Systematics of freycinetia in West New Guinea
SYSTEMATICS OF FREYCINETIA
IN WEST NEW GUINEA
NURHAIDAH IRIANY SINAGA
DEPARTMENT OF BIOLOGY
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
DECLARATION BY CANDIDATE
I hereby declare that this dissertation on Systematics of Freycinetia in West New
Guinea is my own work and effort directed by supervisory commission and has not been
submitted in any forms to any other universities for my award. The information sources
from the published or unpublished references of other authors have been acknowledged
and written in the references at the end of this dissertation.
Bogor, March 2011
Nurhaidah Iriany Sinaga
G 36 1040051
ABSTRACT
NURHAIDAH IRIANY SINAGA. Systematics of Freycinetia Gaudh. in West New
Guinea (supervised by Rita Megia, Alex Hartana, Mien A. Rifai and Ary
Prihardhyanto Keim).
Freycinetia Gaud. in West New Guinea had been investigated. For about 4.000 specimen
collections were studied using morphological, ecological and molecular data of cp DNA
sequence trnL-F, atpB-rbcL and trn-L . The result showed that in West New Guinea
92 species occur. All species can be divided into 4 groups namely F. macrostachya, F.
funicularis, F. oblanceolata and F. angustissima groups. The F. macrostachya group
have 36 species and 12 species are prepare as a new species namely F. aurihasteta
Sinaga, F. brevipedicelata Sinaga, F. clavata Sinaga & Utteridge, F . circuita Sinaga,
F.concordia Sinaga, F. frutaspiralica Sinaga, F. frutonumerata Sinaga, F. mandacana
Sinaga, F.megaauriculata Sinaga & Utteridge, F. stigmabeliata
Sinaga, F.
ultrapedicelata Sinaga and F. yapenina Sinaga. The F. funicularis group have 17
species and 9 species are prepare as a new species namely F. ayawasa Sinaga, F.
fusiforma Sinaga & A.P. Keim, F. hasteta Sinaga, F. imbristigma Sinaga & A.P. Keim,
F. magnoareola Sinaga, F.millikenus Sinaga & A.P. Keim, F. nervoauricula Sinaga,
F. spiroaxilla Sinaga & A.P. Keim and F. simpliaxillata. The F. oblanceolata group
have 31 species and 9 species are considerated as a new species namely F. albaauria
Sinaga, F. batantaensis Sinaga, F. decendata Sinaga, F. folitenella Sinaga, F. frutasolla
Sinaga, F. broccoareola Sinaga,
F. iriana Sinaga, F. manokwariana Sinaga and F.
verstegii Sinaga. The last group is F. angustissima group that have 8 species but only
one as a new species namely F. pauciberria Sinaga. The F. macrostachya and F.
oblanceolata groups living from forest beach to 1700 m asl but F. macrostachya group
are found alone in secondary forest. The F. funicularis and F. angustissima groups prefer
living on the mountain area and both of them are found on 3000 m asl. 35 species has
limited distribution it is only found in Papua but 52 species occurs both in Papua &PNG
and 5 species are widely distribution throughout Malesia and Australia. Analysis
phylogenetic through PAUP program using both morphological and molecular data
support the groups with CI (Coefficient Index) = 0.7, HI (Homoflashy Index) = 0.3 for
morphology characters and CI =0.930; 0.868; 0.890 and HI = 0.07; 0.132; 0.110 for
molecular data by trnL-F; atpB- rbcL and trn-L sequencing of CP DNA. According to
morphological data F. macrostachya group is a primitive one, It is followed by F.
funicularis group, F. oblanceolata and F. angustissima groups as an advance one. But in
the trnL-F sequencing cpDNA data F. macrostahya group as a primitive one separate
from 3 other derivate groups and this result are supported by ecological situation. On
the other hands, atpB–rbcL sequencing strongly support the groups but advance one is
not belong to the F. angustissima group anymore but F. funicularis group, The same way
is found in the trn-L sequencing data. Therefore trnL-F sequencing had given more
conservative characters but atpB-rbcL showed both conservative and advance characters
while trn-L had given more advance characters.
Key words: Freycinetia, West New Guinea, systematics, morphological, ecological and
molecular data.
ABSTRAK
NURHAIDAH IRIANY SINAGA. Sistematika Freycinetia di Nugini Barat dibimbing
oleh Rita Megia, Alex Hartana, Mien A. Rifai dan Ary Prihardhyanto Keim.
Freycinetia Gaud. di Nugini Barat telah diteliti. Sebanyak kurang lebih 4000 lembar
spesimen herbarium telah dipelajari, menggunakan data morfologi, ekologi dan
molekular dengan sekuensing kloroplast DNA yakni sekuens trnL-F; atpB-rbcL ; dan
trn-L .Hasil penelitian menunjukan bahwa di Nugini Barat terdapat sebanyak 92 jenis
Freycinetia . Jenis-jenis Freycinetia ini terdiri atas 4 kelompok yakni kelompok F.
macrostachya, F. funicularis, F. oblanceolata dan kelompok F. angustissima. Kelompok
F. macrostachya memiliki 36 jenis dengan 13 jenis baru yaitu F. aculeata Sinaga, F.
aurihastata Sinaga, F. brevipedicelata Sinaga,
F. clavata Sinaga & Utteridge, F
.circuita Sinaga, F.concordia Sinaga, F.frutaspiralica Sinaga, F. frutonumerata Sinaga,
F. mandacana Sinaga, F.megaauriculata Sinaga & Utteridge, F. stigmabeliata Sinaga,
F. ultrapedicelata Sinaga dan F. yapenina Sinaga. Kelompok F. funicularis memiliki
17 jenis dengan 9 jenis baru yaitu F. ayawasa Sinaga, F. fusiforma Sinaga &
A.P.Keim, F. hastatus Sinaga, F. imbristigma Sinaga & A.P. Keim , F. magnoareola
Sinaga, F. millikenus Sinaga & A.P. Keim, F. nervoauricula Sinaga, F. spiroaxilla
Sinaga & A.P. Keim dan F. simpliaxillata. Kelompok F. oblanceolata memiliki 31
jenis dengan 10 jenis baru yakni F. albaauria Sinaga, F. batantaensis Sinaga, F.
decendata Sinaga, F. folitenella Sinaga, F. frutasolla Sinaga, F .gunungmejensis
Sinaga, F. broccoareola Sinaga, F. iriana
Sinaga, F. manokwariana Sinaga dan
F.verstegii Sinaga. Kelompok F. angustissima memiliki 8 jenis dengan 1 jenis baru
yaitu F. pauciberria Sinaga. Kelompok F. macrostachya dan F. oblanceolata hidup di
hutan pantai hingga ketinggian 1700 m dpl. Sementara kelompok F. funicularis dan F.
angustissima menyenangi tempat tinggi, hingga ketinggian 3000 m dpl. Sebanyak 35
jenis hanya ditemukan di Nugini Barat, namun terdapat 52 jenis yang juga hidup di
PNG dan 5 jenis memiliki persebaran luas hingga ke Malesia bahkan Australia.
Keempat kelompok ini didukung hasil analisis filogenetik dengan program PAUP
berdasarkan data morfologi dan molekular, dengan nilai CI (Coefficient Index)= 0.7, HI
(Homoflashy Index) = 0.3 untuk data morfologi dan nilai CI =0.930; 0.868; 0.890 and
HI = 0.07; 0.132; 0.110 untuk data molekular sekuensing DNA kloroplast berturutturut
trnL-F; atpB-rbcL; dan trn-L. Berdasarkan data morfologi kelompok F.
macrostahya adalah kelompok primitif, dikuti kelompok F. funicularis,, F. oblanceolata
dan F. angustissima. Tetapi sekuensing trnL-F lebih memisahkan kelompok primitif
dari ketiga turunannya dan data sekuensing ini sangat didukung oleh data ekologis. Di
sisi lain, sekuensing atpB-rbcL mendukung keempat kelompok ini, namun kelompok F.
angustissima tidak lagi sebagai yang lebih maju melainkan kelompok F. funicularis,
keadaan yang sama terjadi dalam sekuensing trn-L. Dengan demikian trnL-F lebih
banyak menggambarkan karater yang konservatif sementara atpB-rbcL menggambarkan
keduanya baik sifat konservatif maupun sifat yang lebih maju dan trn-L sekuensing lebih
menggambarkan sifat yang lebih maju.
Kata Kunci: Freycinetia, Nugini Barat, sistematika, morfologi, ekologi dan molekular
data.
SUMMARY
Freycinetia was firstly described by Gaudichaud in 1824. Currently there are about
200 species recognized. The centre of distribution of this genus is in New Guinea, where
at least 60 species have been recognized (Stone 1976; 1983). This number is much
higher compare to Borneo (24) and the Malay Peninsula (8). The majority of herbarium
specimens of Freycinetia from New Guinea are deposited only in few Herbaria such as
Herbarium Bogoriense (BO) in Bogor-Indonesia, Herbarium Manokwariense (MAN) in
Manokwari-Indonesia, Herbarium Lae in Lae-Papua New Guinea, the National
Herbarium of the Netherlands in Leiden (L)-the Netherlands, and Herbarium Kewensis in
Kew-UK. Unfortunately, due to the incomplete nature of the specimens most are labelled
as unknown species. From time to time deposit specimens increase in numbers but
many of them are still unidentified. Therefore, it need taxonomical study to find out how
many species of New Guinea Freycinetia especially in the west area.
First serious molecular study of the family was proceeded by Callmander et al.
(2003). Unfortunately, the infrageneric classification of the genus is still unresolved. So
the aims of this study were to unravel the diversity of Freycinetia in the western part of
New Guinea, revision of the genus particularly the infrageneric classification and
knowing the species distribution in western part of New Guinea in relation with New
Guinea as a whole.
Method of the study using both morphological and molecular approach. The
research was carried out from Jully 2005 until Maret 2009. Taking and collecting
specimens were done at several area in west New Guinea such as Timika (2005),
Manokwari ( 2006), Sarmi (2006) and Jayapura (2006, 2008), followed by ecological
observation in the field and morphological observation in herbarium B0 (2006, 2008),
MAN (2006), LAE (2006), L (2008) and K (2009). Molecular work using sequence
non coding cpDNA: trnL-F, trn-L and atpB – rbcL was done at Plant Biology and
Biotechnology Laboratorium IPB (Bogor Agricultural University) on August to October
2008 and van der Klauw Molecular Laboratorium in Leiden on November 2008 until
January 2009.
The result showed that 92 species of Freycinetia occur in the area. All species can be
divided into 4 groups namely F. macrostachya, F. funicularis, F. oblanceolata and
F. angustissima groups.
The F. macrostachya group have robust habit and are high climbers. Stem robust,
hard, rarely branched, 2-4 cm diameter. Leaf imbricate, linear or lanceolate, margin with
sharp and hard spines, 20-120 cm long, 1.5-7 cm wide; cauline leave colorfull in
common, basically leaf with yellow, orange, red or mixed color, consist of 2-3 whorls;
Infructescence terminal, ternate, sometimes with 4, 8 to 12 cephalia, spirally to umbelly
arranged. Prophyll bracts never found. Cephalium large, 3-20 cm long, 2-6 cm wide.
Berries numerous, thin, stout; stigmatic remains 1-3 or 5-6, rarely 8 or 12, one species
with 32. Seeds linear or ellipse, usually longitudinally arranged.
The F. funicularis group have leafless stem from basal part up to the half part of the
stem, laterally branching, and leaves found only on the terminal part of stem. Stem 2-3
cm diameter. Leaf semi imbricate, linear or lanceolate, margin usually with sharp and
hard spines, 13-50 cm long, 1.5-3 cm wide; Prophyll bracts consist of 4 to more than 8
whorls; bracts mostly consist 3 parts exterior, middle and interior bracts Infructescence
axillary, ternate or rarely quaternate, umbelly arranged. Cephalium oblong or cylindrical,
3-10 cm long, 1.5-3 cm wide. Berries few, a berry usually formed through fusion of
smaller berries giving an impression of a larger berry; stigmatic remains 5-8, rarely 3, 12,
14 or 18, stigmatic position usually changes from transversal to longitudinal, some
species with marginal stigmatic remains. Seed usually flat, transversally arranged.
The F. oblanceolata group possess slender stem, 0.5-2 cm diameter, numerously
branching, branch short, less than 50 cm long. Leaf non imbricate, elliptical to oblong
and oblanceolate, margin with spines on terminal and basal parts only, 2-15 cm long, 0.54 cm wide; Cauline leaves usually green; Prophyll only in both terminalia and axillary
inflorescence, if present only consist of 2 whorls; Infructescence usually terminal, rarely
both terminal and axillary. Cephalium minute, 2-7 cm long, 1-3 cm wide. Berry
numerous or few, some species with flat tips, fusion or self-changing areola present;
stigmatic remains 1-3 or 5-6. Seed linear or oblong-flat, usually arranged transversally,
in some species longitudinally
The F. angustissima group have slender stem, less than 0.5 cm in diameter, usually
numerously branched, branch short, less than 20 cm long. Leaves pseudo-rosette in
appearance, each linear or lanceolate, in some members observed elliptical, margin with
spines on terminal and basal parts, 2-8 cm long, 0.2-1 cm wide. Prophyll bracts never
found. Infructescence terminal, usually binate, rarely single or ternate, arranged in umbel.
Cephalium small, 2-4 cm long, 1-2 cm wide. Berry usually few in numbers, some
species with flat-tipped berries; stigmatic remains 1 or 2, rarely 3, self-changing areola
present giving variation in stigmatic areola. Seeds linear, transversally or longitudinally
arranged.
The F. macrostachya group have 36 species. The previous species have 23 species.
These are F. aculeata Sinaga, F. andajensis
Martelli, F. arfakiana Martelli,
F. archboldiana Merr. & Perry, F. bicolor B.C.Stone, F .excelsa F. Muell., F. fibrosa
Martelli, F. formosula Huynh, F. impundens B.C. Stone, F. Klosii Ridl., F. lacinulata
Kanehira, F. macrostachya Martelli, F. marginata Blume, F. palida Huynh, F. plana
Huynh, F. percostata Merr. & Perry, F. pseudoinsignis Warb., F. rosellana Huynh,
F. solomonensis B.C. Stone, F. trachypoda Merr. & Perry, F. tessellata
Merr. &
Perry, F. undulate Merr. & Perry, and F. whitmorei B.C. Stone. Twelve species are
regarded as new species namely F. aurihasteta Sinaga, F. brevipedicelata Sinaga, F.
clavata Sinaga & Utteridge, F. circuita Sinaga, F. concordia Sinaga, F. frutaspiralica
Sinaga, F. frutonumerata Sinaga, F. mandacana Sinaga, F. megaauriculata Sinaga &
Utteridge, F. stigmabeliata Sinaga, F. ultrapedicelata Sinaga and F. yapenina Sinaga.
The F. funicularis group have 17 species. The previous 8 species are F. cryptocarpa
Kanehira, F. erythospatha Merr. & Perry, F. funicularis Merr.,, F.gibbsiae Rendle,
F. lateriflora Ridll., F. pleurantha Merr. & Perry, F. rhodospatha Ridll., and
F. sterrophylla Merr. & Perry. Nine species are regarded as new species namely F.
ayawasa Sinaga, F. fusiforma Sinaga & A.P. Keim, F. hastata Sinaga, F. imbristigma
Sinaga & A.P. Keim, F. magnoareola Sinaga, F.millikenus Sinaga & A.P. Keim,
F. nervoauricula Sinaga, F. spiroaxilla Sinaga & A.P. Keim and F. simpliaxillata
Sinaga.
The F. oblanceolata group have 31 species. The previous species are 22 species
namely F. biroi Warb., F. brasii Merr. & Perry, F. chartacea Huynh, F .concolor
Huynh, F. ellipsoidalis Merr. & Perry, F. flaviceps Rendl., F. forbesii Ridl.,
F .gunungmejensis Sinaga, F. hagenicola Huynh, F. inermis Ridl., F. lalokiensis Huynh,
F. lagenicarpa Warb., F. lenifolia Huynh, F. linearifolia Merr. & Perry, F. marantifolia
Hemsl., F. obtusiacuminata Huynh, F. oblanceolata Martelli, F. oreophila Merr. &
Perry, F. rectangularis Kanehira, F. scandens Gaudich., F. tenuifolia Huynh, and
F. tenuis Solms. Nine species are regarded as new species namely F. albaauria Sinaga,
F. batantaensis Sinaga, F. decendata Sinaga, F. folitenella Sinaga, F. frutasolla Sinaga,
F. broccoareola Sinaga, F. iriana Sinaga, F. manokwariana Sinaga and F. verstegii
Sinaga.
The F. angustissima group have 8 species and 1 species are regarded as a new
species namely F. pauciberria Sinaga. The previous species here are F. angusta Huynh,
F. angustissima Ridl., F. brachyclada Huynh, F. linearis Merr. & Perry, F. polyclada
Merr. & Perry, F. pseudoangustisimma Huynh and F. stenophylla Warb.
F. macrostachya & F. oblanceolata groups living from forest beach to 1700 m
above sea level. But F. macrostachya group are found alone in secondary forest. on
the other hand, the F. funicularis and F. angustissima groups prefer living on mountain
area and both of them are found up to 3000 m asl. Despite share with F. angustissima
group, the F. funicularis group also share habitat with F. oblanceolata group, especially
in the forest beach where F. angustissima
group never is found, but this group has
ability living on the wet area. West New guinea Freycinetia have 35 species that are
limited distribution, because these species are only occur in the area. But West New
Guinea Freycinetia share 52 species with PNG, and 5 species remains are widely
distribution species that spread through out Malesia region until Australia.
Analysis phylogenetic through PAUP program using both morphological and
molecular data support the groups with CI (Coefficient Index) = 0.7, HI (Homoflashy
Index) = 0.3 for morphology characters and CI =0.930; 0.868; 0.890 and HI = 0.07;
0.132; 0.110 for molecular data by trnL-F; atpB-rbcL and trn-L sequencing of
Chloroplast DNA. According to morphological data
F. macrostachya group is a
primitive one, It is followed by F. funicularis, F. oblanceolata and F. angustissima
group as an advance one. But in the trnL-F sequencing cpDNA data F. macrostachya
group as a primitive one separate from 3 other derivate groups and this data strongly
support ecological situation. On the other hands, atpB–rbcl sequencing strongly support
the groups but advance one is not belong to the F. angustissima group anymore but F.
funicularis groups, it is same way to the trn-L sequencing data. Therefore trnL-F
sequencing had given more conservative characters but atpB-rbcL showed both
conservative and advance characters while trn-L had given more advance characters.
Copyright @2011 Bogor Agricultural University
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SYSTEMATICS OF FREYCINETIA
IN WEST NEW GUINEA
NURHAIDAH IRIANY SINAGA
Dissertation
Submitted as a part of requirement for the fulfillment of Doctor Degree in Biology
DEPARTMENT OF BIOLOGY
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
ii
Examiners in comprehensive examination:
Dr. Nunik Sri Aryanti, M.Si
Department of Biology, Faculty of Mathematics and Natural Science, Bogor Agricultural
University, Bogor, Indonesia
Dr. Ruliyana Susanti
Herbarium Bogoroense, Botany Division,
Research Centre for Biology
Institute of Science, Cibinong, Bogor, Indonesia
Examiners in final examination:
Dr. Tatik Chikmawati, M,Si
Department of Biology, Faculty of Mathematics and Natural Science, Bogor
Agricultural University, Bogor, Indonesia
Dr. Rugayah
Herbarium Bogoroense, Botany Division,
Research Centre for Biology
Institute of Science, Cibinong, Bogor, Indonesia
iii
iv
PREFACE
The main object of the research conducted at November 2005 to January 2009 is
Freycinetia in West New Guinea. This study emphasis four different topics namely the
morphological characters, ecological and distribution, chloroplast DNA sequencing and
description of 92 species of New Guinean Freycinetia. A part of morphological
approach i.e. The Stigma and Areola in the Western New Guinean Species of
Freycinetia Gaud. (Pandanaceae; Freycinetoideae) submmited to the 8 th International
Flora Malesiana Symposium in Singapore and Two New Species of Freycinetia
(Pandanacea) from Manokwari, West Papua have been published in Reinwardtia 13:2
(2010) 183 - 187. Part two: The Ecology and Distribution of Freycinetia Gaud.
(Pandanaceae; Freycinetoideae) in The Indonesian New Guinean have also been
published in Reinwardtia 13:2 (2010) 189 -197.
I am extremely grateful to the advisory committees who directed, advised and read
critically the manuscript, Dr. Rita Megia, DEA (Department Biology IPB), Prof. Alex
Hartana (Department Biology IPB), Prof. Mien A. Rifai (Herbarium Bogoriense, LIPI)
and Dr. Ary .P. Keim (Herbarium Bogoriense, LIPI).
Profound thanks and appreciation to Dr. Eko B. Waluyo and all staff of Herbarium
Bogoriense especially Dr. Rugayah and Prof. Dr. Elizabeth Widjaja; Prof. Dr. Peter van
Welzen, Dr. Barbara Gravendell, Marcel Eurings and Dr. Michael Stech (Leiden
University, NHN Herbarium, Nederland); Dr. Rogier de Kok, Dr. William Bekker and
Dr. Timothy Utteridge (Kew Herbarium, UK); Roy Banka (Lae Herbarium, PNG); Ir.
Jack Wanggai, M.Si and staff (Herbarium Manokwariense) and Dr. Fenny Ismoyo
(Papua University); Ir. Pratita Puradyatmika, M.M (PT. Freeport Indonesia) for any kind
of help during my research.
I gratefully acknowledge the BPPS Scholarship, Sandwich Program and Litdas
from Director General of Higher Education (Dikti), Departement of National Education
Republic Indonesia. I thanks Ir. J.P. Karafir M,Sc (Rector Unipa); Dr. Hasyim (Dean
Math and Science Faculty of IPB) and Dr. Deddy Duryadi DEA (Head of Program Study
Biology Post Graduate School IPB).
I am indebted to all staff and my colleagues in Papua University, my field assistant:
Gasper Taudufu, Christhopere Diaz and Tinus Iwanggin and my friends at Subprogram
Plant Taxonomy IPB: Dr. Iris Rengganis, Dr. Nursahara Pasaribu, Dr. Sri Endarti, Dr.
Puji Widodo, Dr. Fitmawati, Dr. Himah Rustiami, also my brothers: Dr. Soaloon
Sinaga, Kristian Sinaga, SKM, Benjamin Sinaga, S.Tp and my sisters: Sesniwati Sinaga,
SE and Bertha Sinaga, S.Sos with their families. Finally, I give my honor to my beloved
parent: Kasiaman Sinaga and Johana Taudufu.
Bogor, March 2011
Nurhaidah I.Sinaga
v
CURRICULUM VITAE
Nurhaidah Iriany Sinaga was born on 6 January 1969, in Karubaga Papua, as the
second daughter of Kasiaman Sinaga and Johana Taudufu. She graduated from
Forestry Department at Cenderawasih
University in 1991. She studied plant
taxonomy and got M.Si in Biology from IPB in 2001. In 2004, she got scholarship
for study at Program Study of Biology IPB from BPPS Dikti (Directorate General
of Higher Education, National Education Department). Since 1995, she has been
lecturer at the Departement of Forestry, University of Papua.
vi
TABLE OF CONTENT
Page
LIST OF TABLES
LIST OF FIGURES
I. INTRODUCTION
Purpose of The Study
II. METHOD OF THE STUDY
Morphological work……………………………………………………………….
Molecular work……………………………………………………………………
III.RESULTS AND DISCUSSIONS
A. MORPHOLOGICAL CHARACTERS
Stem..........................................................................................................................
Leaves.......................................................................................................................
Spines......................................................................................................................
Prophyll Leaves........................................................................................................
Auricle......................................................................................................................
Cauline Leaves.........................................................................................................
Prophyll Bracts........................................................................................................
Bracts.......................................................................................................................
Peduncle...................................................................................................................
Pedicle......................................................................................................................
Staminate Flower......................................................................................................
Pistillate Flower........................................................................................................
Stigma and Areola...................................................................................................
Cephalia..................................................................................................................
Berries......................................................................................................................
Seed.........................................................................................................................
x
xi
1
3
4
4
4
8
8 - 26
8
9
11
11
13
14
15
15
17
17
17
19
19
22
23
25
B. PHYLOGENETIC ANALYSIS OF FREYCINETIA
Relationship and Evolution of the Freycinetia........................................................
27 - 38
29
C. ECOLOGY AND DISTRIBUTION
Ecology of Western New Guinean Freycinetia........................................................
Distribution of Western New Guinean Freycinetia.................................................
39 - 48
39
41
D. MOLECULAR STUDY OF FREYCINETIA
cpDNA Sequences of trnL-F ..................................................................................
cpDNA Sequence of atbB-rbcL...............................................................................
cpDNA Sequence of trn-L.......................................................................................
Evolution of Western New Guinean Freycinetia....................................................
49 - 64
49
54
59
62
E. DESCRIPTION OF THE FREYCINETIA IN WEST NEW GUINEA
Key to Groups
65 - 205
vii
The F. macrostachya Group ...................................................................................
The F. funicularis Group .........................................................................................
The F. oblanceolata. Group .....................................................................................
The F. angustissima Group......................................................................................
Key to Species of F. macrostachya Group
Species of F. macrostachya Group.........................................................................
F. aculeata Sinaga....…………………………………………………………
F. andajensis Martelli……… ………………………………………………
F. arfakiana Martelli………………………………………………………..
F. archboldiana Merr.&Perry……………………………………………….
F. aurihastata Sinaga………………………………………………………..
F. bicolor B.C.Stone………………………………………………………….
F. brevipedicelata Sinaga……………………………………………………..
F. clavata Sinaga & Utterdige……………………………………………….
F. circuita Sinaga……………………………………………………………
F. concordia Sinaga………………………………………………………….
F. excelsa F. Muell………………………………………………………......
F. formosula Huynh .……………………………………………………….
F. fibrosa Martelli ……………………………………………………………
F. frutaspiralica Sinaga……………………………………………………. .
F. frutonumerata Sinaga…………………………………………………… ..
F. impundens Stone…………………………………………………………..
F. klosii Ridl,…………………………………………………………………
F. lacinulata Kanehira. ………………………………………………………
F. macrostachya Martelli…………………………………………………….
F. marginata Blume…………………………………………………………
F . mandacana Sinaga ……………………………………………………….
F. megaauriculata Sinaga & Utteridge………………………………………
F. pallida Huynh……………………………………………………………..
F. plana Huynh……………………………………………………………. .
F. percostata Merr. & Perry………………………………………………….
F. pseudoinsignis Warb……………………………………………………….
F. rosellana Huynh……………………………………………………………
F. solomonensis B.C. Stone…………………………………………………..
F. stigmabeliata Sinaga…………………………………………………….. .
F. trachypoda Merr. & Perry….……………………………………………. .
F. tessellata Merr. & Perry……………………………………………….
F. undulata Merr. & Perry………………………………………………….
F. ultrapedicelata Sinaga……………………………………………………..
F. whitmorei B.C. Stone……………………………………………………..
F. yapenina Sinaga………………………………………………………….
65
66
67
68
69
73
73
74
75
76
76
78
78
80
81
82
83
84
85
85
87
88
89
89
91
92
93
94
95
96
97
98
98
99
100
101
102
103
104
105
106
viii
Key to Species of F. funicularis Group
Species of F. funicularis Group ................................................................................
F. ayawasa Sinaga ………………………………………………………….
F. cryptocarpa Kanehira……………………………………………………..
F. erythospatha Merr. & Perry………………………………………………
F. funicularis Merr…………………………………………………………..
F. fusiforma Sinaga & Keim ………………………………………………..
F. gibbsiae Rendle ………………………………………………………….
F. hastata Sinaga ……………………………………………………………
F. imbristigma Sinaga & Keim…………………………………………… ..
F. lateriflora Ridll. …………………………………………………………..
F. magnoareola Sinaga………………………………………………………
F. millikenus Sinaga & Keim ………………………………………………
F. nervoauricula Sinaga ……………………………………………………..
F. pleurantha Merr. & Perry ………………………………………………..
F. rhodospatha Ridll…………………………………………………………
F. sterrophylla Merr. & Perry……………………………………………….
F. spiroaxilla Sinaga & Keim..………………………………………………
F. simpliaxillata Sinaga……………………………………………………...
120
122
123
124
125
127
128
129
130
131
132
134
135
136
137
138
139
140
Key to Species of F. oblanceolata Group
Species of F. oblanceolata Group...............................................................................
F. albaauria Sinaga…………………………………………………………
F. batantaensis Sinaga …………………………………………………… ..
F. biroi Warb. …………………………………………………………… .
F. brasii Merr. & Perry……………………..………………………………
F. broccoareola Sinaga……………..………………………………………
F. chartacea Huynh ………..……………………………………………… ..
F. concolor Huynh…………………………………………………………..
F. desendata Sinaga………………………………………………………….
F. ellipsoidalis Merr.& Perry ……………………………………………….
F. flaviceps Rendle…………………………………………………………..
F. folitenela Sinaga …………………………..…………………………….
F. forbesii Ridl,…………………………………………………………….
F. frutasolla Sinaga ….…………………………..……………………… .
F. gunungmejensis Sinaga……………………………………………………
F. hagenicola Huynh ………………………………………………………
F. inermis Ridl. …………………………………………………………….
F. iriana Sinaga ……………………………………………………………
F. lalokiensis Huynh ………………………………………………………
F. lagenicarpa Warb. .....................................................................................
F. lenifolia Huynh ..........................................................................................
F. lineariafolia Merr. & Perry……………………………………………….
152
154
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
173
173
174
F. marantifolia Hemsl…………………………………………………..….
175
ix
F. manokwariana Sinaga……………………………………………………...
F. obtusiacuminata Huynh……………………………………………………
F. oblanceolata Martelli …………………………………………………….
F. oreophila Merr. & Perry…………………………………………………..
F. rectangularis Kanehira……………………………………………………
F. scandens Gaudich. ………………………………………………………
F. tenuifolia Huynh ……………………………………………………….
F. tenuis Solms …………………………………………………………….
F. versteegii Sinaga ………………………………………………………….
176
177
178
179
180
181
182
183
184
Key to Species of F. angustissima Group
Species of F. angustissima Group..............................................................................
F. angusta Huynh ………………………………………………………….
F. angustissima Ridl. ……………………………………………………….
F. brachyclada Huynh ……………………………………………………
F. linearis Merr. & Perry ………………………………………………….
F. pauciberria Sinaga……………………………………………………… .
F. polyclada Merr. & Perry ………………………………………………
F. pseudoangustisimma Huynh …………………………………………….
F. stenophylla Warb…………………………………………………………..
195
196
196
197
198
199
200
201
202
203
IV. GENERAL DISCUSSION AND CONCLUSSION ..............................................
A. GENERAL DISCUSSION......................................................................................
B. CONCLUSSION.....................................................................................................
206 - 210
206
208
REFERENCES..............................................................................................................
APPENDIX...................................................................................................................
211 - 214
215
Aligment of nucleotides sequences of trnL-F spacer of the Chloroplast DNA Freycinetia
Aligment of nucleotides sequences of atpB-rbcL spacer of the Chloroplast DNA Freycinetia
Aligment of nucleotides sequences of trn-L spacer of the Chloroplast DNA Freycinetia
The Ecology and Distribution of Freycinetia Gaud. (Pandanaceae; Freycinetoideae) in The
Indonesian New Guinea
x
LIST OF TABLES
page
1. Primer Using for Sequensing …………………………………………………..…….5
2. Material for Sequencing DNA......................................................................................6
3. Morphological characters for PAUP analysis………………. ………………..…. 32
4. Distribution of the members of the infrageneric groups suggested …………………40
in this current study (A = secondary forest; B = primary beach forest;
C = wet land area)
5. Species distribution of Freycinetia in West New Guinea according to the three .....45
diffrences location
6.Variation of sequence length, AT, GC content of trnL-F intergenic spacer
In Freycinetia............................................................................................................... 50
7. Deletion, insertion and subtitution on DNA sequence of of trnL-F intergenic spacer
some Freycinetia.........................................................................................................51
8.Variation of sequence length, AT, GC content of atpB-rbcL intergenic spacer
in Freycinetia.................................................................................................................55
9. Deletion, insertion and subtitution on DNA sequence of atpB-rbcL intergenic spacer
some Freycinetia........................................................................................................56
10.Variation of Sequence length, AT, GC content of trn-L intergenic spacer
in Freycinetia...............................................................................................................59
11.Deletion, insertion and subtitution on DNA sequence of of trn-L intergenic spacer
some Freycinetia........................................................................................................60
xi
LIST OF FIGURES
Page
1. The fourth kinds of leaves arrangement of the Freycinetia………………..
9
2. The surface of the lamina of Freycinetia ……………………………………
10
3. The shape and colour of the prophyll leaves ……………………………….
12
4. The variation of the auricle …………………………………………………… 12
5. The variation of cauline leaves ………………………………………………..
14
6. The prophyll and variation of the bracts……………………………………… 16
7. The staminate flowers……... ………………………………….…………….
18
8. The pistillate flowers………………….………………………………………
18
9. The variation of the stigmas…………………………………………………..
21
10. The variation of the areola ……………….…………………………………
22
11. The variation and density of the berry ……………………………………..
24
12. The variation of the seed……………………………………………………
26
13. Phylogenetic tree show four groups of West New Guinean Freycinetia…
37
14. Phylogenetic tree show four groups of West New Guinean Freycinetia…
38
15. Distribution Map of the F. angustissima Group………………………….
42
16. Distribution Map of the F. funicularis Group……………………………...
42
17. Distribution Map of the F. oblanceolata Group………………………….
44
18. Distribution Map of the F. macrostachya Group ……….........................
44
19. Strict consensus of the 500 equally most parsimonious trees obtained in …… 54
maximum parsimony analysis of the trnL-F sequences data.
20. Strict consensus of the 500 equally most parsimonious trees obtained in …… 58
xii
maximum parsimony analysis of the atpB-rbcL sequences data.
21. Strict consensus of the 500 equally most parsimonious trees obtained in…… 62
maximum parsimony analysis of the trn -L sequences data.
22. New Species of F. macrostachya Group
1. F. aurihastata Sinaga………….…………..………………………………
2. F. brevipedicelata Sinaga…………. ………….………………………….
3. F. clavata Sinaga & Utterdige………… ..………………………………
4. F. circuita Sinaga ………….……………………………………………
5. F. concordia Sinaga………… .………………………………………….
6. F. frutaspiralica Sinaga…………. .……………………………………..
7. F. frutonumerata Sinaga………….. .……………………………………..
8. F. mandacana Sinaga………….…………………………………………
9. F. megaauriculata Sinaga & Utteridge………….. ..……………………..
10. F. stigmabeliata Sinaga…………… .……………………………………
11. F .ultrapedicelata Sinaga…………. ..……………………………………
12. F. yapenina Sinaga…………. ..…………………………………………
108
109
110
111
112
113
111
112
113
114
115
116
23. New Species of F. funicularis Group
1. F. ayawasa Sinaga…………..……………………………………………
2. F. fusiforma Sinaga & A.P.Keim…………...……………………………
3. F. hastata Sinaga………….……………………………………………. .
4. F. imbristigma Sinaga & A.P. Keim................. …..…………..…………
5. F. magnoareola Sinaga…………. .………………………………………
6. F. millikenus Sinaga & A.P. Keim…………..…………………………...
7. F. nervoauriculata Sinaga …………....………………………………….
8. F. spiroaxilla Sinaga & A.P. Keim …………. .………………………….
9. F. simpliaxillata Sinaga …………. ……………………………………...
142
143
144
145
146
147
148
149
150
24. New Species of F. oblanceolata Group
1. F. albaauria Sinaga, ………… .…………………………………………
2. F. batantaensis Sinaga………….……………………………………….
3. F. broccoareola Sinaga………….. ..…………………………………….
4. F. desendata Sinaga…………. ..…………………………………………
5. F. folitenela Sinaga………………….……………………………………
6. F. frutasolla Sinaga…………. .………………………………………….
8. F. iriana Sinaga………….………………………………………………
9. F. manokwariana Sinaga………….. ……………………………………..
186
187
188
189
190
191
192
193
25. New Species of F. angustissima Group
1. F. pauciberria Sinaga………….. ………………………………………..
205
xiii
Title of Dissertation
Name
Registration Number
Program Study
:
:
:
:
Systematics of Freycinetia in West New Guinea
Nurhaidah Iriany Sinaga
G361040051
Biology
Approved by
1. Advisory Committees
Dr. Rita Megia, DEA
(Chairman)
Prof. Dr. Ir. Alex Hartana
(Member)
2. Program Study of Biology
Prof. Dr. Mien A. Rifai
(Member)
Dr. Ary P. Keim
(Member)
3. Post Graduate School
Dr. Dedy Duryadi Solihin, DEA
(Head)
Dr. Ir. Dahrul Syah, MSc. Agr
(Dean)
Date of Examination: 23 Dec 2010
Date of Graduation…………
I. INTRODUCTION
The Pandanaceae has four genera namely Pandanus, Freycinetia, Sararanga and
Martelidendron. Freycinetia was described in 1824 by the French botanist Gaudichaud.
Until now approximately 200 species present in the world. But this genus is not very
widespread and the real home of Freycinetia is in Malaysia, Indonesia, Philippines,
Formosa, Okinawa, New Guinea and surrounding islands, Australia and and New
Zealand. The centre of Freycinetia distribution is certainly from the Philippines to New
Guinea (Stone, 1970). The New Guinea islands according to Stone (1976, 1983) may be
have 60 species, while in comparison Malaya has 8 species and Borneo has 24 species.
The genus are climbing pandan except F. arborea in Hawai . Freycinetia prefer
living near rivers or in humid areas especially under canopy. Freycinetia could be
found from the sea level to 2400 m asl . It is restricted in their habitat. For example F.
elegantula and F. pseudoangustisimma that are found only in mountains area but only
few species could be found both in the low land and mountain area for example F.
funicularis. The limitation area of occupation may related to the agent of distribution.
Unlike Pandanus with used win as a distribution agent, Freycinetia seeds are spread by
mammals and birds . That is why most species have colourfull beauty bracts to attract
animals to visit the flower.
According to International Plant Nomenclature Index for about 135 names of
Freycinetia present in New Guinea. Perhaps some are synonymous name, but it show
that many species occurs in the area. From eastern New Guinea, more species were
published by Warburg (1900), Marteli (1910), Merril & Perry (1940), Stone (1963) and
Huynh (1996 - 2003). Huynh mentioned about 87 new species into 8 parts journals of
New Guinea Freycinetia . Few of the species are also found in the western area. Stone
(1968) and Huynh (1997) also published many species from Salomon island and few of
them present in Papua too like F. salomonensis Stone and F. plana Huynh. Some
botanists have been published Freycinetia species from Western New Guinea. Solms
et.al in 1883, Martelli in 1910, Rendle in 1917), followed by Ridley in 1916 to 1922,
Merril and Perry in 1940 and Kanehira & Hatusima in 1941. Next, in 1976 Schumann
& Lauterbach mentioned 1 species from Jayapura,
followed by Huynh in 2002 who
2
published 1 new species from Tembagapura. Therefore they all found totally 36
species from Papua and 25 species were new species.
Many specimen from Papua are collected and put in Herbarium Manokwariense
(Man) in Manokwari, Bogoriense (BO) in Bogor, Lae in Morobe PNG, NHN (L) in
Leiden, Nederland and Kew (K) in England . Most of them were put as unknown
species. For example in BO, from 200 specimens only 42 specimens are known species.
How many species exactly occur in Papua need a taxonomical work to find out.
Inter species classification of Freycinetia were published first by Warburg in 1900
which about 2 sections and it was followed by Stone (1968) who mentioned about 15
sections. Warburg worked based on number of stigma namely Oligostigma section
with 1-3 stigmas and Pleiostigma with 3 or more stigmas.
Stone mentioned section
based on raceme and umbel inflorescence and the umbel one were divided for terminal
versus lateral or both lateral and terminal inflorescence. Next the terminal were divided
again based on
pseudopetiole, cephalia, berries, stigma and auricle. However, some
specimens have possibilities to put under 2 section. like F. archboldiana
Other taxonomy problems
are
sinonim name of Freycinetia.
Kanehira and
Hatusima in 1940 mentioned F. inouei as a new species from Dalman in Nabire, West
New Guinea. They mentioned that the species have closely related to F. archboldiana but
different species. Stone in 1972 put the F. inouei as sinonim of F. archboldiana.
Some botanist like Martelli in 1910; Merril & Perry in 1940; Kanehira in 1941 and
Stone in 1967 said that F.angustissima were sinonim to F. stenophylla Warb. and F.
polyclada Merr.& Perry. But, Huynh in 1999 mentioned the three species above were
different species. He used both morphological and anatomical characters. Huynh said
that in morphological characters F. angustissima and F. polyclada were showed their
differences. He also said that the stigma of F. angustisimma is 2 0r 3 rarely 1 or 4 but
stigma and areola is distinctly girdled and F. stenophylla has 2 stigmas with sitgma and
areola is un-distinctly girdled. Actually, our preliminary study showed that stigma and
areola in west New Guinean Freycinetia have many variation and many specimens with
unidentified label have the variation.
Solving all problems above need taxonomy work also phylogenetic analysis to make
sure about the sections. In Freycinetia
species, phylogenetic analysis even used
3
morphological approach never been done. Cox already made cladistic analysis based on
morphological approached but only in generic level. Further, Callmander et. al (2003)
produced phylogenetic trees based on cp DNA data on generic levels too.
Some Freycinetia species without flowers and good auricle looks similar like F.
palida and closely unknown species. For this species information from molecular data
will guide to understanding what species is, is it same or different species. However, for
build a good phylogenetic classification it need comprehensive approached.
According to FWI (Forest Watch Indonesia) report in 2004, deforestation happen in
Indonesia, with acceleration of degradation 3 times foot ball area in every minutes.
Consequently
some
species include Freycinetia
may
disappeared
without ever
identified .
Freycinetia have much utilities . All parts of this plant could be used, for example
hanging root for nail; stem for lance, traditional rain clothes; leaves for roof, and
handicraft like basket and bag, also for perfume; spadix and cephalia as foods; and
bracts for cooling drink and juice. To get more benefit for human life, more researches
need start from basic study. Therefore the taxonomic research were done using main
morphological information and molecular data as a second approach.
Purposed of the Study
The purposed of the West New Guinean Freycinetia study were:
1. To know the diversity of Freycinetia in West New Guinea
2. To obtain delimitation of the genus and species concepts
3. To build phylogenetic classification of Freycinetia based on morphological and
molecular data.
4. To know distribution of West New Guinea Freycinetia
4
I. INTRODUCTION
The Pandanaceae has four genera namely Pandanus, Freycinetia, Sararanga and
Martelidendron. They all occur in the old tropicals, from Africa to India, Indo-China to
Japan, Malaysia to Sumatera until New Guinea area include some islands surroundings
like Salomon islands and also present in Australia. Two genera namely . Martelidendron
and Sararanga have limited in both species number and distribution. Martelidendro has
6 species found only in Madagaskar and some surrounding is
IN WEST NEW GUINEA
NURHAIDAH IRIANY SINAGA
DEPARTMENT OF BIOLOGY
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
DECLARATION BY CANDIDATE
I hereby declare that this dissertation on Systematics of Freycinetia in West New
Guinea is my own work and effort directed by supervisory commission and has not been
submitted in any forms to any other universities for my award. The information sources
from the published or unpublished references of other authors have been acknowledged
and written in the references at the end of this dissertation.
Bogor, March 2011
Nurhaidah Iriany Sinaga
G 36 1040051
ABSTRACT
NURHAIDAH IRIANY SINAGA. Systematics of Freycinetia Gaudh. in West New
Guinea (supervised by Rita Megia, Alex Hartana, Mien A. Rifai and Ary
Prihardhyanto Keim).
Freycinetia Gaud. in West New Guinea had been investigated. For about 4.000 specimen
collections were studied using morphological, ecological and molecular data of cp DNA
sequence trnL-F, atpB-rbcL and trn-L . The result showed that in West New Guinea
92 species occur. All species can be divided into 4 groups namely F. macrostachya, F.
funicularis, F. oblanceolata and F. angustissima groups. The F. macrostachya group
have 36 species and 12 species are prepare as a new species namely F. aurihasteta
Sinaga, F. brevipedicelata Sinaga, F. clavata Sinaga & Utteridge, F . circuita Sinaga,
F.concordia Sinaga, F. frutaspiralica Sinaga, F. frutonumerata Sinaga, F. mandacana
Sinaga, F.megaauriculata Sinaga & Utteridge, F. stigmabeliata
Sinaga, F.
ultrapedicelata Sinaga and F. yapenina Sinaga. The F. funicularis group have 17
species and 9 species are prepare as a new species namely F. ayawasa Sinaga, F.
fusiforma Sinaga & A.P. Keim, F. hasteta Sinaga, F. imbristigma Sinaga & A.P. Keim,
F. magnoareola Sinaga, F.millikenus Sinaga & A.P. Keim, F. nervoauricula Sinaga,
F. spiroaxilla Sinaga & A.P. Keim and F. simpliaxillata. The F. oblanceolata group
have 31 species and 9 species are considerated as a new species namely F. albaauria
Sinaga, F. batantaensis Sinaga, F. decendata Sinaga, F. folitenella Sinaga, F. frutasolla
Sinaga, F. broccoareola Sinaga,
F. iriana Sinaga, F. manokwariana Sinaga and F.
verstegii Sinaga. The last group is F. angustissima group that have 8 species but only
one as a new species namely F. pauciberria Sinaga. The F. macrostachya and F.
oblanceolata groups living from forest beach to 1700 m asl but F. macrostachya group
are found alone in secondary forest. The F. funicularis and F. angustissima groups prefer
living on the mountain area and both of them are found on 3000 m asl. 35 species has
limited distribution it is only found in Papua but 52 species occurs both in Papua &PNG
and 5 species are widely distribution throughout Malesia and Australia. Analysis
phylogenetic through PAUP program using both morphological and molecular data
support the groups with CI (Coefficient Index) = 0.7, HI (Homoflashy Index) = 0.3 for
morphology characters and CI =0.930; 0.868; 0.890 and HI = 0.07; 0.132; 0.110 for
molecular data by trnL-F; atpB- rbcL and trn-L sequencing of CP DNA. According to
morphological data F. macrostachya group is a primitive one, It is followed by F.
funicularis group, F. oblanceolata and F. angustissima groups as an advance one. But in
the trnL-F sequencing cpDNA data F. macrostahya group as a primitive one separate
from 3 other derivate groups and this result are supported by ecological situation. On
the other hands, atpB–rbcL sequencing strongly support the groups but advance one is
not belong to the F. angustissima group anymore but F. funicularis group, The same way
is found in the trn-L sequencing data. Therefore trnL-F sequencing had given more
conservative characters but atpB-rbcL showed both conservative and advance characters
while trn-L had given more advance characters.
Key words: Freycinetia, West New Guinea, systematics, morphological, ecological and
molecular data.
ABSTRAK
NURHAIDAH IRIANY SINAGA. Sistematika Freycinetia di Nugini Barat dibimbing
oleh Rita Megia, Alex Hartana, Mien A. Rifai dan Ary Prihardhyanto Keim.
Freycinetia Gaud. di Nugini Barat telah diteliti. Sebanyak kurang lebih 4000 lembar
spesimen herbarium telah dipelajari, menggunakan data morfologi, ekologi dan
molekular dengan sekuensing kloroplast DNA yakni sekuens trnL-F; atpB-rbcL ; dan
trn-L .Hasil penelitian menunjukan bahwa di Nugini Barat terdapat sebanyak 92 jenis
Freycinetia . Jenis-jenis Freycinetia ini terdiri atas 4 kelompok yakni kelompok F.
macrostachya, F. funicularis, F. oblanceolata dan kelompok F. angustissima. Kelompok
F. macrostachya memiliki 36 jenis dengan 13 jenis baru yaitu F. aculeata Sinaga, F.
aurihastata Sinaga, F. brevipedicelata Sinaga,
F. clavata Sinaga & Utteridge, F
.circuita Sinaga, F.concordia Sinaga, F.frutaspiralica Sinaga, F. frutonumerata Sinaga,
F. mandacana Sinaga, F.megaauriculata Sinaga & Utteridge, F. stigmabeliata Sinaga,
F. ultrapedicelata Sinaga dan F. yapenina Sinaga. Kelompok F. funicularis memiliki
17 jenis dengan 9 jenis baru yaitu F. ayawasa Sinaga, F. fusiforma Sinaga &
A.P.Keim, F. hastatus Sinaga, F. imbristigma Sinaga & A.P. Keim , F. magnoareola
Sinaga, F. millikenus Sinaga & A.P. Keim, F. nervoauricula Sinaga, F. spiroaxilla
Sinaga & A.P. Keim dan F. simpliaxillata. Kelompok F. oblanceolata memiliki 31
jenis dengan 10 jenis baru yakni F. albaauria Sinaga, F. batantaensis Sinaga, F.
decendata Sinaga, F. folitenella Sinaga, F. frutasolla Sinaga, F .gunungmejensis
Sinaga, F. broccoareola Sinaga, F. iriana
Sinaga, F. manokwariana Sinaga dan
F.verstegii Sinaga. Kelompok F. angustissima memiliki 8 jenis dengan 1 jenis baru
yaitu F. pauciberria Sinaga. Kelompok F. macrostachya dan F. oblanceolata hidup di
hutan pantai hingga ketinggian 1700 m dpl. Sementara kelompok F. funicularis dan F.
angustissima menyenangi tempat tinggi, hingga ketinggian 3000 m dpl. Sebanyak 35
jenis hanya ditemukan di Nugini Barat, namun terdapat 52 jenis yang juga hidup di
PNG dan 5 jenis memiliki persebaran luas hingga ke Malesia bahkan Australia.
Keempat kelompok ini didukung hasil analisis filogenetik dengan program PAUP
berdasarkan data morfologi dan molekular, dengan nilai CI (Coefficient Index)= 0.7, HI
(Homoflashy Index) = 0.3 untuk data morfologi dan nilai CI =0.930; 0.868; 0.890 and
HI = 0.07; 0.132; 0.110 untuk data molekular sekuensing DNA kloroplast berturutturut
trnL-F; atpB-rbcL; dan trn-L. Berdasarkan data morfologi kelompok F.
macrostahya adalah kelompok primitif, dikuti kelompok F. funicularis,, F. oblanceolata
dan F. angustissima. Tetapi sekuensing trnL-F lebih memisahkan kelompok primitif
dari ketiga turunannya dan data sekuensing ini sangat didukung oleh data ekologis. Di
sisi lain, sekuensing atpB-rbcL mendukung keempat kelompok ini, namun kelompok F.
angustissima tidak lagi sebagai yang lebih maju melainkan kelompok F. funicularis,
keadaan yang sama terjadi dalam sekuensing trn-L. Dengan demikian trnL-F lebih
banyak menggambarkan karater yang konservatif sementara atpB-rbcL menggambarkan
keduanya baik sifat konservatif maupun sifat yang lebih maju dan trn-L sekuensing lebih
menggambarkan sifat yang lebih maju.
Kata Kunci: Freycinetia, Nugini Barat, sistematika, morfologi, ekologi dan molekular
data.
SUMMARY
Freycinetia was firstly described by Gaudichaud in 1824. Currently there are about
200 species recognized. The centre of distribution of this genus is in New Guinea, where
at least 60 species have been recognized (Stone 1976; 1983). This number is much
higher compare to Borneo (24) and the Malay Peninsula (8). The majority of herbarium
specimens of Freycinetia from New Guinea are deposited only in few Herbaria such as
Herbarium Bogoriense (BO) in Bogor-Indonesia, Herbarium Manokwariense (MAN) in
Manokwari-Indonesia, Herbarium Lae in Lae-Papua New Guinea, the National
Herbarium of the Netherlands in Leiden (L)-the Netherlands, and Herbarium Kewensis in
Kew-UK. Unfortunately, due to the incomplete nature of the specimens most are labelled
as unknown species. From time to time deposit specimens increase in numbers but
many of them are still unidentified. Therefore, it need taxonomical study to find out how
many species of New Guinea Freycinetia especially in the west area.
First serious molecular study of the family was proceeded by Callmander et al.
(2003). Unfortunately, the infrageneric classification of the genus is still unresolved. So
the aims of this study were to unravel the diversity of Freycinetia in the western part of
New Guinea, revision of the genus particularly the infrageneric classification and
knowing the species distribution in western part of New Guinea in relation with New
Guinea as a whole.
Method of the study using both morphological and molecular approach. The
research was carried out from Jully 2005 until Maret 2009. Taking and collecting
specimens were done at several area in west New Guinea such as Timika (2005),
Manokwari ( 2006), Sarmi (2006) and Jayapura (2006, 2008), followed by ecological
observation in the field and morphological observation in herbarium B0 (2006, 2008),
MAN (2006), LAE (2006), L (2008) and K (2009). Molecular work using sequence
non coding cpDNA: trnL-F, trn-L and atpB – rbcL was done at Plant Biology and
Biotechnology Laboratorium IPB (Bogor Agricultural University) on August to October
2008 and van der Klauw Molecular Laboratorium in Leiden on November 2008 until
January 2009.
The result showed that 92 species of Freycinetia occur in the area. All species can be
divided into 4 groups namely F. macrostachya, F. funicularis, F. oblanceolata and
F. angustissima groups.
The F. macrostachya group have robust habit and are high climbers. Stem robust,
hard, rarely branched, 2-4 cm diameter. Leaf imbricate, linear or lanceolate, margin with
sharp and hard spines, 20-120 cm long, 1.5-7 cm wide; cauline leave colorfull in
common, basically leaf with yellow, orange, red or mixed color, consist of 2-3 whorls;
Infructescence terminal, ternate, sometimes with 4, 8 to 12 cephalia, spirally to umbelly
arranged. Prophyll bracts never found. Cephalium large, 3-20 cm long, 2-6 cm wide.
Berries numerous, thin, stout; stigmatic remains 1-3 or 5-6, rarely 8 or 12, one species
with 32. Seeds linear or ellipse, usually longitudinally arranged.
The F. funicularis group have leafless stem from basal part up to the half part of the
stem, laterally branching, and leaves found only on the terminal part of stem. Stem 2-3
cm diameter. Leaf semi imbricate, linear or lanceolate, margin usually with sharp and
hard spines, 13-50 cm long, 1.5-3 cm wide; Prophyll bracts consist of 4 to more than 8
whorls; bracts mostly consist 3 parts exterior, middle and interior bracts Infructescence
axillary, ternate or rarely quaternate, umbelly arranged. Cephalium oblong or cylindrical,
3-10 cm long, 1.5-3 cm wide. Berries few, a berry usually formed through fusion of
smaller berries giving an impression of a larger berry; stigmatic remains 5-8, rarely 3, 12,
14 or 18, stigmatic position usually changes from transversal to longitudinal, some
species with marginal stigmatic remains. Seed usually flat, transversally arranged.
The F. oblanceolata group possess slender stem, 0.5-2 cm diameter, numerously
branching, branch short, less than 50 cm long. Leaf non imbricate, elliptical to oblong
and oblanceolate, margin with spines on terminal and basal parts only, 2-15 cm long, 0.54 cm wide; Cauline leaves usually green; Prophyll only in both terminalia and axillary
inflorescence, if present only consist of 2 whorls; Infructescence usually terminal, rarely
both terminal and axillary. Cephalium minute, 2-7 cm long, 1-3 cm wide. Berry
numerous or few, some species with flat tips, fusion or self-changing areola present;
stigmatic remains 1-3 or 5-6. Seed linear or oblong-flat, usually arranged transversally,
in some species longitudinally
The F. angustissima group have slender stem, less than 0.5 cm in diameter, usually
numerously branched, branch short, less than 20 cm long. Leaves pseudo-rosette in
appearance, each linear or lanceolate, in some members observed elliptical, margin with
spines on terminal and basal parts, 2-8 cm long, 0.2-1 cm wide. Prophyll bracts never
found. Infructescence terminal, usually binate, rarely single or ternate, arranged in umbel.
Cephalium small, 2-4 cm long, 1-2 cm wide. Berry usually few in numbers, some
species with flat-tipped berries; stigmatic remains 1 or 2, rarely 3, self-changing areola
present giving variation in stigmatic areola. Seeds linear, transversally or longitudinally
arranged.
The F. macrostachya group have 36 species. The previous species have 23 species.
These are F. aculeata Sinaga, F. andajensis
Martelli, F. arfakiana Martelli,
F. archboldiana Merr. & Perry, F. bicolor B.C.Stone, F .excelsa F. Muell., F. fibrosa
Martelli, F. formosula Huynh, F. impundens B.C. Stone, F. Klosii Ridl., F. lacinulata
Kanehira, F. macrostachya Martelli, F. marginata Blume, F. palida Huynh, F. plana
Huynh, F. percostata Merr. & Perry, F. pseudoinsignis Warb., F. rosellana Huynh,
F. solomonensis B.C. Stone, F. trachypoda Merr. & Perry, F. tessellata
Merr. &
Perry, F. undulate Merr. & Perry, and F. whitmorei B.C. Stone. Twelve species are
regarded as new species namely F. aurihasteta Sinaga, F. brevipedicelata Sinaga, F.
clavata Sinaga & Utteridge, F. circuita Sinaga, F. concordia Sinaga, F. frutaspiralica
Sinaga, F. frutonumerata Sinaga, F. mandacana Sinaga, F. megaauriculata Sinaga &
Utteridge, F. stigmabeliata Sinaga, F. ultrapedicelata Sinaga and F. yapenina Sinaga.
The F. funicularis group have 17 species. The previous 8 species are F. cryptocarpa
Kanehira, F. erythospatha Merr. & Perry, F. funicularis Merr.,, F.gibbsiae Rendle,
F. lateriflora Ridll., F. pleurantha Merr. & Perry, F. rhodospatha Ridll., and
F. sterrophylla Merr. & Perry. Nine species are regarded as new species namely F.
ayawasa Sinaga, F. fusiforma Sinaga & A.P. Keim, F. hastata Sinaga, F. imbristigma
Sinaga & A.P. Keim, F. magnoareola Sinaga, F.millikenus Sinaga & A.P. Keim,
F. nervoauricula Sinaga, F. spiroaxilla Sinaga & A.P. Keim and F. simpliaxillata
Sinaga.
The F. oblanceolata group have 31 species. The previous species are 22 species
namely F. biroi Warb., F. brasii Merr. & Perry, F. chartacea Huynh, F .concolor
Huynh, F. ellipsoidalis Merr. & Perry, F. flaviceps Rendl., F. forbesii Ridl.,
F .gunungmejensis Sinaga, F. hagenicola Huynh, F. inermis Ridl., F. lalokiensis Huynh,
F. lagenicarpa Warb., F. lenifolia Huynh, F. linearifolia Merr. & Perry, F. marantifolia
Hemsl., F. obtusiacuminata Huynh, F. oblanceolata Martelli, F. oreophila Merr. &
Perry, F. rectangularis Kanehira, F. scandens Gaudich., F. tenuifolia Huynh, and
F. tenuis Solms. Nine species are regarded as new species namely F. albaauria Sinaga,
F. batantaensis Sinaga, F. decendata Sinaga, F. folitenella Sinaga, F. frutasolla Sinaga,
F. broccoareola Sinaga, F. iriana Sinaga, F. manokwariana Sinaga and F. verstegii
Sinaga.
The F. angustissima group have 8 species and 1 species are regarded as a new
species namely F. pauciberria Sinaga. The previous species here are F. angusta Huynh,
F. angustissima Ridl., F. brachyclada Huynh, F. linearis Merr. & Perry, F. polyclada
Merr. & Perry, F. pseudoangustisimma Huynh and F. stenophylla Warb.
F. macrostachya & F. oblanceolata groups living from forest beach to 1700 m
above sea level. But F. macrostachya group are found alone in secondary forest. on
the other hand, the F. funicularis and F. angustissima groups prefer living on mountain
area and both of them are found up to 3000 m asl. Despite share with F. angustissima
group, the F. funicularis group also share habitat with F. oblanceolata group, especially
in the forest beach where F. angustissima
group never is found, but this group has
ability living on the wet area. West New guinea Freycinetia have 35 species that are
limited distribution, because these species are only occur in the area. But West New
Guinea Freycinetia share 52 species with PNG, and 5 species remains are widely
distribution species that spread through out Malesia region until Australia.
Analysis phylogenetic through PAUP program using both morphological and
molecular data support the groups with CI (Coefficient Index) = 0.7, HI (Homoflashy
Index) = 0.3 for morphology characters and CI =0.930; 0.868; 0.890 and HI = 0.07;
0.132; 0.110 for molecular data by trnL-F; atpB-rbcL and trn-L sequencing of
Chloroplast DNA. According to morphological data
F. macrostachya group is a
primitive one, It is followed by F. funicularis, F. oblanceolata and F. angustissima
group as an advance one. But in the trnL-F sequencing cpDNA data F. macrostachya
group as a primitive one separate from 3 other derivate groups and this data strongly
support ecological situation. On the other hands, atpB–rbcl sequencing strongly support
the groups but advance one is not belong to the F. angustissima group anymore but F.
funicularis groups, it is same way to the trn-L sequencing data. Therefore trnL-F
sequencing had given more conservative characters but atpB-rbcL showed both
conservative and advance characters while trn-L had given more advance characters.
Copyright @2011 Bogor Agricultural University
Copyright are protected by low;
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2. It is prohibited to republish and reproduce all or part of this dissertation without the
written permission from Bogor Agricultural University.
SYSTEMATICS OF FREYCINETIA
IN WEST NEW GUINEA
NURHAIDAH IRIANY SINAGA
Dissertation
Submitted as a part of requirement for the fulfillment of Doctor Degree in Biology
DEPARTMENT OF BIOLOGY
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
ii
Examiners in comprehensive examination:
Dr. Nunik Sri Aryanti, M.Si
Department of Biology, Faculty of Mathematics and Natural Science, Bogor Agricultural
University, Bogor, Indonesia
Dr. Ruliyana Susanti
Herbarium Bogoroense, Botany Division,
Research Centre for Biology
Institute of Science, Cibinong, Bogor, Indonesia
Examiners in final examination:
Dr. Tatik Chikmawati, M,Si
Department of Biology, Faculty of Mathematics and Natural Science, Bogor
Agricultural University, Bogor, Indonesia
Dr. Rugayah
Herbarium Bogoroense, Botany Division,
Research Centre for Biology
Institute of Science, Cibinong, Bogor, Indonesia
iii
iv
PREFACE
The main object of the research conducted at November 2005 to January 2009 is
Freycinetia in West New Guinea. This study emphasis four different topics namely the
morphological characters, ecological and distribution, chloroplast DNA sequencing and
description of 92 species of New Guinean Freycinetia. A part of morphological
approach i.e. The Stigma and Areola in the Western New Guinean Species of
Freycinetia Gaud. (Pandanaceae; Freycinetoideae) submmited to the 8 th International
Flora Malesiana Symposium in Singapore and Two New Species of Freycinetia
(Pandanacea) from Manokwari, West Papua have been published in Reinwardtia 13:2
(2010) 183 - 187. Part two: The Ecology and Distribution of Freycinetia Gaud.
(Pandanaceae; Freycinetoideae) in The Indonesian New Guinean have also been
published in Reinwardtia 13:2 (2010) 189 -197.
I am extremely grateful to the advisory committees who directed, advised and read
critically the manuscript, Dr. Rita Megia, DEA (Department Biology IPB), Prof. Alex
Hartana (Department Biology IPB), Prof. Mien A. Rifai (Herbarium Bogoriense, LIPI)
and Dr. Ary .P. Keim (Herbarium Bogoriense, LIPI).
Profound thanks and appreciation to Dr. Eko B. Waluyo and all staff of Herbarium
Bogoriense especially Dr. Rugayah and Prof. Dr. Elizabeth Widjaja; Prof. Dr. Peter van
Welzen, Dr. Barbara Gravendell, Marcel Eurings and Dr. Michael Stech (Leiden
University, NHN Herbarium, Nederland); Dr. Rogier de Kok, Dr. William Bekker and
Dr. Timothy Utteridge (Kew Herbarium, UK); Roy Banka (Lae Herbarium, PNG); Ir.
Jack Wanggai, M.Si and staff (Herbarium Manokwariense) and Dr. Fenny Ismoyo
(Papua University); Ir. Pratita Puradyatmika, M.M (PT. Freeport Indonesia) for any kind
of help during my research.
I gratefully acknowledge the BPPS Scholarship, Sandwich Program and Litdas
from Director General of Higher Education (Dikti), Departement of National Education
Republic Indonesia. I thanks Ir. J.P. Karafir M,Sc (Rector Unipa); Dr. Hasyim (Dean
Math and Science Faculty of IPB) and Dr. Deddy Duryadi DEA (Head of Program Study
Biology Post Graduate School IPB).
I am indebted to all staff and my colleagues in Papua University, my field assistant:
Gasper Taudufu, Christhopere Diaz and Tinus Iwanggin and my friends at Subprogram
Plant Taxonomy IPB: Dr. Iris Rengganis, Dr. Nursahara Pasaribu, Dr. Sri Endarti, Dr.
Puji Widodo, Dr. Fitmawati, Dr. Himah Rustiami, also my brothers: Dr. Soaloon
Sinaga, Kristian Sinaga, SKM, Benjamin Sinaga, S.Tp and my sisters: Sesniwati Sinaga,
SE and Bertha Sinaga, S.Sos with their families. Finally, I give my honor to my beloved
parent: Kasiaman Sinaga and Johana Taudufu.
Bogor, March 2011
Nurhaidah I.Sinaga
v
CURRICULUM VITAE
Nurhaidah Iriany Sinaga was born on 6 January 1969, in Karubaga Papua, as the
second daughter of Kasiaman Sinaga and Johana Taudufu. She graduated from
Forestry Department at Cenderawasih
University in 1991. She studied plant
taxonomy and got M.Si in Biology from IPB in 2001. In 2004, she got scholarship
for study at Program Study of Biology IPB from BPPS Dikti (Directorate General
of Higher Education, National Education Department). Since 1995, she has been
lecturer at the Departement of Forestry, University of Papua.
vi
TABLE OF CONTENT
Page
LIST OF TABLES
LIST OF FIGURES
I. INTRODUCTION
Purpose of The Study
II. METHOD OF THE STUDY
Morphological work……………………………………………………………….
Molecular work……………………………………………………………………
III.RESULTS AND DISCUSSIONS
A. MORPHOLOGICAL CHARACTERS
Stem..........................................................................................................................
Leaves.......................................................................................................................
Spines......................................................................................................................
Prophyll Leaves........................................................................................................
Auricle......................................................................................................................
Cauline Leaves.........................................................................................................
Prophyll Bracts........................................................................................................
Bracts.......................................................................................................................
Peduncle...................................................................................................................
Pedicle......................................................................................................................
Staminate Flower......................................................................................................
Pistillate Flower........................................................................................................
Stigma and Areola...................................................................................................
Cephalia..................................................................................................................
Berries......................................................................................................................
Seed.........................................................................................................................
x
xi
1
3
4
4
4
8
8 - 26
8
9
11
11
13
14
15
15
17
17
17
19
19
22
23
25
B. PHYLOGENETIC ANALYSIS OF FREYCINETIA
Relationship and Evolution of the Freycinetia........................................................
27 - 38
29
C. ECOLOGY AND DISTRIBUTION
Ecology of Western New Guinean Freycinetia........................................................
Distribution of Western New Guinean Freycinetia.................................................
39 - 48
39
41
D. MOLECULAR STUDY OF FREYCINETIA
cpDNA Sequences of trnL-F ..................................................................................
cpDNA Sequence of atbB-rbcL...............................................................................
cpDNA Sequence of trn-L.......................................................................................
Evolution of Western New Guinean Freycinetia....................................................
49 - 64
49
54
59
62
E. DESCRIPTION OF THE FREYCINETIA IN WEST NEW GUINEA
Key to Groups
65 - 205
vii
The F. macrostachya Group ...................................................................................
The F. funicularis Group .........................................................................................
The F. oblanceolata. Group .....................................................................................
The F. angustissima Group......................................................................................
Key to Species of F. macrostachya Group
Species of F. macrostachya Group.........................................................................
F. aculeata Sinaga....…………………………………………………………
F. andajensis Martelli……… ………………………………………………
F. arfakiana Martelli………………………………………………………..
F. archboldiana Merr.&Perry……………………………………………….
F. aurihastata Sinaga………………………………………………………..
F. bicolor B.C.Stone………………………………………………………….
F. brevipedicelata Sinaga……………………………………………………..
F. clavata Sinaga & Utterdige……………………………………………….
F. circuita Sinaga……………………………………………………………
F. concordia Sinaga………………………………………………………….
F. excelsa F. Muell………………………………………………………......
F. formosula Huynh .……………………………………………………….
F. fibrosa Martelli ……………………………………………………………
F. frutaspiralica Sinaga……………………………………………………. .
F. frutonumerata Sinaga…………………………………………………… ..
F. impundens Stone…………………………………………………………..
F. klosii Ridl,…………………………………………………………………
F. lacinulata Kanehira. ………………………………………………………
F. macrostachya Martelli…………………………………………………….
F. marginata Blume…………………………………………………………
F . mandacana Sinaga ……………………………………………………….
F. megaauriculata Sinaga & Utteridge………………………………………
F. pallida Huynh……………………………………………………………..
F. plana Huynh……………………………………………………………. .
F. percostata Merr. & Perry………………………………………………….
F. pseudoinsignis Warb……………………………………………………….
F. rosellana Huynh……………………………………………………………
F. solomonensis B.C. Stone…………………………………………………..
F. stigmabeliata Sinaga…………………………………………………….. .
F. trachypoda Merr. & Perry….……………………………………………. .
F. tessellata Merr. & Perry……………………………………………….
F. undulata Merr. & Perry………………………………………………….
F. ultrapedicelata Sinaga……………………………………………………..
F. whitmorei B.C. Stone……………………………………………………..
F. yapenina Sinaga………………………………………………………….
65
66
67
68
69
73
73
74
75
76
76
78
78
80
81
82
83
84
85
85
87
88
89
89
91
92
93
94
95
96
97
98
98
99
100
101
102
103
104
105
106
viii
Key to Species of F. funicularis Group
Species of F. funicularis Group ................................................................................
F. ayawasa Sinaga ………………………………………………………….
F. cryptocarpa Kanehira……………………………………………………..
F. erythospatha Merr. & Perry………………………………………………
F. funicularis Merr…………………………………………………………..
F. fusiforma Sinaga & Keim ………………………………………………..
F. gibbsiae Rendle ………………………………………………………….
F. hastata Sinaga ……………………………………………………………
F. imbristigma Sinaga & Keim…………………………………………… ..
F. lateriflora Ridll. …………………………………………………………..
F. magnoareola Sinaga………………………………………………………
F. millikenus Sinaga & Keim ………………………………………………
F. nervoauricula Sinaga ……………………………………………………..
F. pleurantha Merr. & Perry ………………………………………………..
F. rhodospatha Ridll…………………………………………………………
F. sterrophylla Merr. & Perry……………………………………………….
F. spiroaxilla Sinaga & Keim..………………………………………………
F. simpliaxillata Sinaga……………………………………………………...
120
122
123
124
125
127
128
129
130
131
132
134
135
136
137
138
139
140
Key to Species of F. oblanceolata Group
Species of F. oblanceolata Group...............................................................................
F. albaauria Sinaga…………………………………………………………
F. batantaensis Sinaga …………………………………………………… ..
F. biroi Warb. …………………………………………………………… .
F. brasii Merr. & Perry……………………..………………………………
F. broccoareola Sinaga……………..………………………………………
F. chartacea Huynh ………..……………………………………………… ..
F. concolor Huynh…………………………………………………………..
F. desendata Sinaga………………………………………………………….
F. ellipsoidalis Merr.& Perry ……………………………………………….
F. flaviceps Rendle…………………………………………………………..
F. folitenela Sinaga …………………………..…………………………….
F. forbesii Ridl,…………………………………………………………….
F. frutasolla Sinaga ….…………………………..……………………… .
F. gunungmejensis Sinaga……………………………………………………
F. hagenicola Huynh ………………………………………………………
F. inermis Ridl. …………………………………………………………….
F. iriana Sinaga ……………………………………………………………
F. lalokiensis Huynh ………………………………………………………
F. lagenicarpa Warb. .....................................................................................
F. lenifolia Huynh ..........................................................................................
F. lineariafolia Merr. & Perry……………………………………………….
152
154
154
155
156
157
158
159
160
161
162
163
164
165
166
167
168
169
170
171
173
173
174
F. marantifolia Hemsl…………………………………………………..….
175
ix
F. manokwariana Sinaga……………………………………………………...
F. obtusiacuminata Huynh……………………………………………………
F. oblanceolata Martelli …………………………………………………….
F. oreophila Merr. & Perry…………………………………………………..
F. rectangularis Kanehira……………………………………………………
F. scandens Gaudich. ………………………………………………………
F. tenuifolia Huynh ……………………………………………………….
F. tenuis Solms …………………………………………………………….
F. versteegii Sinaga ………………………………………………………….
176
177
178
179
180
181
182
183
184
Key to Species of F. angustissima Group
Species of F. angustissima Group..............................................................................
F. angusta Huynh ………………………………………………………….
F. angustissima Ridl. ……………………………………………………….
F. brachyclada Huynh ……………………………………………………
F. linearis Merr. & Perry ………………………………………………….
F. pauciberria Sinaga……………………………………………………… .
F. polyclada Merr. & Perry ………………………………………………
F. pseudoangustisimma Huynh …………………………………………….
F. stenophylla Warb…………………………………………………………..
195
196
196
197
198
199
200
201
202
203
IV. GENERAL DISCUSSION AND CONCLUSSION ..............................................
A. GENERAL DISCUSSION......................................................................................
B. CONCLUSSION.....................................................................................................
206 - 210
206
208
REFERENCES..............................................................................................................
APPENDIX...................................................................................................................
211 - 214
215
Aligment of nucleotides sequences of trnL-F spacer of the Chloroplast DNA Freycinetia
Aligment of nucleotides sequences of atpB-rbcL spacer of the Chloroplast DNA Freycinetia
Aligment of nucleotides sequences of trn-L spacer of the Chloroplast DNA Freycinetia
The Ecology and Distribution of Freycinetia Gaud. (Pandanaceae; Freycinetoideae) in The
Indonesian New Guinea
x
LIST OF TABLES
page
1. Primer Using for Sequensing …………………………………………………..…….5
2. Material for Sequencing DNA......................................................................................6
3. Morphological characters for PAUP analysis………………. ………………..…. 32
4. Distribution of the members of the infrageneric groups suggested …………………40
in this current study (A = secondary forest; B = primary beach forest;
C = wet land area)
5. Species distribution of Freycinetia in West New Guinea according to the three .....45
diffrences location
6.Variation of sequence length, AT, GC content of trnL-F intergenic spacer
In Freycinetia............................................................................................................... 50
7. Deletion, insertion and subtitution on DNA sequence of of trnL-F intergenic spacer
some Freycinetia.........................................................................................................51
8.Variation of sequence length, AT, GC content of atpB-rbcL intergenic spacer
in Freycinetia.................................................................................................................55
9. Deletion, insertion and subtitution on DNA sequence of atpB-rbcL intergenic spacer
some Freycinetia........................................................................................................56
10.Variation of Sequence length, AT, GC content of trn-L intergenic spacer
in Freycinetia...............................................................................................................59
11.Deletion, insertion and subtitution on DNA sequence of of trn-L intergenic spacer
some Freycinetia........................................................................................................60
xi
LIST OF FIGURES
Page
1. The fourth kinds of leaves arrangement of the Freycinetia………………..
9
2. The surface of the lamina of Freycinetia ……………………………………
10
3. The shape and colour of the prophyll leaves ……………………………….
12
4. The variation of the auricle …………………………………………………… 12
5. The variation of cauline leaves ………………………………………………..
14
6. The prophyll and variation of the bracts……………………………………… 16
7. The staminate flowers……... ………………………………….…………….
18
8. The pistillate flowers………………….………………………………………
18
9. The variation of the stigmas…………………………………………………..
21
10. The variation of the areola ……………….…………………………………
22
11. The variation and density of the berry ……………………………………..
24
12. The variation of the seed……………………………………………………
26
13. Phylogenetic tree show four groups of West New Guinean Freycinetia…
37
14. Phylogenetic tree show four groups of West New Guinean Freycinetia…
38
15. Distribution Map of the F. angustissima Group………………………….
42
16. Distribution Map of the F. funicularis Group……………………………...
42
17. Distribution Map of the F. oblanceolata Group………………………….
44
18. Distribution Map of the F. macrostachya Group ……….........................
44
19. Strict consensus of the 500 equally most parsimonious trees obtained in …… 54
maximum parsimony analysis of the trnL-F sequences data.
20. Strict consensus of the 500 equally most parsimonious trees obtained in …… 58
xii
maximum parsimony analysis of the atpB-rbcL sequences data.
21. Strict consensus of the 500 equally most parsimonious trees obtained in…… 62
maximum parsimony analysis of the trn -L sequences data.
22. New Species of F. macrostachya Group
1. F. aurihastata Sinaga………….…………..………………………………
2. F. brevipedicelata Sinaga…………. ………….………………………….
3. F. clavata Sinaga & Utterdige………… ..………………………………
4. F. circuita Sinaga ………….……………………………………………
5. F. concordia Sinaga………… .………………………………………….
6. F. frutaspiralica Sinaga…………. .……………………………………..
7. F. frutonumerata Sinaga………….. .……………………………………..
8. F. mandacana Sinaga………….…………………………………………
9. F. megaauriculata Sinaga & Utteridge………….. ..……………………..
10. F. stigmabeliata Sinaga…………… .……………………………………
11. F .ultrapedicelata Sinaga…………. ..……………………………………
12. F. yapenina Sinaga…………. ..…………………………………………
108
109
110
111
112
113
111
112
113
114
115
116
23. New Species of F. funicularis Group
1. F. ayawasa Sinaga…………..……………………………………………
2. F. fusiforma Sinaga & A.P.Keim…………...……………………………
3. F. hastata Sinaga………….……………………………………………. .
4. F. imbristigma Sinaga & A.P. Keim................. …..…………..…………
5. F. magnoareola Sinaga…………. .………………………………………
6. F. millikenus Sinaga & A.P. Keim…………..…………………………...
7. F. nervoauriculata Sinaga …………....………………………………….
8. F. spiroaxilla Sinaga & A.P. Keim …………. .………………………….
9. F. simpliaxillata Sinaga …………. ……………………………………...
142
143
144
145
146
147
148
149
150
24. New Species of F. oblanceolata Group
1. F. albaauria Sinaga, ………… .…………………………………………
2. F. batantaensis Sinaga………….……………………………………….
3. F. broccoareola Sinaga………….. ..…………………………………….
4. F. desendata Sinaga…………. ..…………………………………………
5. F. folitenela Sinaga………………….……………………………………
6. F. frutasolla Sinaga…………. .………………………………………….
8. F. iriana Sinaga………….………………………………………………
9. F. manokwariana Sinaga………….. ……………………………………..
186
187
188
189
190
191
192
193
25. New Species of F. angustissima Group
1. F. pauciberria Sinaga………….. ………………………………………..
205
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Title of Dissertation
Name
Registration Number
Program Study
:
:
:
:
Systematics of Freycinetia in West New Guinea
Nurhaidah Iriany Sinaga
G361040051
Biology
Approved by
1. Advisory Committees
Dr. Rita Megia, DEA
(Chairman)
Prof. Dr. Ir. Alex Hartana
(Member)
2. Program Study of Biology
Prof. Dr. Mien A. Rifai
(Member)
Dr. Ary P. Keim
(Member)
3. Post Graduate School
Dr. Dedy Duryadi Solihin, DEA
(Head)
Dr. Ir. Dahrul Syah, MSc. Agr
(Dean)
Date of Examination: 23 Dec 2010
Date of Graduation…………
I. INTRODUCTION
The Pandanaceae has four genera namely Pandanus, Freycinetia, Sararanga and
Martelidendron. Freycinetia was described in 1824 by the French botanist Gaudichaud.
Until now approximately 200 species present in the world. But this genus is not very
widespread and the real home of Freycinetia is in Malaysia, Indonesia, Philippines,
Formosa, Okinawa, New Guinea and surrounding islands, Australia and and New
Zealand. The centre of Freycinetia distribution is certainly from the Philippines to New
Guinea (Stone, 1970). The New Guinea islands according to Stone (1976, 1983) may be
have 60 species, while in comparison Malaya has 8 species and Borneo has 24 species.
The genus are climbing pandan except F. arborea in Hawai . Freycinetia prefer
living near rivers or in humid areas especially under canopy. Freycinetia could be
found from the sea level to 2400 m asl . It is restricted in their habitat. For example F.
elegantula and F. pseudoangustisimma that are found only in mountains area but only
few species could be found both in the low land and mountain area for example F.
funicularis. The limitation area of occupation may related to the agent of distribution.
Unlike Pandanus with used win as a distribution agent, Freycinetia seeds are spread by
mammals and birds . That is why most species have colourfull beauty bracts to attract
animals to visit the flower.
According to International Plant Nomenclature Index for about 135 names of
Freycinetia present in New Guinea. Perhaps some are synonymous name, but it show
that many species occurs in the area. From eastern New Guinea, more species were
published by Warburg (1900), Marteli (1910), Merril & Perry (1940), Stone (1963) and
Huynh (1996 - 2003). Huynh mentioned about 87 new species into 8 parts journals of
New Guinea Freycinetia . Few of the species are also found in the western area. Stone
(1968) and Huynh (1997) also published many species from Salomon island and few of
them present in Papua too like F. salomonensis Stone and F. plana Huynh. Some
botanists have been published Freycinetia species from Western New Guinea. Solms
et.al in 1883, Martelli in 1910, Rendle in 1917), followed by Ridley in 1916 to 1922,
Merril and Perry in 1940 and Kanehira & Hatusima in 1941. Next, in 1976 Schumann
& Lauterbach mentioned 1 species from Jayapura,
followed by Huynh in 2002 who
2
published 1 new species from Tembagapura. Therefore they all found totally 36
species from Papua and 25 species were new species.
Many specimen from Papua are collected and put in Herbarium Manokwariense
(Man) in Manokwari, Bogoriense (BO) in Bogor, Lae in Morobe PNG, NHN (L) in
Leiden, Nederland and Kew (K) in England . Most of them were put as unknown
species. For example in BO, from 200 specimens only 42 specimens are known species.
How many species exactly occur in Papua need a taxonomical work to find out.
Inter species classification of Freycinetia were published first by Warburg in 1900
which about 2 sections and it was followed by Stone (1968) who mentioned about 15
sections. Warburg worked based on number of stigma namely Oligostigma section
with 1-3 stigmas and Pleiostigma with 3 or more stigmas.
Stone mentioned section
based on raceme and umbel inflorescence and the umbel one were divided for terminal
versus lateral or both lateral and terminal inflorescence. Next the terminal were divided
again based on
pseudopetiole, cephalia, berries, stigma and auricle. However, some
specimens have possibilities to put under 2 section. like F. archboldiana
Other taxonomy problems
are
sinonim name of Freycinetia.
Kanehira and
Hatusima in 1940 mentioned F. inouei as a new species from Dalman in Nabire, West
New Guinea. They mentioned that the species have closely related to F. archboldiana but
different species. Stone in 1972 put the F. inouei as sinonim of F. archboldiana.
Some botanist like Martelli in 1910; Merril & Perry in 1940; Kanehira in 1941 and
Stone in 1967 said that F.angustissima were sinonim to F. stenophylla Warb. and F.
polyclada Merr.& Perry. But, Huynh in 1999 mentioned the three species above were
different species. He used both morphological and anatomical characters. Huynh said
that in morphological characters F. angustissima and F. polyclada were showed their
differences. He also said that the stigma of F. angustisimma is 2 0r 3 rarely 1 or 4 but
stigma and areola is distinctly girdled and F. stenophylla has 2 stigmas with sitgma and
areola is un-distinctly girdled. Actually, our preliminary study showed that stigma and
areola in west New Guinean Freycinetia have many variation and many specimens with
unidentified label have the variation.
Solving all problems above need taxonomy work also phylogenetic analysis to make
sure about the sections. In Freycinetia
species, phylogenetic analysis even used
3
morphological approach never been done. Cox already made cladistic analysis based on
morphological approached but only in generic level. Further, Callmander et. al (2003)
produced phylogenetic trees based on cp DNA data on generic levels too.
Some Freycinetia species without flowers and good auricle looks similar like F.
palida and closely unknown species. For this species information from molecular data
will guide to understanding what species is, is it same or different species. However, for
build a good phylogenetic classification it need comprehensive approached.
According to FWI (Forest Watch Indonesia) report in 2004, deforestation happen in
Indonesia, with acceleration of degradation 3 times foot ball area in every minutes.
Consequently
some
species include Freycinetia
may
disappeared
without ever
identified .
Freycinetia have much utilities . All parts of this plant could be used, for example
hanging root for nail; stem for lance, traditional rain clothes; leaves for roof, and
handicraft like basket and bag, also for perfume; spadix and cephalia as foods; and
bracts for cooling drink and juice. To get more benefit for human life, more researches
need start from basic study. Therefore the taxonomic research were done using main
morphological information and molecular data as a second approach.
Purposed of the Study
The purposed of the West New Guinean Freycinetia study were:
1. To know the diversity of Freycinetia in West New Guinea
2. To obtain delimitation of the genus and species concepts
3. To build phylogenetic classification of Freycinetia based on morphological and
molecular data.
4. To know distribution of West New Guinea Freycinetia
4
I. INTRODUCTION
The Pandanaceae has four genera namely Pandanus, Freycinetia, Sararanga and
Martelidendron. They all occur in the old tropicals, from Africa to India, Indo-China to
Japan, Malaysia to Sumatera until New Guinea area include some islands surroundings
like Salomon islands and also present in Australia. Two genera namely . Martelidendron
and Sararanga have limited in both species number and distribution. Martelidendro has
6 species found only in Madagaskar and some surrounding is