Mots clés syzygium, Myrtaceae, Indochine, Cambodge, Laos, Viêtnam, révision taxonomique, lectotypification. Key words



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179

ADANSONIA, sér. 3 • 2015 • 37 (2) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. 

www.adansonia.com

MOTS CLÉS

Syzygium,

Myrtaceae,

Indochine,

Cambodge,

Laos,


Viêtnam,

révision taxonomique,

lectotypification.

KEY WORDS

Syzygium,

Myrtaceae,

Indochina,

Cambodia,

Laos,


Vietnam,

taxonomy revision,

lectotypification.

Soh W.-K. & Parnell J. 2015. — A revision of Syzygium Gaertn. (Myrtaceae) in Indochina (Cambodia, Laos and Vietnam). 



Adansonia, sér. 3, 37 (2): 179-275

http://dx.doi.org/10.5252/a2015n2a1



ABSTRACT

The genus Syzygium (Myrtaceae) is revised for Indochina (Cambodia, Laos and Vietnam). The spe-

cies key, descriptions, typification, ecology, conservation status, phenology, vernacular names, usages, 

distribution maps, selected herbarium images and list of specimens examined are given. A total of 

fifty-six species are recognised in Indochina. Eight species including Syzygium bokorense W.K.Soh & 

J.Parn., S. chantaranothaianum W.K.Soh & J.Parn., S. crassiflorum Merr. & L.M.Perry, S. cucphuongense 

W.K.Soh & J.Parn.S. glomerulatum (Gagnep.) Merr. & L.M.Perry, S. harmandii (Gagnep.) Merr. & 

L.M.Perry, S. pierrei (Gagnep.) Merr. & L.M.Perry and S. tonkinense (Gagnep.) Merr. & L.M.Perry 

are confined to Indochina. 63 lectotypes were designated and 23 previously accepted species names 

are now synonyms under Syzygium. Vietnam harbours the largest number of species (49), followed 

by Laos (30) and Cambodia (27). Syzygium attopeuense (Gagnep.) Merr. & L.M.Perry and Syzygium 

corticosum (Lour.) Merr. & L.M.Perry are new records for Thailand.

RÉSUMÉ

Révision du genre Syzygium Gaertn. (Myrtaceae) en Indochine (Cambodge, Laos et Viêtnam).

Le genre Syzygium (Myrtaceae) est révisé pour l’Indochine (Cambodge, Laos et Vietnam). La clé, les 

descriptions, la typification, l’écologie, le statut de conservation, la phénologie, les noms locaux, les 

usages, les cartes de répartition, des images sélectionnées dans un herbarium et une liste des spécimens 

examinés de l’espèce sont inclus. On trouve un total de cinquante-six espèces en Indochine. Huit 

espèces, dont Syzygium bokorense W.K.Soh & J.Parn., S. chantaranothaianum W.K.Soh & J.Parn., 



S. crassiflorum Merr. & L.M.Perry, S. cucphuongense W.K.Soh & J.Parn.S. glomerulatum (Gagnep.) 

Merr. & L.M.Perry, S. harmandii (Gagnep.) Merr. & L.M.Perry, le S. pierrei (Gagnep.) Merr. & 

L.M.Perry et S. tonkinense (Gagnep.) Merr. & L.M.Perry ne peuvent être trouvées qu’en Indochine. 

63 lectotypes ont été reconnus et 23 noms d’espèces acceptés sont maintenant synonymes sous « Syzy-



gium ». Le Viêtnam abrite le plus grand nombre d’espèces (49), suivi du Laos (30) et du Cambodge 

(27). Syzygium attopeuense (Gagnep.) Merr. & L.M.Perry et Syzygium corticosum (Lour.) Merr. & 

L.M.Perry sont maintenant des espèces recensées pour le Cambodge.

Wuu-Kuang SOH 

John PARNELL

Botany Department, School of Natural Sciences,  

Trinity College Dublin (Republic of Ireland)

sohw@tcd.ie

jparnell@tcd.ie

Published on 31 December 2015



A revision of Syzygium Gaertn. (Myrtaceae) in Indochina 

(Cambodia, Laos and Vietnam)

180

ADANSONIA, sér. 3 • 2015 • 37 (2)

Soh W.-K. & Parnell J.

INTRODUCTION



Syzygium Gaertn. is the largest genus in Myrtaceae with 

c. 1200 species of mostly medium to large trees occurring in 

the tropics and subtropics from India to the Pacific Islands, 

and found in a diverse range of habitats from sea shore to 

montane forest (Parnell et al. 2007). The highest concentra-

tion of species and morphological variation is found in the 

Southeast Asian region. The geographical range of Syzygium 

is from Africa through southern Asia, Malesia and Australia 

to the Pacific Islands. The historical basis and the classifica-

tion problems have been extensively reviewed and discussed 

by many authors (Schmid 1972; Craven 2001; Parnell et 



al. 2007). 

François Gagnepain, who was the first botanist to revise 

the Indochinese Syzygium under Eugenia L. s.l., initially 

published all the twenty-eight new species in Notulae Sys-



tematicae (Gagnepain 1917-1918) and later published the 

completed work in the Flore générale de l’Indochine (Gagne-

pain 1920). His work encompassed fifty-five species from 

French-Indochina, which corresponds to the currently rec-

ognised political boundaries in Laos, Cambodia, Vietnam, 

Thailand (part of Mekong basin) and South China (Zhan-

jiang). Merrill & Perry (1938b) later updated the revision 

based on partly original material supplied by Gagnepain and 

additional new material from Indochina, mainly collected 

by Pételot. In their synopsis (Merrill & Perry 1938b), all the 

Indochinese species were transferred from Eugenia L. s.l. to 

Syzygium Gaertn., the species nomenclature was updated, 

eight new species described and new records added. 

Since Gagnepain’s revision in 1920, the generic concept of 

Eugenia-Syzygium has changed and the current revisionary 

works of Syzygium in the neighbouring countries includ-

ing Thailand (Chantaranothai & Parnell 2002) and China 

(Chen & Craven 2007) have suggested several new records 

for Indochina. This is not surprising, since the Flore générale 

de l’Indochine project was prematurely initiated consider-

ing at the time many areas in Indochina were botanically 

unexplored. Since then, there have been many new botani-

cal collections from previously inaccessible localities. This 

coupled with the resurgence of botanical work over the last 

three decades in Cambodia, Laos and Vietnam has generated 

a significant amount of new information. Therefore it is now 

timely to revise Syzygium Gaertn. in Indochina (Cambodia, 

Laos and Vietnam).

C

haraCters



 

useful


 

in

 



identifiCation

 

and



 

genera


  

that


 

Can


 

be

 



Confused

 

with



 S

yzygium

Generally, Syzygium can be recognised easily by the simple, 

opposite, gland dotted (punctilate or pustulate) and exstipu-

late leaves with intramarginal veins. At species level, the 

vegetative characters that are useful for identification include 

the nature and colour of the twig, the leaf arrangement, the 

petiole length and the blade shape, size, colour and venation. 

There are a few reproductive characters useful for identi-

fying species. These are the inflorescence type and length, 

the hypanthium shape and size, the presence or absence of 

pedicel, the calyptrate or free sepals and to a limited extent, 

the fruit shape and size. At the infrageneric level, the hyp-

anthium shape, the ovule arrangement and placentation, 

the anther orientation and the presence or absence of seed 

intercotyledonary intrusion are useful for identifying sub-

generic groups.

In having opposite leaves and intramarginal veins, herbarium 

specimens of Syzygium are often confused with other genera 

such as Eugenia, Garcinia L., Memecylon L. and Rubiaceae Juss. 

All the taxa mentioned above, except for Eugenia, do not have 

a punctilate or pustulate leaf surface (oil glands). Garcinia has 

leaves with petiole bases that clasp the twig while Rubiaceae 

has interpetiolar stipules. Memecylon usually has a shiny and 

wrinkled leaf, and the stamens are crescent-shaped. Eugenia 

has hairy vegetative shoots and inflorescences, and few flow-

ers, from one to three per inflorescence.

MATERIAL AND METHODS

Most of the taxonomic publications relevant to Indochina 

and neighbouring regions were consulted. For the present 

treatment, dried plant specimens acquired from different 

herbaria (A, BK, BKF, BM, CPNP, E, GH, HN, K, KEP, 

L, NY, P, TCD, U and VFU) were examined. In addition 

to this, images and references downloaded from online 

herbaria were also used (A, GH, K, MO, NY and US). In 

this study, over 4000 specimens from Indochina and other 

regions were studied; either on loan or in situ in the vari-

ous herbaria. The study presented here deals with revisional 

work of a number of different species within Syzygium. In 

practice this means that herbarium specimens of the same 

phenotypic assemblage were sorted and grouped together. 

These groups were then either associated with an existing 

type specimen, which is linked to a particular taxon name. 

After the herbarium specimens were sorted into groups, their 

morphological characters were examined. Dichotomous keys 

based on vegetative and reproductive macromorphological 

characters were constructed. 

Unless indicated, all macromorphological measurements 

and observations, including colour, texture and shape are 

recorded from herbarium specimens. When no holotype 

or lectotype is available, then a lectotype is designated or a 

neotype selected by following the guidelines in the Interna-

tional Code of Botanical Nomenclature (McNeill et al. 2006). 

Discussion for each instance of this is found in the relevant 

species notes. Species synonyms were gathered from relevant 

publications, notably the World Checklist of Myrtaceae online 

database (Govaerts et al. 2015) and personal observations 

from herbarium specimens. When possible, the synonyms 

listed in this account were verified by examining the types 

or voucher specimens. In some instances the synonyms were 

excluded when they were found not to conform with the 

species studied here. We applied the IUCN Red List Catego-



ries and Criteria ver. 3.1 (IUCN 2001) in assessing the con-

servation status for each species. About 60% of herbarium 

specimens used in this study are more than fifty years old 


181 

Revision of Syzygium (Myrtaceae) in Indochina

ADANSONIA, sér. 3 • 2015 • 37 (2)

(1861-1959) and since then, the land-use, demography and 

forest cover in Indochina has drastically changed. Therefore 

the conservation status given is possibly overoptimistic and 

an underestimation of the current level of threat.

All specimens examined from Indochina are listed. Speci-

mens from countries outside Indochina are listed only for 

species with a narrow distributional range. The information 

from herbarium label were databased using BRAHMS ver. 

6.04. The species distributions in Indochina were mapped 

using DIVA-GIS ver. 6. The coordinates for mapping species 

distribution were gathered from herbarium labels, gazetteers, 

maps and NGA Geonet Names Server database (

http://earth-

A

B

C



D

F

ig



. 1. — Autographs of selected plant collectors or botanists relevant to Indochina: A, Gagnepain F.; B, Harmand J.; C, Pierre L.; D, Thorel C.

182

ADANSONIA, sér. 3 • 2015 • 37 (2)

Soh W.-K. & Parnell J.

info.nga.mil/gns/html/index.html

).

 The names of major areas 



(provinces) follow the standard used in the Flore du Cambodge 

du Laos et du Viêtnam publication. Distributions outside of 

Indochina were obtained from herbarium specimens and rel-

evant literature. List of examined specimens were organised 

alphabetically in the following sequence: country, major area, 

collector’s name and numerically by collecter’s number. The 

distributions of commercially cultivated species (Syzygium 



aqueum (Burm. f.) Alston, S. aromaticum (L.) Merr. & 

L.M.Perry, S. malaccense (L.) Merr. & L.M.Perry and S. sa-



marangense (Blume) Merr. & L.M.Perry) were not mapped. 

s

peCies



 

ConCept


infrageneriC

  

and


 

infraspeCifiC

 

ClassifiCation



For practicality in species identification, a morphological 

species concept (Sneath & Sokal 1973; Burger 1975; Stuessy 

2009) was employed in this study. The phylogenetic based 

infrageneric classification of Syzygium proposed by Craven & 

Biffin (2010) was adopted for this study. This implies that 

the genera Acmena DC. and Cleistocalyx Blume are now cir-

cumscribed in the genus Syzygium. Within Syzygium, four 

out of a total of six subgenera are currently documented in 

Indochina; Acmena (DC.) Craven & Biffin, Perikion Cra-

ven & Biffin, Sequestratum Craven & Biffin and Syzygium.

Gagnepain (1917; 1920) recognised five sections within 

Syzygium (excluding Acmena) based on the number of fer-

tile locule and the number of petals and their coherence. In 

accordance with Henderson (1949), we do not find the loc-

ule and petal characters taxonomically useful in delimiting 

subgeneric groups nor species. Therefore we do not accept 

Gagnepain’s infrageneric classification of Syzygium.

Infraspecific rank is recognised if a taxon shows a clear geo-

graphical distinction or a small amount of overlap and where 

morphological distinction is minor. In the present treatment, 

we did not recognise any infraspecific ranks for the following 

reasons: firstly, due to the paucity of the Indochinese collec-

tions and the lack of geographical and ecological information, 

it is difficult to ascribe allopatric or peripatric speciation with 

much confidence; secondly, there exists complex patterns of 

variation in some widespread species (e.g. Syzygium antisep-

ticum (Blume) Merr. & L.M.Perry, S. borneense (Miq.) Miq., 

S. claviflorum (Roxb.) Wall ex Steudel, S. cumini (L.) Skeels 

and S. hancei Merr. & L.M.Perry) that do not show clear mor-

phological discontinuities, therefore making identification of 

infraspecific ranks difficult or impossible. A comprehensive 

study focusing on each species or group of species is desir-

able. This should entail detailed phenotypic documentation, 

field observations and population genetic study. In the case of 

rheophytic species, those that has been formally recognised as 

distinct species from their close allies are maintained here in 

this study until further evidence is available to decide on their 

status (e.g. Syzygium abortivum (Gagnep.) Merr. & L.M.Perry 

and S. tsoongii (Merr.) Merr. & L.M.Perry). Morphological 

similarity among rheophytes is a result of convergent evolution; 

allopatric (ecological or geographical) evolution is thought to 

be the main mechanism of speciation for rheophytes (Van 

Steenis 1981; Mitsui et al. 2007). Because of this, ecological 

and geographical criteria are essential in deciding infraspecific 

rank. Unfortunately, these two crucial pieces of information 

are often lacking for Indochinese species. All formally recog-

nised infraspecific names are listed in the list of synonyms to 

highlight morphological variation within the respective species 

and brief comments on our observations are given. 

l

eCtotypifiCation



 

of

 i



ndoChinese

 S



yzygium

In his protologue, Gagnepain (1917-1918) usually cited one 

or more gatherings (i.e. plants from particular localities) and 

he did not indicate any as type. These gatherings are usually 

in duplicates and are mostly kept in P (some duplicates are 

found in BM, E, K, L & NY). Under article 9.2 of Interna-



tional Code of Botanical Nomenclature (McNeill et al. 2006), 

all of these specimens are available for lectotypification. In 

the case where the species have been ‘lectotypified’ by later 

workers, there is often no annotation reflect in this on the 

specimens and there are duplicates and undifferentiated sheets 

at P. Under article 9.15, second step lectotypification to narrow 

down to a single specimen is possible and undertaken herein. 

All the material deposited in P has been annotated by Gag-

nepain on the Herb. Museum Paris label (in addition to the 

Herb. Museum Paris label, the collector’s label is sometimes 

present). The difficulty lies in locating the material that was 

cited in the protologue. The specimen numbers indicated in 

the protologue are in fact usually species numbers and rarely 

collection numbers. Therefore gatherings from different locali-

ties often have the same specimen number, as they are filed 

within the same species pile (pers. obs. in P; Sovanmoly Hul 

pers. comm., April 2009). This is very obviously the case in 

Pierre’s and Thorel’s collections. Furthermore, the labels on 

duplicates are often incomplete, sometimes without locality, 

dates, specimen number or notes. However, usually among a 

set of duplicates, there is one specimen with a complete and 

original label by the collector and it is therefore essential to 

be able to recognise the collector’s handwriting (Fig. 1A-D). 

We reproduce these samples herein because the problems we 

found in respect of Syzygium are duplicated for other groups 

and because samples of the collectors’ handwritings are not 

readily available. Pierre’s Herbarium contains much of Har-

mand’s collection and, as a result, there are sometimes two 

specimen numbers on one herbarium sheet – we cite such 

specimens as, for example, Pierre 3290 (Harmand 1172). In 

some cases the developmental stage of the specimens coupled 

with the label information are good indicators as to whether 

or not they belong to the same gathering. A specimen is dif-

ferent if it has a different collection date and/or locality (the 

number could be absent or similar).

For Indochina, when lectotypification is required we have 

used two criteria to guide selection of lectotype. Firstly, in 

general we have selected the material in best physical con-

dition, but have also, when possible, selected herbarium 

specimens with Delpy’s or Gagnepain’s drawings attached, 

because the details in these drawings were used extensively 

by Gagnepain (1917-1918) in his species descriptions. In 

the Myrtaceae collections in P, Delpy’s drawings are usually 

attached to the respective herbarium specimens. 


183 

Revision of Syzygium (Myrtaceae) in Indochina

ADANSONIA, sér. 3 • 2015 • 37 (2)

SYSTEMATICS

Genus Syzygium Gaertn.

De Fructibus et Seminibus Plantarum 1: 166, t. 33 (1788), nom. 

cons. — Eugenia subg. Syzygium (Gaertn.) Wight, Illustrations 

of Indian Botany 2: 12, 15 (1841). — Jambosa sect. EuSyzygium 

Miq., Flora van Nederlandsch Indië 1: 447 (1855). — Eugenia sect. 



Syzygium (Gaertn.) M.R.Henderson, The Gardens’ Bulletin Singa-

pore 12: 17 (1949). — Type: Sri Lanka (Syzygium caryophyllaeum 

Gaertn.) (lecto-, WAG![WAG0002464]), designated by McVaugh, 



Taxon 5: 164 [1956]).

s

ynonymy



 

list


For generic synonyms see Farr & Zijlstra (1996-onwards) and 

Craven & Biffin (2010).

d

esCription



Tree to 35 m tall or shrub 1-3 m tall. Indumentum mostly 

absent, rarely present. Oil glands present in almost all organs, 

punctilate or pustulate on surface. 

Twig 

Terete, subangular, angular (3- or 4-angled) or winged when 

mature, 2-5 mm diameter, stout or slender, surface usually 

smooth, sometimes flaky or fissured, colour whitish, greyish, 

pale brownish to dark brownish, contrasting or not to leaf 

or petiole colour. 



Leaves

Usually opposite (decussate) or subopposite, sometimes alter-

nate or verticillate in whorls of three, petiolate or subsessile, 

2.5-50 × 2-9 cm, 1.5 to 12 times as long as wide, elliptic, 

ovate, obovate, lanceolate to oblong, base attenuate, cune-

ate, rounded to cordate, margin flat or recurved inward, 

apex usually acute or acuminate, sometimes rounded, rarely 

cuspidate or mucronate, acumen absent or present, 0.3-2 cm 

long, sometimes deflexed, blade chartaceous, subcoriaceous to 

coriaceous, upper surface usually punctilate, rarely pustulate, 

lower surface always pustulate, cork warts (black dots) always 

present, leaf yellowish brown, pale to dark brownish or green-

ish, always paler below, sometimes shiny on both surfaces 

A

D



bs

m

m



m

om

E



F

B

C



F

ig

. 2. — Leaf venation type in Syzygium. Abbreviations: m, intramarginal vein; om, outer intramarginal vein; bs, basal secondary veins. Drawing by the authors.



184

ADANSONIA, sér. 3 • 2015 • 37 (2)

Soh W.-K. & Parnell J.

or just the lower surface; midrib often sunken above, rarely 

flat, mostly smoothly raised below, rarely angled; secondary 

veins 7-45 per side, 0.5-20 mm apart, 30-60 degrees from 

midrib, prominent or faint, raised or sunken above, always 

raised below, narrowly (Fig. 2A) or widely spaced (Fig. 2B, C) 

and connecting to the prominent inner intramarginal veins, 

sometimes 1-3 basal secondary veins (acrodromous veins; Fig. 

2D) arcuate and not connecting but often extending to form 

outer intramarginal veins (Fig. 2C), sometimes intramarginal 

veins absent and all secondary veins arcuate (Fig. 2E) or 

looped (Fig. 2F)to form pinnate-like venation; intermediate 

secondaries present or absent, one to two in between second-

aries, prominent or faint; tertiary veins prominent or faint, 

reticulate, subscalariform or scalariform; intramarginal veins 

frequently present and always prominent, 0.5-5 mm from 

leaf margin, straight or scalloped, sometimes smaller outer 

intramarginal veins present (Fig. 2C); petiole 0.1-3 cm, 1/4-

1/140 of blade length, 0.5-5 mm diameter, slender or stout, 

brownish to dark brownish, to blackish, constrasting or not 

to blade colour. 

Inflorescence

Terminal or axillary on leafy twig or on leafless twig near 

proximal end, 1-17 cm long, solitary, racemose or paniculate-

cymose (first to fourth order branching), flowers few to many, 

one to 300, spreading or in compact heads; axes angular or 

terete, usually smooth, rarely flaky; bracts and bracteoles 

persistent or caducous, 0.3-2 × 0.3-2 mm, triangular, elliptic 

or lanceolate. 



Flowers

Sessile or pedicellate, pedicel 0.05-2 cm long, hypanthium 

1-20 × 0.3-20 mm, obconic, pyriform, elongate or clavate, 

usually smooth, sometimes glaucous, corrugate or warty, 

pseudostalk distinct or indistinct, 1-13 mm long; sepals 4-5, 

free or calyptrate (dome-shaped), triangular, ovate, semior-

bicular or orbicular, 0.3-10 × 0.6-18 mm, equal or unequal 

in size, if unequal outer lobes smaller than inner lobes; petals 

4-6, rarely 8, free or coherent, 1-17 × 1-17, semiorbicular to 

orbicular; stamens many, outer stamens 1-30 mm long; anther 

0.2-2 mm long, mostly oblong and sacs parallel, dehiscence 

by longitudinal slits or wider than long and sacs divaricate, 

dehiscence by apical slits (only in Subg. Acmena), connective 

gland conspicuous or inconspicuous; style 0.8-40 mm long; 

ovary inferior, 2 locules, rarely 3, all fertile, placenta mostly 

axile-median, rarely axile-apical (only in Subg. Acmena), 

ovules 5-35 per locules, arranged in two longitudinal rows 

or radiating. 



Fruit

A berry, 0.5-7 cm long, pyriform, globose, depressed globose, 

ellipsoid to obloid, surface smooth to rugose; with apical cav-

ity, deep or shallow, surrounded by calyx ring, with crown-

like persistent sepal lobes or undulating rim (formed from 

base of caducous sepal lobes or small sepals ring); seed 1-2, 

subglobose or ellipsoid, 0.35-3 cm, testa loosely or closely 

adhering to pericarp, cotyledons free or undivided, inter-

cotyledonary intrusion present or absent, if present ramified 

from the side or apex.

K

eys


 

to

 



speCies

 

of



 S

yzygium

 

Gaertn. 



in

 i

ndoChina



Two keys to the species of Syzygium in Indochina are provided using vegetative characters either in combination with fruit-

ing or flowering characters.

Key A. — This key is based mainly on vegetative and flowering characters. Unless indicated, all observed characters are based 

on dried herbarium specimens. The species positions in the key are arranged to reflect as much as possible their phenetic and 

hypothesised phylogenetic affinities.

1.  Anther sacs divergent; placentation axile-apical  ............. 2. S. acuminatissimum (Blume) DC. (Subg. Acmena)

—  Anther sacs parallel; placentation axile-median  ............................................................................................  2

2.  Hypanthium clavate or elongate with long pseudostalk, ovules in two longitudinal rows  .............................  3

—  Hypanthium obconic or pyriform, ovules irregularly arranged, radiating  ........................  12 (Subg. Syzygium)

3.  Hypanthium glaucous; petiole dark brownish to blackish, contrasting to blade colour  ...... 4 (Subg. Sequestratum)

—  Hypanthium not glaucous; petiole brownish, not contrasting to blade colour  .............................................  7

4.  Inflorescence racemose, short (2-2.5 cm long), few flowered (2-4)  ......  6. S. araiocladum Merr. & L. M. Perry 

—  Inflorescence paniculate, first to second order branching, long (3-7 cm long), many flowered (30-100)  ......  5

5.  Hypanthium abruptly constricting to pseudostalk; leaf apex with distinct acumen  ....................................   

 ......................................................................................................................... 40. S. odoratum (Lour.) DC.

—  Hypanthium gradually narrowing to pseudostalk; leaf obtuse or acute, without distinct acumen  .................  6

6.  Leaf blade elliptic to ovate, 1 to 3.5 times as long as wide  .......  4. S. antisepticum (Blume) Merr. & L.M.Perry

—  Leaf blade lanceolate, 3.5 to 5 times as long as wide  ......................  54. S. tsoongii (Merr.) Merr. & L.M.Perry

7.  Hypanthium wall not fibrous  ...........................................................................................  8 (Subg. Syzygium)

—  Hypanthium wall fibrous  ................................................................................................... 9 (Subg. Perikion)



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ADANSONIA, sér. 3 • 2015 • 37 (2)

8.  Flower sessile  ..............................................................................  8. S. attenuatum (Miq.) Merr. & L.M.Perry

—  Flower pedicellate  ........................................................................... 7. S. aromaticum (L.) Merr. & L.M.Perry 

9.  Leaf blade subsessile, petiole very short (0.05-0.1 cm); leaf base cordate, sometimes rounded  ........................   

 ...............................................................................................11. S. boisianum (Gagnep.) Merr. & L.M.Perry 

—  Leaf blade petiolate, petiole long (0.3-0.6 cm); leaf base cuneate or attenuate  ...........................................  10

10.  Leaf blade lanceolate  ............................................................... 1. S. abortivum (Gagnep.) Merr. & L.M.Perry

—  Leaf blade elliptic, oblong-elliptic, ovate to obovate  ..................................................................................  11

11.  Leaf blade small, 4-8.5 × 1-4 cm; petiole short, 0.15-0.2 cm  ..... 16. S. championii (Bentham) Merr. & L.M.Perry

—  Leaf blade large, 6-22 × 1.5-7.5 cm; petiole long, 0.3-0.6 cm  .......... 18. S. claviflorum (Roxb.) Wall ex Steude

12.  Hypanthium large, ≥ 1 cm long; inflorescence racemose ............................................................................  13 

—  Hypanthium small, < 1 cm long; inflorescence paniculate  .........................................................................  24

13.  Leaves verticillate, arranged in whorls of three at twig-end  ........................................................................  14

—  Leaves in pairs, opposite or subopposite at twig-end  ..................................................................................  16

14.  Inflorescence terminal, rarely axillary on leafy twig  ............. 23. S. diospyrifolium (Wall. ex Duthie) S.N.Mitra 

—  Inflorescence axillary on leafless proximal end of twig  ...............................................................................  15

15.  Leaf blade large, 15-35 × 6-18 cm  ................................................................ 25. S. formosum (Wall.) Masam.

—  Leaf blade small, 11-18 × 1.5-2.5 cm  ...................................  29. S. harmandii (Gagnep.) Merr. & L.M.Perry

16.  Inflorescence axillary on leafless proximal end of twig  .................... 35. S. malaccense (L.) Merr. & L.M.Perry

—  Inflorescence terminal or axillary on leafy twig  ..........................................................................................  17

17.  Leaf blade > 20 cm long  ............................................................................................................................  18

—  Leaf blade ≤ 20 cm long  ............................................................................................................................  19

18.  Tertiary venation ladder-like (scalariform); leaf blade oblong elliptic, 3 to 4 times as long as wide  ..................   

 ............................................................................................ 36. S. megacarpum (Craib) Rathakr. & N.C.Nair

—  Tertiary venation net-like (reticulate); leaf blade lanceolate, 5 to 9 times as long as wide  ................................   

 ............................................................................................................. 20. S. crassiflorum Merr. & L.M.Perry

19.  Basal secondary veins arcuate, not connecting to each other  ......................................................................  20

—  Basal secondary veins connecting to each other  .........................................................................................  21

20.  Leaf subsessile, petiole ≤ 2 mm long; leaf base cordate; hypanthium apex abruptly constricted to pseudostalk; 

fruit turbinate (in vivo)  ................................................................................... 5. S. aqueum (Burm. f.) Alston

—  Leaf petiolate, petiole > 2 mm long; leaf base cuneate, rounded or slightly cordate; hypanthium gradually constricted 

to pseudostalk; fruit pyriform to subglobose (in vivo)  ................ 46. S. samarangense (Blume) Merr. & L.M.Perry

21.  Petiole < 3 mm long  ..................................................................................................................................  22

—  Petiole ≥ 3 mm long  ..................................................................................................................................  23

22.  Leaf base cuneate; tertiary veins reticulate  ................................... 42. S. pierrei (Gagnep.) Merr. & L.M.Perry

—  Leaf base cordate; tertiary veins scalariform to subscalariform  ......  17. S. chantaranothaianum W.K.Soh & J.Parn.

23.  Leaf apex gradually narrowing without distinct acumen; hypanthium greenish (in vivo)  ...........................   

 ................................................................................................................................32. S. jambos (L.) Alston

—  Leaf apex with distinct acumen; hypanthium reddish-green (in vivo) ..........................................................   

 ....................................................................................................47. S. siamense (Craib) Chantar. & J.Parn.

24.  Inflorescence axillary on leafless proximal end of twig  ...............................................................................  25

—  Inflorescence terminal or axillary on leafy twig  ..........................................................................................  29

25.  Sepals calyptrate; intramarginal veins absent, all secondary veins arcuate  ..........  37. S. nervosum A.Cunn. ex DC.

—  Sepals free; intramarginal veins present  ......................................................................................................  26

26.  Basal secondary veins connect to adjacent secondary veins to form intramarginal veins  .............................  27

—  Basal secondary veins arcuate and not connecting to adjacent secondary veins to form intramarginal veins  .....  28

27.  Twig terete; secondary veins ≥ 20 pairs per side, narrowly spaced (2-5 mm)  ............. 22. S. cumini (L.) Skeels 

—  Twig angular (4-angled) and winged; secondary veins < 20 pairs per side, widely spaced (5-15 mm)  ..............   

 ........................................................................................................50. S. tetragonum (Wight) Wall. ex Walp.


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28.  Petals free; leaf base slightly unequal; leaf blade brownish  ...........................  43. S. polyanthum (Wight) Walp.

—  Petals coherent; leaf base equal; leaf blade greenish  ......................................  10. S. balsameum (Wight) Walp.

29.  Inflorescence hairy  ....................................................................................................................................  30

—  Inflorescence glabrous  ...............................................................................................................................  31

30.  Leaf blade hairy; twig brownish not contrasting to leaf colour; secondary veins widely spaced, ; tertiary veins 

subscalariform  ..........................................................................................  55. S. vestitum Merr. & L.M.Perry

—  Leaf blade glabrous; twig whitish contrasting to leaf colour; secondary veins narrowly spaced (1-3 mm); tertiary 

veins reticulate  ................................................................................ 33. S. levinei (Merr.) Merr. & L.M.Perry

31.  Sepals calyptrate  .............................................................................  38. S. nigrans (Gagnep.) Craven & Biffin

—  Sepals free  .................................................................................................................................................  32

32.  Inflorescence short, ≤ 3 cm long  ................................................................................................................  33

—  Inflorescence long, > 3 cm long  .................................................................................................................  35

33.  Leaf blade lanceolate, 5 to 8 times as long as wide  ............................. 48. S. sterrophyllum Merr. & L.M.Perry

—  Leaf blade elliptic to obovate, 1.5 to 3 times as long as wide  ......................................................................  34

34.  Leaf blade large (usually > 3 cm long); leaf dark greenish brown  ..................  28. S. hancei Merr. & L.M.Perry

—  Leaf blade small (≤ 3 cm long); leaf light brownish  ....................................... 15. S. buxifolium Hook. & Arn.

35.  Flower pedicellate  ......................................................................................................................................  36

—  Flower sessile  .............................................................................................................................................  41

36.  Inflorescence unbranched  .................................................................. 3. S. angkae (Craib) Chantar. & J.Parn.

—  Inflorescence branched at least once  ..........................................................................................................  37

37.  Basal secondary veins arcuate, not connecting to adjacent secondary veins  ................................................  38

—  Basal secondary veins connecting to adjacent secondary veins  ...................................................................  40

38.  Twig whitish, contrasting to leaf colour; leaf blade 3 to 4 times as long as wide  ..............................................   

 ............................................................................. 56. S. zimmermannii (Warb. ex Craib) Merr. & L.M.Perry

—  Twig brownish, not contrasting to leaf colour; leaf blade 2 to 3 times as long as wide  ................................  39

39.  Tertiary veins reticulate; secondary veins 7-15 mm apart; peduncle surface flaky  .......................................   

 ............................................................................................................ 30. S. hemisphericum (Wight) Alston

—  Tertiary veins subscalariform; secondary veins 15-20 mm apart; peduncle smooth  .........................................   

 ....................................................................................................................31. S. imitans Merr. & L.M.Perry 

40.  Leaf blade small (< 10 cm long), elliptic, oblong-elliptic or ovate, base cuneate and slightly attenuate; midrib 

smoothly raised below…  ............................................................... 34. S. lineatum (DC.) Merr. & L.M.Perry

—  Leaf blade large (≥ 10 cm long), obovate, base strongly attenuate; midrib angled below  .................................   

 ...................................................................................................... 24. S. fastigiatum (Bl.) Merr. & L.M.Perry

41.  Inflorescence with flowers clustering at branch-ends, compact; branch-ends with 7-13 flowers  ..................  42

—  Inflorescence not compact at branch-ends; branch-ends with 3 flowers  .....................................................  50

42.  Intramarginal veins absent, all secondary veins arcuate  ...............................................................................   

 ..................................................................................... 26. S. glomerulatum (Gagnep.) Merr. & L.M.Perry

—  Intramarginal veins present  .......................................................................................................................  43

43.  Secondary veins sunken above  ............................................................................ 13. S. borneense (Miq.) Miq.

—  Secondary veins raised above  .....................................................................................................................  44

44.  Hypanthium large, > 5 mm long; sepals unequal, inner sepals larger than outer sepals  ..............................  45

—  Hypanthium small, ≤ 5 mm long; sepals equal  ..........................................................................................  46

45.  Secondary veins 8-12 per side, 8-10 mm apart  ...................................................  27. S. grande (Wight) Walp.

—  Secondary veins 20-25 per side, 2-5 mm apart  ..................  41. S. pachysarcum (Gagnep.) Merr. & L.M.Perry

46.  Twigs whitish, contrasting to leaf colour  ....................................................................................................  47

—  Twigs brownish, not contrasting to leaf colour  ..........................................................................................  48

47.  Leaf blade elliptic or obovate, one to two times as long as wide …  ........................... 22. S. cumini (L.) Skeels

—  Leaf blade lanceolate, 2.5 to six times as long as wide…  ................  45. S. ripicola (Craib) Merr. & L.M.Perry


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ADANSONIA, sér. 3 • 2015 • 37 (2)

48.  Leaf apex mucronate  .......................................................................  21. S. cucphuongense W.K.Soh & J.Parn.

—  Leaf apex blunt , acute or acuminate  .........................................................................................................  49

49.  Secondary veins narrowly spaced, 2-3 mm; leaf apex without acumen  ........................................................   

 ................................................................................................. 51. S. thorelii (Gagnep.) Merr. & L.M.Perry

—  Secondary veins widely spaced, 5-8 mm; leaf apex with distinct acumen  ........................................................   

 ...............................................................................................53. S. tonkinense (Gagnep.) Merr. & L.M.Perry

50.  Leaf subsessile, petiole very short, ≤ 2 mm long, 1/25-1/60 of blade length; base cordate  ...............................  51

—  Leaf petiolate, petiole long, ≥ 3 mm, 1/7-1/22 of blade length; base cuneate or attenuate…  ..........................  52

51.  Twig whitish, contrasting to leaf colour…  ..................................  14. S. bullockii (Hance) Merr. & L.M.Perry

—  Twig dark brownish, not constrasting to leaf colour…  ............................ 12. S. bokorense W.K.Soh & J.Parn.

52.  Leaf blade lanceolate, 5 to 8 times as long as wide  .................. 9. S. attopeuense (Gagnep.) Merr. & L.M.Perry

—  Leaf blade elliptic, ovate, obovate to oblong, 1.5 to 4 times as long as wide  ...............................................  53

53.  Twig strongly angular (4-angled) and winged  .............................  44. S. praecox (Roxb.) Rathkr. & N.C. Nair

—  Twig terete  ................................................................................................................................................  54

54.  Secondary veins narrowly spaced, ≤ 2 mm  ..................................  49. S. syzygioides (Miq.) Merr. & L.M.Perry

—  Secondary veins widely spaced, ≥ 4 mm  ....................................................................................................  55

55.  Hypantium small, ≤ 2.5 mm long  .............................................  19. S. corticosum (Lour.) Merr. & L.M.Perry 

—  Hypanthium large, > 3 mm long  ...............................................................................................................  56

56.  Leaf blade dark dark brown to blackish  .........................................  52. S. thumra (Roxb.) Merr. & L.M.Perry

—  Leaf blade yellowish brown  ...............................................................  39. S. oblatum (Roxb.) Wall. ex Steudel

Key B. — This key is based mainly on vegetative and fruiting characters. Unless indicated, all observed characters are based on 

dried herbarium specimens. Syzygium chantaranothaianumS. fastigiatumS. glomerulatumS. harmandiiS. pierreiS. thumra

S. tonkinense and S. zimmermannii were excluded from this key because their fruits were not seen.

1.  Seed with intercotyledonary intrusion  .........................................................................................................  2

—  Seed without intercotyledonary intrusion  ....................................................................................................  6

2.  Fruit depressed globose  .......................................................................... 2. S. acuminatissimum (Blume) DC.

—  Fruit ellipsoid to obovoid  ................................................................................................... 3 (Subg. Perikion)

3.  Leaf blade subsessile, petiole very short, 0.05-0.1 cm; leaf base cordate, sometimes rounded .....................   

 ............................................................................................ 11. S. boisianum (Gagnep.) Merr. & L.M.Perry 

—  Leaf blade petiolate, petiole long, 0.3-0.6 cm; leaf base cuneate or attenuate  ...............................................  4

4.  Leaf blade lanceolate ................................................................ 1. S. abortivum (Gagnep.) Merr. & L.M.Perry

—  Leaf blade elliptic, oblong-elliptic, ovate to obovate .....................................................................................  5

5.  Leaf blade small, 4-8.5 × 1-4 cm; petiole short, 0.15-0.2 cm  ...... 16. S. championii (Bentham) Merr. & L.M.Perry

—  Leaf blade large, 6-22 × 1.5-7.5 cm; petiole long, 0.3-0.6 cm  .......... 18. S. claviflorum (Roxb.) Wall ex Steude

6.  Fruit diameter > 4 cm long  ..........................................................................................................................  7

—  Fruit ≤ 4 cm long  ......................................................................................................................................  15

7.  Leaves verticillate, arranged in whorls of three at twig-end  ........................... 25. S. formosum (Wall.) Masam.

—  Leaves in pairs, opposite or subopposite at twig-end  ....................................................................................  8

8.  Inflorescence axillary on leafless proximal end of twig  .................... 35. S. malaccense (L.) Merr. & L.M.Perry

—  Inflorescence terminal or axillary on leafy twig  ............................................................................................  9

9.  Leaf blade > 20 cm long  .....................................................  36. S. megacarpum (Craib) Rathakr. & N.C.Nair

—  Leaf blade ≤ 20 cm long  ............................................................................................................................  10

10.  Basal secondary veins arcuate, not connecting to each other  ......................................................................  11

—  Basal secondary veins connecting to each other  .........................................................................................  12



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11.  Leaf subsessile, petiole ≤ 2 mm long; leaf base cordate; fruit turbinate (in vivo)  ..... 5. S. aqueum (Burm. f.) Alston

—  Leaf petiolate, petiole > 2 mm long; leaf base cuneate, rounded or slightly cordate; fruit pyriform to subglobose 

(in vivo)  ...............................................................................  46. S. samarangense (Blume) Merr. & L.M.Perry

12.  Secondary veins narrowly spaced, ≤ 2 mm apart  .........................  49. S. syzygioides (Miq.) Merr. & L.M.Perry

—  Secondary veins widely spaced, > 2 mm apart  ...........................................................................................  13

13.  Leaf drying pale brownish, coriaceous  ................................................................  27. S. grande (Wight) Walp.

—  Leaf drying greenish, subcoriaceous to chartaceous  ....................................................................................  14

14.  Leaf apex gradually narrowing without distinct acumen  ..........................................  32. S. jambos (L.) Alston

—  Leaf apex with distinct acumen  ....................................................  47. S. siamense (Craib) Chantar. & J.Parn.

15.  Fruit distinctly whitish  .............................................................................................  16 (Subg. Sequestratum)

—  Fruit not distinctly whitish, greenish, yellowish or reddish  ........................................................................  19

16.  Infructescence racemose, short, 2-2.5 cm long  ...................................  6. S. araiocladum Merr. & L. M. Perry 

—  Infructescence paniculate, first to second order branching, long, 3-7 cm long  ...........................................  17

17.  Leaf apex with distinct acumen  .......................................................................... 40. S. odoratum (Lour.) DC.

—  Leaf obtuse or acute, without distinct acumen  ...........................................................................................  18

18.  Leaf blade elliptic to ovate, 1 to 3.5 times as long as wide  .......  4. S. antisepticum (Blume) Merr. & L.M.Perry

—  Leaf blade lanceolate, 3.5 to 5 times as long as wide  ......................  54. S. tsoongii (Merr.) Merr. & L.M.Perry

19.  Leaves verticillate, arranged in whorls of three at twig-end  ..........................................................................   

 ........................................................................................ 23. S. diospyrifolium (Wall. ex Duthie) S.N.Mitra

—  Leaves in pairs, opposite or subopposite at twig-end  ..................................................................................  20

20.  Leaf blade > 20 cm long  ......................................................................  20. S. crassiflorum Merr. & L.M.Perry

—  Leaf blade ≤ 20 cm long  ............................................................................................................................  21

21.  Infructescence hairy  ..................................................................................................................................  22

—  Infructescence glabrous  .............................................................................................................................  23

22.  Leaf hairy; twig brownish not contrasting leaf colour; secondary veins widely spaced; tertiary veins subscalari-

form  .........................................................................................................  55. S. vestitum Merr. & L.M.Perry

—  Leaf glabrous; twig whitish contrasting to leaf colour; secondary veins narrowly spaced (1-3 mm); tertiary veins 

reticulate  ......................................................................................... 33. S. levinei (Merr.) Merr. & L.M.Perry

23.  Infructescence axillary on leafless proximal end of twig  .............................................................................  24

—  Infructescence terminal or axillary on leafy twig  ........................................................................................  28

24.  Intramarginal veins absent, all secondary veins arcuate  ................................ 37. S. nervosum A.Cunn. ex DC.

—  Intramarginal veins present  .......................................................................................................................  25

25.  Basal secondary veins connecting to adjacent secondary veins to form intramarginal veins  ........................  26

—  Basal secondary veins arcuate and not connecting to adjacent secondary veins to form intramarginal veins  .....  27

26.  Twig terete; secondary veins ≥ 20 pairs per side, narrowly spaced, 2-5 mm  ............... 22. S. cumini (L.) Skeels

—  Twig angular (4-angled) and winged; secondary veins < 20 pairs per side, widely spaced, 5-15 mm ................   

 ........................................................................................................ 50. S. tetragonum (Wight) Wall. ex Walp.

27.  Leaf base slightly unequal; leaf blade brownish  ...........................................  43. S. polyanthum (Wight) Walp.

—  Leaf base equal; leaf blade greenish  ..............................................................  10. S. balsameum (Wight) Walp.

28.  Infructescence short, ≤ 3 cm long  ..............................................................................................................  29

—  Infructescence long, > 3 cm long  ...............................................................................................................  31

29.  Leaf blade lanceolate, 5 to 8 times as long as wide  ............................. 48. S. sterrophyllum Merr. & L.M.Perry

—  Leaf blade elliptic to obovate, 1.5 to 3 times as long as wide  ......................................................................  30

30.  Leaf blade larger, usually > 3 cm long; leaf dark greenish brown  ..................  28. S. hancei Merr. & L.M.Perry

—  Leaf blade smaller, ≤ 3 cm long; leaf light brownish  ...................................... 15. S. buxifolium Hook. & Arn.

31.  Infructescence unbranched  ................................................................ 3. S. angkae (Craib) Chantar. & J.Parn.

—  Infructescence branched at least once  ........................................................................................................  32



189 

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ADANSONIA, sér. 3 • 2015 • 37 (2)

1. Syzygium abortivum (Gagnep.) Merr. & L.M.Perry



Journal of the Arnold Arboretum 19: 101 (1938); P.H.Hô, An Il-

lustrated Flora of Vietnam (Câyco Viêtnam) 2: 45, fig. 3722 (1992); 

K.D.Nguyen, (Myrtaceae) Checklist of Plant Species of Vietnam 2: 900 

(2003); M.F.Newman et al.A Checklist of the Vascular Plants of Lao 

PDR: 244 (2007). — Eugenia abortiva Gagnep., Notulae Systematicae 3: 

316 (1917); Gagnep. in Lecomte, Flore générale de l’Indochine 2: 802, 

fig. 86 (1920). — Type: Laos, Viangchan [Vien-chang, Vien-tiane], fl., 

Thorel s.n. [2245] (lecto-, P![

P00589305

], here designated; isolecto-, 

P![


P00589306

]). — Remaining former syntypes: Laos, between Xieng-

kuoang and Pak-lay, fl., Thorel s.n. [2245] (P!); Thailand, Lakhon 

Nakhon Panom, fl., Thorel s.n. [2245] (P![

P00589304

]) (see Note).

a

dditional



 

material


 

examined


. — 

: sites -> default -> files -> articles -> pdf
files -> Parodontologie
files -> Бу хмао-югры
files -> Schedule b financial Contributions for Transition in the
files -> Guidance note on transferring contributions from one un agency to another for the purpose of programmatic activities
pdf -> Adansonia 37 (2), 2015 Index des nouveautés taxonomiques et nomenclaturales
pdf -> Laboratoire de Phanérogamie, Muséum national d’Histoire naturelle
pdf -> RÉsumé Description de Syzygium guehoi Bosser & Florens (Myrtaceae), nouvelle espèce de l’île Maurice affine de S. cymosum (Lam.) Dc var cymosum des Mascareignes, et de la var montanum J. Guého & A. J. Scott de la Réunion. Abstract
pdf -> A new species of Syzygium (Myrtaceae) from the Kalakkad-Mundanthurai Tiger Reserve in Peninsular India Madepalli Byrappa Gowdu viswanathan


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