Artabotrys (Annonacea) in East Malesia

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ARTABOTRYS (ANNONACEAE) IN EAST MALESIA

TRIASTINURMIATININGSIH

THE POST GRADUATE PROGRAM

BOGOR AGRICULTURAL UNIVERSITY

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LETTER OF STATEMENT

I express that thesis entitling :

ARTABOTRYS (ANNONACEAE) IN EAST MALESIA

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ABSTRAK

Penelitian taksonomi marga Artabotrys di Malesia Timur dilakukan berdasarkan pada ciri-ciri morfologi spesimen herbarium dari Herbarium Bogoriense (BO), Herbarium Celebense (CEB), Herbarium Manokwari (MAN), National Herbarium of Papua New Guinea (LAE) dan koleksi hidup di Kebun Raya Bogor. Hasil penelitian menunjukkan terdapat 13 jenis, 9 jenis diantaranya

telah dikenal sebelumnya yaitu A. camptopetala Diels, A. gracilis King, A. hexapetalus (L.f.) Bhandari, A. inodorus Zipp., A. macranthus Holthuis, A. speciosus Kurz., A. suaveolens Blume, A. sumatranus Miq., dan A. venustus King. Empat jenis lainnya diusulkan sebagai jenis baru yaitu A. desmidantha Diels ex Triasti., A. longipetalus Triasti., A. palmatiflorus Triasti. dan A. trichofolius Triasti.. Kunci identifikasi dan deskripsi dari masing- masing jenis telah dibuat. Artabotrys suaveolens mempunyai persebaran yang luas sedangkan A.camptopetala, A. desmidantha, A. longipetalus, A. inodorus, A. palmatiflorus dan A. trichofolius persebarannya terbatas. New Guinea merupakan kawasan yang paling tinggi keanekaragaman jenisnya di Malesia Timur.

Analisa hubungan kekerabatan dengan Cyathocalyx sumatranus sebagai outgroup menghasilkan satu kladogram parsimony dengan panjang 26 langkah dan menunjukkan 3 kelompok besar. Kelompok I adalah Cyathocalyx sumatranus, kelompok kedua adalah Artabotrys macranthus yang paling dekat dengan outgroupnya. Sedangkan kelompok III disatukan oleh karakter synapomorf no 2 (panjang daun), 3(hook), 11(bentuk daun buah), 14(jumlah bakal biji) dan 15 (letak biji).

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ABSTRACT

Studies on the taxonomy of the genus Artabotrys R. Br. ( Annonaeceae ) in East Malesia were based on morphological characters by examining a large number of Herbarium Bogoriense (BO), Celebense (CEB), Manokwari (MAN), Departemen of Forest Lae (LAE) and living plants of Bogor Botanical Garden. There are eight species which have been known previously: A. camptopetala Diels, A. gracilis King., A. hexapetalus (L.f.) Bhandari, A. inodorus Zipp., A. macranthus Holthuis, A. speciosus Kurz., A. suaveolens Blume, A. sumatranus Miq. and A. venustus King. Four new species are proposed in this treatment : A. desmidantha Diels ex Triasti., A. longipetalus Triasti., A. palmatiflorus Triasti., and A. trichofolius Triasti.. An identification key and detailed discription are provided for taxa. Artabotrys suaveolens Blume is widely distributed. While A. camptopetala, A. desmidantha, A. longipetalus, A. inodorus, A. palmatiflorus and A. trichofolius have a limited distribution. New Guinea is the area with the highest diversity of Artabotrys species in East Malesia.

A phylogenetic analysis with Cyathocalyx sumatranus as outgroup, shows three major groups. The first group is Cyatocalyx sumatranus. The second group is A. macranthus which has the closest relationship with the outgroup. The third group separated from the other group by the synapomorph characters 2 (leaf length), 3 (hook), 11 (carpel shape), 14 (number of ovule) and 15 (position of seed).

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ARTABOTRYS (ANNONACEAE) IN EAST MALESIA

TRIASTINURMIATININGSIH

Thesis submitted

As partial fulfillment requireme nt for the Master Degree In Plant Taxonomy

THE POST GRADUATE PROGRAM BOGOR AGRICULTURAL UNIVERSITY

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Title : Artabotrys (Annonacea) in East Malesia Name : Triastinurmiatiningsih

Reg. Number : G351040041

Certified by

Supervisor Committee

Dr. Sri Sudarmiyati Tjitrosoedirdjo, M. Sc. Dr. Rugayah, M. Sc. Chairman Member

Acknowledged by

Head of Study Program of Biology Dean of Graduate School

Dr. Ir. Dedi Duryadi Solihin, DEA Prof. Dr. Ir. Khairil Anwar Notodiputro, M.Sc.

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CURRICULUM VITAE

The author was born in Ngawi on 22th January 1969. The third daughter from four children of Mr. Soebari and Mrs. Pudji Asti

She passed her Elementary School (SD) in 1981, Junior High School (SMP) in 1984 and Senior High School (SMA) in 1987, all in Ngawi. She attended the Faculty of Biology, Gadjah Mada University, Yogyakarta and graduated in 1992.

Since 1994 she was employed by Department of Biology, Faculty of Mathematic and Natural Science, Pakuan University (UNPAK), Bogor as candidate and promoted as lecturer in 1995.

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ACKNOWLEDGEMENT

The author realizes that this thesis will never be completed without assistance of many colleagues and friends who have contributed in various ways. I would like thank to Dr. Sri Sudarmiyati Tjitrosoedirdjo and Dr. Rugayah for their valuable advices and guidance in completing the thesis. I am grateful to DIKTI and UNPAK Bo gor for the financial support for a master degree at the graduate school IPB.

I would like to express my gratitude to Dr. Eko Baroto Waluyo Head of the Herbarium Bogoriense (BO) and to other Herbaria: National Herbarium of Papua New Guinea (LAE), Herbarium Celebense (CEB) and Herbarium Manokwari (MAN) for grating me to examine their specimens for studies. In particulary I am grateful to Prof. Dr. Soedodo Hardjoamidjojo Head of UNPAK Bogor; Dr. Khairil Anwar Notodiputro Director of Graduate School Bogor Agricultural university; Dr. Dedi Duryadi Program coordinator of the biology program studies, Prof. Dr. Mien A. Rifai Program coordinator taxonomy sub program studies.

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TABLE OF CONTENTS

8. A. palmatiflorus Triasti. ... 28

9. A. speciosus Kurz... 28

10.A. suaveolens Blume ... 29

11.A. sumatranus Miq. ... 30

12.A. trichofolius Triasti. ... 31

13.A. venustus King ... 32

CONCLUSION... 40

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LIST OF TABLE

Table Page

1. Distribution of Artabotrys in East Malesia and the other place in

Malesia ...8

2. Morphological characters and character states used in phylogenetic Analysis...9

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ARTABOTRYS (ANNONACEAE) IN EAST MALESIA

TRIASTINURMIATININGSIH

THE POST GRADUATE PROGRAM

BOGOR AGRICULTURAL UNIVERSITY

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ABSTRAK

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ABSTRACT

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Certified by

Acknowledged by

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ACKNOWLEDGEMENT

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TABLE OF CONTENTS

8. A. palmatiflorus Triasti. ... 28

9. A. speciosus Kurz... 28

10.A. suaveolens Blume ... 29

11.A. sumatranus Miq. ... 30

12.A. trichofolius Triasti. ... 31

13.A. venustus King ... 32

CONCLUSION... 40

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LIST OF TABLE

Table Page

Malesia ...8

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LIST OF FIGURE

Figure Page

1. Forms of Artabotrys leaves. shape : a. ovate, b. obovate, c. lanceolate, d&e. oblong, f. ellipse. Apex : a. acute, b. cuspidate, c&f. long acuminate, d&e. shortly acuminate. Base : a, d&f. cuneate, b&c. attenuate, e. rounded. ...5

2. Variation of Artabotrys stamens. 1. filament, 2. theca, 3. apical. a. oblong flat apical prolongation, b. oblong suborbiculate apical prolongation,c. oblong suborbiculate apical prolongation ...6

3. Variation of Artabotrys carpel. 1. stigma; 2. carpel. a. cylindrical carpel bilobed stigma, b. cylindrical carpel clavate stigma, c. cylindrical carpel Ellipsoid stigma, d. ellipsoid carpel, e. ovoid carpel wooly stigma, f.ovoid carpel cup stigma, g.ovoid carpel axe stigma, h. ovoid carpel ovoid stigma ...7

4. Cladogram of Artabotrys...10

5. A. desmidantha Diels ex Triasti.: a. habit, b. sepal, c. petal, d.stamen, e. carpel and stigma (a-e: Docters vanLeuween 11388) ...21

6. Artabotrys gracilis King, a. habit, b. calyx, c. stamen, d. carpel and

stigma (from Nurainas, 2004)...22

7. Artabotrys hexapetalus (L.f) Bhandari. a. habit, b. calyx, c. inner petal, d.outer petal, e. carpel and stigma, f. stamen ( from Nurainas, 2004)……….……..23

8. Artabotrys inodorus Zipp.: a. habit, b.sepal, c. outer petal, d.inner petal e. stamen, f. carpel (a- f: J.S. Womersly 13430)...24

9. Artabotrys longipetalus Triasti.: a. habit, b. leaf, c. sepal, d. outer petal, e.inner petal, f. stamen, g. carpel and stigma (a-g: M. Jacobs 9315) ...25

10. Artabotrys macranthus Holthuis: a. habit, b. petal, c. stamen d. carpel and stigma ( from L. B. Holthuis, 1942) ...26

11. Distribution of Artabotrys in East Malesia. A. camptopetala, A. desmidantha, A. gracilis, A. hexapetalus, A. inodorus, A. longipetalus and A. macranthus .. ...27

12. Artabotrys palmatiflorus Triasti.: a. habit, b.sepal, c. outer petal, d. inner petal, e. stamen, f. carpel (a- f: Ramlanto 444)...33

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14. Artabotrys suaveolens Blume, a. habit, b. sepal, c.outer petal, d. inner petal, e. stamen, f. carpel and stigma (a-c: G. Kjellberg 2752; d-f: Moh. Amir 254) ... ...35

15. Artabotrys sumatranus Miq.: a. habit, b. flower, c. outer petal, d.inner petal, f. carpel and stigma (from Nurainas, 2004) ...36

16. Artabotrys trichofolius Triasti. : a. habit, b. sepal, c. petal, e. stamen, f.carpel. (a-f : Ramadhanil & Ch. H. Scultze 816) ...37

17. Artabotrys venustus King : a. habit, b. stamen, c. carpel and stigma, d. fruit ( from Nurainas, 2002) ...38

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INTRODUCTION

Artabotrys is one of the large genus of Annonaceae family. The genus has more than 100 species distributed in Tropical Africa and Eastern Asia. Several species can be used for drugs, poison and cosmetic (Burkill, 1935). One of them has economically important for parfume namely A. odoratissimus, known as Ylang-ylang oil (Keßler, 1993).

The genus Artabotrys was described almost 187 years ago by Robert Brown, based on A. odoratissimus R. Br. Previously A. odoratissimus was included in genus Unona, Annona, and Uvaria. However, it differs from Unona in petal shape, seed number, seed position and peduncle characters, from Annona in fruit, petal and peduncle characters, and from Uvaria in their peduncle shape, petal and fruits characters (Brown, 1820). Bhandari (1964) reduced A. odoratissimus as the synonym of A. hexapetalus.

Keßler (1993) include Artabotrys in an informal grouping of Xylopiae together with Xylopia, Anaxagorea, Pseduo artabotrys, Cyathocalyx, Drepananthus and Diclinanona based on climbers habitat, peculiar hooked peduncle, bisexual flowers, equal depth of petal, spoon like outer petal, connivent inner petal covering reproductive organ and berry fruits. While Sinclair (1955) placed Artabotrys as a member of sub family Annonoideae tribe Unonae together with Cyathocalyx, Desmos, Monocarpia, Oncodostigma, Meiogyne, Polyalthia, Cananga, Mezzetia and Disepalum. Artabotrys included in the tribe Unoneae based on valvate petal, flat, differentiation of blade and claw, and inner petal covering reproductive organ.

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and A. inodorus ß inodorus from New Guinea. Keßler et. al (2002) said in the checklist of plant in Sulawesi that there are Artabotrys suaveolens, Artabotrys trichopetala and Artabotrys sp. Which have not recorded yet in Moluccas..

The limited data of Artabotrys in East Malesia is due to the lack of intensive explorations, collections and investigations so this is the reason that studies were worshy subject of investigations.

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MATERIAL AND METHODS

Explorations

Exploration was conducted at Celebes and Moluccas for addition of the collected specimens due to the scarse of the previous collections. Exploration and specimen collection were based on Balgooy method (1987)

Morphology

Morphology and distribution of the species were based on dried specimens and spirit- preserved materials which were deposited at LAE, CEB, BO, MAN and the collected specimens in 2006.

Measurement and observation were recorded: locality; general habitat; stem (surface, indument, colour); branches (straight or curved); leaves (shapes, base, apex, midrib, number of nervus lateralis, leaf length, petiole length, surface); inflorescense (shapes, induments, branche, number of flowers); flowers (colour, sepal surface, sepal shape, petal surface, petal shape, length pedicel, shape and number stamen, shape and number carpells), fruitlets shape were observed from the specimens. Terminology used in this studies following Stearn (1988) and Harris & Harris (1994)

Revision

Revision was made using standard preparation guide-lines proposed by Rifai (1976), Vogel (1987), and Maxted ( 1992).

Phylogenetic Analysis

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RESULT AND DISCUSSION

MORPHOLOGY Habit

The habit of Artabotrys are climbers. This is one of an important character to distinguishis from other genus in Annonaceae .

Indumentum

Type of indumentum in the genus are puberulous, pubescent, tomentose and villose. Types of indumentum on young twigs, petiols, and midrib are pubescent or sparsely pubescent, on blade is sparsely pubescent. Indumentum in peduncle is sparsely pubescent, while in flowers are puberulous, pubescent, tomentose and villose.

Branches

Generally their young branches are hairy but A. suaveolens and A. gracilis are glabrous. Most of the Artabotrys species have straight branches such as A. desmidantha, A. gracilis, A. longipetalus, A.palmatiflorus, A. suaveolens, A. sumatranus, A. hexapetalus, A. inodorus, A. longipetalus, A. speciosus, A. venustus, A. venustus. Whereas A. trichofolius and A. macranthus have curved branches.

Leaves

The leaves of the genus have various shape (Fig.1). The shape of blade were elliptic to oblong, oblong, elliptic, ovate, obovate and lanceolate. Most species of Artabotrys have elliptic to oblong leaves but obovate leaves found in A.

sumatranus and A. macranthus, lanceolate or ovate in A. palmatiflorus,

lanceolate or obovate in A. inodorus. Exception is found in A. speciosus and A. trichofolius which only have oblong leaves. Petioles of Artabotrys are usually short but A. palmatiflorus has longer than other species.

Most species of Artabotrys have cuneate leaves base, attenuate found in

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Generally the leaves apex in Artabotrys are short acuminate , but A. inodorus and A. hexapetalus have rather lengthened acuminate apex, while A.suaveolens has

acuminate to cuspidate, and A. palmatiflorus has acute apex.

Fig. 1. Forms of Artabotrys leaves. Shape : a. ovate, b. obovate, c. lanceolate, d & e.oblong, f. elliptic. Apex : a. acute, b. cuspidate, c & f. long acuminate, d & e. shortly acuminate. Base : a, d & f. cuneate, b & c. attenuate, e. rounded

Some species of Artabotrys have shinning upper leaf surface such as A. camptopetala, A. suaveolens, A. venustus and A. gracilis. Generally both of leaf

surfaces are glabrous, except in midrib beneath, but A. trichofolius has hairy at lower surface. The midrib of Artabotrys raised at lower surface, but some species Artabotrys have midrib raised at both surface such as A. sumatranus, A. venustus,

A. macranthus and A. palmatiflorus.

c

a b

3cm

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Flowers

The flowers are bisexual in solitary or fasicle. Peduncles are extra axillary, opposite the leaves, reflexed, hooked, flatten or terete. Calyx consists of 3 sepal valvate, sometimes connate at base with erected or deflected in ovate, deltoid or triangular shape. Corolla consists of 6 petals, valvate in two whorls. It has 3 outer petal, free, fleshy, concave at base with oblong, lanceolate or linear shape. Outer petal longer and broader than inner petals, sometimes different shape. A. sumatranus, A. gracilis, A. speciosus and A. longipetalus have outer petal different from inner petal shape.

The shape of stamen was oblong with the apical prolongation flat, acute or concave. Most of species Artabotrys have stamen oblong with apical prolongation flat but A. sumatranus and A. trichofolius have stamen oblong with apical prolongation acute, while A. speciosus and A. longipetalus have stamen oblong with apical prolongation suborbiculate. They consists of three parts: filament, theca and apical prolongation (Fig. 2).

The carpels have three shapes: ovoid for most species, cylindrical for A. macranthus, A. hexapetalus and A. longipetalus, while ellipsoid found in

A. trichofolius and A. speciosus. Most species have 2-5 carpels, but A macranthus

has 8-13 carpels.

Fig 2. variation of Artabotrys stamens. 1. filament; 2. theca; 3. apical; a. oblong flat apical prolongation; b. oblong acute apical prolongation; c. oblong suborbiculate apical prolongation

The shape of stigma various for all species. A. gracilis cup shape stigma, A. hexapetalus clavate shape, cylindrical bilobed in A. macranthus, ellipsoid

1mm

a b c

1

2

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d

villose in A. desmidantha, ellipsoid in A. longipetalus, ovoid in A. sumatranus and axe shape in A. venustus .

Fig 3. Variation of Artabotrys carpel. 1. stigma; 2. carpel; a. cylindrical carpel bilobed stigma, b. cylindrical carpel clavate stigma, c. cylindrical carpel ellipsoid stigma, d. ellipsoid carpel, e. ovoid carpel villose stigma, f.ovoid carpel cup stigma, g.ovoid carpel axe stigma, h. ovoid carpel ovoid stigma.

Fruits

The fruits of Artabotrys are berry, with various shape: ellipsoid with acute or blunt apex, oblong with blunt apex, and obovoid with acute or blunt apex. Artabotrys hexapetalus has ellipsoid fruit with acute apex. A. suaveolens and A. sumatranus have ellipsoid fruit with blunt apex, while A. inodorus has obovoid fruit with acute apex and A macranthus has oblong fruit with blunt apex.

DIVERSITY

Twelve taxa of Artabotrys are recognized in this study. There are nine species which have been known previously : A. camptopetala,A. gracilis King. A. hexapetalus (L.f.) Bhandari, A. inodorus Zipp., A. macranthus Holthuis, A. speciosus Kurz, A. suaveolens Blume, A. sumatranus Miq. and A. venustus King. Four species are proposed in this treatment: A. desmidantha Diels ex Triasti., A. longipetalus Triasti., A. palmatiflorus Triasti., and A. trichofolius Triasti..

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DISTRIBUTION

Artabotrys is found in all over East Malesian area. Artabotrys suaveolens Blume is widespread in East Malesia and also other places. It has been recorded from Celebes, Moluccas, New Guinea and also in Malay Peninsula, Sumatra, Java, Borneo, Philippines. In East Malesia some of species have limited distribution such as A. trichofolius which only occurs in Celebes, A. palmatiflorus only found in Moluccas, while A camptopetala, A. longipetalus, A. desmidantha and A. inodorus are found in New Guinea.

The region with the highest species diversity in Malesia is New Guinea. There are 9 species in New Guinea (Table 1.). Species of Artabotrys in East Malesia are more diverse than West Malesia and Java. There are 13 species in East Malesia, while in West Malesia has 7 species and 3 species in Java.

Table 1. Distribution of Artabotrys in East Malesia and the other place in Malesia (from several references and own observation of the East Malesian specimens)

East Malesia Other places in Malesia Species

C. New Guinea G. Sumatra (Nurainas, 2004)

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PHYLOGENETIC ANALYSIS OF ARTABOTRYS

A phylogenetic analysis of Artabotrys was carried out based on morphological characters, using Cyathocalix sumatranus as the outgroup. The phylogenetic analysis used 16 morphological characters (Table 2.), which all of them are scored with the primitive state based on the state of the outgroup. Data matrix of those characters which has been used for the analysis shown in Table 3. The analysis result three parsimonious trees with 26 steps length, Consistency index (CI) of 0,65 and Retention index (RI) of 0.70. A single tree, which was selected from three fundamental tree is shown in Figures 4. From 16 characters analysed, 8 of them are apomorph (5 synapomorph, 3 autapomorph). Other characters with Consistency Index lower than 1.00 showed parallel ( characters 6, 7, 8, 9, 10, 13) and reversal (character 0 &11).

Table 2. Morphological characters and character states used in phylogenetic analysis

The cladogram divided the taxa into three groups which consists of group I : Cyathocalyx sumatranus as the outgroup; group II: A. macranthus and group

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A. trichopetalus, A. venustus. A. macranthus is close related with the outgroup might be the most primitive and separated from the ingroup because of character 13 (torus concave).

Fig. 4. Cladogram of Artabotrys in East Malesia

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Table 3. Matrix of morphological characters at the taxa

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TAXONOMY

Generic Description

Artabotrys R. Brown

Artabotrys R. Br. Bot. Reg. 5: 423. 1819; Blume, Fl. Java. 2: 58. 1828; Miquel,

Fl. Ned. Ind. 38. 1860; Hooker, Fl . Brit. India 1 : 53. 1875; King, Jour. As. Soc. Beng. 61: 30. 1893; Ridley, Fl. Mal. Pen. 1: 38. 1922; Sinclair, Gard. Bull. Sing. 14(2) : 146.1955; Backer and Bakhuizen van den Brink Jr., Fl. Java. 1 : 113. 1963. -Type species: Artabotrys hexapetalus (L.f.) Bhandari, Koenig, s. n., India, Madras (Holo: LINN).

Climbers. Lateral branches lenticellate, glabrous to hairy. Leaves: alternate, simple; margin entire; midrib raised or flat at upper surface; lateral nerves pinnate, flat or raised on the lower surface, anastomosing near to the margin; veins scalariform. Inflorescence: peduncle extra – axillary, opposite the leaves, reflexed, hooked, flattened or terete; bract small; often caducous. Flowers: hermaphrodite, fleshy, solitary or fascicle; sepal valvate, 3, erect or deflexed, connate at base; petal 6 in two series, valvate, the blade terete or flat, oblong, lanceolate or linear; outer petal 3, spoon – like at base, free, longer and broader than the inner ones; inner petal 3 remaining attached at their bases when they fall, claws the blade joins it connivent, close to the sexual organs. Torus flat to concave, pillose. Stamen hypogynous, numerous tongue shaped, apex flat, acute or convex. Carpel free, few to many, glabrous or hairy; stigma persistent; ovule 2. Fruit berried, glabrous, puncticulate. Seed 2, erect, collateral , dorsiventral, ventral side flat and dorsal side convex.

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Key to the Species

1.a Lateral branches curved...2 b. Lateral branches straight ...3 2.a. Young twig with long reddish hairs; leaves 17.5-20 x 7-8 cm, lower surface

glabrous; outer petal oblong, tomentose; carpels 8-12, cylindrical bilobed ... ...A. macranthus b. Young twig with short white ha irs; leaves 4-8.5 x 2.5-3.5 cm, lower surface

sparsely pubescent; outer petal ovate, puberulous; carpels 5, ellipsoid bilobed ...A. trichofolius 3.a. Outer petal terete, inner petal terete ...4 b. Outer petal flat, inner petal terete or flat ...6 4.a. Petiole 2-4 mm long, sparsely pubescent; number of flower 3-5; pedicel 2-6

mm long, apical prolongation flat ...5 b. Petiole 6-10 mm long, glabrous; number of flower 20-40; pedicel 15-20 mm long, apical prolongation suborbiculate ...A. palmatiflorus 5.a. Sepal ovate or broadly ovate; petal 6-15 x 1-1.5 mm, apex recurved,

pubescent or sparsely pubescent ...7 b. Sepal lanceolate; petal 16-20 x 1.5-2 mm, apex erected, puberulous ...

...A. inodorus 6.a. Blade coriaceous; sepal ovate, pubescent; outer petal 8-15 mm long,

pubescent; stigma cup shape ...A. suaveolens b. Blade chartaceous; sepal broadly ovate, sparsely pubescent; outer petal 15-20

mm long, puberulous; stigma ovoid shape...A. camptopetala 7.a. Inner petal terete, linear; top of apical prolongation acute or suborbiculate .... ...8 b. Inner petal flat, lanceolate or ligulate; top of apical prolongation flat ...9 8.a. Blade thinly coriaceous, elliptic to oblong or obovate, base cuneate; flowers

3-4; top of apical prolongation acute; carpels 5 ...A. sumatranus b. Blade coriaceous, oblong, base rounded; 5-6 flowers; top of apical

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b.Sepal suborbiculate, apex cuspidate; outer petal ligulate, 5.5-7 cm long ... ...A. longipetalus 10. a. Bract elliptic; pedicel 2-3 mm; torus flat ...11

b. Bract elliptic to oblong; pedicel 15-25 mm; torus convex...12 11.a. Pedicel villose; sepal ovate, 5 x 3 mm; outer petal lanceolate, 15 x 4 mm,

villose; carpels broadly cylindrical; stigma ellipsoid, villose ... ...A. desmidantha b. Pedicel glabrous; sepal broadly ovate, 2 x 2 mm; outer petal linear 3-5 x 1mm, pubescent; carpels broadly ovoid; stigma cup shape, glabrous ... ...A. gracilis 12. a. Leave shining at upper surface, lateral nerves raised on both surface; outer petal oblong 6-10 x 3-5 mm, stigma axe shape ...A. venustus

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Distribution : New Guinea

Habitat and Ecology : primary forest at 5400 feet alt

Specimen examined: New Guinea. Oksapmin, Telefomin subdistrict, West Sepik District, 17-10-1968, E.E. Henty, R. Isgar & M. Galore 41608 (LAE!); Ambunti subdistrict, Waskuk hills, around Langu and Garuka villages, West Sepik District, 29-6-1995, J. Regalado & W. Takeuchi 1441 (LAE!).

2. A. desmidantha Diels ex Triasti., spec.nov. ( Fig. 5)

Folia sub coriacea, glabra, elliptica, 8-12 cm longa, 3.5-6 cm lata, nervus lateribus 6-8 paribus, apica breviter acuminata, basi cuneata. Petioli 2-4 mm longa, sparsissima pubescent. Pedunculi trifidi 2-3 flori; pedicelli villose, 2-3 mm longa, sepal lato-ovata, 5 mm lata, apice acuminata, villose; petala externa lanceolata, 15 mm longa, 4 mm lata, apice acute, utrinque villose; interna angusti-lanceolata. Stamen angusti-oblonga, apical prolongation plana. Carpels 5, lato-cylindrica; stigma ellipsoidea, villose. -Typus : Docters van Leeuwen 1138, Memberamo, New Guinea ( Holo: BO!)

Lateral branches straight, lenticelled, black when dry. Young twigs pubescent. Leaves: petiole 2 – 4 mm long, pubescent; blade thinly coriaceous, glabrous in both surface, pale brown at upper surface, dark brown at lower surface when dry, elliptic to oblong, 8 – 12 (-13) cm x 3.5 – 6 cm; base cuneate; apex short acuminate; midrib flat at upper, raised at lower surface and sparsely pubescent, lateral nerves 6 – 8 pairs, anastomosing 3–6 mm to the margin. Inflorescence: peduncle terete, three branches, sparsely pubescent, with 3 – 5 flowers; bract elliptic, 2 x 1.5 mm, flower bud ovoid, 4 x 3 mm, villose. Flowers: pedicel 2 – 3 mm long, villose; sepal valvate, fleshy, erect, ovate, 3 x 2.5 mm, villose, apex acuminate; outer petals flat, fleshy, lanceolate, 15 x 4 mm, both surface villose, apex acute; inner petals flat, narrower, 13 x 3.5 mm, villose; torus flat; stamen c. 14, oblong, top of apical prolongation flat; carpels 5, broadly cylindrical, 1.5 x 0.5 mm; stigma ellipsoid 1 x 1 mm, villose. Fruit and seed not seen.

Distribution: New Guinea

Habitat and Ecology: primary forest at 60 m alt.

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desmidantha has elliptic to oblong leaf only, terete peduncle with three branches, flat torus, top of apical prolongation flat, stamen 14 and stigma ellipsoid stigma. The indumen of pedicel, sepal, petal and stigma are the most important character for this species. Diels (1932) described Leeuwen specimen as A. desmidantha but it has not published yet. Therefore, this study propose this as a new species. Specimen examined: New Guinea : Memberamo, November 1926, Docters van Leeuwen 11388 (BO!)

3. Artabotrys gracilis King (Fig. 6)

A. gracilis King, Jour. As. Soc. Beng. 16(2): 35. 1893; Ridley, Fl. Mal. Pen. 1: 44. 1922; Ridley, Sarawak Mus. Journ. 1(3): 77. 1913; Sinclair, Gard. Bull. Sing. 2(14): 258. 1955. - Type: G. King 4987, Malay Peninsula, Perak (Holo: L, Isotype: BO!)

Lateral branches straight, lenticelled, glabrous, dark colour when dry. Young twigs glabrous. Leaves: petiole 2-4 mm long, pubescent; blade thinly coriaceous, both surface glabrous, shining at upper surface, brown when dry; elliptic to oblong, 6-12 x 5-7 cm; base cuneate; apex acuminate; midrib flat at upper, raised at lower surface, sparsely pubescent; lateral nerves 7-9 pairs, anastomosing 4-6 mm to the margin, flat on both surface. Inflorescence: peduncle terete, sparsely pubescent, three branches; 5-7 flowers; bract elliptic, 5-7 x 1-2 mm, persistent, pubescent; flower bud broadly ovoid, 1.5 x 2 mm, pubescent. Flowers: pedicel 2-2.5 mm long, glabrous; sepal valvate, erect, ovate, 2 x 2 mm, apex acuminate, outside sparsely pubescent, inside glabrous; outer petal fleshy, flat, linear, 3-5 x 1 mm, apex acute, pale green, tomentose; inner petals slightly narrower, terete, linear, 2-3 x 1 mm; torus flat; stamen 20-24, oblong, 1-2 x 0.5-1mm, top of apical prolongation flat; carpels 5, broadly ovoid, 0,4-0,6 x 0,3-0,4 mm, glabrous; stigma cup shape, glabrous. Fruit and seed not seen.

Habitat & Ecology : Rain forest on hill slope at 300 feet alt.

Field note : Leave glossy dark green above, dull and pale below, flowers pale green

Vernacular name: wasmenyangu (Waskuk), efai(Wagu)

Notes : This species allied to A. suaveolens, mainly in vegetative characters but differs in sepal and outer petal. A. suaveolens has sepal ovate and outer petal terete. Whereas A. gracilis has sepal broadly ovate and outer petal flat.

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4. Artabotrys hexapetalus (L.f.) Bhandari (Fig. 7)

Artabotrys hexapetalus (L.f.) Bhandari, Baileya 12(4): 147.1964. Annona hexapetala L.f., Suppl. 270. 1781; Hort. Kew. 2. ed. 2. 3: 335. 1811.Type: Koenig, s. n., India, Madras (Holo: LINN).

Artabotrys uncinatus (Lam) Merr., Philipp. Journ. Sci. Bot. 7 : 234. 1912; Backer & Bakhuizen van den Brink. Jr., Fl. Java. 1:134. 1963.

Artabotrys odoratissimus R. Br. ex Kew Gawler, Bot. Reg. 5: 423. 1819. Artabotrys hamatus Blume. Fl. Java. Anon. 60. t. 29, 31C.1828; Hook f. & Thoms., Fl. Brit. India 1:54.1872.

Lateral branches straight, lenticelled, glabrous, dark brown when dry. Young twigs sparsely pubescent. Leaves: petiole 2-5 mm, sparsely pubescent; blade thinly coriaceous, both surface glabrous except for the midrib beneath, pale green when dry, elliptic to oblong or lanceolate, 6-16 x 3-7 cm; base cuneate; apex acuminate; midrib flat at upper, raised at lower surface; lateral nerves 8-11 pairs, anastomosing 3-4 mm to the margin, flat on both surface. Inflorescence: peduncle flat, sparsely pubescent, two branches; 1-2 flowers; bract elliptic to oblong, 1-2 x 0.5-1 mm, pubescent; flower bud broadly ovoid. Flowers: pedicel 15-20 mm long, sparsely pubescent; sepal erect to reflexed, thinly fleshy, ovate, 3-8 x 3-6 mm, apex acute, pubescent; petal fleshy, yellow; outer petal flat, lanceolate, 20-30 x 5-10 mm, pubescent, apex acute; inner petal narrower, flat, 20-30 x 3-6 mm; torus convex; stamen c. 29, oblong, top of apical prolongation flat; carpels 5, cylindrical, 2 x 0.5 mm; stigma clavate, glabrous. Fruit obovoid to ellipsoid, 3.5-4 x 1.5 cm, apex mucronate; stalk 20 x 1.5 mm, sparsely pubescent. Seed 2, ellipsoid.

Distribution : Celebes & New Guinea

Habitat & Ecology : Primary and secondary forest at 50-100 m.alt.

Specimen examined : Celebes: Saleier, 28-5-1913, Docters van Leeuwen 1904, 1905 (BO!). Moluccas: Kp. Kiandarat, G. Kilia, 18-8-1938, P. Buwalda 5637 (BO!).

5. Artabotrys inodorus Zipp. (Fig. 8)

Artabotrys inodorus Zipp., Ann. Mus. Bot. Lugd. Bat. ii. 41.1866 Type: Zippelius s.n. (Holo: L)

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lanceolate, 9-13 x 3-6 cm; base cuneate; apex acuminate; midrib flat at upper, raised at lower and sparsely pubescent; lateral nerves 8-12 pairs, anastomosing 4-5 mm to the margin, flat on both surface. Inflorescence: peduncle flat, three branches, sparsely pubescent; 3-5 flowers; bract ovate, 2 mm x 1,5 mm, puberulous outside, glabrous inside; flower bud ellipsoid, 2 mm x 2 mm, puberulous. Flowers: pedicel 3-5 mm long, glabrous; sepal deflect, thinly fleshy, lanceolate, 5 x 3 mm, glabrous inside, puberulous outside, apex long acuminate; outer petals, terete, fleshy, linear, 15-20 x 1 mm, triquetrous, puberulous; inner petals narrower, terete, 15-18 x 1 mm; stamen oblong, top of apical prolongation flat; carpels 3, broadly oblong; stigma not seen. Fruit obovoid, 30-40 x 1.5-2 cm; stalk 0.5 cm long. Seed not seen.

Habitat and Ecology : Atzera Range, near Lae, Morobe District. at 250 feet alt. Field Notes: Flower in short, leaf opposed. Inflorescences, outer whorl of perianth lobes short, triangular, middle and inner whorl elongate, clawed at the base, lobea cream. Fruit olive brown.

Notes : This species similar to A. suaveolens in petal linear. A. inodorus has oblong to lanceolate leaf, peduncle flat, ellipsoid flower bud, lanceolate sepal, petal longer, triquetrous and fruit obovoid. While A. suaveolens has elliptic to oblong leaf, terete-sub terete peduncle, broadly ovoid flower bud, ovate sepal, terete petal and ellipsoid fruit.

Specimen examined: New Guinea.Dagarungu Ridge, Jimni Valley, Western Highland District, 1-9-68, E. E. Henty & H. Streimann 38932 (BO!, LAE!); Atzera range, near Lae, Morobe District, Juni 1961, J. S. Womersly 13420, 13430 (LAE!); Markham river, Morobe Distric, 20 Nov. 1962, T. G. Hartley s.n (LAE!).

6. Artabotrys longipetalus Triasti., spec. nov. (Fig. 9)

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Lateral branches straight, lenticelled, glabrous, black when dry. Young twigs sparsely pubescent. Leaves: petioles 2 –3 mm long, sparsely pubescent, blade coriaceous, both surface glabrous, pale brown at upper surface, dark brown at lower surface when dry, elliptic, 6 –9 x 3 – 4.5 cm; base cuneate; apex short acuminate; midrib flat at upper surface, raised at lower surface and sparsely pubescent; lateral nerves 8 –11 pairs, anastomosing 3 – 4 mm to margin, flat on both surface. Inflorescence: peduncle, sub terete to terete, three branches, sparsely pubescent; 1-3 flowers; bract and flower bud not seen. Flowers: pedicel 1 – 1,5 cm, sparsely pubescent; sepal valvate, fleshy, flat, erect, suborbiculate, 3 x 3mm, sparsely pubescent outside, glabrous inside, apex cuspidate; outer petal valvate, thinly fleshy, flat, ligulate , 55 – 70 x 3 – 3,5 mm, both surface puberulous, apex acute; inner petal triangulate to the middle, flat toward to apex, 40 – 60 x 1 – 3 mm, puberulous, apex acute; torus flat; stamen c.30, oblong, 1 x 0.4 mm, top of apical prolongation suborbiculate; carpels c. 3, cylindrical, 1 mm x 0.5 mm; stigma ellipsoid 0.5 x 0,3 mm, glabrous. Fruit and seed not seen

Habitat and Ecology : old well-drained volcanic soil, primary forest. Alt. 600-700 m

Note : A. longipetalus differs from all other species in elliptic leaf, suborbiculate sepal, cuspidate apex, very long petal, ligulate, inner petal1/3 triquetrous, flat toward to apex and ellipsoid stigma . The species similar to A. sumatranus and A. speciosus in outer petal linear and flat, but differs in elliptic leaf and outer petal longer.

Specimen examined : New Guinea. Mount Basovi, Northen New Guinea, 18-10-1973, M. Jacobs 9315 (BO!, LAE!).

7. Artabotrys macranthus Holthuis (Fig. 10)

Artabotrys macranthus Holthuis, Blumea 5: 180. 1942; Keßler, Rheedea 5: 97. 1995.-Type: H. J. Lam, 3003, Indonesia, Sangi Talaud Islands, Karakelang, Pasir Malap, E. of Lobo (Holo: L; Iso: BO!)

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sparsely pubescent; two branches, 1-2 flowers; bract not seen; flower bud ovoid, 3-4 x 2-3 mm, tomentose. Flowers: pedicel 1-1.2 cm long, tomentose; sepal erect, fleshy, triangular, 10-20 x 5-10 mm, apex acute, both surface tomentose; outer petal fleshy, flat, greenish yellow, oblong, 35-40 x 20-25 mm, pubescent; inner petal narrower, flat, 35-38 x 15-20 mm; torus flat; stamen numerous, oblong, 1 x 0.5 mm, top of apical prolongation flat; carpels 8-12, stigma cylindrical bilobed, glabrous. Fruit oblongoid, 1.5 x 1 cm; stalk 1.3 cm long, pubescent. Seed not seen.

Distribution : Celebes, Moluccas Habitat & Ecology : alt. 60-100 m

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Fig. 5. A. desmidantha Diels ex Triasti.: a. habit, b. leaf, c. sepal, d. outer petal, e. inner petal, f. stamen, g. carpel and stigma (a-g: Docters. van Leuween 11388)

c b

a

f

g

d

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8. Artabotrys palmatiflorus Triasti., spec. nov. (Fig. 12)

Folia sub coriacea, ovata, superiora and inferiora glabra, 7-13 (-15) cm longa, 3-4.5 cm lata, nervus lateribus 8-12 paribus, basi cuneata, apice acuminata, petioli 6-10 mm longa. Pedunculi bifida multiflori; pedicelli 1.5-2 cm longa; sepal lato ovata, 2-2.5 mm longa, 2 mm lata, apice acuta, pubescent; petala externa linearis, 8 mm longa, 0.5 mm lata, interna breviora, utrinque tomentosa; stamina oblonga, apical prolongatus planus, carpella 3 ovoidea, glabra.- Typus: Ramlanto 444. G. Wapaluhu, Liang, Moluccas (Holo: BO!)

Lateral branches straight, lenticelled, black when dry. Young twigs glabrous. Leaves: petiole 6-10 mm long, glabrous; blade thinly coriaceus, both surface glabrous, ovate or lanceolate, 7 –13 (- 15) x 3 – 4.5 cm; base cuneate; apex acute; surface pale brown when dry; midrib flat to slightly raised at upper surface, raised at lower surface, glabrous; lateral nerves 8 – 12 pairs; anastomosing 2 – 5 mm to the margin. Inflorescence: peduncle sub terete to terete, two branches, glabrous, with 20-40 flowers; bract broadly ovate,1 x 1.5 mm, slightly pubescent; flower bud orbicular, 1 x 1 mm, tomentose. Flowers: pedicel 1.5 – 2 cm, slightly pubescent; sepal erect, valvate, fleshy, flat, broadly ovate, 2 – 2.5 x 2 mm, apex acute, sparsely pubescent; outer petals terete, fleshy, linear, 8 mm long, upper part terete, 0.5 mm wide, tomentose, apex acute; inner petals slightly narrower, terete, linear, 7 x 0.5 mm; torus flat; stamen c.26, oblong, 0.5 x 0.4 mm, top of apical prolongation suborbiculate; carpels 3, broadly ovoid, 0.2 – 0.4 x 0.3 mm, groove surface; stigma not seen. Fruit not seen.

Distribution: Moluccas

Habitat and Ecology: Primary forest at 60 m alt.

Notes: A. palmatiflorus is similar to A. suaveolens in petal terete but differs in leaf, number of flower, petal apex and top of apical prolongation. A. palmatiflorus has ovate to lanceolate leaf, chartaceous lamina, 20-40 flowers, tomentose petal, acute petal apex and suborbiculate of apical prolongation.

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9. Artabotrys speciosus Kurz ex Hook.f & Thomson. (Fig. 13)

Artabotrys speciosus Kurz ex Hook. f. & Thomson, Fl. Brit. Ind. i: 55. 1872 -Type : Kurz s.n. Andaman Isl. (Holo: K)

Lateral branches straight, lenticelled, black when dry. Young twigs pubescent. Leaves: petiole 3 – 5 mm long, slightly pubescent; blade coriaceous, oblongs, 13 – 16 x 3.5 – 6.5 cm; base rounded; apex acuminate, dark brown on both side when dry, glabrous on both side; midrib flat at upper surface, raised at lower surface and sparsely pubescent, lateral nerves 10 – 11 pairs, prominent at lower surface, anastomosing 4 – 6 mm to the margin. Inflorescence: peduncle sub- terete to terete, two branches , sparsely pubescent, with 5 – 6 flowers; bract ovate, 2 x 1.5 mm, apex acute, sparsely pubescent; flower bud ovate, 2 x 2 mm, pubesscent. Flowers: pedicel 4 – 6 mm, sparsely pubescent; sepal valvate, fleshy, erect, glabrous inside, pubescent outside, triangular, 2 – 3 x 2 mm, apex acuminate; outer petal valvate, thinly fleshy, flat, lanceolate, 2 – 2.5 x 0.1- 0.3 cm, tomentose; inner petals , terete, linear, 2 – 2.5 x 0.1 cm, tomentose; torus flat; stamen c 20, oblong, 1 x 1 mm, top of apical prolongation acute; carpels 5 , broadly ovoid, 1 – 1.5 x 1 mm; stigma not seen. Fruit ellipsoid, 3-3.5 x 2-2.5 cm. Seed not seen.

Distribution: Moluccas and New Guinea Habitat & Ecology: primary forest. Alt 30-40 m

Field Notes: climber. Leave dark green above light green below. Flowers yellow or yellowish green, with sickening sweet scent. Fruit green

Notes: This species is similar to A. sumatranus in outer petal flat and linear, but differs in leaf, sepal and carpel. A. speciosus has oblong leaf, lateral nerves prominent at lower surface, triangulate sepal and ellipsoid carpels. Whereas A. sumatranus has lanceolate or obovate leaf, lateral nervus flat at lower surface. Specimen examined: Moluccas.P. Wokam, Dasinamalaoe, Aroe islands, 11-5-1938. P. Buwalda 4922 (BO!), New Guinea. Netherlands, 8-5-1954. P. van Royen 3574 (BO!, LAE!)

10. Artabotrys suaveolens Blume (Fig. 13)

Artabotrys suaveolens Blume, Fl. Jav. Anon. 2 : 62. 1828 Tab. 30, 31D; Hook. F. et Th. Fl. Ind. 1: 129. 1855. Fl. Br. Ind. 1 : 55.1872; Boerl., Icon. Bog. 1: 120. 1897. Tab. 39; Ridley, Fl. Malay Penins. 1: 42, 1922; Sinclair, Gard. Bull. Sing. 2(14): 256. 1955. Backer & Bakh, van den Brink Jr. Fl. Java 1:114. 1962; Kessler & Heusden, Rhedeea 3 : 55. 1993.

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Lateral branches straight, lenticelled, dark coloured. Young twig sparsely pubescent. Leaves: petiole 2-4 mm long, sparsely pubescent; blade thinly coriaceous, both surfaces glabrous except for the midrib beneath, shining above, brown when dry, elliptic to oblong, 4.5-9(-12) x 2 - 4 cm; base cuneate; apex acuminate to cuspidate; midrib flat at upper surface, raised at lower surface; lateral nerves 8-10 pairs, prominent at above side, anastomosing, 3-5 mm to the margin. Inflorescence: peduncle terete- sub terete, pubescent, three branches, with 3-5 flowers; bract lanceolate, 1.5 x 0.5 mm, sparsely pubescent outside, glabrous inside; flower bud broadly ovoid, 2 x 1.5 mm, pubescent. Flowers: pedicel 4-6 mm long, sparsely pubescent; sepal valvata, fleshy, erect, ovate, 1.5 - 2 x 1-1.5 mm, glabrous inside, pubescent outside, apex acuminate; outer petals linear toward the apex, terete, pubescent, 7-15 x 0.5 mm; apex rounded; inner petals terete, slender, linear, 7-15 x 0.5 mm, pubescent, apex rounded; torus flat; stamen c. 20-25, oblong, 0.2 x 0.5 mm, top of apical prolongation flat; carpels 2-3, broadly ovoid, 0.4-0.5 mm, glabrous; stigma cup shape. Fruit ellipsoid, 7 x 5 mm, glabrous, stalk 5-10 mm. Seed 2.

Distribution: Celebes, Moluccas, New Guinea

Habitat & Ecology: Primary and secondary forest, at 0-150 m alt

Specimen examined: Celebes. G. Papalia, Wolasi, Ranometo, Kendari, 19-2-1986, Moh. Amir 254 (BO!); Malili, 8-11-1929, G. Kjellberg 2752 (BO!). Moluccas. Halmahera, G. Sembilan, 1-10-1951, Idjan/Mochtar, 290 (BO!); Soelabesi, G. Berberi, Atjih (B. Hulstyin )321 (BO!); New Guinea. Netherlands, 9-6-1959, W. Vink 8567 (BO!, LAE!).

11. Artabotrys sumatranus Miq. (Fig 15)

Artabotrys sumatranus Miquel, Fl. Nederl. Ind. 1:375.1860; Miquel, Ann. Mus. Bot. Lugd. 2: 43. 1860; Backer & Bakhuizen van den Brink. Jr., Fl. Java: 1:114.1963. - Type: J. E. Teysman 4382, Lampung, (Iso: BO!)

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surface, raised lower surface; lateral nerves 7-11 pairs, anastomosing, 3-6 mm to the margin. Inflorescence: peduncle, flat, sparsely pubescent, three branches, 3-4 flowers; bract ova tes, 2-3 x 0.5-1 mm, pubescent outside, glabrous inside; flower bud broadly ovoid, 1.5-2 x 1-1.5 mm, pubescent. Flowers: pedicel 3-5 mm long, sparsely pubescent; sepal erect, sometimes deflexed, valvate, connate at base, ovate, 3-5 x 2-3 mm, apex caudate, glabrous inside, densely pubescent outside; outer petals thinly fleshy, flat toward the apex, densely pubescent, lanceolate, 1.5-2 cm x 1-1.5-2 mm, apex acute; inner petal terete, 8-17 mm long, apex obtuse; torus flat; stamen 20-25, oblong, top of apical prolongation acute; carpels 3, glabrous, free, ovoid, 1-2 x 0.5-1; stigma ovoid. Fruit 2-3, ellipsoid, 3-4 x 2-2,5 cm, stalk 1-2 cm. Seed not seen.

Distribution: New Guinea

Habitat & Ecology: primary & secondary forest. Alt. 200- 4000 feet

Collector Notes: Liana, climbing in trees. Fruit yellow to orange, leaves dark green above and below

Specimen examined: New Guinea. Atzera range, 24-11-1970, J. S. Womersly 43682.

12. Artabotrys trichofolius Triasti., spec. nov. (Fig.16)

Folia coriacea, superiora glabra, inferiora sparsissima pubescent, oblong, 4 – 8,5 cm longa, 2,5 – 3,5 cm lata, nervus lateribus 8 – 9 paribus, basi cuneate, apice breviter acuminata,. Pedunculi bifida uniflori; pedicelli 1-1.5 cm longa, glabra; sepala lato-ovata, 3 mm longa, 3 mm lata, apice acuta, puberulosa; petala externa ovata, 10 mm longa, 4 mm lata, utrinque puberulosa; interna breviora, omnia flava; petala interna angusti ovata; stamina multa, angusti-oblonga, apical production acuta; carpella 3, ovoidea glabra.- Typus : P. Ramadhanil & Ch. H. Scultze, 816, 4-08-02. Donggala, Central Sulawesi (Holo: CEB!).

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peduncle flat, sparsely pubescent, two branches, with one flowers. Flowers: pedicel 1 – 1.5 cm long, glabrous; sepal valvate, free, deltoid, 3 x 3 mm, erect, glabrous inside, puberulous outside, apex acute; petals fleshy, flat, outer petals valvate, ovate, 10 x 4 mm, puberulous outside, glabrous inside; inner petals slightly small, flat, ovate, 10 x 3 mm; torus flat; stamen c 17, oblong, 1.5 x 0.5– 0.8 mm; top of apical prolongation acute; carpels 5, ovoid, 2x1 mm, bilobus; stigma not seen. Fruit and seed not seen

Distribution: Celebes.

Habitat and Ecology: Margin forest of grand forest Park Palu. Alt. 208 m.

Notes: A. trichofolius differs from other species in oblong leaf (4-8.5 x 2.5-3.5 cm), hairy lower surface of leaf, deltoid sepal, ovate petal, top of apical prolongation acute, and carpels ovoid, bilobed.

Specimen examined: Donggala, Biromaru, Pombewe Paneki, Palu, Central Sulawesi, 4-8-2002, P. Ramadhanil dan Ch. H. Schultze, 816 (CEB!)

13. Artabotrys venustus King. (Fig. 17.)

Artabotrys venustus King. Jour. As. Soc. Beng, 61: 32. 1893; Ridley, Fl. Mal. Pen. 1: 40. 1922; Sinclair, Gard. Bull. Sing. 2(14): 252. 1955.

Type: King 6499, Malay Peninsula, Perak, Larut ( Isotype: BO !)

Lateral branches straight, lenticelled, brown when dry. Young twig sparsely pubescent. Leaves: petiole 3 – 5 mm long, sparsely pubescent; blade coriaceous, both surfaces glabrous, shinning at upper surface, yellowish – brown when dry; elliptic to oblong, 9 –15 x 3 –6 cm; base cuneate; apex short acuminate; midrib raised on both surface; lateral nerves 7 – 9 pairs, thin, raised on both surface, anastomosing, 4 – 5 mm to the margin. Inflorescence: peduncle sub terete, two branches, sparsely pubescent, with 3-5 flowers; bract and flower bud not seen. Flowers: pedicel 2 – 2.5 cm, glabrous; sepal erect to deflect, ovate, 3–4 x 2–3 mm, apex caudate, tomentose outside; outer petal fleshy, flat, oblong 6-10 x 3-5 mm; apex acute, greenish yellow, tomentose; inner petal narrower, flat, oblong, 6-10 x 2-4 mm; stamen not seen; carpels about 6, ovoid, 1-2 x 0.5-1 mm, narrowly ovoid; stigma axe shape. Fruit not seen.

Distribution : New Guinea Habitat & Ecology : primer forest

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Fig. 14. Artabotrys suaveolens Blume : a. habit, b. leaf, c. sepal,

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CONCLUSION

The recent study indicated that there were 13 species Artabotrys occure in East Malesia. Nine species have been known previously A. camptopetala Diels,

A. gracilis King, A. hexapetalus (L.f.) Bhandari, A. inodorus Zipp., A. macranthus Holthuis, A. speciosus Kurz, A. suaveolens Blume, A. sumatranus

Miq. and A. venustus King, four species will be proposed as a new species in this treatment: A. desmidantha Diels ex Triasti., A. longipetalus Triasti., A. palmatiflorus Triasti., A. trichofolius Triasti. An analysis of the distribution of these species indicated that A. suaveolens Blume are widely distributed. Whereas A. desmidantha, A. longipetalus, A. inodorus, A. palmatiflorus and A. trichofolius have limited distribution. New Guinea is the area with the highest diversity of the species in East Malesia.

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ARTABOTRYS (ANNONACEAE) IN EAST MALESIA

TRIASTINURMIATININGSIH

THE POST GRADUATE PROGRAM

BOGOR AGRICULTURAL UNIVERSITY

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ABSTRAK

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ABSTRACT

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Certified by

Acknowledged by

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ACKNOWLEDGEMENT

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TABLE OF CONTENTS 8. A. palmatiflorus Triasti. ... 28 9. A. speciosus Kurz... 28 10.A. suaveolens Blume ... 29 11.A. sumatranus Miq. ... 30 12.A. trichofolius Triasti. ... 31 13.A. venustus King ... 32 CONCLUSION... 40

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LIST OF TABLE

Table Page

Malesia ...8

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LIST OF FIGURE

Figure Page

1. Forms of Artabotrys leaves. shape : a. ovate, b. obovate, c. lanceolate, d&e. oblong, f. ellipse. Apex : a. acute, b. cuspidate, c&f. long acuminate, d&e. shortly acuminate. Base : a, d&f. cuneate, b&c. attenuate, e. rounded. ...5

2. Variation of Artabotrys stamens. 1. filament, 2. theca, 3. apical. a. oblong flat apical prolongation, b. oblong suborbiculate apical prolongation,c. oblong suborbiculate apical prolongation ...6

3. Variation of Artabotrys carpel. 1. stigma; 2. carpel. a. cylindrical carpel bilobed stigma, b. cylindrical carpel clavate stigma, c. cylindrical carpel Ellipsoid stigma, d. ellipsoid carpel, e. ovoid carpel wooly stigma, f.ovoid carpel cup stigma, g.ovoid carpel axe stigma, h. ovoid carpel ovoid stigma ...7

4. Cladogram of Artabotrys...10

5. A. desmidantha Diels ex Triasti.: a. habit, b. sepal, c. petal, d.stamen, e. carpel and stigma (a-e: Docters vanLeuween 11388) ...21

6. Artabotrys gracilis King, a. habit, b. calyx, c. stamen, d. carpel and

stigma (from Nurainas, 2004)...22

7. Artabotrys hexapetalus (L.f) Bhandari. a. habit, b. calyx, c. inner petal, d.outer petal, e. carpel and stigma, f. stamen ( from Nurainas, 2004)……….……..23

8. Artabotrys inodorus Zipp.: a. habit, b.sepal, c. outer petal, d.inner petal e. stamen, f. carpel (a- f: J.S. Womersly 13430)...24

9. Artabotrys longipetalus Triasti.: a. habit, b. leaf, c. sepal, d. outer petal, e.inner petal, f. stamen, g. carpel and stigma (a-g: M. Jacobs 9315) ...25

10. Artabotrys macranthus Holthuis: a. habit, b. petal, c. stamen d. carpel and stigma ( from L. B. Holthuis, 1942) ...26

11. Distribution of Artabotrys in East Malesia. A. camptopetala, A. desmidantha, A. gracilis, A. hexapetalus, A. inodorus, A. longipetalus and A. macranthus .. ...27

12. Artabotrys palmatiflorus Triasti.: a. habit, b.sepal, c. outer petal, d. inner petal, e. stamen, f. carpel (a- f: Ramlanto 444)...33

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14. Artabotrys suaveolens Blume, a. habit, b. sepal, c.outer petal, d. inner petal, e. stamen, f. carpel and stigma (a-c: G. Kjellberg 2752; d-f: Moh. Amir 254) ... ...35

15. Artabotrys sumatranus Miq.: a. habit, b. flower, c. outer petal, d.inner petal, f. carpel and stigma (from Nurainas, 2004) ...36

16. Artabotrys trichofolius Triasti. : a. habit, b. sepal, c. petal, e. stamen, f.carpel. (a-f : Ramadhanil & Ch. H. Scultze 816) ...37

17. Artabotrys venustus King : a. habit, b. stamen, c. carpel and stigma, d. fruit ( from Nurainas, 2002) ...38