See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/306019334

Leaf anatomy of Cinnamomum Schaeffer (Lauraceae) with

special reference to oil and mucilage cells

Article  in  Blumea journal of plant taxonomy and plant geography · January 1992

CITATIONS READS

12 204

3 authors, including:

Els Bakker Arnout F Gerritsen

Leiden University Medical Centre Genmab
41 PUBLICATIONS   666 CITATIONS    35 PUBLICATIONS   1,195 CITATIONS   

SEE PROFILE SEE PROFILE

All content following this page was uploaded by Els Bakker on 10 August 2016.

The user has requested enhancement of the downloaded file.

BLUMEA 37 (1992) 1-30

Leaf anatomy of Cinnamomum schaeffer (Lauraceae)

with special reference to oil and mucilage cells

M.E. Bakker A.F. Gerritsen & P.J. van der Schaaf

Rijksherbarium/Hortus Botanicus, Leiden, The Netherlands

Summary

The morphology and distribution patterns of oil and mucilage cells in the leaf of 150 species of Cin-

described. Idioblasts in the and the spongy parenchyma.

namomum are are always present palisade
both oil and cells in species either oil or mucilage cells
Usually mucilage occur; some are present.

Both of idioblasts suberized wall The idioblasts species in size/

types possess a layer. vary between

and number. Variations in the distribution partly be explained by the

shape, stainability pattern can

of the oil and mucilage cells.

proposed homology

Other leaf anatomical characters are also mentioned, such as lamina and cuticle thickness, bundle

sheath extensions, sclerification of the epidermal, the palisade, and the parenchyma cells,
spongy

number of palisade layers, presence or

absence of a hypodermis, the indumentum and papillate ab-

axial epidermal cells, and the venation pattern.

Most species differ from each other in only one or few leaf anatomical characters (including oil

and mucilage cells). A combinations in character distribution were observed. However,

great many

the distribution in approximately one-fifth of all studied species deviated largely from the
pattern

typical leaf anatomical character distribution in the of Cinnamomum

pattern occurring majority

species. There was a maximum of seven differing features out of the sixteen features studied. Within

this almost all neotropical Cinnamomum species are included. The latter species lack scleri-
group

fied epidermal cells and almost all have penninerved instead of the generally occurring triplinerved
leaves.

Cluster analyses based on all leaf anatomical features studied revealed that the distribution pat-

terns of the oil and mucilage cells play a significant part in the grouping of the species. Therefore

oil and mucilage cells possess at least some diagnostic value within the genus Cinnamomum. The

systematic significance of oil and mucilage cells at the infrageneric level remains uncertain for lack

of a detailed infrageneric classification of Cinnamomum for comparison with the idioblast distribu-

tion patterns.

Introduction

In a previous study on oil and mucilage cells in species of the genus Annona (Bakker
& Gerritsen, 1992) we explored the systematic value of these idioblasts. Since the

distribution of the idioblasts showed large variation within of species it was

groups

concluded that oil and mucilage cells had little or no taxonomic value below the

genus level. This can partly be understood in the light of the proposed homology of

oil and mucilage cells (Bakker & Gerritsen, 1989, 1990, 1992; Bakker et al., 1991).
2 BLUMEA VOL. No. 1992
37, 1,

In the the diagnostic (discriminating) and systematic value of oil and

present paper

mucilage cells will be investigated for the genus Cinnamomum Schaeffer.

Cinnamomum is a genus with about 200 accepted species (Kostermans, 1964,

1986,1988). A number of neotropical species previously placed in other genera such

and Phoebe have been transferred Cinna-

as Oreodaphne, Persea, Laurus, Ocotea, to

momum (Kostermans, 1961, 1988; Van der Werff, 1991a, b). The not neotropical
Malesian Australia, Asia, the Pacific Islands
species are present in the area, and

1961, 1968, 1969, 1970a, b, 1983, 1986, 1988; Hyland, 1989). The
(Kostermans,

genus is especially known for the production of cinnamon and camphor (Pax, 1889;
The of the bark dates from
Hegnauer, 1966; Kostermans, 1986). use (cinnamon) an-

cient times (see Kostermans, 1986). Nowadays, the plants are also used as medicine

and in the cosmetic and perfume industry.

Leaf anatomical descriptions of Cinnamomum species are scarce. General notes

Pax Solereder (1899), Metcalfe & Chalk (1950) and Metcalfe

were given by (1889),
Perrot (1891) described the histology of oil and mucilage cells in species of
(1987).
Lauraceae, including five Cinnamomum species. Later only two detailed leaf ana-

tomical studies were published for a restricted number of Cinnamomum species

(Santos, 1930; Marlier-Spirlet, 1946).

The genus Cinnamomum is poorly understood and poses

many
taxonomic diffi-

culties. The two sections distinguished by Meissner (1864): Malabathrum Meissn.

and Camphora Nees, are still accepted nowadays as the only subdivision of the

genus (Kostermans, 1986). The sections, besides their different smells, also can be

distinguished by bud morphology and leaf

arrangement (Kostermans, 1986).

In the the distribution of oil and mucilage cells and other leaf ana-
present paper
tomical features are described for 150 species of Cinnamomum. A comparison of the

leaf anatomical character distribution in the species is carried of cluster

out by means

analysis and the results are discussed in the light of the existing classification in order

to evaluate the diagnostic and/or systematic value of the oil and mucilage cells below

the
genus level.

MATERIAL AND METHODS

Material and fixation

The 207 specimens of the 150 species of Cinnamomum studied are listed in Table 1.

In this study leaves from herbarium vouchers, kept in Leiden, were boiled in water

until they permanently sank in water, and then were stored in a FAPA-solution.

Fresh leaves of C. burmanni, C. camphora, C. daphnoides, C. iners, C. kina-

baluense, C. loureirii, C. obtusifolium, C. parthenoxylon, C. sericeum, C. tamala,

C. verum, and C. spec., grown in greenhouses of several Botanical Gardens in

Western Europe (in Table 1 marked with an asterisk), were collected and fixed

in FAPA-solution. Fresh leaves from C. burmanni (Leiden, Java), C. camphora

(Berkeley, Bogor), C. iners (Wageningen, Bogor), C

i . kinabaluense (Bogor), C.
and C. fixed in modified
parthenoxylon verum (Delft) were
Karnovsky fixative fol-
lowed by OSO4 and embedded in Epon (see Bakker & Gerritsen, 1989).
 M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Leaf anatomy of Cinnamomum 3

Table 1. List of material examined of Cinnamomum.

All material is herbarium material located in L.

altissimum Kosterm.: Chelliah FRI 6543, Malaysia — amoenum (Nees) Kosterm.: Smith &

Klein 11260, Brazil —

angustitepalum Kosterm. 1: Kostermans 12974, Borneo —
angusti-

2: Brain anak Tada S 16253, Borneo archboldianum Allen 1: Craven & Schodde
tepalum —

1239, New Guinea —

archboldianum 2: bb. 25066, New Guinea —
archboldianum 3: Koster

BW 6854, New Guinea —

archboldianum 4: Streimann & Stevens LAE 54854, New Guinea —

archboldianum 5: Foreman & Vinas LAE 60127, New Guinea —

archboldianum 6: Schodde

& Craven 5014, New Guinea —

archboldianum 7: Aet & Idjan 394, New Guinea —
archbold-

ianum 8: Carr 14814, Papua New Guinea —

archboldianum 9: Carr 13253, Papua New Guinea

—
archboldianum 10: Stevens LAE 54769, New Guinea—- aromaticum Nees: Schewe s.n., s.l. —

aureo-fulvum Gamble: Wong Khoon Meng FRI 32230, Malaysia —

baileyanum (F. Muell.)

Francis: Hyland 7547, Australia —

bejolghota (Ham.) Sweet: Schewe s.n., Asia —

bintulense

Kosterm.: Ding Hou 349, Borneo —

bodinieri Gamble/Lev.: Beauvais 132, China —

brevipedun-
culatum Chang: Ching-en Chang 6643, Taiwan —
bullatum Kosterm.: Womersley NGF 11335, New

Guinea —
burmanni (Nees) Blume 1: Schewe s.n., Japan —
burmanni 2: de Vriese & Teijs-

mann s.n., Sumatra —

burmanni 3: de Vriese & Teijsmann s.n., Java —
burmanni 4: Boer-

lage 1888, Java —

burmanni 5: Keith For. Dep. No. 5999, Borneo —
burmanni 6: Perrottet

s.n., Philippines —
burmanni 7: Teijsmann s.n., Celebes —
burmanni 8: Schmutz 2836,

Lesser Sunda Islands burmanni 9: Iboet Lesser Sunda Islands burmanni 10: Kos-
—

514, —

termans s.n., Java —

burmanni 11*: Hortus Botanicus Leiden, The Netherlands —
burmanni

12*: Baas s.n., Cibodas, Java —

calciphilum Kosterm.: Paul et al. S 37382, Borneo — cam-

bodianum Poilane 23247, Cambodia 1: d'Aleizette

Lec.: —

camphora (L.) Presl 558m, Japan

—
camphora 2*: Bot. Gardens Kew No. 00073 12261, U.K. —
camphora 3*: Royal Bot.

Garden Edinburgh No. 69 6417, U.K. —

camphora 4*: Bot. Garden Techn. Univ. Delft, The

Netherlands 5*: Bot. Garden Utrecht No. The Netherlands

—

camphora 73GR00538, — cam-

6*: Van der Werff s.n.,

phora Berkeley, U.S.A. —

camphora 7*: Bot. Garden Bogor No. XXB

112, Java camphora 2*: Baas Cibodas, Java 9*: Gardens

—
s.n., —

camphora variegata Bot.

Kew No. 00073 12262, U. K. cappara-coronde Blume: Kostermans 25596, Ceylon

— —

caryo-

phyllus (Lour.) Moore: Bordeneuve (Chevalier 40969), Indochina —

celebicum Miq.: bb. 5555,

Celebes —

chittagongense Kosterm.: King 249, India —

cinereum Gamble: Corner Ma-
s.n.,

laysia citriodorum Thw.: Kostermans 23455, Ceylon clemensii Allen: New

— —

Eyma 5093,
Guinea —
cordatum Kosterm.: Whitmore FRI 20470, Malaysia —
coriaceum Cammerl.: bb.

14079, Borneo —
corneri Kosterm.: Carson SAN 28012, Borneo —
crassinervium Miq.: Puasa,
Borneo
B.N.B. For. Dep. 3159, —
crispulum Kosterm.: Poilane 21913, Indochina —
cubense

(Nees) Kosterm.: Sintenis 1036, Puerto Rico culitlawan (L.) Kosterm.: Blackw.t.?
—

Labill.,

391, Moluccas —
curvifolium (Lour.) Nees: Poilane 32.126, Indochina —
cuspidatum Miq. 1:

Suppiah FRI 11901, Malaysia —

cuspidatum 2: de Wilde & de Wilde-Duyfjes 12889, Sumatra

damhaense Kosterm.: Chevalier 37468, Asia daphnoides Sieb. & Zucc. 1: Koidzumi
— —

s.n.,

Eur./Asia —

daphnoides 2*: Bot. Gardens Kew No. 00073 12263, U.K. —

deschampsii

Gamble: Corner SFN 31000, Singapore —

dewildei Kosterm.: bb. 2796, Sumatra —
doeder-

leinii Engl.: Fosberg 37185, Ryukyu Islands —

dubium Nees 1: Kostermans 24095, Ceylon —

dubium 2: s.n., Java —

durifolium Kosterm.: Poilane 9030, Asia —

ebaloi Kosterm.: Ebalo

1198, Philippines —

effusum (Meissn.) Kosterm.: Sousa 7876, Mexico —

elephantinum Kos-

ellipticifolium Kosterm.: Balus? Burma elon-

term.: B6jaud 742m, Cambodia —

12288, —

West Indies
gatum (Vahl) Kosterm.: Curtiss 309, (Central America) —
englerianum Schewe:
Ledermann 9806, New Guinea —
eugenoliferum Kosterm.: bb. 32907, New Guinea —
fouil-
4 BLUMEA VOL. 37, No. 1, 1992

(Table 1 continued)

loyi Kosterm.: Endert 3840, Borneo —

frodinii Kosterm.: Henty & Katik NGF 49456, New Guinea

—
gigaphyllum Kosterm.: Telussa BW 5161, New Guinea —
glaucescens (Wall.) Drury: Poilane

1272, Asia Kosterm.: Stocks s.n., Asia grandiflorum Kosterm.: Hoogland 5052,
—

goaense —

New Guinea —
griffithii Meissn. 1: J.C. 1641, Malaysia —
griffithii 2: Kostermans 5283,

Borneo —
iners Reinw./Blume 1: Pierre 5170, Asia —
iners 2: Kostermans? s.n., Java —

iners 3*: Bot. Gard. Agricult. Univ. Wageningen No. 72 PTO 1275, The Netherlands —
iners

4*: Bot. Gard. Bogor No. XXB74, Java —

insulari-montanum Hay.: Kao 9834, Taiwan —

javanicum Blume 1: bb. 2729, Sumatra javanicum 2: Java kami Kosterm.: Frodin
—

s.n., —

NGF 28255, New Guinea —

keralaense Kosterm.: Kostermans 26283, South India —
kinabalu-

ense Heine 1: Aban Gibot SAN 66838, Borneo —

kinabaluense 2*: Bot. Gard. Bogor No. XXA

85,Java —
kunstleri Ridley: Poilane 31019, Indochina —
laubatii F. Muell.: Irvine 1668, Aus-

tralia —
ledermannii Schewe: Powell UPNG 1639, Papua New Guinea —
litseafolium Thw.:

Davidse 8455, Ceylon —

longitubum Kosterm.: Poilane 29260, Asia —
loureirii Nees 1:

Yoshida 2201, Japan —

loureirii2*: Royal Bot. Gard. Edinburgh No. 100 029, U. K. —
lucens

Meebold 14876, Burma Hook, f.: Kostermans 24534, South India —

Miq.: —

macrocarpum

Kostermans 1230, Moluccas Kosterm.: Poilane 29510,

macrophyllum Miq.: —
magnifolium
Indochina —
mairei Lev.: s.n., Asia —
malabaricum Garc.?: Ridsdale 495, Asia —
malabath-

rum
(Burm.f.) Blume: Kostermans 26119, South India —

malayanum Kosterm.: Kochummen

FRI 16657, Malaysia —

melastomaceum Kosterm.: Poilane 29636, Indochina —
melliodo-

rum Kosterm.: Sayers NGF 24238, New Guinea —

mendozae Kosterm.: Mendoza PNH 42310,

Philippines mercadoi Vid.: Merrill Sp. Blanc. No. 758, Philippines Kos-
— —

microcarpum

term.: Martin S 38157, Borneo —

microphyllum Ridley: Whitmore FRI 15582, Malaysia —

mollissimum Hook, f.: Corner SFN 30885, Malaysia — montanum (Sw.) Berchth. & Presl:

Krug & Urban 5259, Jamaica —

myrianthum Merr.: Conklin PNH 37897, Philippines —

myr-

tifolium Kosterm.: Chevalier 35.993, Cambodia —

nalingway Kosterm.: Kostermans 270,

Asia nooteboomii Kosterm.: bb. 6628, Sumatra Nees *: Bot. Gard.

— —

obtusifolium Royal

Edinburgh No. 35 0159, U. K. —

oliveri F. M. Bailey: Tadman s.n., Australia—- osmeophleum
Kan.: Kao 9650, Taiwan —
ovalifolium Wight: Hoogland 11550, Ceylon —
ovatum Allen:

Jean 38.397, Indochina pachyphyllum Kosterm.: Sinclair & Kish bin Salleh SFN
—

39922,

Malaysia paiei Kosterm.: Paie & Mamit S 29332, Borneo Kosterm.: Martelino
— —

panayense

& Edafio BS 35653, Philippines parthenoxylon Meissner *: Bot. Gard. Bogor No.
—

XXB41,

Java perrottetii Meissn.: Kostermans 25803, India piniodorum Schewe: BW

— —

Versteegh
3919, New Guinea —
podagricum Kosterm.: Sayers NGF 21699, New Guinea —
poilanei

Kosterm.: Poilane 33772, Indochina S

—
politum Miq.: Smythies 7803, Borneo —

polyadel-

phum (Lour.) Kosterm.: Poilane 28504, Indochina Kosterm.: de Wilde &

—

porphyrospermum

de Wilde-Duyfjes 12984, Sumatra Kosterm. 1: Bodinier China

—

porrectum (Roxb.) 1118, —

porrectum 2: Burkill KEP 76692, Malaysia —

porrectum 3: Rahmat si Toroes 4694, Sumatra

propinquum F.M. Bailey: Hyland 6577, Australia pseudopedunculatum Hay.: Tuyama

—
—

s.n., Bonin Island —

psychotrioides (H.B.K.) Kosterm.: Tokatia a/1541, Mexico —

racemo-

sum Kosterm.: Shea & Minjuk SAN 76182, Borneo reticulatum

—

Hay.: Chang 3771, Taiwan

rhynchophyllum Miq. 1: Thorenaar 52, Sumatra 2: Clemens

rhynchophyllum 22194,
—
—

Borneo rhynchophyllum 3: Sinanggui SAN 56217, Borneo 4: Gibot

— —

rhynchophyllum
SAN 37019, Borneo —

rhynchophyllum 5: Kostermans 4559, Borneo —

rhynchophyllum

6: de Wilde & de Wilde-Duyfjes 15545, Sumatra 7: Forbes

—

rhynchophyllum 2969, Sumatra

rhynchophyllum 8: Ogata KEP 105031, Malaysia rhynchophyllum 9: Grashoff 733,

—
—

Sumatra riedelianum Kosterm.: Riedel Brazil rigidum

—
s.n., —

Gillespie: Webster & Hild-

roth 14198, Fiji riparium Gamble: Ridsdale 628, South India Kosterm.: Kos-
— —

rivulorum

termans 27831, Ceylon Kosterm.: Ridsdale SMHI 264, Philippines scalarinerve

—

rupestre —
 M.E. A.F. Gerritsen & P.J. van der Schaaf: Leaf of Cinnamomum 5
Bakker, anatomy

(Table 1 continued)

Kosterm.: Chevalier 38.371, Indochina —

scortechinii Gamble: Poilane 11143, Asia —
sellowi-

anum (Nees & Mart.) Kosterm.: Hatschbach 17399, Brazil —

sericans Hance: Pierre 5170, Asia —

sericeum Siebold *: Royal Bot. Gard. Edinburgh No. 69 6418, U.K. —

sessilifolium Kan.:

Hosakawa 5954, Ponape. Pacific —

simondii Lec.: Chevalier 32075, Indochina —

sinharajense

Kosterm.: Kostermans 27486, Ceylon —

sintok Blume: Kostermans s.n., Java —
solomonense

Allen: Waterhouse Solomon Islands soncaurium Kosterm.: Hansen & Smiti-

129A-B, —

(Ham.)
nand 12882, Thailand —

spec.*: Royal Bot. Gard. Edinburgh No. 69 6352, U. K. —

subavenop-

sis Kosterm.: Boschproefstation Cel/IV-94, Celebes —

subavenium Miq. 1: Put 3526, Thailand

—
subavenium 2: Kostermans 37, Sumatra —
subcuneatum Miq.: Lorzing 11289, Sumatra —

subpenninervum Kosterm.: Poilane 9680, Asia —

sulphuratum Nees: Hook. f. & Thomson

2617, Asia —
tahyanum Kosterm.: Othman etal. S 41520, Borneo —
tamala (Ham.) Th. Nees &

Eberm. 1: Tatemi Shimizu et al. T-20932, Thailand —

tamala 2*: Bot. Gardens Kew No. 00073

12264, U. K. —
tampicense (Meissn.) Kosterm.: s.n., Mexico —
tazia 'Hamilton' Hook, f.: Bot.

Garden Calcutta s.n., Asia —

tepalinum Kosterm.: Koelz 29958, India —
tonkinense (Lec.)
Chev.: Poilane 32261, Indochina —
trichophyllum Quis. & Merr.: Meijer 9759, Celebes —

triplinerve (R. & P.) Kosterm.: Dombey 201, Peru —

tsoi Allen: Poilane 24801, Annam —

vaccinifolium Kosterm.: Sleumer & Vink BW 14254, New Guinea —

velutinum Ridley:
Whitmore FRI12116, Malaysia —
verum Presl 1: Hasskarl s. n., Japan —
verum 2: Wight 2511,
India —
verum 3: Sedies? s.n., Mauretania —
verum 4*: Bot. Gard. Berlin-Dahlem, Germany —

verum 5*: Bot. Gard. Techn. Univ. Delft, The Netherlands —

verum 6*: Bot. Gard. Agric. Univ.

Wageningen, The Netherlands —

vesiculosum (Nees) Kosterm.: Klein 3.269, Brazil —
vimi-

neum Nees: Loh FRI 19170, Malaysia —

virens R.T. Baker: Jones 2693, Australia —
wightii
Meissn.: Kostermans 25731, India —
xanthoneurum Blume: Versteegh & Vink BW 8309, New

Guinea —
xylophyllum Kosterm.: Kostermans 12981, Borneo —
zollingerii (de Lukm.) Kos-

term. 1: Walker et al. 6059, Okinawa Island —

zollingerii 2: Fosberg 38155, Ryukyu Islands.

Sectioning, staining and microscopy

The methods used for sectioning and staining of the leaves are identical to those

described for Annona leaves in Bakker & Gerritsen (1992). Thick (30 pm) transverse

sections of FAPA-fixed leaves stained for oil with Sudan IV

were or Chrysoidin/
Acridin red. Mucilage was stained with Alcian Blue. For suberin the Berberin and

Anilin Blue fluorescent staining method was applied. Ultrathin and 1 pm sections

of Epon-embedded material examined with transmission electron

were microscopy

(TEM) and light microscopy (LM) respectively (see Bakker & Gerritsen, 1992; Bakker

et al., 1991).
Some leaf fragments were air-dried, gold-sputtered and examined with the scan-

ning electron microscope (SEM).

Examination and measurements

In the 30 pm sections the number of oil and/or mucilage cells in the

palisade and/
or spongy parenchyma was counted and calculated per mm leaf width. In the same

sections the lamina thickness measured and the number (per leaf and
was mm width)

length of hairs was determined (see Bakker & Gerritsen, The

1992). presence/ab-
sence of an
indumentum was verified by examination of the leaves with a stereo mi-

croscope and partly with a scanning electron microscope (SEM).

6 BLUMEA VOL. 37, No. 1, 1992

Phenetic analysis
Phenetic analyses were carried out with the help of NTSYS-pc, version 1.30

(Rohlf, 1986), a for numerical taxonomic and multivariate analy-

computer program

ses. Each analysis consists of a two-step procedure, i.e., the computation of the over-

all similarity among

taxa and the representation thereof by a phenogram. In the first

datamatrix, based Table 2, is used calculate the

step a on to (dis)similarity among

expressed Manhattan distances. In the second the

taxa as
step, creating a phenogram,

pairwise distances among taxa serve as input for the analysis, using UPGMA (Un-

weighed Pair-Group Method using arithmetic Averages) as a cluster criterion.

RESULTS

Oil and mucilage cells

30 pm sections -

Light microscopy

Sudan IV —
Oil cells were recognized by the stained contents (red, orange, pink,

brown; depending on the species). The oil is as small irregular bodies (Fig.
present

1A), clustering globules (Fig. IB), a distinctly stained mass (Fig. 1C), or a homo-

geneous oil drop. The oil drop may be visibly attached to the wall by a cupule (Fig.

ID). Other oil cells did not contain oil but could be distinguished by the presence of a

cupule (Fig. IE) and/or a characteristic fold in the cell wall (Fig. IE, F). These oil

cells had a glossy pink Mucilage cells were unstained and empty. The
appearance.

suberized layer was visible as a thin red line in the oil and cells.
mucilage

Chrysoidinlacridin red —
The oil cells were recognized as described for the

Sudan IV-staining, but generally the idioblasts were less clear because of the
strong

background staining. Mucilage cells were unstained (empty) or faintly yellowish.

Alcian Blue —
Oil cells stained glossy greenish-blue or were empty (Fig. 2A) and

could be distinguished from the mucilage cells by a thick dark blue cell wall, sometimes

containing cupule, by the ring-like

a or appearance of the cytoplasm. In mucilage cells

the retained mucilage distinctly stained blue and sometimes extruded from the cells

(Fig. 2A, B, D). The mucilage often showed (circular) striation (Fig. 2C). The presence

of elder pith fragments (used to clamp leaf fragments for sectioning) sticking to the

sections also indicated the presence of mucilage in the leaf. In control samples the muci-

lage was stained less densely and generally spread out over the section as a
result of

the dissolution of the in water, the

mucilage thereby obscuring mesophyll (Fig. 2D).

30 pm sections -
Fluorescence microscopy
The cell walls of oil and mucilage cells showed autofluorescence of the suberized

wall layer. When stained for suberin (Berberin) this layer appeared
specifically as a

thin yellow/whitish thin line in the cell wall. This reaction has been describ-
staining
ed earlier for oil and cells of Cinnamomum burmanni (Bakker al„
mucilage et 1991).

1 pm sections -

Light microscopy
Both idioblast types were easily distinguished from each other by their overall

cytoplasmic composition. Oil cells were also characterized by the presence of a

cupule. Mucilage cells showed the distinctly purple-stained mucilage and/or the

characteristic pattern of the cytoplasmic strands.

 M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Leaf anatomy of Cinnamomum 7

Transmission electron microscopy

The suberized wall layer was already observed against the inner side of the cell

wall in a young oil cell in the leaf of Cinnamomum camphora (Fig. 3A) and turned

out to be always present in mature oil (Fig. 3D) and mucilage cells. The composition
of the of the oil and mucilage cells in the leaf is similar that de-
cytoplasm very to

scribed for the idioblasts in the shoot apex of C. burmanni (Bakker & Gerritsen, 1989;
Bakker et al., 1991). In a very young oil cell, present in the palisade parenchyma,
electron dense oil droplets and characteristic plastids were present in the cytoplasm

(Fig. 3B). The plastids lacked thylakoids but contained starch, dark globules and

very small white globules (Fig. 3B). In another young oil cell in the palisade paren-

chyma a thickened part of the inner wall layer was observed: the cupule base (Fig. 3C).
In mature oil cells in leaves of different Cinnamomum species a cupule was generally
observed (Fig. 3D), showing the cupule base from which the cupule projects into

the cytoplasm surrounding the oil cavity. These observations are identical to those

described earlier for cupules in oil cells of C. burmanni (Bakker & Gerritsen, 1989;
Bakker et al., 1991) and Annona muricata (Bakker & Gerritsen, 1990).

Shape and size

In most species the oil cells are oblong-ovoid in the palisade parenchyma (Fig.

1A, C, D) and more or less globular in the spongy parenchyma (Fig. IB, E, F). The

cells ovoid in the and

mucilage generally are
palisade parenchyma (Fig. 2A, B, D)
globular in the spongy parenchyma (Fig. 2B, C). The sizes of the idioblasts vary per

species (Figs. 1A-F & 2A-D). In general mucilage cells are larger than oil cells.

Distribution

Oil and/or mucilage cells are always present in both the palisade and the spongy

parenchyma (Table 2). Most species both oil and mucilage cells in the leaves.
possess

The majority of these species contain the two idioblast in both the palisade and
types
the spongy parenchyma. The exclusive of oil cells in the leaf occurred in c.
presence

17% of the species and only 5 species contained mucilage cells in the
exclusively
leaves (Cinnamomum archboldianum [no. 4 and 5], C. magnifolium, C. nalingway,

C. rivulorum and C. sinharajense).

In the spongy parenchyma idioblasts are usually located against the lower side of

the palisade parenchyma (Fig. 2A, B) and against the abaxial epidermal cells.

Of 18 species of which more than 1 specimen was studied 11 were constant for

the distribution of the oil and mucilage cells in the palisade and spongy parenchyma

(C. angustitepalum [2 specimens], C. burmanni [12], C. cuspidatum [2], C. daph-

noides [2], C. javanicum [2], C. loureirii [2], C. porrectum C.
[3], rhynchophyllum

[9], C. subavenium [2], C. tamala [2] and C. verum [6]). The other species showed

variation in their distribution Small variations concern the presence/absence

patterns.
of oil or mucilage cells in one of the mesophyll layers (C. griffithii [2] and C. zollin-

gerii [2]). Larger variations occur when part of the specimens lacks one type of idio-
blast iC. archboldianum [10], C. camphora [9], C. dubium and C. iners
[2] [4]).
Cinnnamomum kinabaluense [2] showed different distribution
two patterns (Table 2).
The frequency of the idioblast types in the leaf was found to be highly variable
between species (Table 2). Between specimens of the minor vari-
same species only
ations occurred in the number of idioblasts. The maximum number in the
palisade
8 BLUMEA VOL. 37, No. 1, 1992
 M.E. A.F. Gerritsen & P.J. der Schaaf: Leaf anatomy of Cinnamomum 9
Bakker, van

Fig. 2. Transverse 30 thick sections of Cinnamomum species. A—D: Mucilage cells stained with
µm

Alcian Blue. Light microscopy. All x 345 A: C. virens. Mucilage cell in the palisade
—

paren-

chyma showing extrusion of the stained mucilage from the cell into another cell. Also note the

neighbouring unstained oil cell (oc) in the

spongy parenchyma. —
B: C. nalingway. Mucilage cells

in the palisade and spongy both extruding mucilage into the intercellular
parenchyma, spaces of the

parenchyma. Note the darker stained mucilage in the centre of the cells. C: C. vaccini-
spongy —

folium. Mucilage cell in the parenchyma. Note the circular striations in the mucilage. D:
spongy —

C. magnifolium. Mucilage cell in the palisade parenchyma. Note the less densely stained mucilage
extruded from the cell and spread out over the section.

Fig. 1. Transverse 30 thick sections of Cinnamomum species. A—F: Oil cells stained with
µm

Sudan IV. Light microscopy. All x 725.— A: C. cubense. Oil cell in the palisade parenchyma

showing loose pieces of stained oil. B: C. crispulum. Oil cell in the filled
—

spongy parenchyma
with a mass of globules of stained oil. C: C. vesiculosum. Oil cell the lower layer of the
— in
pali-

sade parenchyma filled with a heterogeneous oil drop. —

D: C. tampicense. Oil cell in the palisade

parenchyma filled with a large oil drop. —

E: C. brevipedunculatum. Empty oil cell in the
spongy

parenchyma showing a cupule (arrow) and a fold in the cell wall (arrowhead). —
F: C. vimineum. Large

oil cell in the parenchyma showing the fold in the cell wall.
empty spongy
10 BLUMEA VOL. 37, No. 1, 1992

3. Cinnamomum 8. A—D: Transmission electron microscopy. A: Detail of the cell wall

Fig. camphora
—

of a
very young
oil cell in which only a suberized wall layer (s) has been deposited between the primary

cell wall (w) and the cytoplasm (c); X

58,200. B: Detail of the cytoplasm of oil cell depicted in
young
—

(st), dark globules (arrowheads), and small white globules

A. Note the specific plastids containing starch

Note the oil globule (o) in the cytoplasm; 27,250.

(small arrows). The plastids lack thylakoids. large X —

C: Detail of the cell wall of a young oil

cell showing the development of a
cupule base (cb) deposited

against the suberized wall The base is the thickened of the inner wall which is
layer (s). cupule part layer
the whole oil cell; 27,250. D: Detail of
deposited against the suberized layer throughout X — a developed

cupule in an
older oil cell in the palisade parenchyma. Note the cupule base (cb) from which the cupule

(arrows) projects into the cytoplasm and surrounds the oil cavity (oc); x
16,930.
proper

Fig. 4. Transverse 30 thick sections of leaves of Cinnamomum species. A—F: Stained with Sudan IV.
µm

Light microscopy. — A: C. vaccinifolium. Extremely thick, densely stained cuticle (17 µm) showing an irreg-

ular outer
surface; x 400. — B: C. glaucescens. Adaxial epidermis with large non-sclerified epidermal cells.

Note the cutinized (stained) anticlinal walls; x 400. —

C: C. wightii. Vertically transcurrent lower order

vein with bundle sheath extensions; x 400. D: C. velutinum. Detail of the uppermost
sclerenchymatous —

of vertically transcurrent lower order vein showing the continuation of the sclerified bundle sheath
part a

cells with the sclerified adaxial epidermal cells; x 1,000. —

E: C. sintok. Layer of aligned sclerified outer

periclinal walls of the palisade cells underneath the small epidermal cells; x 400. — F: C. sericeum. Hypo-
dermis underneath the adaxial epidermis. Note the sclerenchymatous cells in this layer (arrow); x 400.
(hy)
M.E. A.F. Gerritsen & P.J. der Schaaf: Leaf of Cinnamomum 11
Bakker, van anatomy
 12 BLUMEA VOL. 37, No. 1992
1,

c
e + + + + + + + + + + + +
+ + + + + +
+ + + + +
+ + +
+ + + + +
+ +
+
+ + f + + + + t

v
V t p
P p
p
t t t t P
p
t P
p
t t p
p pp t
t p
P p
p P
p P
p
t t
t I
t tt pp P
p t
t

3s
3
s

3cs
3cs 2
s
2s 2s Is
2s Is 2s
4t 2s
4l 3t
3t 3s
3
s
2s 2i
2i
ha
ha
1

-It -lc -lc* t—+lc2- -It 13-4 +lc-It 3—+lc-* +lc—-± - 1±lc-It 13-It —It-lc* It3 1
ab. f
f 5
5 22 ?? -
-
2 5
5 1

- 3
4 - 3 _ - - - -
1

- 3 _ - 1
3

-
-
1
1
3s
3s
ha

species. ad.
ad. f 1 - - - - - - _ _ - _ _ - - - - - - - - - - - -
-

Cinamou
+
pa
pa _ - - - - - + -
+++ - ±
+ ± -
-± - ± ± ± +
± ± - - - - - -

-
-

+ ±
+ +
+ +- ± ±
± +
+ + -± ±
+ + ± +
+ +
+ +
+ ± +
+
-

s + - + - - ± -- - t - - - + ± +
s
-

-
in
—1—
3- lc lc lc* lc*- lc*lc* lc* lc*

1-
It 3- lc lc It It 1-1- lc It lc lc 1- 1- 1- lc It It It It It It

features
p
P

hy
hy _ - - - - - - - - - - - - - - - - - - - - - - - -
-

-
++ ++++
++ +- ++++ ++++ ++
++

an tomical
ep +
+
-
+
+
+- +
+ +
+ +
+ ++
-
+
+ ++ +
+ +
+ - - - +
+ +
+ +
+ +
+ +
+ +
+ -

cu
cu 2
2 2
2 2
2 3
3 2 2
2 2
2 3
3
2
2 2
2 2
2 2
2 2
2 2
2 2 2
2 2
2 1
1
1
1 1
1
1 2 2
2
1
1 2 2 2
2 2
2

leaf
of t 2
2 3
3 4
4 4
4 2 2 3
3 3
3
3
3 4
4 2
2 3
3 3
3 3
3 2 4
4
3
3
2
2 2
2 2
2 2 3
2 3
3
2
2 3
3 2
2 3
3 3
3

Distrbuion muc
s
s

P
2
2 - 4 3
3
3 3 4
4 P
P 2
2 2
2 - 2
2 4
4 3
3 3 4
4 3
3 P 2
P 2 - - 3 2
2 2
2 2
2 2 3
3 P

P 2 - 4 4
4 4 4 P 4 2
2 - 3 3 4 3 4 4 2
P 2 2 4
4 2
2 4 2
2 P
4 4 4 1
1 4 3 3 4 3 4 4 - - 1
1

2.

Table
3
3 2 P 2
2 2 P 3
3 2
2 - 4
4 4
4 3 2
2 3 3
3 2
2 P 4 3 3
3 4 4
4 3 2
2 4 4
4 P
P
s
s -

oil
Oi
P 2 P 2 2 2 - 2
2 4
4 P 3 4 4 2
2 - 3 3
3 P
P
-
1 1
1 1 1 1
_ 4
4 - 4
4 2 2 4
4 2 3 4 4
-

-10

1-1-33,6,-610- 5
5
6

5,
1 2 4,
4,

3,
3,

brevipdunclatmwn bulatum burmani caliphlum cambodianum camphora camphora camphora camphora cap r-conde caryophylus celbicum chitagones cinerum citrodrum clemensi
8
8 9

1-12 2, 7, 4,
4,

altis mu m amoenum angustliepaum angustiepalum archboldianum archboldianum aromaticum aureo-fulhvwmn baileyanum bejolghota bintulens bodineir
1

species
 M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Cinnamomum 13
Leaf anatomy of

e + +
+ + +
+ +
+ +
+ + +
+ + + + + +
+ + + + +
+ + + + + + + + +
+ + + + + +
+
+ + + + + + + + + + + + + + + + +

v
V t t t p
P t t t
t t t
t t t t t PP t P
P t
t t t
t P t t

1
li
1
2t 4s 2s 3s 2s 2s
3s 2s 2s 2s
2s 3s li 2s
3s 2s 2s Is
2s Is
ha
ha

ab.
ab. f
f _ - _ _ _ _ 1
_ 11 4 3 _ 3
3 2 3 -
1
12s _ _ 3 - 4 - 2 2 2 _ 2
2

-It— -lc* 2 -It lc* -1 lc*3It—-1 2It 3-lc 3-It - -It2_1 lc* 3-2lc*
1
4s 3s
ha
ha

ad.
ad. f - - — _ _ _ _
1- _ _ _ _ _ _ - _ - _ - - - -
1
1
_ _ _
_

-
pa +
+ - _ _ ±
+ _ _ _ _ +
± _ +
± ±
± - _ _ - +
+
++
++ ±
+ - - - - +
+ -
_

-
_lt±
s +
+ ±
+ +
+ +
+ ±
± ±
± ±
± +
+ +
+ _ + ± ±
± +
+ ± ±
+ - ±
± ±
± ±
± - -- - ±
± +
+ ± ±
± - ±
± ±
±

1--
hy
P
p 2t2t

_
It ItIt

- -
lc* lc* 2c*2c* lc*lc*

_ - _ _
It

-
lc*

-
lc

-
_2c±
2c

-
lc*

-
It

_
It

_
_ic*+
lc*

-
It

-
lc

-
It 2c2c

-
It

-
1-

-
2c*2c* 1- lc*lc*

- - -
It It

- -
lc*

-
1-

-
lc*lc* lc*

- -
-

ep

cu
cu
+
±

3=
3= 2
2
+
+

2
2
+
+ +
+

2
2 2
2
+
+

2
2
_

2
2
+
+

2
2
+
+

2
2
+
+

2
2
+
+

3
3
+

1
1
+
+

2
2
±
±

2
2
+
+

i
1
+
+

2
2
+
+

2
2
+
+

2
2
+
+

3=
3=
- +
+

2
2
2+-
+

2 2
2
-

2
2
+
+

2
2
+
+

2
2
+
+

l
1
+
+

2
2
+
+

1
-

2
2
+
+

1
1
+
+
tt
t

s
s
4
4

4
3
3

4
33

-
4 3
3

4
2
2

2
2

-
2
2

4
2
2

3
2
2

2
2
2

J
4
4

1
3
3

2 2
3 3

2
22

-
2
2

2
4
4

2
3

3
3

-
3
3

4
3
3

4
2
2

-
3
3

5
3
3

4
22

-
2
2

4
3
3

4
3
-

2
2
2

muc
P
p
44 34 54 22
4 3 - 5 2 3
3
44 23 42 ?? 11 32 12 22
4 2 4 ? 1
3 1 2
22 42 43
2 4 4 2
2
44 54 1- 45 44
4 5 1 4 4 -
54 44
5 4 -
42 42
4 4
-

s 1 2
2 5 2 4 4 3 2
2 4 S 3 3 2 4 4 4 1
2 3 22
1
4 3 2 4 3 3 5 3 3

oil
oi
P
P
21
2
25 -2 14 54 23 24 45 43 43 12 34 24 34 11 12 43
2 - 1 5 2 - 2 4 4 4 1 3 2 3 1
1 4 - 1
11 54 53 12 44 33 13 55
5 5 1 4 3 1 5 -
23
2
-

2 2
1,2
1, 1,2
1,

cordatum coriaceum corneri crasinervum crispul m cubense culitaw n curvifolum cuspidatum damhaens e daphnoides deschampsi dewildei doe rlein dubium dubium durifolium eebalbaloi ef usum el phanti um eliptcfolium elongatum englerianum eug noliferum fouiloyi frodini gi aphylum glauces ns goaense grandiflorum
1
1
2

species
 BLUMEA VOL. No. 1992
14 37, 1,

e
e + + + + + + +
+ + + + +
+ +
+ + + + +
+ +
+ +
+ + + + + + + + +
+ + + +
+ + + + + + +

v
I t I t t t t t t t 1
t t t t t t t 1 t t t t t t t t t t t t t t
v

3i 2i 2s 3s
3s 3s 5s 2s
2s 4t 3t 3s
3t 3s 3s 2t 2s 2s
2t 3s Is 2t
ha 1

ab. f _ 3 3 2 - 4 - _ 4
4 3 2 _ 3 _ -
-3 2 2 3 1
3 - 3
3 2
2 _ - 1

-
I t
-t 3*-±2lc 4-2 3-It2 1—3- ltc-3±2 *- 3-It -12sIt -t 1
1
4i
4i 2s 3i
3i
ha
ha

ad.
ad. f
f _ - - - - - 3
3 - - - - - - - - - _ _ - 1 1
1 _ - -
-

-
1

-
-
pa _ - ±
+ ± - ±
± - - - - +
± - - - ± - - - - - ±
± - - - - -
-

-
-

s
s ±
± ±
±
±- - - +
+ ±
± ±
± ±
+ - +
+ +
+ ±
± - - +
+ - +
+ +
+ +
+ ±
± ±
± ±
± ±
± - +
+ +
+ +
+ +
+

lclc
lc*lc* lc* lclc lc ItIt 2c
lc*lc* 2c2c 1- lt
lc*lc* It
lc*lc* lclc ltc lc lc 2c* lc*lc* It
lc*lc* It It 1-1- lc*lc* It It It 2t2t
P
p

hy
hy
_
- - - - - - - - - - - - - - - - - - -
- - - - -

-
-

+ + +
ep
ep +
± +
+ +
+ ++ +
+ +
+ +
+ +
+ ++ - +
+ +
+ ++ +
+ +
+ +
+ +
+ ++
++
t + +
+ ++ +
+ +
+ +
+
++
+ ++
++
+
-

32+
cu
cu 1
1
1
1 2
2 2
2 2
2 2
2 3 2
2 3
3 2
2 2
2 2
2 2
2 2
2 2
2 2
2 2
2 2
2 2
2 2 2
2 2
2 2
2 1
1 2
2 2
2 2
2 2
2 2
2
-2-
t 2
2
-3t-
2
2 2
2 33 3
3 2
2 4 3
3 4 2 3
3 2
2 2
2 4 3 2
2 2
2 3
3 4
4 3 3 22 3
3 1
1 4
4 3 2
2 2
2 33

s
s 2 3 2 - 4
1

4 4 4 - 1 2 3 2 2 2 1 2 2 - - 3 P 5
5 3 2 2 -
3

muc
P
p
32 43 22 44 41 54 44 44
3 4 2 - 4 4 5 4 4 -
11 42 53 32 43 22 32 21 32 22
1 4 5 3 4 2 3 2 3 2 - -
43 PP 43 32 32
4 P 5
5 4 3 3 -

- 2 2 1 2 - 2 P 2
2 3 4 4 3
s
s 4 3 3 4 4 5 3 3 3 4 3 2 4 2 4 4

oil
oi
P
p
24 1- 32 42 13 23 34 24 -1 45
2 1 3 4 1 2 3 2 - 4 1
13 13 23 34 23 22 44 22
1 2 3 2 2 4 2 - 3
34 24 12 -p 23 24 14 43
2 1 - 1
1 2 2 1 4
-

conti ued)
insular-motnu javnaicum keralense kinabluens kinabluen.s kunstleri laubati ledrmani litseafolium longitubmu loureiri
2 1 2

1,
1, 2

macro pum macrophylum magnifoluim mairei malbricum malbthrum mal yanum melasto mceu meliodrum mendozae mercadoi microarpum
1
1 2
2
1,2
1,

2
1-3
1-3 ■ 4
4

(Table species grif ithi grif thi hi iners iners kami /ul ce/in.?s
 M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Leaf of Cinnamomum 15
anatomy

e
e + + +
+ + +
+ + + +
+ + + +
+ +
+ +
+ + +
+ +
+ +
+ +
+ + +
-t- + + + +
+ +
+ +
+ +
+ + +
+ +

v
V — t t
P
p t t t t P
P t t t t t
t t t t
t t P
p t P
P t t t

s s
2s 3s 2s
3s 2s 2s 2s
2s 2s
2s 2t 4i
4i 3s 2s
2
2s
2s 3i
3i
3
ha 1

ab. 4 3 4 - - 3 3 - - - 4
4 - 1 - - 3 - 22 - - - - -1-1 - - 22 -
-3
f

-
Is

-It34- It53- lc* -+ —lct-I - 13It- -±lc*It- lc t- lc -132

1
li
li 4t
4t
ha

ad.
ad. f 2
2 - - - 1 1 - - - 3 - - - - - - - - - - - - - - - - - - -
-

pa _ +
+ + - - +
+ - + - ± +
+ ±
+
++
++ ±
± - +±
++
++ - - - - ± -
-±-± - - - - -±
-±
± ±

-
+ + ± ± - X ± - ± ± + - + - + + - ± - - ± - - - +
s
s ± ±
± ±
± ± ± ± ± ± ± ± ± + ±
+ + + + ± + ±

ItIt It 2c*2c* llIt lc*lc* It lclc

lc* lc* lc
lc*lc* lc ItIt ll ItIt 1- ltIt It lc* lc* lc*lc* ItIt lc* It Ic- lc 2c*2c* 1- lc lc lc*lc*
P
p

hy _ - - - - - - - - - - - - - - - - - - - - - - - - - - - -
-

ep + +
+ +
+ +
+ ++*+ + + +
+ +
+
++ ++f+
++ + +
+ - +
+ +
+ +
+ +
+ +
+ +
+ +
+ - +
+ ±
+ -
++
++ +
+
++
++
+ - +
+ + + +

cu
cu 2
2 2
2
2
2 2
2 2 2
2 2 1
1
2
2 2
2 2
2 2
2 1
1
2
2 2
2
1
1 2
2 2
2
2
2 2
2 2
2 2
2 2 2
2 2
2 3
3 2
2
1
1 2
2 1
1
-
-t t
t 2 2
2 2
3 22 2
2 3
3 3
3 3
3 2
2 33 3
3 2
2 44 3
3 3
3 33 3 2
2 3 2
2 2
2 2
2 3
3 3
3 3 3 3
3 4 2 3
3

2
1

2 - 1
1
4
4 - 3 4 P 4 -
s 4 2 5 2 2 4 - 2 4 3 2 2 4 2 3 3
-

muc
P
p
24 52 42 21 22 22 24
2 - 2 2 2 -
44 - 4
43 22 22 44 42 11 43 43 21
2 2 4 4 2 3
1- 43 54 PP 54
4 5 P 5 -
-
1
1
5 4 2 5 1 4 4 4 1

2 4 3 2 2 P 5
5
s
s
5 4 3 4 4 - 5 4 4 4 4 33 4 4 4 4 3 3 2 3 1 3 3 2

oil
oi

P
p
45 44 33 24 -4
4 4 3 2 - - 3
35 34 44 14 34
3 4
1
3 1
14 24 14 34 23 12 33 -2 -3 -1 24
2 1
3 2 1 3 - - - 2 2
2 3
32 43 22 pp 12
4 2 P 1 4
4
-

1-10

microphylum molismu montaum myrianthum myrtifolum nalingway notebomi obtlusifolum oliveri osm.eophleu.mrn ovalifoulim ovatum pachyp lum panyens parthenoxyln peroteti pinod rum podagricum poilanei politum polyadelphum porhy spermu porectum proinqum pseudo nclatum psychotri des racemosu m reticulatmu rhyncopylum
1-3
1-3

species paiei
 16 BLUMEA VOL. 37, No. 1992
1,

c
e +
+ +
+ + +
+ +
+ +
+ + + +
+ +
+ + +
+ + + + + +
+ + + +
+ +
+ +
+ + + +
+ +
+ +
+ +
+
+

t t t t t t t t t t
V
v Pp t t P
p t t t t t t t t P
P t t P
P P

1
2s 4i
4i 3s
3
3s
s
3cs 3s
i

li
1
3s
3
s
2t 2s ?
3i
3i 2i
2i 3s
ha 1

ab.
ab. - 44
3 _ 2 - _ - 3 22 2
1? - 2 - _ 2 _ _ 2
f 2 - 4 3 3 3 _ 3 1
2

_
-

-
I t _-
-It 2lc-+4It±_3 -2c*It+ -_+lc It -lc2 2- lc* It 1lc*t 2
—- l c *
1 ?

ha

ad. f
f - _ - _ _ - - _ _ _ 1
1? _ _ _ - - - _ - _ _ _ -
_ - _ -
_

pa - +
+ - + + +
+ ±
+ + - - ± - _ _ ±
± - -
-+
-+ _ - +
+ - - - _ - _ +
+

s
s +
+ ±
± ±
± - - - _ ±
+ - - ±
± _ +
± ±
± +
+ ±
± ±
± -
±-
+- _ ±
± ±
+ +
+ - - - +
+ - +

It-
P
p

hy
hy
lc*
lc* It

-
lc*lc*

-
lc

-
lc*
lc*

-
It

-
-lc*-
lc* 2c*

- -
It

-
1-
1-

+
+
lt

-
lc lclc

- -
It

-
lc*

-
lc

-
lc*

-
lc lc

- _
It It

- -
It It

- -
lc*

-
1-

-
lt

-
It

_it
-
lc* lc*

-
_ic*+
-

++-
ep

cu
cu 1
1_
_

2
2
+
+

2
2
±
±

2
++
++

2
++

1
1
-_ +
+

2
+

3-
3
-

2
2
+
+

1
1
+
±

2
2
+
+

1
1
-

2
+
+

1
+
+

1
++

2
+
+

1
1
++

2
2
+
+

2
2
+
+

2
2
+
++
+

2
+
+

2
+
+

1
+
+

2
2
+
+

1
1
-

2
+
+ +
+

1
i 2
2
+
+

1
l
+
2t-
3 3 22 2 4 2 3 3 3 3 3 2 4 3 2 2 3 2 2 2 3 3 2 2 2
2 2 3 2 2
2
t
3 3 4 2 3 3 3 3 3 2 4 3 2 3 2 2 2 3 3 2 3 2

s
s 3 - 4 4 4 4 -
1

2 4 - 4 1 3 2 2 2
1
1 2 4 2 - 2 1 2 3 3 2

muc
muc
P
p
43
4 - 3
34 54 14 24
5 1 2
-
21 22 44 54 21 43 32 22
2
2 4 - 5
2
4 3 2 - 1
1
42 34 22
4 3 2 -
12 41 22 43 23 42
1
4
2
4
2
4

3 5 4 - 2 4 3 2 2 - 2 - 2 4 1 4 4 4 4 4 4 2 2 3
s
s 3 3 4 3 4 4 4

oil
oi

P
p
33 -5 34
3 3 -
22 24 33 52 42 23 23 44
2 2 3 5 4 2 2 4 -
22 1- 22 23 24
2
1
2 2 2 2
2
11
1
- 3
34 34 24 24 24 22 23
3 2 2 2 - 2 2
-

conti ued)
2

1,

subaveniops subavenium subcnealtum subpeni rveum sulphuratm tahijanum tamala tampicense tepalinum lotnkiens e trichopylum
2

1,2

ried lianum rigidum riparium rivulorum rupestre scalrinevre scortechin selowianum sericans sericeum se ilfoum simondi sinhar jens sintok solmones soncaurim spec.
1,
2

(Table species tamala tazia

 M.E. A.F. Gerritsen & P.J. van der Schaaf: Leaf of Cinnamomum 17
Bakker, anatomy

=
1
4 = =
e
e +
+ +
+ + +
+ + +
+ +
+ +
+ + +
+ +
+ +
+ +
+ Pi- ab-
?

periclnal
2
pa

V
v t t
t t t t P tt P
P t tt t l
t tt
absent;
=
hairs/m ; 50
pm; pm;
300
intermdiate less)
or
veins:
sclerifed. papilae. outer
1-2
<

=
>

=
i
=

(more
order
frequncy;
1

domed
s = 5

-— - -=
2s
2s 3s It
It 3s It
It = =

thin-waled;
3
ha
1
1
3

length: pm;
t

lower
sclerifed; club-shaped hairs; hair/m ; 20 -30
ab. -2-2

distinct;
22 2 2

lowly
f _ - 3 2 - - - - - -
=
f =

1 =
the
= t

of
±
= vein
0.1-1
hairs/m ;

-2t 32-lc* It- 2I -t lc - -—= —-p —s

ha
ha
1

weakly flat; ++
=
(solid);
abaxial type. main ext nsions
4

=
=
ad.
ad.
pm;
papilae;
f _ - - - - - - - - - - - 2
-

=
1

hair

10 -20 thick-waled curly

=

width; 5-10
±

sclerifed; epidermis:
ha
sheath
pcniervd;
=
ab.
leaf 5

dome-shaped hairs; hair/m hairs/m ;

pa ++ - - - +
± - - - - - -
=
-
-

=
=
s
c

bundle
abaxial present.
3
not =

pm;
thicknes ;
_ +
+ + - ± - +
+ ± +
+ +
+ - -
s
s ± ±

1-2+
=
-

adaxial
undetrmined;
=
=

venation: plincrved. transcur nt

+
+
0.1
P
p
2- 2t 2c 2c* lc* 2c* It It 2
1
It ItIt lc lc
chyma: papilate walls; ha
=
<

=
2-5
=
50-10 wall sent;
=
hy =
_ - - - - - - - - - - -
ad.
-

v e

ep _ +
+ +
+ +
+ +
+ - +
+ +
+ +
+
++
+
++ ++
++ ++ +
+ = = =
=
not ++ not
cell/ paren-
2- spongy periclinal anticlnal
p 4 4

=
=
cu 4= pm; pm;
sclerifed; presnt.
1
2
2 4 2
2
1

1
2 2
2
1
1 2
2 2 2
2 2
2 2.
2

0.1-1
cels/m ;
=

8-13 cells; spongy

3-layerd;
s

20 -30 outer and

=

2
2
= =

parenchyma; width; 5-10 indstinct,

=
2 3
3 4
4 3
3 2 33 2
2 3 3
3 2
2 3
3 3 3
3

epidermis distinctly
t

sclerifed periclnal
3
+
=
=

pm;
3
3

leaf
=

pm;
absent;
moderately 2-layer d; outer sclerif d.
5

type 3-8
adaxial
s 2 2 3 - - 2 5 3 3

and
1
s
_ 4 3 3
=

mucmuc 22 42 44 43 42 35 43 -1 43 palisade
cel/mm cels/m ; idoblast 10 -20
=

thicken d thicken d total y

P
p _ 2 3 - 4 - 4 =
4 4 3 •- 4 3 4 =

pm;
hypodermis:
2
=
ep =
p = 3

0.1 ? +

2-5
unilayer d;
2
<

cells; <

pm;
= with layer)
sclerifed;
4 2 2 4 4 2 4 2
3 4 4 4 3 3

surface. with
s =

mucilage absent; cels/m ; undet rmined;

= 1
=

oil 54 12 23 32 44 44 14 22 23 14 23 12 1 4

100 hy
layer)
layer) (upper
=
P 4 2 2 2

thicknes : ir egular slightly

1 1 1 1 1
p 5 2 3 4
1

<

(upper
=
2:

sclerif d. parenchyma:
1
t

(upper
= =

Table muc 1-2 frequency thicknes : cuticle

=

= ±
=
c
=

=
sclerifed;
parenchyma;
of = pm.
pm;
sclerifed; strongly palisade sclerifed; wall; walls
1 2
cells; 3
c*

vacinfoliwunm veluintum verum vesiculosmu vimnieum virens wighti xanthoeurm xylophylum zolingeri zolingeri Explan tion present, lamina 300 adaxial
1-6 13

triplinerve mm;
oil
> > =

species tsoi
tsoi oil
=

t
=
=

cu
18 BLUMEA VOL. 37, No. 1, 1992

parenchyma was 8 cells/mm leaf width for oil cells (<C. camphora 7) and 10 cells/

mm
leaf width for mucilage cells (C. psychotrioides). In the parenchyma the
spongy

maximum number was 11 oil cells (C. C. rhynchophyllum 8) and 8 mucilage cells/mm
leaf width ( C. englerianum). The frequency of both idioblast types together in the leaf

ranges from 0.5 to over 17 idioblasts per mm

leafwidth (iC. englerianum).

Other leaf anatomical characteristics (Table 2)

Lamina thickness —
Four categories of lamina thickness were distinguished

(Table 2). Most species are between 100 and 300 pm thick. Only one species (C.

malayanum ) is less than 100 pm thick. Sixteen species have a very thick lamina (over
300 pm; Table 2).

Lower order veins —

The lower order veins are vertically transcurrent and pos-

sess
sclerified bundle sheath extensions, showing birefringence and (auto-)fluores-

cence (Fig. 4C; Table 2). Only in C. riedelianum such bundle sheath extensions were

lacking, and the veins were embedded in the mesophyll.

Cuticle —
The cuticle (including the cuticular layer) stained bright to dark red,

with Sudan IV. Four classes in the cuticle thickness

pink or orange were recognized

(Table 2). Most species had a cuticle thickness between 3 and 8 pm; 28 species had

thinner cuticles; C. has a cuticle only 2 pm thick; 9 species had thick cu-
tampicense
ticles of which the cuticle of C. vaccinifolium showed the maximum value of 17 pm

(Fig. 4A). The outer cuticle surface is almost always flat. In 3 species, however, an

irregular surface was observed; C. cordatum, C. effusum and C. vaccinifolium (Fig.

4A; = in Table 2).

Adaxial and abaxial epidermis In species the adaxial cells
—
most epidermal are

small, square rectangular in shape (Fig. 4A), and show various of bi-
to degrees

refringence and (auto)fluorescence of the cell walls which indicate sclerification (Fig.

4C, D). Some species showed strong sclerification. Other species possessed large

epidermal cells which were hardly sclerified (Fig. 4B; Table 2). The adaxial epider-
mal cells had cutinized anticlinal walls (Fig. 4B). Generally, the cells of the abaxial

epidermis show flat periclinal outer walls (Table 2). Some species have lowly domed

outer periclinal epidermal walls (Fig. 5A; Table 2). Other species possess papillae

varying from dome-shaped (Fig. 5B; + in Table 2) to club-shaped (Fig. 5C; ++ in

Table 2). Generally, the cells of the abaxial epidermis show sclerification identical to

that in the adaxial cells.

occurring epidermal

Hypodermis A hypodermis occurred in 1 C. sericeum

—

only species: (Table 2).

Some of the hypodermal cells were sclerified (Fig. 4F).

Fig. 5. Transverse 30 thick sections of Cinnamomum species. A—G: Stained with Sudan IV.
µm

Light microscopy. —
A: C. bodinieri. Abaxial epidermis with lowly domed outer periclinal walls;
435. B: Abaxial of the leaf with C:
x —

C. cubense. epidermis dome-shaped papillae; x 435. —

C. abaxial of the leaf

bejolghota. Papillate epidermis showing more or less club-shaped papillae;

x 435. —
D: C. wightii. Short, thin-walled appressed hair on the abaxial surface of the leaf; x 435.

—
E: C. bintulense. Long, broad, thin-walled hair on the abaxial surface of the leaf; x 175. —
F:

C. fouilloyi. Very thick-walled erect hair on the abaxial surface of the leaf; x 435. —
G: C. sellow-

ianum. Curly, thick-walled hairs on the abaxial surface of the leaf; x 435.
M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Leaf anatomy of Cinnamomum 19
 BLUMEA VOL. 37, No. 1, 1992
20

6. electron micrographs of Cinnamomum leaves. A: C. aureo-fulvum. Dense align-

Fig. Scanning —

260. B: C. sellowianum. Curly erect hairs; 260.

ed hairs: x
— x

Almost all species studied possessed a distinctly 1-layered palisade

Mesophyll —

The presence of recorded in 19

parenchyma. a 2-layered palisade parenchyma was

species (Table 2). Only in C. amoenum a 3-layered palisade parenchyma was ob-

served. In many species the palisade cells (of the upper layer in case of a multi-layered

palisade) were (more or less) entirely sclerified (c* en t in Table 2). In the partially

sclerified cells the sclerified parts in the anticlinal walls had tapering ends on the

lower side of the cells when seen in transverse section (Fig. 4D, E). In
approximate-
20% of the species only the upper periclinal wall was thickened and sclerified,
ly
thus forming a
kind of cap (Fig. 4D; c in Table 2). Where sclerification of the pali-
sade cells (and adaxial epidermal cells) was present a layer of 'fused' sclerified caps

was mostly apparent underneath the epidermis (Fig. 4E). Only 13 species had non-

sclerified palisade cells (Fig. 4B; Table 2).

In about two-thirds of the species studied a slight to distinct sclerification was ob-

served in the cells of the spongy parenchyma (Table 2). A subepidermal layer against
the abaxial epidermis mostly showed distinct sclerification.

Trichomes —
Trichomes were absent in about 50% of the species (Table 2). In

the other species solitary unicellulartrichomes were Most of these species

present.
bore trichomes on their abaxial surface only. The frequencies varied from few to

abundant (Fig. 6A, B; Table 2). The length of these hairs varied from c. 30 pm (e.g.

C. wightii, Fig. 5D) to over 250 pm (e.g. C. kinabaluense 2, Table 2; Fig. 5E). In

only ten species trichomes were also present on the adaxial side of the leaf, usually in

low The of the adaxial trichomes varied from 50

frequencies (Table 2). length c. pm

to 250 pm (Table 2). The trichomes were of different erect (Fig. 5F), appres-
types:
sed (Fig. 5D) or curly (c in Table 2; e.g. C. sellowianum, Figs. 5G, 6B). In most

cases the hairs were thick-walled (Fig. 5F, G). Some species showed thin-walled

hairs (Fig. 5D, E) or hairs with an intermediate wall-thickness.

 M.E. A.F. Gerritsen & P.J. van der Schaaf: Leaf of Cinnamomum 21
Bakker, anatomy

Venation pattern —
In Cinnamomum the triplinerved leaf (subpalmate venation

generally (t in Table 2). The be described acrodromal.

pattern) occurs nervature can as

The main vein and two (sub)basal lateral veins are conspicuous and in some cases

even an additional lateral vein pair is distinct. However, some species have penni-
nerved (pinnately nerved) leaves (p in Table 2).

General leaf in Cinnamomum

anatomy
The lamina thickness ranges from 100 to 300 pm. The upper cuticle is 3-8
pm

thick and is flat. The adaxial epidermal cells are small, rectangular to
square in trans-

verse section and possess more or less sclerified walls. These walls are continuous

with the sclerenchymatous bundle sheath extensions of the vertically transcurrent

lower order veins. There is no hypodermis. The palisade parenchyma consists of one

layer of cells of which the outer periclinal wall is more or less thickened and sclerified,
thus forming a cap. In species the sclerified caps in the
most aligned outer periclinal
wall form a kind of layer underneath the adaxial epidermis. The anticlinal walls of the

palisade cells show sclerification. The cells of the also show

spongy parenchyma
sclerification. Oil and/or mucilage cells in the and spongy
are always present palisade

parenchyma. The outer periclinal walls of the abaxial epidermal cells are more or less

flat and sclerified. Hairs, if present, are unicellular, thick-walled, and mostly occur

on the abaxial side. The majority of the species differed from this in
general pattern

only one or a few features. However, about one-fifth of all studied deviated
species
more strongly with a maximum number of seven out of the sixteen features studied.

Phenetic analysis
All leaf anatomical features presented in Table 2 coded for cluster
were analysis
(Table 3a) and datamatrix for the calculation of the
a was generated similarity among
the species/specimens. Several
groups of leaf anatomically identical species were

entered as single species (Table 3b). The first mentioned of each of these
species
was used in the analysis. The resulting phenogram of the cluster
groups analysis
based upon Manhattan distances is presented in Figure 7.

Ten arbitrary clusters can be recognized: cluster 1: C. altissimum- -iC. durifolium;

cluster 2: C. aromaticum- - C. pachyphyllum; cluster 3: C. bullatum- C
I . melastoma-

ceum; cluster 4: C. bodinieri

- C. cluster 5: C. and C.
malayanum; angustitepalum

bejolghota; cluster 6: C. cubense C. montanum; cluster 7: C. cordatumt-iC. kina-

I

baluense 1\ cluster 8: C. archboldianum 4-1C. nalingway; cluster 9: C. amoenum

i-

C. triplinerve, and cluster 10: C. bintulense

-<C. Most Cinnamomum
, ovalifolium.

species are clustered at low distance values, especially clusters 1 3 which

to (Fig. 7)
include two-thirds of all studied. The
species neotropical Cinnamomum species (n in

Fig. 7) are especially present within the clusters 6 and which show deviation
9, more

(higher Manhattan distance) from the in the upper of the

species placed part pheno-

gram (Fig. 7).

When analyzing the variation in idioblast distribution within the clusters it
patterns
was striking that three clusters for the idioblasts
were constant (Table 4). The species/
specimens of cluster 8 were characterized by the absence of oil while the spe-
cells,
cies in the clusters 9 and 10 lacked mucilage cells. In the clusters 3-7 1 2
only or

species/specimens deviated from the general pattern within a cluster (see Table 4).
22 BLUMEA VOL. 37, No. 1, 1992

Figure 7

*
altissimum

archboldianum 1-10*
*
brevipedunculatum

cambodianum *

solomonense

propinquum
*
baileyanum
burmanni

subpenninervum
cinereum

calciphilum
*
durifolium
*
aromaticum

*
caryophyllus

aureo-fulvum

griffithii

ellipticifolium

englerianum

subaveniopsis

zollingerii
coriaceum

goaense

pachyphyllum
*
bullatum

clemensii

trichophyllum
*
curvifolium
*
macrocarpum

velutinum

wightii
*
doederleinii

*
microphyllum

spec.

macrophyllum

rupestre

melastomaceum

*
bodinieri (n)

camphora 1*

porrectum 1,2

dewildei *

parthenoxylon
sericeum

malayanum
 M.E. A.F. Gerritsen & P.J. der Schaaf: Leaf of Cinnamomum 23
Bakker, van
anatomy

(Fig. 7 continued)

1
angustitepalum

angustitepalum 2

bejolghota
cubense n

effusum n

elongatum n

montanum n

cordatum

*
kami

daphnoides 1

daphnoides 2*

kinabaluense 1

archboldianum 4*

rivulorum

sinharajense
archboldianum 5

nalingway

amoenum n

vesiculosum n

sellowianum n

*
camphora 7, 8

kinabaluense

riedelianum n

triplinerve n

*
bintulense

*
corneri

microcarpum

mollissimum

4
camphora
camphora 9
*
damhaense

simondii

*
obtusifolium

riparium

ovalifolium

Fig. 7. Phenetic cluster analysis of Cinnamomum species based a datamatrix

on
derived from Table 2.

*
The scale on the X-axis reflects Manhattan distances between species or groups of species; refers

to groups of identically coded species listed in Table 3b (n =

neotropical species).
24 BLUMEA VOL. 37, No. 1, 1992

Table 3a. Coding characters of Table 2 for cluster analysis.

1. Oil cells in lamina: 0 =

1 =
absent.
present;

2. Mucilage cells in lamina: 0 = 1 = absent.

present;

3. Oil cells in palisade parenchyma: 0 =

absent; 1 =
present.

4. Oil cells in parenchyma: 0 absent; 1

spongy
= =
present.

5. Mucilage cells in palisade parenchyma: 0 =

absent; 1 =
present.

Mucilage cells in parenchyma: 0 absent; 1

6. spongy
= =
present.

7. Lamina thickness: 0 = < 300 pm; 1 = > 300 pm.

8. 0 < 8 pm; 1 8
Cuticle thickness: = = > pm.

Adaxial epidermal cells: 0 slightly sclerified; 1 distinctly

9. = not or = or

10. Adaxial hypodermis: 0 absent; 1 [strongly sclerified,

= =
present.

11. Palisade 0=1 1 2 3 layers.

parenchyma layers: layer; = or

12. Palisade parenchyma cells: 0 = not

sclerified; 1 = more or less sclerified.

Spongy parenchyma cells: 0 lightly sclerified; 1 = sclerified.

13. = not or

14. Adaxial 0 absent; 1

papillae: = =
present.

15. Adaxial hairs: 0 =

absent; 1 =
present.

16. Abaxial hairs: 0 =

absent; 1 =
present.

17. Venation: 0 =
triplinerved; 1 =
penninerved.

18. Bundle sheath extensions: 0 =

absent; 1 =
present.

Table 3b. Groups of identically coded Cinnamomum species.

C. C.
C. altissimum; C. crassinervium; chittagongense; politum.
C. archboldianum 1-3,6-10; C. cappara-coronde; C. celebicum; C. cuspidatum 1,2; C. laubatii;

C. loureirii 1; C. mendozae; C. mercadoi; C. panayense; C. xylophyllum.

C. archboldianum C. magnifolium.
4;

C. aromaticum; C. perrottetii; C. scoriechinii; C. subavenium 2; C. sulphuratum.

C. baileyanum; C. citriodorum.

C. bintulense; C. fouilloyi

C. bodinieri; C. porrectum 3; C. psychotrioides; C. tampicense.

C. brevipedunculatum; C. elephantinum; C. myrtifolium; C. ovatum; C. poilanei; C. polyadelphum;

C. rigidum; C. tepalinum.
C. bullatum; C. javanicum; C. loureirii2.

C. burmanni; C. oliveri; C. virens.

C. cambodianum; C. crispulum; C. culitlawan; C. eugenoliferum; C. insulari-montanum; C. kera-

laense; C. longitubum; C. myrianthum; C. osmeophleum; C. piniodorum; C. porphyro-

C. C. racemosum; C. reticulatum; C. sessilifolium; C. sintok; C.
spermum; soncaurium;
C.
tamala 1,2; tazia; C. lonkinense; C. verum p.p.; C. xanthoneurum ; C. zollingerii 2.

C. camphora 1; C. camphora 2-6; C.

porrectum 1, 2.
C. camphora 7,8; C. glaucescens.
C. caryophyllus; C. deschampsii; C. dubium 2; iC. ebaloi; C. frodiniv; C. gigaphyllum; C. grandi-

florum; C. iners 1-3; C. ledermannii; C. lucens; C. malabathrum; C. podagricum; C. scala-

rinerve; C. sericans; C. subavenium 1; C. subcuneatum; C. verum

p.p.

C. corneri; C. tahijanum; C. vimineum.

C. curvifolium; C. kunstleri.
C. damhaense; C. dubium 1; iC. iners 4; C. mairei; C. C. paiei; C. rhynchophyllum
malabaricum; p.p.

C. daphnoides 2; C. vaccinifolium.
C. dewildei; C. griffithii 1.

C. doederleinii; C. melliodorum.

C. durifolium; C. litseafolium.

C. kami; C. pseudopedunculatum.

C. C. tsoi.
macrocarpum;

C. microphyllum; C. nooteboomii.

C. obtusifolium; C. rhynchophyllum p.p.

 M.E. Bakker, A.F. Gerritsen & P.J. der Schaaf: Cinnamomum 25
van
Leaf anatomy of

The general idioblast distribution pattern in the clusters 1-4 was that oil and mucilage

cells are in both layers. Clusters 1 and 2 showed more variation in the idio-
present
blast distribution than clusters 3 and 4. About 80% of the species within the former

clusters possessed the general idioblast distribution. The remaining species/specimens

within these clusters only lack one idioblast type in one
of the
mesophyll layers.
The idioblast characters are more constant than the other leaf anatomical characters

such as the presence or

absence of sclerified epidermis, palisade and spongy paren-

chyma cells, the layering of the palisade parenchyma, the lamina and cuticle thick-

ness, and the venation pattern (Table 4). The combination of the idioblast distribution

pattern and other constant leaf anatomical characters can be used to distinguish the

clusters (Fig. 7) from each other (Table 4).

DISCUSSION

The leaf anatomical features studied confirm and greatly extend the existing
(Table 2)

knowledge on Cinnamomum leaf anatomy (Santos, 1930; Marlier-Spirlet, 1946).

These features, including the idioblasts, did not show obvious correlations enabling
the recognition of species groups. In order to some insight in the overall similari-
get
ties of the individual species/specimens of Cinnamomum cluster
analyses were ap-

plied. A tentative cladistic analysis was not carried out, as the number of taxa is out

of with the number of features. Moreover, such

proportion as compared an analysis

yielded already unsatisfactory trees in a study on oil and mucilage cells in Annona

(Bakker & Gerritsen, 1992) in which only c. 40 taxa were used.

Some Cinnamomum species of which more than one specimen was studied ended

up in different clusters (Fig. 7), e.g. C. archboldianum in the clusters 1 and 8; C.

in the clusters 4, 9 and 10, and C. kinabaluense in the clusters 7 and 9.

camphora
However, slightly different specimens of most other species ended up in the same

clusters, e.g. C. angustitepalum and C. daphnoides.

In the clusters 1 and 2 the idioblast distribution show the largest variation
patterns

(Table 4). However, these variations are less obvious than in the idioblast distribu-

tion in leaves of Annona (Bakker & Gerritsen, 1992). The other leaf anatomical fea-

useful in discriminating these clusters. In the clusters 3-7

tures were not only one or

two species show minor idioblast variations. Cluster 3 always possesses mucilage
cells in both mesophyll layers and oil cells in the spongy parenchyma. The remaining
leaf anatomical features were not discriminating. Species in cluster 4 can be recog-

nized by the combination of the following features; presence of both idioblast

types
in both layers, non-sclerified epidermal cells, and a weakly or not sclerified
spongy

parenchyma. Cluster 5 does not

possess a discriminating leaf anatomical character

distribution. Cluster 6 lacks mucilage cells in the spongy parenchyma and is addi-

tionally characterized by the possession of non-sclerified epidermal cells and

spongy

parenchyma, a sclerified palisade parenchyma and leaves.

two-layered penninerved
Cluster 7 is characterized by the presence of oil and cells in both
mucilage layers, a

thick lamina and cuticle, a two-layered sclerified palisade parenchyma and tripli-
 26 BLUMEA VOL. 37, No. 1, 1992

Table 4. Constant leaf anatomical features per cluster.

1 feature oil muc

1 c e P s pa ad ab v

P s P s

cluster 1

+* +* 1+
1 + < +
-
_

2 +* + < + 1 - t

3 + + + < +

4 +* + + + < < -
1 _

_* +* > + 1+
5 + + -

6 + + + < -
2+ -

7 +* + + + > > 2+ -
t

8 - -
+ + < 1 -

9 + + - - < -
-

10 + + - -
< +

Legend:

Clusters: for species composition of each cluster see Figure 7 and Table 3b.

Features: oil: p = oil cells in palisade parenchyma

s = oil cells in spongy parenchyma

mucilage cells in
muc: p
=
palisade parenchyma
s =
mucilage cells in spongy parenchyma

+ -

present; -
= absent

+* =
present, but absent in one species
-* but in
=
absent, present one species

1 =
lamina thickness

< = lamina thickness < 300 gm

> = lamina > 300 gm

c = cuticle thickness

< = cuticle thickness < 8 gm

> = cuticle > 8

pm

e = adaxial epidermis
+ =
sclerified; -
= non-sclerified

p =
palisade parenchyma
1 = uni- layered; 2 =
two-layered
+ = sclerified; -
= non-sclerified

s =
spongy parenchyma
+ =
distinctly sclerified; -
=
weakly or non-sclerified

pa =
papillae; -
= absent

ad = adaxial hairs; -
= absent

ab = abaxial hairs

v = leaf venation

t =
triplinerved; p =
penninerved
 M.E. Bakker, A.F. Gerritsen & P.J. van der Schaaf: Leaf anatomy of Cinnamomum 27

veined leaves. The distribution of the idioblasts is constant in the following

pattern
three clusters (Table 4). Cluster 8 lacks oil cells and clusters 9 and 10 are
charac-

terized by the absence of mucilage cells.

At least in the latter three clusters the idioblast distribution pattern is the discrimi-

nating factor. Therefore, oil and mucilage cells possess some diagnostic value, gen-

erally in combination with other leaf anatomical features.

A classification of Cinnamomum species was presented by Meissner (1864). He

placed 54 Cinnamomum species in two sections: Malabathrum and Camphora (see

also No literature is available the infrageneric taxonomy
Pax, 1889). more recent on

of Cinnamomum (Kostermans, 1986). In an to correlate the leaf anatomical

attempt

characters of the species (Table 2) with the existing classification, we analyzed the

position of the species in the phenogram (Fig. 7). In the present paper 23 out of the

42 species included in section Malabathrum (Meissner, 1864) are discussed. Of these

species 18 are placed in the upper part of the phenogram (Fig. 7: clusters 1-3). The

five other placed in cluster 7 and 10. All these species sclerified
species are possess

epidermal cells and/or palisade cells. Only two out of the twelve species belonging

to section Camphora (Meissner, 1864) were studied here (C. camphora and C. par-

thenoxylon). These species are placed in the clusters 4, 9 and 10 (Fig. 7) and lack

sclerified epidermal and palisade cells (Table 2). A few other not neotropical species
C. clemensii, C. glaucescens, C. kinabaluense 2, C. C.
(C. bodinieri, malayanum,
C. simondii) also sclerification
porrectum and lack (Table 2) and are almost all locat-

ed in the same clusters. Because of the low number of species studied in section Cam-

phora little can be concluded about characters on sectional level, but from the fact that

both sections have species in cluster 10 it is evident that there is no clear leaf anatom-

ical distinction between them.

Of the 11 neotropical species in the 10 transferred from other

present study, were

genera such as: Phoebe (C. cubense, C. effusum, C. elongatum), Oreodaphne (C.
C. C. Laurus
amoenum, tampicense, vesiculosum), (C. montanum, C. triplinerve),
Ocotea (C. psychotrioides) and Persea (C. riedelianum). Almost all neotropical spe-

cies are placed in the clusters 6 and 9 (Fig. 7). Furthermore C. cubense, C. effusum
and C. elongatum (cluster 6) are closely related as is the case
with C. amoenumi and

C. vesiculosum (cluster 9). The first three species were included in the genus Phoebe

and the latter two in Oreodaphne.

It is striking that nine of these species have leaves (Table 2; not C. sel-
penninerved
lowianum and C. triplinerve). Of the remaining 139 Cinnamomum species only 15

showed this type of leaf venation (Table 2). Triplinerved and penninerved leaves

have been reported before in Cinnamomum In ad-

(Klucking, 1987; Hyland, 1989).

dition, all neotropical species possessed non-sclerified epidermal cells (Table 2).

Further, eight of eleven neotropical species showed a 2- or 3-layered palisade paren-

chyma against eleven of the remaining not neotropical species (Table 2). Therefore, it

can be concluded that the neotropical species show more differences from the
general

occurring leaf anatomical pattern in Cinnamomum than does the majority of the

species studied. Richter (1981) already recognized a South American group of Cin-

namomum species based wood and bark

on anatomy.
28 BLUMEA VOL. 37, No. 1, 1992

The presence of oil cells, and to a lesser extent also of mucilage cells, is characteristic

Metcalfe & 1950; Metcalfe,

for the Lauraceae (Pax, 1889; Solereder, 1899; Chalk,

1987). In the oil and/or mucilage cells in Cinna-

present study are always present

momum
in both mesophyll layers of the leaf (Table 2). In Annona (Annonaceae) the

idioblasts were always present in the spongy parenchyma only (Bakker & Gerritsen,

and of oil and

1992). The variation in number, staining reaction, size/shape mucilage

cells in Cinnamomum was

similar to that found in Annona species (Bakker & Gerrit-

1992). The different staining colours and/or appearances of the oil cells in dif-
sen,

ferent species (Fig. 1) have been attributed to the different compositions of oil in the

The presence of oil and cells in leaves

cells (Perrot, 1891; Kurata, 1952). mucilage

of the neotropical Cinnamomum species studied, which formerly were placed in

other genera, are in agreement with the literature concerning the leaf anatomy of

American Lauraceae, including Phoebe, Persea, Ocotea, and Oreodaphne (Petzold,

1907).

species of which in the than studied

Eleven present study more one specimen was

for their idioblast distribution pattern. However, other

were constant seven
species
(C. archboldianum, C. camphora, C. dubium, C. griffithii, C. iners, C. kinabalu-

ense, C. zollingerii) showed variable distribution patterns (Table 2). It is not

likely
that these specimens were determined incorrectly. The variation in idioblast distribu-

of the of idioblasts
tion can partly be explained by the suggested homology two types

1991; also below). Moreover, it is that different ecologi-

(Bakker et al., see possible
cal conditions also played in the variation found in C. camphora, C. iners, and
a part
C. kinabaluense. It is not known which genetic and/or ecological triggers play a

and seasonal variations have been reported effect the number of

part. Ontogenetic
to

oil and mucilage cells in the camphor tree, C. camphora (Shirasawa, 1903). Leaf oil

composition varied between varieties of C. cassia (Cheng et al., 1989). The varia-

tions used to of C. al., and

were identify types camphora (Shi et 1989) were applied
in numerical chemotaxonomical studies in Cinnamomum & Ex-
(Tao Zhong, 1988).

perimental studies under varying conditions are needed in order to explain the appar-

ent variations in idioblast distribution in Cinnamomum species.

Since inffageneric classifications within Cinnamomum, based upon other charac-

ters such as macromorphology and pollen morphology, are very scarce, the idioblast

distributionpatterns could not be correlated with diversity patterns in other character-

istics. Therefore little can be concluded about the systematic significance of the oil

and mucilage cells.

In the literature oil and mucilage cells are mentioned as occurring together and were

from
supposed to develop one type into the other (Tschirch, 1889,1914; Janssonius,

1926, 1934; West, 1969). Baas & Gregory (1985) and Gregory & Baas (1989) crit-

ically discussed this phenomenon and further research. In earlier papers

suggested
described the ultrastructure and development of oil and cells of Cinna-
we
mucilage
momum (Bakker & Gerritsen, 1989; Bakker et al., 1991) and Annona (Bakker &

Gerritsen, 1990), especially to investigate the possible homology of both idioblast

found that suberized wall layer,

types. We a typical plastids, an extraplasmatic space

for
storage of the secreted product, and a cupule were present in both oil and mucilage
cells (Bakker al., 1991), which indicated of both idio-
et a homologous development
 M.E. Bakker, A.F. Gerritsen & P.J. der Cinnamomum
van Schaaf: Leaf anatomy of 29

blast In addition, the first zone of deposited mucilage in mucilage cells resem-
types.
bled the inner wall layer in oil cells. The present observations on oil and mucilage
cells in a large number of Cinnamomum species also support this homology. A suber-

ized always detected in both idioblast the light microscopical and

layer was types at

ultrastructural level.

CONCLUSIONS

Oil and/or mucilage cells are always present in both mesophyll layers in all Cinna-

momum species. The idioblasts show variations in shape/size, stainability, number,

and distribution The latter variation can partly be understood by the
pattern. proposed

homology of both idioblast types. Oil and mucilage cells possess some diagnostic
value in combination with other leaf anatomical characters. Because of the lack of

comparative classification of Cinnamomum, the remains

a
systematic significance
questionable. Further studies on the classification of Cinnamomum are necessary to

evaluate the systematic value of oil and mucilage cells at the infrageneric level.

ACKNOWLEDGEMENTS

We thank Dr L. Goosen-de Roo (Botanical Laboratory, Leiden) for valuable comments and critical

reading of the manuscript. The curators of the Hortus Botanicus in Leiden and Utrecht, of the Botan-

ical Gardens of the Agricultural University Wageningen and the Technical University Delft (The
Netherlands), of the Botanical Garden of Berlin-Dahlem (Germany), of the Botanical Gardens of Kew

and the Royal Botanical Garden Edinburgh (U. K.) for kindly providing fresh leaves of several Cin-

species, Dr. P. Baas for collecting several species in botanical in Java

namomum
gardens (Indonesia),
and Dr. H. van der Werff (Missouri Botanical Garden, U.S.A.) for collecting leaves of C. camphora
in Berkeley. This work was supported by the Foundation for Fundamental Biological Research

(BION) which is subsidized by the Netherlands Organization for Scientific Research NWO (grant nr.

811-440-023 to M.E. Bakker).

REFERENCES

& M. Gregory. 1985. A of oil cells in the with their

Baas, P., survey dicotyledons comments on re-

placement by and joint occurrence with mucilage cells. Isr. J. Bot. 34: 167-186.

Bakker, M.E., & A.F. Gerritsen. 1989. A suberized layer in the cell wall of mucilage cells in Cin-

namomum. Ann. Bot. 63: 441-448.

Bakker, M.E., & A.F. Gerritsen. 1990. Ultrastructure and of oil idioblasts in Annona
development
muricata L. Ann. Bot. 66: 673-686.

Bakker, M.E., & A.F. Gerritsen. 1992. Oil and mucilage cells in Annona and their
(Annonaceae)

systematic significance. Blumea 36: 411-438.

Bakker, M.E., A.F. Gerritsen & J.P. van der Schaaf. 1991. Development of oil and mucilage cells

in Cinnamomum burmanni. An ultrastructural Acta Bot. Neerl. 40: 339-356.

study.

Cheng, B., Y. Xu, X. Yu, J. Ding & J. Zhou. 1989. The introduction and cultivation of Cinnamo-

mum cassia in Yunnan Province. Acta Bot. Yunnanica 11: 433-439.

Gregory, M., & P. Baas. 1989. A of mucilage cells in the

survey vegetative organs of the dicotyle-
dons. Isr. J. Bot. 38: 125-174.

Hegnauer, R. 1966. Chemotaxonomie der Pflanzen 4. Birkhauser

Verlag, Basel, Stuttgart.
Hyland, B.P.M. 1989. A revision of Lauraceae in Australia (excluding Cassytha). Aust Bot.
Syst.
2: 135-367.
 BLUMEA VOL. 37, No. 1, 1992
30

Janssonius, H.H. 1926. Mucilage cells and oil cells in the woods of the Lauraceae. Trop. Woods 6:

3-4.

Janssonius, H.H. 1934. Mikrographie des Holzes der auf Java vorkommenden Baumarten. Band 5.

Brill, Leiden.

Klucking, E.P. 1987. Leaf venation patterns. Vol. 2. Lauraceae. 216 pp., 149 plates. Cramer, Ber-

lin, Stuttgart.

A.J.G.H. 1961. The world species of Cinnamomum Trew. (Lauraceae). Rein-

Kostermans, new

wardtia 6: 17-24.

A.J.G.H. 1964. Bibliographia Lauracearum. 1450 P.T. Djulie 'ArchipeT, Bogor.

Kostermans, pp.

A.J.G.H. 1968. Materials for a revision of Lauraceae I. Reinwardtia 7: 291-356.

Kostermans,

Kostermans, A.J.G.H. 1969. Materials for a revision of Lauraceae II. Reinwardtia 7: 451-536.

Kostermans, A.J.G.H. 1970a. A new Bornean species of Cinnamomum Sch. Reinwardtia 8: 13-

15.

Kostermans, A.J.G.H. 1970b. Materials for a revision of Lauraceae III. Reinwardtia 8: 21-196.

A.J.G.H. 1983. The South Indian species of Cinnamomum Schaeffer (Lauraceae)

Kostermans,

Bull. Bot. Surv. India 25: 90-133.

Kostermans, A.J.G.H. 1986. A monograph of the Cinnamomum Schaeff. (Lauraceae). Part 1.

genus

Ginkgoana6: 1-171.

Kostermans, A.J.G.H. 1988. Materials for a revision of Lauraceae V. Reinwardtia 10: 439-469.

Kurata, T. 1952. Microscopic observation on oil cell of camphor tree. J. Jap. For. Soc. 34: 232-234.

Marlier-Spirlet, M.L. 1946. Sur quelques epidermes de feuilles de Cinnamomum L. Bull. Jard. Bot.

Brux. 17: 266-305.

Meissner, C.F. 1864. In: A. de Candolle, Prodromus systematis naturalis regni vegetabilis. Vol. 15,

522 Masson & Sons, Paris.

pp.

Metcalfe, C.R. 1987. Anatomy of the dicotyledons. 2nd ed., Vol. III. Magnoliales, Illiciales and

Laurales. 224 Oxford Science Publ.

pp.

Metcalfe, C.R., & L. Chalk. 1950. Lauraceae. In: Anatomy of the dicotyledons. 2: 1145-1156.

F. 1889. Lauraceae. In: A. Engler & K. Prantl, Naturliche Pflanzenfamilien 3 (2): 106-114.
Pax,

Perrot, E. 1891. Contribution l'etude histologique des Lauracees. Thesis, Paris.

Petzold, V. 1907. Systematisch-anatomische Untersuchungen iiber die Laubblatter der amerikani-

schen Lauraceen. Bot. Jahrb. 38: 445-474.

Engl.
Richter, H.G. 1981. Anatomie des sekundaren Xylems und der Rinde der Lauraceae. Sonderb. Natur-

wiss. Vereins Hamburg 5. Parey, Hamburg.

Rohlf, F.J. 1986. NTSYS-pc. Numerical and multivariate
taxonomy analysis systematics. Applied
Biostatistics Inc., Setauket, New York.

Santos, J.K. 1930. Leaf and bark structure of some cinnamon trees with special reference to the

Philippine species. Philipp. J. Sci. 43: 305-365.

Shi, W., W. He & G. Wen. 1989. Study on chemical constituents of the essential oil and classifi-

cation of from Cinnamomum camphora. Acta Bot. Sinica 31: 209-214.

types

Shirasawa, H. 1903. fiber Entstehung und Verteilung des Kampfers im

Kampferbaume (Bull. Coll.

Agric. Tokyo 5: 373-401). See: Just's Bot. Jahrber. 38 (1): 1373-1374 (1913).
Solereder, H. 1899. Laurineae. In: Systematische Anatomie der Dicotyledonen: 791-796. Verlag
Enke, Stuttgart.

Tao, G., & Y. Zhong. 1988. Study on numerical chemotaxonomy of Cinnamomum in Hubei Prov-

ince. Acta Phytotaxon. Sinica 26: 409-417.

Tschirch, A. 1889. Angewandte Pflanzenanatomie I. Urban and Schwarzenberg, Vienna.

Tschirch, A. 1914. Die Membran als Sitz chemischer Arbeit. Verhandl. Schweiz. naturf. Gesellschaft.

II. Teil, Sektion fur Botanik: 178-188.

Werff, H. van der. 1991a. A key to the of Lauraceae in the New World. Ann. Missouri Bot.
genera

Gard. 78: 377-387.

Werff, H. van der. 1991b. New species of Lauraceae from Ecuador and Peru. Ann. Missouri Bot.

Gard. 78: 409-423.

West, W.C. 1969. of oil cells in the woody Ranales. Bull. Torrey Club 96: 329-344.
Ontogeny BoL

View publication stats