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The Natural History Journal of Chulalongkorn University 7(1): 35-45, May 2007 

©2007 by Chulalongkorn University 



 

Contributions to the Leaf Anatomy and Taxonomy of Thai 

Myrtaceae 

 

 



C

HORTIP 

KANTACHOT, P

RANOM 

CHANTARANOTHAI

*

AND 

A

CHRA 

THAMMATHAWORN

 

 

 



Applied Taxonomic Research Center, Department of Biology, Faculty of Science, 

Khon Kaen University, Khon Kaen 40002, Thailand.

 

 



 

A

BSTRACT

.– Leaf anatomical studies of 28 taxa in 12 genera of Thai 

Myrtaceae were investigated to determine their taxonomic relevance. 

Fourteen characteristics could be used for the species identification: unifacial 

or bifacial leaves, shape of the midrib; margin and petiole, presence or 

absence of trichomes, hypodermis, stomatal types, epidermal cell wall, 

number of spongy layers, presence or absence of bundle sheath extension, 

shape of vascular bundle at midrib and petiole, presence or absence of 

sclereid at midrib and crystal types. Leaf anatomical data support 

morphological evidence for separating taxa at the generic level except for 

Cleistocalyx and Syzygium. 

 

K



EY WORDS

leaf anatomy; taxonomy; Myrtaceae 

 

 



I

NTRODUCTION

 

 

The family Myrtaceae with its ca. 100 



genera and 3,000 species is distributed 

mainly in the tropics and subtropics. 

Fourteen genera and 115 species have been 

enumerated in Thailand (Parnell & 

Chantaranothai, 2002).  

Anatomical studies for various genera in 

Myrtaceae have been carried out by many 

workers, including Metcalfe & Chalk 

(1957), Carr & Carr (1969), Wilson & 

Waterhouse (1982), Hussin et al. (1992) 

and Haron & Moore (1996). Metcalfe & 

Chalk (1957) recognized four stomatal 

types in Myrtaceae: anomocytic, 

anisocytic, cyclocytic and paracytic. Most 

species have unicellular uniserriate 

trichomes. Hussin et al. (1992) studied leaf 

anatomy of Eugenia from the Malay 

Peninsula and concluded that the following 

characteristics were useful for species 

identification: stomatal type, leaf shape in 

transverse section (TS), shape of midrib 

bundle, sclerenchyma sheath, cutinization 

of the outer wall, presence or absence of 

sclereid, idioblast, hypodermis, columnar 

epidermal cells, solitary crystals, number 

of palisade layers, shape of vascular strand, 

sclerenchyma sheath and sclereid in ground 

tissued of the petiole. The aims of the 

present paper are to describe the 

comparative leaf anatomy of the Thai 

* Corresponding author: 

Tel: (6643)-342-908 

Fax: (6643)-364-169 

E-mail: pranom@kku.ac.th 



NAT. HIST. J. CHULALONGKORN UNIV. 7(1), MAY 2007 

36

Myrtaceae and to assess the taxonomic 



value of these characteristics. 

 

M



ATERIALS AND 

M

ETHODS



 

 

Fresh leaves of 28 taxa of 12 genera 



were collected. The specimens were fixed 

in 70% alcohol or 70% FAA, dehydrated 

in tertiary butyl alcohol series, sectioned on 

a rotary microtome and stained in safranin 

and fast green. TS of midribs, leaf margins 

and the middle parts between the midrib 

and margin of the lamina were made. For 

epidermal studies, samples were prepared 

by mechanical scraping at the midway 

between the base and apex of the lamina, 

stained in safranin and then mounted in 

DePeX artificial mounting medium. The 

leaf anatomical data of the taxa examined 

are complied and presented in Tables 1 and 

2, Figures 1 to 3 and Keys. Photographs 

were taken with the aid of an Olympus BH

light microscopy. 



 

R

ESULTS



 

 

General leaf anatomical description 



Lamina

 

Epidermal cells: in surface view 

epidermal cells on both surfaces are 

circular, ovate, square, triangular to 

polygonal; anticlinal walls of adaxial and 

abaxial cells are straight (Figs. 1A, 1B), 

curved (Fig. 1C) and undulate (Fig. 1D), 

adaxial cells larger than abaxial cells and 

variable shape in TS. 

Stomata: occurring on both surfaces or 

confined to abaxial surface: anomocytic 

(Fig. 1E), paracytic (Figs. 1F, 1G) and 

anisocytic (Fig. 1H).  



Trichomes: absent in all species except 

unicellular, uniserriate trichomes present in 



D. parviflorum var. parviflorum,  M. 

cajuputi,  P. guajava,  R. dumetorum,  Rh. 

tomentosa and T. burmanica var. 

rufescens 

Hypodermis: absent in all species but 

present in adaxial with 1-3 rows and the 

third row often becoming palisade-like 

(Figs. 2D, 2F) in P. guajava, S. cuminiS. 



malacense and S. ripicola 

Mesophyll: palisade cells and spongy 

cells different in shape: palisade cells 1 - 3 

layers; spongy cells 3 - 15 layers (Fig. 

2G). Palisade and spongy cells have similar 

shape in B. frutescens and E. camaldu-

lensis. Sclereid cells present only in S. 

jambos, S. malaccense and S. samarangense 

var.  samarangense. Oil cavities located 

close to both surfaces, globular, and lined 

with epithelial-like cells (Figs. 2E, 2F).  



Vascular bundle: collateral bundles 

without  bundle sheath extension  except 

(Figs. 2B, 2F) in E. camaldulensis,  Eu. 

uniflora,  Rh. tomentosa,  S. cumini and  S. 

ripicola. Fibre caps with only fibre cells 

present in all species except in S. aqeum



S. aromaticumS. formosumS. jambosS. 

malaccense,  S. megacarpum,  S. samaran-

gense var. samarangense and S. siamense 

which have both fibre cells and sclereid 

cells.  

Crystals: all species present druses (Fig. 

2H) and prisms only in mesophyll except S. 



ripicola has druses in hypodermis and 

mesophyll. 



Midrib  TS:  outline in adaxial surface 

concave, flat or convex. Abaxial surface 

convex to keeled. Vascular bundle is 

circular, heart-shaped or U-shaped, arms 

incurved or straight; bundle sheath with 

fibre or parenchyma (Figs. 3A-3D). 

Crystals: druses and prisms present in 

ground tissue. 

 


KANTACHOT ET AL. – LEAF ANATOMY AND TAXONOMY OF THAI MYRTACEAE 

37

T



ABLE 

1. Leaf epidermal characters of the studied taxa. 

 

Surface view

 

Adaxial cell wall

 

Abaxial cell wall

 

Species



 

Stomatal 

type

 

S C U S C U 



Trichome

 

Baeckea frutescens 

PA  + + - + + - 



Callistemon citrinus 

AN  + + - + -  - 



Cleistocalyx  nervosum var. nervosum 

PA  -  - + -  - + 



Decaspermum parviflorum var. parviflorum 

AN  -  - + -  - + 



Eucalyptus camaldulensis 

AN  + -  - + -  - 



Eugenia uniflora 

AN  - + -  - + - 



Melaleuca cajuputi 

AN  + -  - + -  - 



Psidium guajava 

AN  + - - - + - 



Rhodamnia dumetorum 

AN  -  - + + + - 



Rhodomyrtus tomentosa 

AN  + -  - + -  - 



Syzygium albiflorum 

AN  -  - + -  - + 



S. aqeum 

PA  -  - + -  - + 



S. aromaticum 

AN  -  - + - + - 



S. cinereum 

AN  -  - + -  - + 



S. claviflorum 

AN  - + -  - + - 



S. cumini 

ANI, 

PA -  - + -  - + 





S. diospyrifolium 

AN  -  - + -  - + 



S. formosum 

PA  -  - + -  - + 



S. jambos 

PA  -  - + -  - + 



S. laetum subsp. jugorum 

ANI, 


PA + + -  - + - 



S. malaccense 

PA  - + -  - + + 



S. megacarpum 

PA  - + -  -  - + 



S. ripicola 

ANI, 

PA - + -  -  - + 





S. samarangense var. samarangense 

PA  -  - + -  - + 



S. siamense 

PA  -  - + -  - + 



S. winitii 

PA  - + -  -  - + 



S. zimmermanii 

PA  -  - + -  - + 



Tristaniopsis burmanica var. rufescens 

AN  - + - + + - 

 

AN = anomocytic, ANI = anisocytic, C = curved, PA = paracytic, S = straight, U = undulate, + = present, - = 



absent 

Petiole TS 

Outline: adaxial of petiole in TS flat

concave (Fig. 3E), convex or V-shaped 

(Fig. 3F). Abaxial surface circular or 

ovate.  



Ground tissue:  collenchyma present 

below epidermis with some oil cavities; 

parenchyma present mainly in ground 

tissue.  



Trichome: unicellular, uniserriate 

trichomes present only in D. parviflorum 

var. parviflorumM. cajuputiP. guajava

R. dumetorum,  Rh. tomentosa and T. 

burmanica var. rufescens 

Vascular bundle: collateral with 

circular or ovate, heart-shaped or U-

shaped; bundle sheath cells with fibre or 

parenchyma. 



Crystals: Idioblast present in ground 

tissue underneath epidermal cells. Phloem 

parenchyma cells with druses or prisms 

(Fig. 3G).

 

 

Key to Genera Based on Anatomy of 



Lamina and Petiole 

 

1. Leaves unifacial..……...............…... 2 



-  Leaves bifacial......……….…............ 5 

2. Trichomes on the abaxial surface of 

lamina present...…............ Melaleuca 

-  Trichomes on both surfaces of lamina 

absent.………………….………………... 3 


NAT. HIST. J. CHULALONGKORN UNIV. 7(1), MAY 2007 

38

 



 

F

IGURE 

1. Adaxial (A-D) and abaxial (E-H) surfaces view. A) B. frutescens, B) M. cajuputi, C) S. jambos, D) S. laetum 

subsp. jugorum, E) S. albiflorum, F) S. formosum, G) S. jambos, H) S. ripicola

 

 

 



KANTACHOT ET AL. – LEAF ANATOMY AND TAXONOMY OF THAI MYRTACEAE 

39

 



 

F

IGURE 

2. Transverse section of margin and lamina. A) C. citrinus, B) E. camaldulensis, C) S. laetum subsp. jugorum

D) S. malaccense, E) M. cajuputi, F) S. cumini, G) S. diospyrifolium, H) druses in hypodermis of S. ripicola. 

 

 

 



 

NAT. HIST. J. CHULALONGKORN UNIV. 7(1), MAY 2007 

40

 



 

F

IGURE 

3. Transverse section of midrib and petiole. A) E. camaldulensis, B) Eu. uniflora, C) P. guajava, D) S. winitii

E) S. aromaticum, F) S. claviflorum, G) prism in M. cajuputi, H) trichomes in M. cajuputi. 

 

 

 



KANTACHOT ET AL. – LEAF ANATOMY AND TAXONOMY OF THAI MYRTACEAE 

41

3.  Vascular bundle in midrib heart-shaped. 



Stomata in the abaxial surface present... 

.......……….................... Eucalyptus 

-  Vascular bundle in midrib circular or 

ovate. Stomata in the both surfaces 

present………………….…................ 4 

4. Stomata anomocytic……..... Callistemon 

- Stomata paracytic...……......... Baeckea 

5. Trichomes present.....……............... 6 

-  Trichomes absent.....………............ 10 

6. Hypodermis present..………..... Psidium 

-  Hypodermis absent.........………....... 7 

7. Bundle sheath extension in collateral 

bundle of lamina present.. Rhodomyrtus 

-  Bundle sheath extension in collateral 

bundle of lamina absent.......……...... 8 

8. Anticlinal walls in abaxial surface deeply 

undulate........………..... Decaspermum 

-  Anticlinal walls in abaxial surface 

straight or shallowly curved............. 9 

9. Druses present. Anticlinal walls in 

adaxial surface deeply undulate………… 

…………………………….….. Rhodamnia 

-  Druses and prisms present. Anticlinal 

walls in adaxial surface curved..…….... 

...……………................. Tristaniopsis 

10. Vascular bundle in petiole circular….... 

.………………………………...... Eugenia 

-  Vascular bundle in petiole heart-shaped 

or U-shaped…. Cleistocalyx & Syzygium 

 

Key to Species Based on Anatomy of 



Lamina and Petiole of Syzygium 

 

1. Bundle sheath extension present……... 2 



-  Bundle sheath extension absent…….... 3 

2. Outline of midrib in adaxial surface 

concave...……………………… S. cumini 

-  Outline of midrib in adaxial surface flat 

.....……………................. S. ripicola 

3. Outline of petiole in adaxial surface V-

shaped......………......... S. claviflorum 

-  Outline of petiole in adaxial surface not 

V-shaped......………………………...... 4 

4. Stomata anomocytic…….................. 5 

-  Stomata paracytic or paracytic and 

anisocytic.........…………………....... 8 

5. Anticlinal walls in abaxial surface 

curved...........………... S. aromaticum 

-  Anticlinal walls in abaxial surface 

undulate.................………............ 6 

6. Vascular bundle in petiole U-shaped ….. 

.......……............……... S. albiflorum 

-  Vascular bundle in petiole heart-shaped.. 

…………...…………………………....... 7 

7. Vascular bundle in midrib U-shaped…... 

……………………..………… S. cinereum 

-  Vascular bundle in midrib heart-shaped 

.....……………......... S. diospyrifolium 

8. Hypodermis present…….. S. malaccense 

-  Hypodermis absent...……................ 9 

9. Stone cells in vascular bundle of midrib 

present..........……………………......10 

-  Stone cells in vascular bundle of midrib 

absent........…………................... 15 

10.  Outline of midrib in adaxial surface 

concave……………………………….... 11 

-  Outline of midrib in adaxial surface flat 

........……………..………………….... 13 

11. Druses present. Anticlinal walls in 

adaxial surface shallowly curved.……... 

..………………........... S. megacarpum 

-  Druses and prisms present. Anticlinal 

walls in adaxial surface deeply undulate 

...………………………................... 12 

12.  Vascular bundle in midrib U-shaped.... 

..………………………………… S. aqeum 

-  Vascular bundle in midrib heart-shaped 

..........…………....... S. samarangense 

13. Spongy cells arranged in less than 10 

layers.………………………………..... 14 

-  Spongy cells arranged in more than 10 

layers..........………...... S. formosum 

14. Outline of petiole in adaxial surface 

convex...........……......... S. siamense 



NAT. HIST. J. CHULALONGKORN UNIV. 7(1), MAY 2007 

42

-  Outline of petiole in adaxial surface 



concave…......……............. S. jambos 

15. Anticlinal walls in adaxial surface 

straight or shallowly curved........... 16 

- Anticlinal walls in adaxial surface 

deeply undulate.................. S. winitii 

16. Druses present. Anticlinal walls in 

abaxial surface curved.....………...... 

………………..S. laetum subsp. jugorum 

-  Druses and prisms present. Anticlinal 

walls in abaxial surface undulate.……... 

………………............. S. zimmermanii

 

 



D

ISCUSSION AND 

C

ONCLUSIONS



 

 

The Myrtaceous plants uniquely share a 



suit of morphological and leaf anatomical 

features. Tables 1 and 2 show that certain 

characteristics have been useful in the 

identification of the Thai Myrtaceae. Three 

types of stomata are observed as in the 

work of Hussin et al. (1992): anomocytic, 

paracytic and anisocytic. Haron & Moore 

(1966) show the anomocytic and paracytic 

types for the majority of Myrtaceae. The 

anomocytic or paracytic type was found to 

be the most common occurring in all 

 

T

ABLE 

2. Anatomical characters of the studied species in transverse section. 

 

Lamina 



Species 

Unifacial Bifacial 

Number of 

hypodermis 

layer 

Midrib 

Vascular 

bundle shape 

Petiole 

Vascular 

bundle shape 

Crystal 

type 

Baeckea frutescens 

+ + 




D,P 

Callistemon citrinus 

+ + 




D,P 

Cleistocalyx nervosum var. nervosum 

- + 






Decaspermum parviflorum var. parviflorum 

- + 






Eucalyptus camaldulensis 

+ - 


2  D,P 



Eugenia uniflora 

- + 


1  D,P 



Melaleuca cajuputi 

+ - 






Psidium guajava 

- +  + 


(2) 





Rhodamnia dumetorum 

- + 






Rhodomyrtus tomentosa 

- + 






Syzygium albiflorum 

- + 






S. aqeum 

- + 


2  D,P 



S. aromaticum 

- + 






S. cinereum 

- + 






S. claviflorum 

- + 






S. cumini 

- +  + 


(1) 





S. diospyrifolium 

- + 






S. formosum 

- + 


2  D,P 



S. jambos 

- + 


3  D,P 



S. laetum subsp. jugorum 

- + 






S. malaccense 

- + + 


(2-3) 





S. megacarpum 

- + 






S. ripicola 

- +  + 


(1) 





S. samarangense var. samarangense 

- + 


2  D,P 



S. siamense 

- + 






S. winitii 

- + 






S. zimmermanii 

- + 


3  D,P 



Tristaniopsis  burmanica var. rufescens 

- + 


3  D,P 



1 = circular & ovate, 2 = heart-shaped, 3 = U-shaped, D = druse, P = prism, + = present, - = absent 

KANTACHOT ET AL. – LEAF ANATOMY AND TAXONOMY OF THAI MYRTACEAE 

43

studied taxa except in S. cuminiS. laetum 



subsp. jugorum and S. ripicola which have 

both anisocytic and paracytic stomata in the 

same leaf. 

The stomatal complex are dispersed 

randomly over the abaxial surface, except 

in  B. frutescens,  C. citrinus and M. 



cajuputi. Only C. citrinus and M. cajuputi 

have a stomatal type with a non-prominent 

T-piece cutinization at the pole of the guard 

cell. Haron & Moore (1999) suggested a 

non-prominent T-piece cutinization type 

was found in the New World species. 

Epidermal cells in TS are circular or 

square-like in shape. Hypodermis is 

presented in four species: P. guajava, S. 

cuminiS. malaccense and S. ripicola. The 

anticlinal walls of both adaxial and abaxial 

epidermal cells as seen in surface view 

vary from straight to undulate. Cutin 

occurs on the leaf surfaces, the form of 

which varies from species to species being 

more distinct in B. frutescens,  C. citrinus 

and  E. camaldulensis. All species have 

globular oil cavities in mesophyll that are 

underneath both epidermis in agreement 

with the study of Carr & Carr (1969). The 

outline of midrib in TS in the adaxial 

surface ranges from flat, convex or 

concave while the abaxial one varies from 

convex to obtuse, angled to keeled. This is 

consistent with the work of Hussin et al. 

(1992). The leaf margin in TS is straight, 

directed downwards or re-curved, and the 

tip is acute, obtuse or acuminate. The 

shape of the midrib and petiole vascular 

strand is circular, heart-shaped or U-shaped 

but the ends of the strand in Cl. nervosum 

var.  nervosum and S. malaccense are 

deflexed and broken into smaller bundles. 

The bundle sheath extension in the lamina 

appears in five species: E. camaldulensis



Eu. unifloraRh. tomentosaS. cumini and 

S. ripicola. Fibre cap in lateral veins occur 

in all species. Sclereids are observed in S. 



aqeum,  S. aromaticum,  S. formosum,  S. 

jambosS. malaccenseS. megacarpumS. 

samarangense var. samarengense and S. 

siamense. They are at the midrib and 

petiole vascular bundle which Hussin et al

(1992) observed only in E. helferi (S. 

helferi). Druses and prisms occur in the 

ground tissue of the lamina and petiole, 

especially  S. ripicola present  druses in 

hypodermis and ground tissue. Metcalfe & 

Chalk (1957) and Hussin et al. (1992) 

reported only druses in the majority of the 

Myrtaceous plant. The occurrence of 

unifacial and bifacial leaf types appears to 

be a good diagnostic characteristic for the 

generic level. Most of them are bifacial 

leaf, but the unifacial leaf is found in B. 

frutescens,  C. citrinus,  E. camaldulensis 

and M. cajuputi

Using a combination of anatomical 

characteristics, it is possible to construct a 

key to study the species. Only two genera, 

Cleistocalyx and Syzygium, could not be 

separated. Both are morphologically 

different, especially in flower, the former 

has a fused calyx while the latter has free 

sepals (Parnell and Chantaranothai 2002). 

Further study is needed to elucidate their 

status. 

Comparative leaf anatomical character-

istics of the closely related species show 

that these characters are particularly 

important in discrimination the genus 

Syzygium, especially in the following pairs 

of species:  



Syzygium albiflorum and S. zimmer-

manii: the former has anomocytic stomatal 

type, druses present only in mesophyll, tip 

of leaf margin is obtuse and adaxial surface 

of petiole is flat. The latter has paracytic 

stomatal type, druses and prisms present in 


NAT. HIST. J. CHULALONGKORN UNIV. 7(1), MAY 2007 

44

mesophyll, tip of leaf margin is acute and 



adaxial surface of petiole is concave. 

Syzygium aqeum differs from S. sama-

rangense var. samarangense in its acute tip 

of leaf margin, outline of petiole in adaxial 

is concave; vascular bundle in midrib is U-

shaped. The tip of leaf margin  in  S. 



samarangense var. samarangense is 

obtuse, outline of petiole in adaxial is 

slightly convex, vascular bundle in midrib 

is heart-shaped. 



Syzygium diospyrifolium and S. 

formosum:  S. diospyrifolium has anomo-

cytic stomatal type, only druses present in 

mesophyll, two layers of palisade cells, 8 - 

10 layers of spongy cells, outline of midrib 

in adaxial is concave and sclereid absent. 

S. formosum has paracytic stomatal type, 

druses and prisms present in mesophyll, 

one layer of palisade cells, 11 - 15 layers 

of spongy cells, outline of petiole in adaxial 

is flat and sclereid present. 

Syzygium jambos and  S. siamense: the 

former has both druses and prisms in 

mesophyll and outline of petiole in adaxial 

is concave. The latter has only druses and 

outline of petiole in adaxial is convex. 

Syzygium ripicola and S. winitii:  S. 

ripicola has both paracytic and anisocytic 

stomatal types, one layer of hypodermis, 

bundle sheath extension present, tip of leaf 

margin is acute, outline of midrib in 

adaxial is flat and petiole is V-shaped. The 

S. winitii has paracytic, stomatal type, 

hypodermis and bundle sheath extension 

absent, tip of leaf margin is obtuse and 

outline of midrib and petiole in adaxial are 

concave. 

The leaf anatomy features alone are 

insufficient for the delimitation of some 

generic groups. In general, it can be 

supported that the features have consider-

able systematic value and give additional 

support to the distinctness of the species. 

However, like all other taxonomic 

evidence, leaf anatomical characteristics 

must be interpreted with circumspection. 

 

S

PECIMENS 



E

XAMINED


 

 

All specimens examined are deposited at 



KKU (Khon Kaen University Herbarium); 

Baeckea frutescens L., C. Kantachot 43. – 

Callistemon citrinus (Curtis) Skeels, C. 

Kantachot 213. – Cleistocalyx nervosum 

(DC.) Kesterm. var. nervosum,  C. Kanta-



chot 190. – Decaspermum parviflorum 

(Lam.) A.J.Scott var. parviflorum,  C. 



Kantachot 159. – Eucalyptus camal-

dulensis Dehnh., C. Kantachot 166. – 

Eugenia uniflora L.,  C. Kantachot 120. – 

Melaleuca cajuputi Powell, C. Kantachot 

72. – Psidium guajava L., C. Kantachot 

132. – Rhodamnia dumetorum (Poir) Merr. 

& L.M.Perry, C. Kantachot 15. – Rhodo-



myrtus tomentosa (Aiton) Hassk., C. 

Kantachot 49. – Syzygium albiflorum 

(Duthie & Kurz) Behadur & R.C.Guar, C. 



Kantachot 105. – S. aqeum (Burm.f.) 

Alston, C. Kantachot 202. – S. aromaticum 

(L.) Merr. & L.M.Perry, C. Kantachot 

194. – S. cinereum (Kurz) Merr & 

L.n.Perry,  C. Kantachot 185. – S. clavi-

florum (Roxb.) A.M.Cowan & Cowan, C. 

Kantachot 152. – S. cumini (L.) Skeels, C. 

Kantachot 29. – S. diospyrifolium (Wall. 

ex Duthie) S.N.Mitra, C. Kantachot 117. – 



S. formosum (Wall.) Masam., C. 

Kantachot 22 & C. Kantachot 34. – S. 

jambos (L.) Alston, C. Kantachot 207. – S. 

laetum (Buch.-Ham.) Grandhi subsp. 

jugorum (Craib) Chantar. & J.Parn., C. 

Kantachot 203. – S. malaccense (L.) Merr. 

& L.M.Perry, C. Kantachot 58. – S. mega-



carpum (Craib) Rathakr. & N.C.Nair, C. 

Kantachot 65. – S. ripicola (Craib) Merr. 

KANTACHOT ET AL. – LEAF ANATOMY AND TAXONOMY OF THAI MYRTACEAE 

45

& L.M.Perry, C. Kantachot 21. – S. 



samarangense (Blume) Merr. & L.M.Perry 

var. samarangenseC. Kantachot 195. – S. 



siamense (Craib) Chantar. & J.Parn., C. 

Kantachot 106. – S. winitii (Craib) Merr. 

& L.M.Perry, C. Kantachot 119. – S. 



zimmermanii (Warb.) Merr. & L.M.Perry, 

C. Kantachot 204. – Tristaniopsis burma-

nica (Griff.) Peter G.Wilson & 

J.T.Waterh. var. rufescens J.Parn. & Nic 

Luhadha, C. Kantachot 23. 

 

A



CKNOWLEDGEMENTS

 

 



We would like to thank Dr. David A. 

Simpson and Dr. Kenneth Courten

a

y for 


assistance with English language. This 

work was supported by the TFR/BIOTEC 

Special Program for Biodiversity Research 

and Training grant BRT T_145029. 

 

L

ITERATURE 



C

ITED


 

 

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Haron, N.W. and Moore, D.M. 1966. The 

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phology in the genus Eugenia L. Botanical 

Journal of the Linnean Society, 120: 256-

277. 


Hussin, K.H., Cutler, D.F. and Moore, D.M. 

1992. Leaf anatomical studies of Eugenia L. 

(Myrtaceae) species from the Malay 

Peninsula. Botanical Journal of the Linnean 

Society, 110: 137-156. 

Metcalfe, C.R. and Chalk, L. 1957. Anatomy 

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Parnell, J. and Chantaranothai, P. 2002. 

Myrtaceae. In: Santisuk, T. and Larsen, K. 

(Eds.). Flora of Thailand, vol. 7, part 4, 

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review of the genus Tristania  R.Br. 

(Myrtaceae): a heterogeneous assemblage of 

5 genera. Australia Journal of Botany, 30: 

413-446. 



 

Received: 22 November 2006 

Accepted: 11 May 2007 

 

 


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