The Species of Polyalthia in Berau Regency, East Kalimantan

THE SPECIES OF POLYALTHIA IN BERAU REGENCY,
EAST KALIMANTAN

BY :

DlAN AKBARlNI
NRP: 98285

POST GRADUATE PROGRAM
BOGOR AGRICULTURAL UNIVERSITY
2002

ABSTRAK

Dian Akbarini. Jenis Polyalfhia di Kabupaten Berau, Kalimantan Timur. Dibimbing
oleh: Prof. Dr. Ir. Edi Guhardja, M.Sc., Prof. Dr. Mien A. Rifai dan Dr. Paul J. A.
Kessler.

Penelitian marga Polyalthia Blurne ( Annonaceae) telah dilakukan di
Kabupaten Berau. Marga ini terdiri dari lima belas jenis (Polyalthia berauensis,
Polyalthia cauliflora, Polyalthia dicfyoneura, Polyalthia glauca, Polyalthia igniflora,

Polyalthia jenkensii, Polyalthia laferiflora, Polyalfhia microtus, Polyalthia oblonga,
Polyalthia obliqua, Polyalthia rumphii, Polyalthia sclerophylla, Polyalthia
sumafrana, Polyalthia tipuliflora, dan Polyalthia sp. A ). Polyalthia berauensis
diusulkan sebagai suatu jenis baru.

ABSTRACT

Dian Akbarini. The Species of Polyalthia in Berau Regency, East Kalirnantan.
Supervised by : Prof. Dr. Ir. Edi Guhardja, M. Sc., Prof. Dr. Mien A. Rifai, and Dr.
Paul J . A. Kessler.

The study of the genus Polyalthia Blurne (Annonaceae) in Berau Regency
is presented. The genus comprises of 15 species (Polyalthia berauensis,
Polyalthia cauliflora, Polyalthia dictyoneura, Polyalthia glauca, polyalthia igniflora,
Polyalthia jenkensii, Polyalthia lateriflora, Polyalthia microtus, Polyalthia oblonga,
Polyalthia

obliqua, Polyalthia rumphii, Polyalthia sclerophylla, Polyalthia

sumatrana, Polyalthia tipuliflora, and Polyalthia sp. A). Polyalthia berauensis is

proposed as a new species.

Surat Pernyataan

Dengan ini saya menyatakan bahwa tesis yang berjudul:
THE SPECIES OF POLYALTHIA IN BERAU REGENCY,
EAST KALIMANTAN
Adalah benar merupakan hasil karya saya

sendiri dan belum pernah

dipublikasikan. Semua sumber data dan informasi yang digunakan telah
dinyatakan secara jelas dan dapat diperiksa kebenarannya.

Bogor, Mei 2002

NRP: 98285

THE SPECIES OF POLYALTHIA IN BERAU REGENCY,
EAST KALIMANTAN


BY :
DlAN AKBARlNl
NRP: 98285

A Thesis Submitted for the Degree of Master
at
Post Graduate Program Bogor Agricultural University

Department of Biology
POST GRADUATE PROGRAM
BOGOR AGRICULTURAL UNIVERSITY
2002

Tittle of Thesis

: The Species of Polyalthia in Berau Regency, East
Kalimantan

Name


: Dian Akbarini

NRP

: 98285

Study Program

: Biology

Approved

1.Supervisor Committee

Prof. Dr. Ir. Edi Guhardia, M.Sc.
Chairman

Dr. Paul J.A. Kessler
Supervisor


Prof. Dr. Mien A. Rifai
Supervisor

f Biology Program

Day of Graduation

: May, 04, 2002

Graduate Program

CURRICULUM VITAE

Dian Akbarini was born on June, 12, 1975 in Pangkalpinang, as the first
daughter from four children, from father H. M. Azro'i

Kosim and mother Hj.

Nur'aini.

In 1987, she graduated from SD Negeri No. 9 Tanjungpandan, in 1990
she graduated from SMP Negeri 1 Pangkalpinang, and in 1993 she graduated
from SMA Negeri 2 Pangkalpinang. She graduated from Biology Department of
the Faculty of Mathematics and Natural sciences, Padjadjaran University in
Jatinangor, 1998.
In 1998, she registered as a post graduated student at Bogor Agricultural
University.

ACKNOWLEDGEMENT

Alharndulillah.
I would like to thanks to my supervisors : Prof. Dr. Ir. Edi Guhardja, M.Sc,
(chairman of the supervisor committee), Prof. Dr. Mien A. Rifai and Dr. Paul J.A.
Kessler (members of the supervisor committee) for their supervising this study.
My thanks are due to Dr. Rugayah, M.Sc and Dra. Tutie Djarwaningsih,
M.Si for her support and comment of this manuscript and to Dr. lrawati head of
Herbarium Bogoriense who permitted this study to be executed in the Herbarium
Bogoriense. Miss Rina and Herbarium staff to provided some references which
are related to this study. Ir. Teguh Triono, M.Sc. and Yelita, S.Si.


helped to

make photographs, Mr. Sobari assisted in making the illustrations and Mr. Arif
Hidayat helped in compiling specimens from East Kalimantan.
I would like also to thanks all post graduate students of Plant Taxonomy.
IPB '98 for their support.
Finally to my parents, for their moral support and financial support of my
study; my brothers for their moral support and last but not least, to my husband
Ade Kelana Risfiardy, ST., for his moral support, his understanding and deep
patient.

TABLE OF CONTENTS
ABSTRAK

....................................................................................................ii

ABSTRACT

....................................................................................................
111


CURRICULUM VITAE

................................................................................vi

ACKNOWLEDGEMENTS ............................................................................vii
TABLE OF CONTENTS ............................................................................
LIST OF TABLE AND FIGURES

viii

............................................................... x

INTRODUCTION ..................................... ....................................................1
MATERIAL AND METHOD

........................................................................ 2

RESULT AND DISCUSSION ...................................................................
3


. . .

Geographical Distr~but~on
...............................................................
Floristic Analysis of Polyalthia in Berau Regency

............................

3

6

Morphological Characters ............................................................ 7
Habit

............................................................................................8

Twig ............................................................................................


7

Leave

.................................... .. ..................................................

Flower

..........................................................................................
8

Stamen

.........................................................................................9

7

Carpel ...........................................................................................
9


. . ........................................................................................9
Carp~d~a
. .
Key and Descript~ons.........................

.
.
...........................................11

Key to Species

.............................................................................12

Polyalthia berauesis ....................................................................

14

Polyalthia cauliflora .......................................................................

15

Polyalthia dictyoneura ................................................................... 17
Polyalthia glauca ..........................................................................

19

. . .
Polyalthra rgnrflora ........................................................................ 21
Polyalthia jenkensii

........................................................................
23

Polyalthia lateriflora ....................................................................... 25
. .
Polyalthra mrcrotus

........................................................................
26

Polyalthia obliqua ................................ ....

.................................. 28

Polyalthia oblonga ......................................................................... 29
Polyalthia rumphii ............................. ............................................. 30
Polyalthia sclerophylla ..................................................................

32

Polyalthia sumatrana ...................................................................

33

Polyalthia tipuliflora

35

Polyalthia sp.A
CONCLUSION

............................................................................. 37

............................................................................................
40

REFERENCES ........................................................................................ 4

1

LIST OF TABLES AND FIGURES
TABLES
1. The Distribution of ThePolyalthia in Berau Regency .................................

4

2. The Polyalthia in Berau Regency based on Sist & Saridan (1998)

..

and Akbar~n~
(2002) .................................................................................

FIGURES
1. Distribution of Polyalfhia in Berau Regency ............................................
2 . The various size and shape of flowers Polylathia ...................................

3. Stamen Polyalfhia .................................................................................
4 . The carpidia shapes of Polyalthia ...........................................................

5 . Polyalthia berauensis .............................................................................
6. Polyalthia cauliflora...............................................................................
. .
7 . Polyalthra drctyoneura ...........................................................................

8. Polyalthia glauca ..................................................................................

. . .

9. Polyalthra rgn~flora.................................................................................

. .
10. Polyalthra jenkensii .................................................................................
11. Polyalthia lateriflora ................................................................................

. .
12. Polyalthra rnrcrofus .................................................................................
13. Polyalthia obliqua ...................................................................................
14. Polyalthia rurnphii ...................................................................................
15. Polyalthia sclerophylla ..........................................................................
16. Polyalthia surnatrana ..............................................................................

. . .

................................................................................
17. Polyalthia t~pulrflora
18. Polyalthia sp.A ........................................................................................

6

The genus was first published by Blume (1829) based on Polyalthia
subcordata Blume. Polyalthia Blume is one of the largest genera in Annonaceae,
containing over 100 species and has wide distribution. The generic name
Polyalthia is derived from the Greek words polus- much and althein - to heal,
probably because it was much used as native medicine (Corner, 1940).
Hildebrandt (1952) enummerated one species (Polyalthia g1auca)in
Berau (Borneo). Kessler and Heusden (1993) recorded 7 species (Polyalthia
glauca, Polyalthia sumatrana, Polyalthia lateriflora, Polyalthia rumphii, Polyalthia
flagellaris, Polyalthia cauliflora and Polyalthia microtus) in Balikpapan

-

Samarinda area, East Kalimantan. Argent et.al. (1998) described 4 species
(Polyalthia sumatrana, polyalthia glauca, Polyalthia rumphii, Polyalthia lateriflora
and Polyalthia xanthopetala) in Central Kalimantan. Sist and Saridan (1998)
described

11 species Polyalthia in this Labanan area, 5 of them identified

(Polyalthia beccarii, Polyalthia glauca, Polyalthia lateriflora, Polyalthia rumphii,
and Polyalthia sumatrana ), and others were unidentified. Recently, Johnson and
Murray (1999) recorded 4 new species of Polyalthia from Borneo (Polyalthia
endertii, Polyalthia igniflora, Polyalthia montis-silam and Polyalthia tipuliflora ).
A study of Polyalthia from Berau Regency is undertaken as part of a
general floristic study conducted there. The aim of this study is to inventorize, and
to make identification key by using morphological characters. This information
can be used for development and conservation of Polyalthia in the future.

MATERIAL AND METHOD

This research was conducted in the Herbarium Bogoriense (BO), from
January2000-June2000.
Research materials were
Bogoriense (BO,

herbarium specimens from

Herbarium

Herbarium Wanariset (WAN) and living collection in Bogor

Botanical Garden.
This research used the descriptive method, as described by Leenhouts
(1968), Rifai (1976), de Vogel (1987) and Maxted (1992), : select all herbarium
specimens of Polyalthia from Berau regency, sort the specimens into groups
according to their overall resemblances, gather the literature, identify and
describe the herbarium specimens, analyse the data, deliniate the taxa, construct
an identification key, the nomenclature and to write the report.

RESULT AND DISCUSSION
Based on herbarium specimen, there are 15 species which can be
recognized, namely; Polyalthia berauensis, P. cauliflora, P. dictyoneura, P.
glauca, P. igniflora, P. jenkensii, P. lateriflora, P. microtus, P. obliqua, P. oblonga,
P. rumphii, P. sclerophylla, P. sumatrana, P. tipuliflora and Polyalthia sp. A.
Polyalthia berauensis is proposed as a new species.
Geographical Distribution
The genus is distributed throughout the Old World Tropics and particularly
numerous in South East Asia (Sinclair, 1955).
In this study, the species are found in logged over forest, primary forest,
low hills, logged over dipterocarp forest, limestone rocks, secondary forest and
logged over mixed dipterocarp forest. (Table. 1 and Fig. 2).

Table 1. The Distribution of The Polyalfhfa in Berau Regency

Floristic Analysis of Polyalthia i n Berau Regency

In this floristic study, there are 15 species which can be recognized based
on the specimen deposited in herbarium Bogoriense (table 2).
Table. 2

. The Polyalthia i n Berau regency based o n Sist & Saridan (1998)

and Akbarini (2002)

Morphological Characters
Habit
This character is important to recognized in species level. Species of
Polyalthia in Berau regency are either treelets or trees. The treelets are P.
berauensis, P. cauliflora, P. igniflora, P. microtus, P. rumphii, P. tipuliflora and
Polyalfhia sp.A. The trees species are P. dicfyoneura, P. glauca, P. jenkensii, P.
laferiflora, P. oblonga, P. obliqua, P. sclerophylla and P. sumatrana.

Twig
The twigs are circular, sometimes lenticelled (P. cauliflora, P. dictyoneura,
P. jenkensii, P. lateriflora, P.sclerophylla and Polyalthia sp.A). The twigs are
glabrous and hairy. lnduments type in this genus is simple hair. The twigs are
hairy in P. berauensis, P. cauliflora, P. jenkensii, P. microtus, P. oblonga, P.
rumphii and P. tipuliflora.

Leave
Some species of Polyalthia can be recognized by their leaves. Their
shapes are narrow ovate, oblong, broadly lanceolate, elliptic, spathulate or
oblanceolate. Generally acute or rounded base. Polyalthia obliqua is specific at
base, cuneate. Their apexes are various, apex, acuminate or caudate. The size
of leaves varies from 4.6 crn to 50 cm. The biggest size is at P. lateriflora ( 7.7
crn- 50 crn). Polyalthia glauca and P. sumatrana differ from others, because they
have glaucous of their below surface, and their secondary veins unclear and
difficult to be counted. Polyalthia microtus has leaves with rise in furrow midrib in
upper surface and this is a specific character of this species. Polyalthiajenkensii
has densely hairs on both side.

Flower
The flower is important character to recognize in species level.
Calyx

Calyx in this genus has 3 sepals, imbricate, usually triangular, sometime
broadly triangular (Polyalthia jenkensii), and fleshy. It has simple hairy in the
outside and the margin. The size is from 0.5 mrn to 3 cm. The apex of sepal is
acute.
Corolla
The corolla consists of two whorls, inner petal (3) and outer petal (3). The
inner petal and outer petal are various in size or shape (fig. 2), equal or unequal.
The petal usually hairy in outside and margin. The shape of inner petal and outer
petal are elliptic, narrow ovate and ovate, with acute apex. The position of the
inner petal to outer petal is valvate.

-

b. P. microtus. c. P. berauensis. d. P. dictyoneura. e. P. sumatrana.
lateriflora. g. P. rumphii.

f. P.

Stamen
Type of the stamen in Polyalthia is uvarioid (Fig. 3). Uvariod stamen is
broad with conspicuos connective stamen ( kessler, 1993).The shape of stamen
is cylindrical. Generally, the size of stamen is 1 mm in length, except P.
berauensis, P. igniflora, P. microtus, P. rumphii P. tipuliflora and Polyalthia sp.A
which are 2 mm in length. lndument in the stamen can be used to recognize the
species;Polyalthia microtus differ from other species, because this species has

Carpel
Generally, cylindrical shaped. The carpels are covered by simple hairs,
and the size is mostly less than 2 mm in length, cylindrical shaped . The carpels
in this genus are numerous ( more than 20).
Carpidia
The carpidia shapes of the Polyalthia are globose (P. glauca) or ellipsoid
(P.cauliflora, P. dictyoneura, P. jenkensii, P. lateriflora, P. microtus, P. rumphii, P.
sclerophylla and P. sumatrana), with either acute apex or rounded apex. The

apex on carpidia can be used to identify the species (fig.4). Polyalthia rumphii,
has glandular in its carpidia, whereas the other species have no glandular.
Polyalfhia obliqua is yellow colour when dry, whereas the are others black or dark
brown colour when dry.

Fig. 4. The carpidia shapes o f Polynlthia. a. Polyalthia sclerophylla. b. P.

slmiatma. c. P. nrmphii. d. P. cauliflora. e. P. glauca. f. P. obliqua. g. P.
Interrjlorn.
The seed shaped is mostly ellipsoid. The number of seed are 1 or 2

KEYS AND DlSCRlPTlONS
POLYALTHIA

Polyalthia Blume, FI. Jav. Anonac. (1829) 68; Hook.f., FI. Br. Ind. 1 (1875)
62; King, Ann. Roy. Bot. Gard. Cal. (1893) 62; Boerl., Icon. Bog. 1 (1901) 103;
Koord & Valet., Meded Land Platent. ixi. (1903) 2983; Ridley, FI. Malay Pen. 1
(1922) 48; J. Sincl., Gard. Bull. Sing. 14 (1955) 279; Whitmore, Tree FI. Mal. 1
(1972) 85, Bakhuizen & Backer. Jr., FI. Jav. 1 (1963) 106; Kessler, P.J.A. &
Heusden, Rheedea 3.1 (1993) 75; Heusden, Blumea suppl. 7 (1992) 125.
Shrubs, treelets or trees up to 40 m. Bark smooth, lenticelled or
unlenticelled, fibrous, bole circular, straight. Twigs glabrous or hairy, lenticelled or
unlenticelled. Leaves: petioles glabrous or hairy (simple hairy), 1-9 mm in length,
1-9 mm in diameter; lamina oblong, narrow ovate, oblanceolate, broadly
lanceolate; texture carthaceous to coriaceous, sometime shiny above;

base

acute, rounded, or sub cordate; apex acute, acuminate or caudate; margin
entire; glabrous or hairy above and below surface, glaucous or not glaucous in
below surface; midrib above sunken, flat or raised in furrow, midrib in below
surface prominent. Flowers solitary or cluster, axillary, supra axillary, cauliflorous
or ramiflorous. Flowers bisexual; sepals 3, triangular, 0.5 mm- 3 cm, usually hairy
in margin, and outside, valvate, fleshy; petals in two whorls ( inner petals and
outer petals), equal or unequal in size and shaped, elliptic, narrow ovate or ovate,
apex acute, inner petal 3, glabrous outside, fleshy, outer petals 3, glabrous
outside, fleshy. Stamens numerous (more than 20), fleshy, have connective
prolongation (uvarioid stamen), 0.7 mm

- 2mm in length. Carpels numerous

(more than 20), cylindrical. Carpidia: stalked, globose or ellipsoid shape, apex
acute or rounded. Seed 1 to 2.

Distribution: Assam, lndo China, Thailand, Malay Peninsula, Philippines,
Sumatra, Java, Celebes, Borneo, New Guinea, Africa, Australia.

Key To Species of Polyalthia
I.a. Tree. diameter more than 12 cm

.

b. Treelet, diameter less than 12 cm

.................................................

2

.................................................9

2. a. Leaves glabrous above and below surface

........................................

3

..........................PPobl.onga

b. Leaves hairy above and below surface

3. a. Leaves below surface glaucous, secondary veins not distinct

....................4

b. Leaves below surface not glaucous, secondary veins clear

...................5

4. a. Carpidia globose, intrasecondary veins 10-15 mm

..................P. glauca

b. Carpidia ellipsoid, intrasecondary veins 2-5 mm

.............P. sumatrana

5. a. Carpidia globose, seed globose

................................................6

b. Carpidia ellipsoid, seed ellipsoid

................................................
.7

6. a. Leaves coriaceous, midrib in above sunken

b. Leaves charthaceous, midrib in above flat

......................P. sclerophylla

......................P. dictyoneura

................... 8

7. a. Midrib in above sunken, base acute, truncatus or rounded
b. Midrib in above surface flat, base cunete

.......................... .P. obliqua

8. a. Secondary veins 5-7 in pairs, leaves chartaceous, petals ovate

P. jenkensii
b. Secondary veins 7-8 in pairs, leaves coriaceous, petals elliptic ..P. laterifora

9. a. Midrib hairy above
b. Midrib glabrous above

.....................................................................
I0
...................................................................... 11

.

10. a. lndument pale yellow 2-3 mm long

................................. P . berauensis

b.lndument dark brown or light brown, less than 1 mm long
11. a. Leaves hairy above, densely hairy below

......... P. rumphii

........................................

b. Leaves glabrous above, glabrous or sparsely hairy below

12

...................13

12. a. lnner petals 2.5

- 3.9 cm in length, secondary veins 14-21 pairs ........
P. igniflora

b. lnner petals 4.2

- 5-9 cm in length, secondary veins 21-24 pairs .........
P. tipuliflora

13. a. Flowers cauliflorous, stamen less than 2 mm in length
b. Flowers axillary or supraaxillary, stamen 2 mm in length
14. a. Midrib raised and furrowed above, twigs unlenticelled
b. Midrib flat above, twigs lenticelled

P. cauliflora

............14

.... P. microtus

... ... ... .......... Polya1thia.sp.A

1. Polyalthia berauensis Akbarini sp.nov.
Polyalthia berauensis. --Type : Ambriansyah et al. Berau 839. Borneo
(BO).
Folia chartacea, oblonga-lanceolata. 31-34 cm longa, 8.5-9.5 cm lata,
basi cordata, apice acuti-acuminati, newis lateris 20-26 paribus. Petioli 2.4 mm
diametro, 3 mm longi. Pedicello ca. 6 mm longa. Sepalum ca. 5 mm longa.
Interior petalum 2.7 cm longa, exterior petalum 2.5 cm longa. Stamen 2 mm
longa. Typus : Ambransyah etal. Bemu 839. Borneo (Holo: 80).
Treelet 7 m tall, 6 cm in diameter. Bark black, fibrous, rachis and lower
surface of leaves with long pale yellow hairs. Twigs 4-5 mm in diameter, densely
hairy (long hairy, 2-3 mm in length). Leaves: petioles hairy, 2.4 mm in diameter,
3 mm in length; lamina charthaceous, oblong

- lanceolate 31-34 cm by 8.5-9.5

cm; base subcordate, apex acute-acuminate; hairy above and below; midrib flat
above and hairy below, secondary veins flat above, prominent below; secondary
veins 20-26 in pairs. Flowers on the main bole; pedicel 6 mm in length, densely
hairy, like silky hairs; sepals triangular, 5 mm in length, hairy; petals elliptic,
creamy white, densely hairy, inner petals 2.7 cm in length, outer petals 2.5 cm in
length. Stamens 2 mm in length. Carpidia not seen.
DISTRIBUTION : Borneo (Berau )
HABITAT AND ECOLOGY : Logged over forest. Altitude : 50 m.
SPECIMEN EXAMINED: Ambriansyah et al. Berau 839.
NOTE: This species differs from other species in its indument. It has long
hairs, 2 - 3 mm which are pale yellow or brown yellow when dried.

Fig. 5. Polyalthia berauensis sp. nov. A. Sepal. B. Inner petal. C. Outer petal. D.
Stamen. E. Flower,

2. Polyalthia cauliflora Hook. f . & Thorns.
Polyalthia cauliflora Hook. f. & Thorns., FI. Ind. 1 (1855) 138; Hook.f.,

FI. Br.

Ind. 1 (1875) 66; Airy Shaw, Kew Bull. (1939) 280. Ridl., FI. Mal. Pen. 1 (1922)
51; Kessler & Heusden, Rhedeea 3, 1 (1993) 78 Fig. 9. 79.
Polyalthia macrorhyncha Miq.. Bot Ann. Mus. Lugd.-Bat. 2 (1865) 14.
Monoon teysmannii Miq., Ann. Mus. Bot. Lugd. - Bat. 2 (1865) 19.

Treelet or tree up to 11 m tall. Bark smooth, black, inner bark thick, hard,
bole straight. Twigs hairy, 0.15 in diameter, lenticelled. Leaves: petioles hairy 1
mm in diameter, 2-5 cm in length; lamina charthaceous, young leaves purple,
ovate to oblong, 4.6

- 14.8 by 0.9 - 3.8 cm; base acute, apex caudate; glabrous

above and hairy below; midrib sunken and glabrous above, hairy and prominent
below; secondary veins sunken and glabrous above, prominent and hairy below,
9 in pairs. Flowers on the main bole, cauliflorous, on twigs, short-tubercled shoot;
pedicel glabrous, 1 mm in diameter, 1-2 cm in length; sepals triangular, 3 mm,
hairy in outside and margin; petals greenish yellow, equal (inner and outer petals)
narrow ovate, 1.6-1.8 cm hairy; stamens 1 mm, creamy yellow. Carpidia: green
when young, stalked 1.1-2.5 cm, ellipsoid, apex acute, 3-7 mm in diameter.
Carpels cream yellow. Seed 2.
DISTRIBUTION : Malay Peninsula, Sumatra, Thailand, Borneo ( Kelai
River near Long Lanuk, Berau)
HABITAT AND ECOLOGY : Logged over forest along road, on slope,
primary forest, undulating area, low hills. Altitude 30 m.
SPECIMENS EXAMINED: Kostermans 21167, Ambriansyah et al. Berau
878, F. Slik, Pono & Rasyadi Berau 547, Kade Sidiyasa et al. Berau 1204, Z.
Arifin Berau 569, Kessler P.J.A; Arbainsyah B 1371, K. Sidiyasa et al. Berau
1255.

Fig. 6. Polyalthia cauliflora A. Flower. B. Carpidia. C. Stamen. D. Sepal
E. Carpel.

3. Polyalthia dictyoneura Diels
Polyalthia dictyoneura Diels, Mitt. Inst. Bot. Hamburg, vii (1927) 78.Type: H. Winkler 1148, West Borneo, Bukit Mehipit, 29.12.1924 (Iso: BO)
Tree 7-20 m tall, 15-20 cm in diameter. Bark smooth, blackish, hooped,
bole tapered.

Twigs 3.5 mm in diameter, lenticelled. Leaves: lamina

chartaceous, oblong, 9.2 -24.8 cm by 3.2 -15.3 cm; base subcordate; apex
acute to acuminate glabrous above and below, midrib flat and glabrous above.

prominent and glabrous below; secondary veins prominent above and below, 1012 in pairs. Flowers

ramiflorous; pedicels 1 mm in length; hairy; sepals

triangular, 2.5 mm in length, hairy; petals when young green, inner petals ovate;
ca. 7 mm in length; outer petals ovate, 6.7-7 mm in length. Stamen and carpel
not seen. Carpidia: stalked 11 mm in length, ellipsoid, apex acute, diameter 5-6
mm.
DISTRIBUTION : Borneo ( along Sungai Sepan at base Mt. Njapa, Kelai
River)
HABITAT AND ECOLOGY: logged over mixed dipterocarp forest, along
river bank, primary forest, on limestone rocks, covered by thick humus. Altitude:
40 m. Flowering : October. Fruiting : October.
FIELD NOTES: Flowers strong sweet smell.
SPECIMENS EXAMINED : Arifin, Z. et al. B 1458, Kessler, P.J.A.;
Arbainsyah B 1345, 51416, 51425. A. Kostermans 21196 (BO).

Fig. 7. Polyalthia dictyoneura. A. Sepal. B. Inner petal. C. Outer petal.
D. Stamen. E. Carpel. E. Lenticelled on twig.
4. Polyalthia glauca Boerl.

Polyalthia glauca Boerl., Icon. Bogor. 1 (1901) 104; Koord. & Valeton,
Bijdr. IX (Meded. PI. Tuin No. LXI) (1908) 285; Whitmore. Tree FI. Mal. 1 (1983)
90. Fig. 10; Kessler & Heusden, Rhedeea 3, 1 (1993) 80. Fig. 10. 81.
Unona merritiiMerr.. Philip. J. Sci: 1, Supp. 1 (1906) 190.
Polyalthia merritii (Merr.) Merr., Philip. J. Sci. 10 (1915) 250.
Polyalthia parkinsonii Hutchinson, Bull. Misc. Inform. (1917) 25.

Tree up to 41 m tall, up to 47 cm in diameter. Bark smooth, light orangegreyish, bolestraight, circular. Twigs glabrous 6 mm in diameter. Leaves: lamina
chartaceous; oblong to broadly lanceolate, 8-15 cm by 3-4 cm; base acute, apex
acuminate -caudate; glabrous above, glaucous below; midrib sunken and
glabrous above, prominent below; secondary veins flat above and below, unclear;
intra secondary veins 1- 1.5 cm. Flowers not seen. Stamen not seen.
Carpidia:when young linght green, ripe dark, stalked 0.7 cm

- 4.2 cm in length,

globose, 1.7-1.9 cm in diameter. Seed 1, 16 mm in diameter.
VERNACULAR NAME : Landur
DISTRIBUTION: Malay Peninsula, Sumatra, Java, Celebes, Philippines,
Borneo (Kampoeng lnaran, Berau).
HABITAT AND ECOLOGY

: Logged over dipterocarp forest. Altitude

75-100 m, Fruiting : October.
SPECIMENS EXAMINED: Arifin, Z; Arbainsyah B1341; Van der Zwaan
600 (00).
NOTE: The species is similar to P. sumatrana in leaf character, which is
glaucous below. However they differ in carpidia shape. Polyalthia glauca has
globose carpidia while Polyalthia sumatrana has ellipsoid carpidia.

Polyalthia. igniflora D.M. Johnson, Contr. University of Michigan
Herbarium. 22.1999.96.--Type:

Clemens & Clemens 32841. Borneo. Sabah

(East Malaysia). Mt. Nunkok, 2500-3500 ft. 13 April 1993 (fl) (Holotype : A.
Isotype: U. Ny, not seen).
Treelet, 3m, 6-23 cm in diameter. Twigs 1.4-5 mm in diameter, thick,
longitudinally ridged or wrinkled, grey-brown to orange brown, glabrous or hispid-

hairy with golden-brown to brown acicular hairs 0.5-0.7 mm in length, eventually
glabrous, occasionally sparsely lenticellate. Leaves: lamina chartaceous,
oblanceolate or oblong-oblanceolate, 18.3-38 cm by 5.1-14 cm, olive-grey ,
usually

paler and tan-colored; base obliquely rounded to subcordate, apex

acute-acurninate; glabrous and sparsely hairy above and below, midrib flat to
sunken but with a longitudinal groove running along either side above and
prominent below; secondary veins sunken above and prominent below.
secondary veins 14-21 in pairs. Flowers internodal, occasionally subopposit to
the leaves, frequently on leafless segments of twigs, pedicels 1-2, arising from a
tubercle, 7-27 mm in length, 1.3-1.8 mrn thick at midpoint; sepals brown, orange
suffused with green or dull orange-yellow, 11-23 mm in length; petals of both
whorls orange, yellow, or dull red, glabrous above, sparsely appressedpubescent in below surface, outer petals 2.1-3.6 cm in length, 6.5-11.5 rnm in
width, inner petals 2.5-3.9 cm in length, 3.5-8 mm in width. Carpidia not seen.

DISTRIBUTION : Borneo (Kelai River near Long Lanuk).
HABITAT : along ridge.
SPECIMENS EXAMINED: Kostermans 21108,21376.

Fig. '9. Polyalthia igniflora. A. Inner petal. 8. Outer petal. C. Stamen.

6. Polyalthia jenkensii Benth. & Hook.
Polyalthia jenkensii Benth & Hook; FI. Br. Ind. 1 (1876) 64; Ridley, Sar.
Mus. Jour. (1911-1913) 81; Whitmore, Tree FI. Mal. (1983) 90. Fig. 10. ---TypeSouth Andarnan, Kurz s.n., (BO).
Tree, up to 20 rn tali, 8-20 crn in diameter. Bark greyish, smooth, hooped.
Twigs hairy, lenticelled. Leaves: petioles 1-2 rnrn in diarnater. 3 rnrn in length,
hairy; lamina charthaceopus, oblong, 7.4-16 crn by 2.3-4 crn; base acute, apex

acuminate-caudate; glabrous above and below; midrib sunken and glabrous
above, prominent and glabrous below; secondary veins prominent and glabrous
above and below, secondary veins 5-7 in pairs. Flowers extraaxillary, sepals
broadly triangular 1 cm in length, hairy; petals broadly ovate, outer petals 3.9- 4
cm in length, inner petals 3.5-3-8 cm hairy. Carpidia ellipsoid. Seed 1.
DISTRIBUTION : Borneo (Berau).
HABITAT AND ECOLOGY: limestone rocks covered by thick, on river
bank of small stream. Altitude: 40 m. Flowering: October.
FIELD NOTES : Petals strong sweet smell, carpidia green.
SPECIMEN EXAMINED: Kessler, P.J.A.; Arbainsyah B 1346.; Arbainsyah
B 1365.
NOTE: This species can be distinguished from the other species by its
leaves, because in its lower surface the cluster of base of secondary veinsare
hairy.

m

7. Polyalthia laferiflora King
Polyalfhia laterflora King, Journ. As. Soc. Beng.

1x1. (1892) ii. 58. , Ann.

Roy. Bot. Gard. Calcutta V, IV (1893) 73.
Tree 12-30 m tall, 10-55 cm in diameter. Bark smooth, sparsely
lenticelled, hooped, greyish black-brownish, inner bark thick, buttresses
spreading, bole straight. Twigs glabrous. Leaves: petioles glabrous, 1.5-2.5 mm
in diameter, 5-9 rnm in length; lamina coriaceous, shiny above, oblong, 7.7-50
cm by 4-12 cm; base rounded, apex acuminate; glabrous above and below
surface; midrib sunken above and prominent below; secondary veins sunken and
glabrous above, prominent and glabrous below, secondary veins 7-18 in pairs.
Flowers solitary or in few- flowered fascicles on short shoots; pedicels hairy, 2.13.4 cm length, sepals triangular, 4mm, hairy; petals greenish yellow, inner petals
and outer petals elliptic, 1-3 cm in length, hairy. Stamens 2 mm in length, stamen
greenish, carpels 2.8 mm, hairy, stigma cream. Carpidia: stalked ca. 4.5 cm in
length, ellipsoid, acute rounded, ca.3 cm length, ca.2 cm in diameter. Seed I .
DISTRIBUTION : Malay Peninsula, Sumatra, Java, Borneo ( Mt. Njapa
and Kelai River).
HABITAT AND ECOLOGY : logged over forest, logged over dipterocarp.
Altitude: 85-100 m. Flowering : October.
SPECIMENS

EXAMINED:

Kostermans

21465,

Kessler,

P.J.A.;

Arbainsyah 81381; Kessler et al. Berau 14, Arifin 2.; Arbainsyah B 1342, Arifin et
al. 81461, Arifin, Z. 81379.

Fig. 11. Polyalthia lateriflora A. Stamen. B. Petal. C.

8. Polyalthia microtus Miq.
Polyalthia microtus Miq., Ann. Mus. Bot. Lugd. Bat. 2 (1865) 13; Kessler &
Heusden. Rhedeea 3, 1 (1993) 80-82.
Treelet 2.5-5 m tall, 4 cm in diameter. Twigs hairy, 3-4 mm in diamater.
Leaves: petiole hairy, 2 mm in diameter, 4-7 mm in length; lamina charthaceous,
elliptic, 20.5-28.7 cm by 6-9.6 cm.; base subcordate, apex acuminate; glabrous
above and below surface; midrib raise in furrow and glabrous above, glabrous

and prominent below; secondary veins glabrous and sunken above, prominent
and glabrous below, secondary veins 14-20 in pairs.

Flowers supraaxillary,

ramiflorous, solitary; pedicels 3-9 mm in length, hairy; sepals yellowish,
triangular, c.2 cm. in length, hairy; petals orange, inner petals elliptic, c.3 cm,
hairy, outer petals c.2.8 cm hairy. Stamen 2mm in length. Carpidia 2-13, stalked
3-13 mm in length, shortly cylindrical to ellipsoid, 7 - 9 mm in length, 4 - 5 mm in
diameter. Seed not seen.
DISTRIBUTION : Borneo (Tanjung Redeb, Kelai River near Long Lanuk.
HABITAT AND ECOLOGY: primary forest, logged over forest, flat pain,
ridge. Altitude : 30-85 m.
SPECIMENS EXAMINED: Kade Sidiyasa et al. Berau 1220, 1230 (BO).
NOTE: This species is easily recognized by midrib character which is
raised and furrowed on the umer surface.

Fig. 12. Po!yalthia microtus A. Sepal. B. Stamen. C. Inner petal. u. Outer petal.

9. Polyalthia obliqua H0ok.f. & Thoms.
Polyalthia. obliqua H0ok.f. & Thoms. FI. Ind. 1 (1855) 138; Miq. FI. Ned.
Ind. (1859) 44; Hook., FI. Br.. Ind. 1. (1875) 67; Ridley, Sar. Mus. Jour. (19111913) 81.
Polyalthia sirnilis Merr. Philip. Jour. Sci. 8 (1913) 371.
Tree up to 10 m tall, 12 cm in diameter.

Bark greyish-white, black,

smooth, hooped, deppled, lenticelled. Twigs glabrous 5-6 mm in diameter,
fissured. Leaves: petioles glabrous, 1.2 mm in diameter, 4-9 mm length; lamina
coriaceous,shiny above, oblong, 9.2

- 17.5 cm by 3.2 - 4.7 cm; base cuneate,

apex acuminate to caudate; glabrous above and below surface; midrib flat and
glabrous above, prominent and glabrous below; secondary veins prominent and
glabrous above and belows, secondary veins 11-15 in pairs. Flowers not seen.
Carpidia: stalked 3 cm, ellipsoid, apex acute, glabrous, 16 mm in diameter. Seed
1.
DISTRIBUTION : Borneo (Berau)
HABITAT: Primary forest.
SPECIMENS EXAMINED: Adriansyah; Boestani, A.N. AA 2197.
NOTE: The species has leaves the edges of which are very slightly
revolute and its carpidia are yellow when dried.

Fig. 13. Polyalthia obliqua
10. Polyalthia oblonga King
Polyalthia oblonga King, Journ. As. Soc. Beng 11, 2 (1892) 65.---Type:
King s.n. Malaysia : Perak (Holo: CAL, not seen ).
Tree, 15 m tall, 12 cm in diameter, bark cracked, hooped. Twigs circular,
fissured, hairy. Leaves: petioles hairy, 3 mm in diameter, 3 mm in length; lamina
charthaceous, oblong to spathulate, 13.6-25.8 cm by 4.9-9 cm; base truncatus,

apex acute; glabrous above and hairy below; midrib sunken and hairy above,
prominent and hairy below; secondary veins sunken and glabrous above,
prominent and hairy below, secondary veins 10-16 in pairs. Flowers and carpidia
not seen.
DISTRIBUTION : Malay Peninsula, Borneo (Berau)
HABITAT AND ECOLOGY: logged over dipterocarp forest. Altitude: 1000
m.
SPECIMEN EXAMINED : Arifin.2.; Arbainsyah B 1332

11. Polyalthia rumphii (Blume) Merr.
Polyalthia rumphii (Blume) Merr. Enum. Philipp. FI. PI. ii (1923) 162;

.

Whitmore, Tree FI. Mal. 1 (1983) 90. Kessler & Heusden, Rheedea 3 1 (1993)
82.
Polyalthia. kunstleri King, Journ. As. Soc. Beng. Pt. Nat. Hist. 61 (1893)
55.
Treelet, up to 8 m tall, up to 12 cm in diameter: Bark fibrous, smooth,
white to black-brownish, bole straight, circular, columnar. Twigs circular, hairy, 4
mm in diameter, fissured. Leaves: petioles hairy, 1-2 mm in diameter, 0.4

- 1 cm

in length; lamina charthaceous, oblong-ovate. 5.4-12.5 cm by 2.4-5.2 cm; base
acute, apex acuminate-caudate; glabrous above and below; midrib sunken and
hairy above, prominent and hairy below; secondary veins flat and glabrous
above, prominent and hairy below, secondary veins 6-12 in pairs. Flowers extra
axillary, cluster; pedicels 0.5-1 mm in diameter; sepals triangular, hairy; petals
green becoming yellowish; inner petals and outer petals unequal, inner petals
elliptic (1.4 cm) , hairy, outer petals (1.2 cm), hairy. Stamen 2 mm in length.
Carpidia: green, stalked 2-5 cm; ellipsoid, 5.5 mm in diameter, Seed 1.

DISTRIBUTION : Sumatra, Java, Amboina, Philippines,Borneo (Berau)
HABITAT AND ECOLOGY: Logged over forest, primary forest and
secondary forest Altitude: 45 m.
SPECIMENS EXAMINED: Ambriansyah; Arifin, Z. Berau 984, Kessler et
al. 282, Kessler et al. 319 (BO).
NOTE: The glands on carpidia surface can be used to recognize this
species.

Fig. 14. Polyalthia rumphii A. Inner petal. B. Outer petal. C. Sepal. D. Carpidia.
E. Stamen.

12. Polyalthia sclerophylla Hook .f. & Th.
Polyalthia sclerophylla Hook. f. & Th., FI. Br. Ind. 1 (1872) 65
Tree.c.10 m up to 15 m tall, 7-12 cm in diameter, bark whitish. Twigs
fissured, glabrous with lenticelled. Leaves: petioles hairy, 2 mm in diameter, 5-9
mm in length; lamina coriaceous, oblong, 13 cm-25 cm by 5.8 cm- 8.5 cm.; base
rounded; apex acute

- acuminate; glabrous above and below surface; midrib

sunken above and prominent below; secondary veins flat and glabrous above,
prorninent and glabrous below, 10-13 in pairs. Flowers not seen.. Carpidia:
stalked 4.1 cm in length, hairy, shortly ellipsoid, diameter 18-20 mm, stalks
monocarpidia 2.8 cm in length, 4 cm in diameter, hairy.
DISTRIBUTION : Berau, PT. lnhutani I logging area, KM 35. along Sungei
Sepan
HABITAT : Over mixed Dipterocarp forest, along river bank
SPECIMEN EXAMINED : Kessler, Arbainsyah 81417
NOTE : This species is close to Polyalthia lateriflofa from its leaves
character, but the leaves of P. sclerophylla are shorter than P. lateriflora.

Fig. 15. Polyalthia sclerophylla

13. Polyalthia sumatrana (Miq.)Kurz
Polyalthia sumatrana (Miq.) Kurz, Sar. Mus. Jour. (1911-1913) 80.; Ridl.,
FI. Mal. Pen. 1 (1922) .52. ; Merrill, Univ. Cal. Publ. Bot. XV. (1924) 67; Airy
Shaw, Kew Bull. (1939) 280; Whitmore. Tree FI. Mal. 1 (1983) 90; Kessler &
Heusden, Rhedeea 3 , l (1993) 82.
Guatteria sumatrana Miq., FI. Ind. Bat. Suppl. (1860) 380.

Monoon sumatranurn Miq. Ann. Mus. Bot. Lugd. Bat. 2 (1865) 19. --Type

: Teijsman s.n., Sumatra occid., Lubu Alang (not seen).
Tree 8-20 m tall, up to 25 cm in diameter. Bole straight, circular, bark
smooth, fibrous, hooped, white, greyish, inner bark thick. Twigs glabrous. Leaves:
petioles glabrous, mm in diameter, 5-12 mm in length, lamina chatthaceous,
oblong to broadly lanceolate, 23.4-28 cm by 4.7-16.6 cm; base acute, apex
acuminate-caudate; glabrous above and below surface; midrib sunken and
glabrous above, prominent below; secondary veins flat and glabrous above and
below; glaucous below, secondary veins not clear. Flowers axillary, subaxillary;
pedicels 1.5-2 cm in length; sepals triangular, hairy; inner and outer petals nearly
same length, 3.7 cm in length, elliptic, hairy. Stamen not seen. Carpidia: red
when young becoming black, stalked c.3.2 cm in length, 2 mm in diameter;
ellipsoid, 13-21 mm in diameter. Seed 1.
DISTRIBUTION : Malay Peninsula, Sumatra, Java, Borneo (Berau PT.
lnhutani I, Berau road to Tj. Redeb, Near Teluk Bayur, Mt. Njapa, G. Lamin,
Labanan).
HABITAT AND ECOLOGY : primary forest, logged over forest, secondary
forest, burned over forest along road, Hill forest, altitude : 20-200 m. Flowering :
February, June, October, November. Fruiting : October.
SPECIMENS EXAMINED: Kessler et al. Berau 135, 191, 325, 331, 892,
Arifin Berau 575, Kostermans 21378, 21560, (60).

Fig. 16. Polyalthia sumatrana A. Sepal. B. Petal

14. Polyalthia tipuliflora D.M. Johnson
Polyalthia tipuliflora D.M. Johnson, Contr. Univ. Michigan Herb. 22 (1999)
22: 99. Type:Kostermans 13902:Borneo:Berouw, Mt. llas Bungaan (1so:BO).

Treelet 3-5 m tall, trunk 5 cm in diameter. Bark smooth. Twigs 4.5-6 mm
in diameter, light brown to grey-brown, densely covered with a pale yellow to pale
brown tomentum formed of matted and twisted hairs 1.5-3 mm long. Leaves:
lamina charthaceous, oblanceolate, 36-6 cm by 11-12 cm, olive grey; base
subcordate; apex acminate; above with long hairy (2 mm)along midrib toward the
base, below surface hairy; midrib sunken above, prominent below, secondary
veins sunkenabove, prominent below, secondary veins 21-24 in pairs. Flowers
arising from woody tubercles on thrunk or leafless twigs, pedicel 5-6 mm by 2.5
mm thick at midpoint, tirsute; sepals 6.5-11,3 mm by 4.5-5.5 m at base,
triangular, caudate, densely hirsute; petals dirty-orange to orange, slightly fleshy,
filiform, sparsely pilose to glabrous in upper surface, sparsely pubescent to
glabrous below surface; outer petals 4.5-6.1 cm by 5-6 mm, acute, subauriculate
at base because of short inflexed claw; inner petals 4.2-5.9 cm by 3-3.5 mm,
acute. Stamens numerous, 2 mm in length, glabrous. Carpidia not seen.
DISTRIBUTION Borneo ( Mt. llas Bungaan)
HABITAT: sand and limestone.
SPECIMEN EXAMINED: Kostermans 13902 (BO)

Fig. 17. Polyalthia tipuliflora A. Flower. B. Stamen.
15. Polyalthia sp. A
Treelet C. 3 m tall, 4 cm in diameter. Bark smooth. Twigs circular, 3 mm in
diameter, glabrous with lenticelled. Leaves: petioles 5-7 mm in length, 2-2.5 mm
in diameter, hairy; lamina charthaceous, oblong-ovate, 10.7-22.1

cm by 3.7-6.9

cm.; base acute, apex acuminate-caudate; glabrous above and below; midrib flat
and glabrous above, prominent and hairy below; secondary veins glabrous and
flat above, prominent below, secondary veins 10-13 in pairs. Flowers axillary;

pedicel 10 - 12 rnrn in length, hairy; sepals triangular, 5 rnm, hairy; petals elliptic,
hairy, inner petals 15 mrn, outer petals 10 mrn. Stamen 2 mm in diameter.
Carpidia not seen.
DISTRIBUTION : Borneo (Berau).
HABITAT AND ECOLOGY: Secondary forest along road ca.2 km from
Labanan Altitude : 10 rn.
FIELD NOTES : Young fruits green, young leaves purple. Flowers whitish
cream.
SPECIMEN EXAMINED : Kessler et al. Berau 309
NOTE: The species resembles to P. rumphii in flower character and P.

dictyoneura in twigs character.The identify of this species is still uncertain
because there is not enough specimen to describe it as a new species.

Fig.18. Polyalthia sp.A. A. Sepal. 8.Inner petal. C. Outer petal. D. Stamen.

CONCLUSION

There are 15 species of Polyalthia from Berau Regency. Polyalfhia
berauensis, P. cauliflora, P. dictyoneura, P. glauca, P. igniflora, P. jenkensii, P.
lateriflora, P. microtus, P. oblonga, P. obliqua, P. rumphii, P. sclerophylla,
P.sumatrana, P. tipuliflora, and Polyalthia sp. A. Polyalthia berauensis

is

proposed as a new species.
The species of Polyalthia is distributed in logged over forest, primary
forest, low hills, logged over dipterocarp forest, limestone rocks, a long ridge,
secondary forest and logged over mixed dipterocarp forest in Berau Regency.

REFERENCES
Argent, G.. A. Saridan, E.J.F. Campbell, P. Wilkie, G. Fairweather, J.T. Hadiah,
D.J. Middletton, C. Pendly, M. Pinard, M. Warwick & K.S. Yulita. 1998.
Polyalthia. In Manual of The Larger and More Important Non Dipterocarp
Trees of Central Kalimantan Indonesia. Vol.1. Forest Research Institute,
Samarinda. p. 68-71.
Blume, C. L. von. 1829. FI. Jav. Anonac. 68. Bruxellis: Suntibus Librariae J.
Frank.
Corner, E. C. H. 1940. Wayside Trees of Malaya. Singapore: Government
Printing Office. P. 135.
Hildebrandt, F. H. 1952. Daftar Nama Pohon-Pohonan " Bulungan dan Berau"
Kalimantan Timur. Dalam Laporan Balai Penyelidikan Kehutanan No. 55.
Balai Penyelidikan Kehutanan Bogor.
Johnson, D. M. & Nancy A. Murray. 1999. Four New Species of Polyalthia
(Annonaceae) from Borneo and Their Relationship to Polyalthia insignis.
Contr. Univ. Michigan Herb. 22 : 95 - 104.
Kessler, P.J.A. 1993. Annonaceae. In Kubitzki, K. et al.(Eds.) Fam. Gen. Vas.
Plants Vol. 2. Springer, Berlin. P. 97 -1 29.
Kessler, P. J. A. & E. C. H. van Heusden. 1993. The Annonaceae of the
Balikpapan Samarinda Area, East Kalimantan Indonesia. Rhedeea 3:
50 - 89.

-

Kochummen, K. H. 1972. Annonaceae. In Whitmore, T.C. (Eds.)Tree Flora of
Malaya. Vol. 1. Kuala Lumpur: Longman. p. 85.
Leenhouts, P.W. 1968. A Guide to the Practice of Herbarium Taxonomy. The
International Bureau for Plant, Taxonomy and Nomenclature of The
InternationalAssociation of Plant Taxonomy, Utrecht. The Netheerland.
Maxted, N. 1992. Towards defining a taxonomic revision methodology. Taxon
41(4): 653 660.
Rifai. M.A. 1976. Sendi-Sendi Botani Sistematika. Lembaga Biologi Nasional LIPI, Bogor.
Sinclair, J. 1955. A Revision of Malayan Annonaceae. Gard. Bull. Sing. 14 (2):
149 - 516.

-

Sist, P. & A. Saridan. 1998. Description of the Primaty Lowland Forest of Berau.
In Bertault, J. G. & K. Kadir (Eds.). Silvicultural Research in A Lowland
Mixed Dipterocarp Forest of East Kalimantan. Jakarta: Cirad. p. 51 94.
Vogel, E.F. de. 1987. Guideline for the Preparation of Revisions. In Vogel (ed.)
Manual Herbarium Taxonomy Theory and Practice.

-