromous
Figure 9.12 Leaf Venation, specialized terminology, after Hickey (1973) and Hickey & Wolf (1975).
 10 Systematic Evidence and Descriptive Terminology

Flower Parts Flower Attachment

anther
stamen
[androecium] filament stigma

style pistil
[gynoecium,
ovary 1- carpels] pedicellate sessile
petal
[corolla]
Flower Sex
perianth
sepal
[calyx] nectary ovules
receptacle
bract
pedicel
staminate pistillate
perfect/bisexual unisexual
Symmetry
Figure 9.13 Flower Parts, Sex, and Attachment.

biradial radial / bilateral /

actinomorphic zygomorphic
Figure 9.14 Flower symmetry types.

A C E

B D F
Figure 9.15 Flower symmetry examples. A-B. Actinomorphic / radial symmetry. A. Five planes of symmetry. B. Six planes of sym-
metry. C. Slight zygomorphy (bilateral symmetry), vertical plane of symmetry. D-E. Strong zygomorphy of all floral parts. F. Asymmetry,
caused by twisting of floral parts.
Chapter 9 Plant Morphology 11
Perianth Cycly / Merosity
inner
outer tepal
petal tepal
corolla

calyx

calyx sepal

uniseriate biseriate dichlamydeous homochlamydeous

pentamerous tetramerous biseriate / trimerous
Figure 9.16 Perianth Cycly and Merosity.

outer inner
tepals tepals calyx

corolla
calyx

A B C

Figure 9.17 Perianth Cycly. A. Biseriate, homochlamydeous, with outer and inner perianth parts (tepals) similar. B. Biseriate,
dichlamydeous, with distinct calyx and corolla. C. Uniseriate, with a single whorl of perianth parts, by default termed a calyx.

Perianth Fusion

petal petal

sepals corolla calyx lobes corolla

lobe lobe

aposepalous aposepalous synsepalous synsepalous

apopetalous sympetalous apopetalous sympetalous
Figure 9.18 Perianth Fusion.
12 Systematic Evidence and Descriptive Terminology

upper (posterior) lip operculum

tube corona

c.s.
throat

lower (anterior) lip tepals

bilabiate calyptrate/ campanulate carinate coronate

operculate

hood

cruciate cucullate (hooded) infundibular disk ligulate / ray

wing
banner petals
constriction

[section]

keel corolla tube

petals sac/
l.s. pouch
papilionaceous personate rotate saccate

calyx spur limb

salverform

claw
corolla spur
spurred tubular unguiculate urceolate

Figure 9.19 Perianth Types/Parts. (c.s. = cross section; l.s. = longitudinal section)

Aestivation

c.s. c.s. c.s. c.s. c.s.

imbricate-alternate convolute/contorted quincuncial valvate involute

Figure 9.20 Perianth Aestivation.
Chapter 9 Plant Morphology 13
Stamen Types / Parts Stamen Arrangement
anther
thecae
filament
laminar antherode
staminal body
disk

filamentous laminar staminodia didynamous tetradynamous didymous

Figure 9.21 Stamen Types and Parts. Figure 9.22 Stamen Arrangement.
Stamen Cycly/Position

antisepalous / antipetalous / diplostemonous obdiplostemonous

alternipetalous alternisepalous
uniseriate biseriate
Figure 9.23 Stamen Cycly (uniseriate or biseriate) and Position.

Stamen Insertion

inserted exserted
Figure 9.24 Stamen Insertion.

Androecial / Stamen Fusion

1 stamen
with filament anthers
point of basally fused filaments connate
filament fused
insertion
9 stamens
petal with connate
filaments

apostemonous epipetalous diadelphous monadelphous syngenesious

Figure 9.25 Stamen Fusion.
14 Systematic Evidence and Descriptive Terminology

Anther Type and Parts theca Anther Attachment

connective microsporangium

line of
dehiscence
c.s.
pollen theca
locule
theca theca
c.s.
dithecal monothecal basifixed dorsifixed subbasifixed dorsifixed basifixed
versatile
Figure 9.26 Anther Parts and Attachment (c.s. = cross section)

longitudinal poricidal transverse valvular

Figure 9.27 Anther Dehiscence Types.

Anther Dehiscence Direction

c.s. c.s. c.s.

extrorse introrse latrorse

staminode
abortive anther

c.s. c.s.
introrse downward
(early in development) (at maturity)
Figure 9.28 Anther Dehiscence Direction. (c.s. = cross section)
Chapter 9 Plant Morphology 15
Carpel Development / Pistil Parts
stigma

style

pistil
ovary wall

locule
ovary
ovule
c.s. funiculus
c.s. c.s. placenta

Figure 9.29 Gynoecium: Carpel Development. (c.s. = cross section; l.s. = longitudinal section)

septum

c.s. c.s. c.s.

c.s. l.s. stipe
carpels 3, locule 1 per carpel carpels 3, locule 1 carpels 2, locule 1 carpels 4, locules 4 carpel 1, locule 1
apocarpous syncarpous unicarpelous
Figure 9.30 Gynoecium Fusion. (c.s. = cross section)

A c.s. c.s. c.s.

B c.s. c.s. c.s.

Figure 9.31 Gynoecium: Carpel Boundaries. (c.s. = cross section)

 Hypanthium Presence / Ovary Position / Perianth-Androecial Position
16 Systematic Evidence and Descriptive Terminology
hypanthium hypanthium hypanthium hypanthium hypanthium
absent present absent present absent

hypanthium
hypanthium

ovary superior ovary inferior ovary half-inferior

hypogynous perigynous epigynous epiperigynous epihypogynous

Figure 9.32 Ovary Position and Perianth/Androecial Position.

column

c.s. c.s.
parietal-septate parietal-axile
c.s. c.s.

l.s. c.s. c.s. l.s. c.s. c.s. c.s. l.s. l.s. l.s.
axile parietal marginal laminar free-central apical / basal apical-axile
pendulous
Figure 9.33 Gynoecium: Placentation. (c.s. = cross section; l.s. = longitudinal section)

terminal lateral gynobasic discoid globose linear plumose

Figure 9.34 Gynoecium: Style Position and Stigma/Stig-

matic Region Type.

peduncle
involucre

simple umbel simple umbel

fascicle glomerule (indeterminate) (determinate)

Figure 9.37 Indeterminate or determinate inflorescences types.

Chapter 9 Plant Morphology 17

X
pedicel
bract
peduncle
X X top view

(prophyll) side view

stem axis

A
simple
dichasium D helicoid cyme

stem axis

X
X
top view

X side view

B E
scorpioid cyme
compound
dichasium

a
b c d e
1 3 5 6 4 2 top view 1 2 3 4 5 6 top view
b d e c a

1 3 5 6 4 2
1 2 3
4
e side view 5 side view
d e
d c
c 6
b b

C F G
a
compound rhipidium drepanium
cyme

Figure 9.35 Determinate (cymose) inflorescences. A-C. Dichasial cymes. A. Simple dichasium. B. Compound dichasium. C.
Compound cyme. D-G. Monochasial cymes. D. Helicoid cyme, showing derivation from compound dichasium by development of one
axis on the same side of the primary axis. E. Scorpioid cyme, showing derivation from compound dichasium by development of one axis
on alternating sides of the primary axis. F. Rhipidium. G. Drepanium. (Terminology after Weberling, 1981.)

rachis

pedicel inflorescence rachis

axis

spike raceme corymb (simple) corymb (compound) panicle

Figure 9.36 Indeterminate inflorescences types.

18 Systematic Evidence and Descriptive Terminology

thyrse verticillaster compound simple corymb of heads

raceme

1o bracts rays
(involucre) 2o bracts
(involucel)
peduncle

compound umbel raceme of spikes panicle of spikelets

(indeterminate)

Figure 9.38 Secondary inflorescences.

female flower
male flowers
spathe male flowers
bract

spadix involucre phyllary

palea lemma
c.s.
thick, flowers
cyathium
fleshy axis

florets

flowers
glumes
compound compound l.s.
l.s. receptacle receptacle
spikelet
catkin / ament spadix hypanthodium (grass) head / capitulum

Figure 9.39 Specialized inflorescence types.

Chapter 9 Plant Morphology 19
hard, thick
pericarp
pericarp
pericarp pericarp
seed
seed
locule
seed coat seed
l.s. l.s. l.s. wing l.s.
achene grain/caryopsis utricle samara nut
Figure 9.40 Fruits: simple, dry, and indehiscent fruit types. (c.s. = cross section; l.s. = longitudinal section)

valves
replum
2 lines of
dehiscence
c.s.
replum c.s.
c.s. valves
2 lines of
dehiscence

1 line of
dehiscence

follicle legume silique silicle

valves lines of transverse suture pores
dehiscence

c.s. c.s.

loculicidal septicidal circumscissile poricidal

capsule capsule capsule capsule capsule
1 line of
dehiscence mericarps
/ unit fruit

two carpels
of pistil
apically
connate
l.s. calyx carpophore

schizocarp of follicles schizocarp of mericarps schizocarp of nutlets

Figure 9.41 Fruits: simple, dry, and dehiscent fruit types. (c.s. = cross section; l.s. = longitudinal section)
20 Systematic Evidence and Descriptive Terminology

exocarp
fleshy perianth/androecium scar withered perianth/
endocarp juice sacs androecium
fleshy
l.s. mesocarp septum hypanthium
fleshy exocarp tissue
berry mesocarp ovary
tissue
hard c.s. c.s. cartilaginous
endocarp leathery l.s. endocarp
leathery
seed exocarp outer pericarp pedicel
l.s.
drupe pepo hesperidium pome
Figure 9.42 Fruits: simple, fleshy fruit types. (c.s. = cross section; l.s. = longitudinal section)
Aggregate Fruits

pericarp

seed seed

l.s.
l.s. l.s.
unit
achene unit endocarp unit
drupe follicle
receptacle calyx
calyx
achenecetum drupecetum follicetum
(Fragaria sp., Strawberry) (Rubus sp., Blackberry) (Magnolia grandiflora, Fl. Magnolia)
Figure 9.43 Fruits: aggregate fruit types. (l.s. = longitudinal section)
Multiple Fruits

remains of
perianth/
unit fruit androecium
flower
bract
bract
seed
l.s.
unit achene
(longitudinal section)

multiple fruit of achenes l.s.

(Platanus sp., Sycamore) sorosis
(Ananas comosus, Pineapple)

scales achenes

involucral
bracts
unit 15 mm
fruits

l.s. compound
receptacle l.s.

syconium bur
(Ficus sp., Fig) (Xanthium sp., Cocklebur)
Figure 9.44 Fruits: multiple fruit types. (l.s. = longitudinal section)
Chapter 9 Plant Morphology 21

prolate discoid

capitate oblate clavate lenticular filiform fistulose fusiform ligulate pilate terete turbinate
Figure 9.45 Seed Parts, endosperm types. (l.s. = longitudinal section)

Shape: Plane

linear narrowly oblong oblong narrowly elliptic elliptic widely elliptic orbicular
12:1 - 6:1 6:1 - 3:1 2:1 - 1.5:1 6:1 - 3:1 2:1 - 1.5:1 1.5:1 - 1.2:1 ~ 1:1

lanceolate lance-ovate ovate widely ovate oblanceolate oblance-ovate obovate

6:1 - 3:1 3:1 - 2:1 2:1 - 1.5:1 1.5:1 - 1.2:1 6:1 - 3:1 3:1 - 2:1 2:1 - 1.5:1

narrowly triangular widely deltate rhombic trullate

triangular 2:1 - 1.5:1 triangular ~ 1:1 2:1 - 1.5:1 2:1 - 1.5:1
6:1 - 3:1 1.5:1 - 1.2:1

acicular ensiform strap-

shaped cordate/ pandurate reniform
falcate spatulate subulate
> 12:1 cordiform

Figure 9.46 Solid (three-dimensional) shapes.

22 Systematic Evidence and Descriptive Terminology

obtuse rounded
> 90

attenuate narrowly cuneate cuneate

< 45 < 45 45 - 90 truncate cordate

petiole

reniform auriculate hastate sagittate oblique peltate

Apex:Note
Figure 9.47 Planar (two-dimensional) shapes. Shape
that bottom of drawings are points of attachment.

acute obtuse truncate

45 - 90 >90 ca. 180

acuminate caudate cuspidate narrowly

< 45 < 45 < 45 acute rounded emarginate retuse
< 45
Figure 9.48 Bases. Note that bottom of drawings are points of attachment.

apiculate aristate cirrhose mucronate mucronulate spinose/

> 3:1, flexuous > 3:1, stiff 1:1 - 3:1 1:1 pungent

Figure 9.49 Apices. Note that bottom of drawings are basal.

base & apex base cuneate base reniform base sagittate base cordate
rounded apex acute apex cuspidate apex truncate apex obtuse

Figure 9.50 Apical processes.

Chapter 9 Plant Morphology 23

entire serrate serrulate doubly dentate denticulate crenate crenulate spinose praemorse ciliate ciliolate filiferous
serrate
Figure 9.51 Shape combinations. Note that the five shapes are all elliptic, but differ in the base and apex shapes.
SHAPE: Division

sinus
lobe

pinnately lobed pinnately cleft pinnately parted pinnately divided sinuate incised
1/8 - 1/4 1/4 - 1/2 1/2 - 3/4 3/4 - ca. 1

palmately lobed palmately cleft palmately parted palmately divided laciniate pectinate
1/8 - 1/4 1/4 - 1/2 1/2 - 3/4 3/4 - ca. 1

3o leaf division
(pinnule)

leaf division/lobe
(pinnule) 2o leaf division
(pinnule)
1o leaf division
(pinna) 1o leaf division 2o leaf division
(1o pinna) (2o pinna)
pinnatifid
bipinnatifid tripinnatifid decompound

Figure 9.52 Margins.

24 Systematic Evidence and Descriptive Terminology

posterior (adaxial)
adaxial (ventral) distal
(apical)
terminal
(apical)

lateral
(axillary)
abaxial (dorsal)
proximal anterior (abaxial)
(basal) basal/radical cauline acrocaulis
general position specialized position
Figure 9.53 Division.

Arrangement

spiral distichous decussate non-decussate whorled

alternate opposite

c.s.
leaf
(of long shoot)
short shoot
(fascicle) bract

equitant fasciculate imbricate valvate secund

Figure 9.54 Position.

Chapter 9 Plant Morphology 25

appressed Orientation
15
ascending
30 appressed, distinct
anthers
45 inclined
30
45
15
horizontal
15 (divergent)
45
30
45 reclined
antrorse retrorse
30 connivent
15 descending reflexed erect pendant/
depressed pendulous
Figure 9.55 Arrangement. (c.s. = cross section)

Posture: Transverse

recurved cernuous squarrose incurved flexuous

Posture: Longitudinal

c.s.

c.s.
c.s. c.s.

cup-shaped
conduplicate revolute involute (abaxially concave)

c.s. Posture: Twisting/Bending

induplicate
c.s.
reduplicate
180o twist

c.s.
c.s.

plicate undulate/repand resupinate geniculate

Figure 9.56 Orientation.

26 Systematic Evidence and Descriptive Terminology

Configuration Vestiture
beard

canaliculate punctate rugose/bullate striate bearded

Epidermal Excrescence

aculeate/prickly farinaceous/scurfy papillate/ muricate ciliate/ comose

verrucate/tuberculate ciliolate

Vestiture

pubescent puberulent tomentose tomentulose lanate/villous

pilose arachnose/arachnoid scabrous hirsute hispid

strigose strigulose sericeous floccose lepidote

Figure 9.57 Posture. (c.s. = cross section)

Trichome Types Bristle Types

unicellular

malpighian uniseriate multiseriate pilate capitate barbed/barbellate plumose

(2-armed) tapering glandular

sessile stalked dendritic peltate uncinate/hooked glochidiate

stellate

Figure 9.58 Surface Features: Configuration, Epidermal Excrescence, and Vestiture.