Daemonorops fissa Complex in West Malesia
DAEMONOROPS FISSA COMPLEX IN WEST MALESIA
DWI PUTRI RAMADHANI
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015
STATEMENT OF RESEARCH ORIGINALITY AND
INFORMATION SOURCE
This is to verify that my thesis entitled: Daemonorops fissa complex in
West Malesia is my own work and never been submitted to any institution before.
All the incorporated data and information are valid and stated clearly in the text,
and listed in the references.
Bogor, May 2015
Dwi Putri Ramadhani
NIM G353120231
SUMMARY
DWI PUTRI RAMADHANI. Daemonorops fissa complex in West Malesia.
Supervised by TATIK CHIKMAWATI and HIMMAH RUSTIAMI.
Daemonorops fissa complex is highly varied in West Malesia, but
information about this complex is limited. The objective of the research was to
reveal species diversity of Daemonorops fissa complex in West Malesia based on
morphological and anatomical characters.
Evaluation of morphological characters was conducted in Herbarium
Bogoriense (BO) Cibinong Science Centre LIPI, Bogor (West Java). A total of
355 herbarium specimens were observed in this study representing all species of
Daemonorops fissa complex. This study explored Daemonorops fissa complex,
including the following: 1) species diversity, (2) distribution, (3) morphological
variation, (4) anatomical variation and (5) phenetic relationships. Data obtained
then was analized descriptively and phenetically.
Species of Daemonorops fissa complex in West Malesia share some
unique characters such as erect peduncle (to ± 2 cm), alternate leaves, length of
involucrum 1 cm, globose fruits, and ruminate endosperm. This complex species
consists of eight species namely, D. angustifolia, D. calicarpa, D. fissa, D.
grandis, D. lewisiana, D. melanochaetes, D. monticola, D. stenophylla.
Daemonorops binnendijkii, D.palembanica and D. trichroa proposed as new
synonymies. Daemonorops binnendijkii and D. trichroa are new synonyms to D.
angustifolia, and D. palembanica is a new synonym to D. melanochaetes. All
species found in lowland rain forest, especially along stream and other wet places
or more open areas, swampy area, ridgetop hill Dipterocarp forest, bamboo forest,
and river side on altitude from 250 m – 1900 m asl.
Morphological variation of Daemonorops fissa complex in West Malesia
could be differentiated into 20 morphological characters. Cluster analysis
separated all species into two main groups with coefficient similarity of 36%.
Group I consisted of D. angustifolia, D. fissa, D. grandis, D. melanochaetes, D.
monticola and D. stenophylla, while group II consisted of D. calicarpa and D.
lewisiana. They were separated from five characters, total height, number of
leaflets, width of leaflets, type of inflorescence and type of spine in leafsheath.
Leaf anatomical characters of paradermal section could not be used to
separate among species within D. fissa complex, but leaf anatomical characters of
transverse section could be used to identify among species. All examined species
of D. fissa complex have three similar characters, tetracytyc stomata, cryptopor
type with sunken position and cell wall of sinous, but they have different pattern
of bulliform cell and palisade tissues. Both characters are considered as two
valuable new characters for supporting morphological data for species
delimitation.
Keywords: anatomy, Daemonorops fissa complex, morphology, West Malesia
RINGKASAN
DWI PUTRI RAMADHANI. Daemonorops fissa kompleks di Kawasan Malesia
Barat. Dibimbing oleh TATIK CHIKMAWATI dan HIMMAH RUSTIAMI.
Daemonorops fissa kompleks memiliki keanekaragaman yang tinggi di
Malesia Barat, tetapi informasi tentang kompleks ini sangat terbatas. Penelitian ini
mempelajari keanekaragaman jenis Daemonorops fissa kompleks di kawasan
Malesia Barat berdasarkan karakter morfologi dan anatomi.
Evaluasi karakter morfologi dilakukan di Herbarium Bogoriense, Pusat
Penelitian Biologi, LIPI dengan mengamati 355 lembar spesimen herbarium yang
mewakili semua jenis Daemonorops fissa kompleks. Kajian yang dilakukan
meliputi 1) keanekaragaman jenis, (2) persebaran, (3) variasi morfologi, (4)
variasi anatomi dan (5) kekerabatan fenetik. Analisis data meliputi analisis
deskriptif dan analisis fenetik.
Jenis-jenis dari Daemonorops fissa kompleks di kawasan Malesia Barat
memiliki karakter bersama yaitu tanpa tangkai bunga atau memiliki tangkai bunga
sampai ± 2 cm, susunan anak daun berseling, panjang daun pembalut 1 cm,
bentuk buah bulat dan bentuk endosperma termamah. Daemonorops fissa
kompleks di kawasan Malesia Barat terdiri dari delapan jenis yaitu D.
angustifolia, D. calicarpa, D. fissa, D. grandis, D. lewisiana, D. melanochaetes,
D. monticola, D. stenophylla. Daemonorops binnendijkii, D.palembanica dan D.
trichroa diusulkan sebagai sinonim. Daemonorops binnendijkii dan D. trichroa
adalah sinonim dari D. angustifolia, sedangkan D. palembanica adalah sinonim
dari D. melanochaetes. Semua jenis ditemukan di berbagai habitat seperti hutan
hujan tropis, tempat terbuka, hutan Dipterocarpaceae, hutan primer, hutan bambu
dan sepanjang aliran sungai dengan ketinggian sekitar 250 m sampai 1900 mdpl.
Variasi morfologi Daemonorops fissa kompleks dapat dibedakan ke dalam
20 karakter yang digunakan untuk menyusun deskripsi dan kunci identifikasi.
Analisis gugus memisahkan semua jenis ke dalam dua kelompok utama pada
koefisien keserupaan 36 %. Kelompok I terdiri dari D. angustifolia, D. fissa, D.
grandis, D. melanochaetes, D. monticola dan D. stenophylla. Kelompok II terdiri
dari D. calicarpa dan D. lewisiana. Kedua kelompok ini memisahkan berdasarkan
karakter tinggi total, jumlah anak daun, lebar anak daun, tipe perbungaan dan
bentuk duri pada batang.
Karakter anatomi daun D. fissa kompleks dari irisan paradermal tidak dapat
digunakan sebagai pembeda jenis dalam D. fissa kompleks, tetapi irisan melintang
dapat digunakan sebagai pembeda jenis. Semua jenis D. fissa kompleks yang
diamati memiliki tiga karakter yang sama, bentuk stomata tetrasitik, stomata
tenggelam dan tepi sel epidermis bergelombang, tetapi memiliki bentuk sel kipas
dan jaringan palisade yang berbeda. Kedua karakter tersebut dijadikan sebagai
karakter baru untuk menguatkan karakter morfologi sebagai batasan jenis D. fissa
kompleks.
Kata kunci: anatomi, Daemonorops fissa kompleks, morfologi, Malesia Barat
Copyright@ 2015, Bogor Agricultural University
All Right Reserved
It is prohibited to cite all or part of this thesis without referring to and
mentioning the source. Citation only permitted for the sake of education,
research, scientific writing, report writing, critical writing or reviewing
scientific problems. Citation does not inflict the name and honour of Bogor
Agricultural University.
It is prohibited to republish and reproduce all part of this thesis without
copyright permission from Bogor Agricultural University.
DAEMONOROPS FISSA COMPLEX IN WEST MALESIA
DWI PUTRI RAMADHANI
A thesis submitted to fulfill one of the requirements for the
Master Degree in Plant Biology Graduate Program
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015
Examiner of the examination:
Dr. Rugayah, M.Sc.
Herbarium Bogoriense, Botany Division,
Research Center of Biology – Indonesia Institute of Science,
Cibinong Science Center, Cibinong
Thesis Title : Daemonorops fissa Complex in West Malesia
Name
: Dwi Putri Ramadhani
NIM
: G353120231
Certified by
Supervisor Commitee
Dr Ir Tatik Chikmawati, MSi
Chairman
Dr Himmah Rustiami, SP, MSc
Member
Approved by
The Head of Plant Biology Graduate Program
The Dean of Postgraduate School
Dr Ir Miftahudin, MSi
Dr Ir Dahrul Syah, MScAgr
mExamination date: 6 February 2015
Graduation Date
PREFACE
I am grateful to my supervisors, Dr. Tatik Chikmawati M.Si., and Dr. Himmah
Rustiami, SP, M.Sc from Herbarium Bogoriense (BO) LIPI for their advice, kindness,
patience and valuable discussion throughout this study.
Many people have helped me through good advice, suggestions and supports.
In particular, I would like to thank Mahya, Gilang, Zulfan and Aini Qomariah for
their help during field, Dian and Hariri for their help in GIS Mapping for this study. I
would like to thank Dr. Sunaryo, Drs. Erlin Rachman, Eka Fatmawati T, S.Si., Ujang
Hapid, Widoyanti and Darius for their help and technical supports during my
anatomical study.
I would also like to thank all people in Herbarium Bogoriense for the
information and facilities when I carried out specimen examinations. I sincerely thank
all my friends of Plant Biology Graduate Program for their friendship.
I express deepest appreciation to Indonesian Government through DIKTI for
providing financial support for my research. Finally, my deepest appreciation is
also to my family, Erlis Suriani (mother) and Djulias Muhar (father), Sitha Muriani
(my sister) and Doni Tri Hariansyah (my brother) who has given me love and moral
supports. Undoubtedly, without them it is going to be impossible for me to finally
finish this study.
Bogor, May 2015
Dwi Putri Ramadhani
CONTENTS
LIST OF TABLES
LIST OF FIGURES
LIST OF APPENDIXES
1 INTRODUCTION
2 MATERIALS ANDMETHODS
Time and Place
Plant Materials
Research Methods
Data Analysis
3 RESULT AND DISCUSSION
Species Diversity
Distribution
Morphological Variations
Anatomical Variations
Anatomical Key to Daemonorops fissa complex in West Malesia
Phenetic Analysis
Taxonomic Treatment
Morphological Key to Daemonorops fissa complex in West Malesia
Species Description of Daemonorops fissa complex in West Malesia
4 CONCLUSION
REFERENCES
CURRICULUM VITAE
vi
vi
vii
1
3
3
3
3
6
6
6
7
8
14
16
17
19
19
20
35
35
38
LIST OF TABEL
1
2
3
Morphological characters of Daemonorops fissa complex in West
Malesia
Morphological variation of Daemonorops fissa complex in West
Malesia
Anatomical characters of Daemonorops fissa complex in West
Malesia
5
12
18
LIST OF FIGURES
1
2
3
4
5
6
7
8
9
10
11
12
13
Part of rattan. [A] Vegetative parts; [B] Generative parts. 1. cirrus, 2.
flagellum, 3. stem bearing a crownshaft, 4. stem bearing open
sheaths, 5. knee, 6. ocrea, 7. rachilla, 8. fruit
Distribution of Daemonorops fissa complex in West Malesia
D.angustifolia ( ), D.grandis ( ), D.calicarpa ( ),D.fissa (
D. lewisiana ( ), D. melanochaetes ( ),D. monticola ( ),
D. stenophylla ( )
Spine types on the leaf sheath of D. fissa complex [A] oblique circle;
[B] scattered
Type of knee Daemonorops fissa complex [A] bulgy; [B] elongatefolded; [C] elongate-flat; [D] elongate inflated.
Density of spine in prophyll of Daemonorops fissa complex
[A] velutinose with triangular spine; [B] velutinose with net-like
spine; [C] setose; [D] net-like spine; [E] triangular spine.
Flower of Daemonorops fissa complex. [A] flower of D.
melanochaetes; [B] Male flower of D. angustifolia; [C] Female
flower of D. melanochaetes; [D] Sterile male flower of D.
melanochaetes. [1] sterile male flower; [2] female flower; [3] male
flower; [4] calyx; [5] corolla; [6] androecium; [7] corolla; [8]
gynoecium; [9] corolla; [10] calyx; [11] empty anther.
Fruit scale surface of Daemonorops fissa complex [A] concave;
[B] flat
Seed surface of Daemonorops fissa complex [A] smooth; [B]
reticulate
Endosperm of Daemonorops fissa complex [A] slightly ruminate;
[B] deeply rumninate
Epidermal cell of Daemonorops [A] abaxial surface; [B] adaxial
surface. [1] stomata; [2] epidermis.
Transverse section of leaf, [1] epidermis; [2] sclerenchym; [3]
bulliform cell; [4] palisade; [5] sponge; [6] tannin; [7] phloem
strands; [8] xylem.
Position of bulliform cell Daemonorops fissa complex [A] between
two vascular bundle; [B] near midvein; [C] midvein.
Type of bulliform cell Daemonorops fissa complex [A] letter V; [B]
curved upward; [C] parallel; [D] curved downward.
4
7
8
9
10
11
11
11
11
14
14
15
16
14 Palisade tissue Daemonorops fissa complex [A] with one layer of
palisade tissue (dorsiventral); [B] with two layer of palisade tissue
(isobilateral). [1] upper palisade; [2] under palisade.
15 Dendogram of Daemonorops fissa complex in West Malesia based
on morphological characters
16 Daemonorops angustifolia (Griff.) Mart. [A] herbarium type
collection SING; [B] female and sterile male flower; [C] fruit; [D]
endosperm; [E] knee; [F] latex; [G] cane.
17 Daemonorops calicarpa (Griff.) Mart. [A] herbarium type collection
K; [B] spine oblique circle in leaf sheaths; [C] flower; [D]
endosperm. [1] sterile male flower; [2] female flower; [3] male
flower.
18 Daemonorops fissa Blume [A] herbarium collection BO; [B]
knee; [C] cirrus; [D] fruit; [E] flower; [F] endosperm. [1] sterile male
flower; [2] female flower; [3] male flower.
19 Daemonorops grandis (Griff.) Mart.[A] herbarium type collection
SING; [B] knee; [C] male flower; [D] endosperm. [1] corolla; [2]
stamen; [3] calyx.
20 Daemonorops lewisiana (Griff.) Mart. [A] herbarium type collection
K; [B] spine in leaf sheaths; [C] female and male flower; [D] seed;
[E] endosperm; [F] transversal veinlets.
21 Daemonorops melanochaetes Blume. [A] herbarium type collection
K; [B] knee; [C] prophyll; [D] flower; [E] endosperm. [1] sterile
male flower; [2] female flower; [3] male flower.
22 Daemonorops monticola (Grift) Mart., [A] herbarium type collection
K; [B] prophyll; [C] spine in leaf sheath; [D] male flower; [E]
flower; [F] endosperm. [1] sterile male flower; [2] female flower; [3]
male flower.
23 Daemonorops stenophylla Becc. [A] herbarium type collection K;
[B] knee; [C] prophyll; [D] fruit.
16
17
22
23
25
27
28
31
33
34
1 INTRODUCTION
Daemonorops is a rattan genus included in the subtribe Calamoideae. It is
the second largest rattan genus after Calamus, which was consisted of about 120
species (Rustiami 2011). The genus Daemonorops was first described by Blume
(1849), based on the type specimen Daemonorops melanochaetes. Daemonorops
was distributed from India and South China through the Malay Archipelago to
New Guinea where was represented by one species. Twenty-two species occur in
Southern Asia (Beccari 1911; Dransfield 1979, 2001; Evan et al. 2001; Hodel
1998). Daemonorops species are highly variable, but they are homoplasious in
many morphological characters (Umapathy et al. 2014).
The greatest morphological variation of Daemonorops was found in Malay
Peninsula, Sumatra and Borneo (Dransfield et al. 2008). West Malesia consisted
of Borneo, Sumatra, Malay Peninsula and Java (Raes & van Welzen 2009). High
diversity in West Malesia is supported by geological conditions and climate
history of the complex as well as the fact that this region is an archipelago with a
variety of obstacles. This condition opens up opportunities for the development of
its type in the isolated region. Rising sea levels during the Pleistocene as well as
the isolated islands causes the distribution patterns of plants including rattan is
unique in the West Malesia (Rustiami 2009).
The species of Daemonorops are mostly confined to primary tropical rain
forest on a great various soils, but some species had narrow ecological
requirements. A few species are a rather more weedy nature, found abundant in
forest habitats with high-light intensities such as riverbanks. One species in
Borneo, Daemonorops longispatha, grows on the landward margin of mangrove
forest. Some species are strictly montane, occurring at altitude up to ca. 2500 m
above sea level (m asl). Several species in Borneo are confined to heath forest, or
to limestone or serpentine rock (Dransfield et al. 2008).
Daemonorops spesies have been recognized having many uses.
Daemonorops melanochaetes is used as raw material for the manufacture
of tables, chairs and household crafts, and ripe fruit is edible (Harada et
al. 2005), D. draco for jernang sap seekers and cane can be used to make
household equipments (Sulasmi et al. 2012). In Jambi, umbut (young
shoots) of D. angustifolia is used for ulcer drugs by Suku Anak Dalam
(Jumiati et al. 2012).
The genus is classified into two sections based on its inflorescence
structure, section Daemonorops and Piptospatha (Beccari 1911; Rustiami et al.
2014). Species within section Daemonorops have concave boat-shaped bracts,
which at anthesis are completely enclosed by the prophyll and split longitudinally
to expose their flowers. All bracts are usually persistent. In contrast to section
Daemonorops, species within section Piptospatha have bracts splitting to the base
and only the lower part of inflorescence rachis is enclosed by the prophyll, and the
subsequent rachis bracts are usually deciduous (Dransfield et al. 2008, Rustiami et
al. 2011, Rustiami 2014).
2
Daemonorops fissa complex was classified into the section Daemonorops
with the sheath upright and directly attaching to the stem by peduncle length of ±
2 cm (pers. com 2013). Based on observed specimen in herbarium and compared
to the previous studies, species concept of Daemonorops fissa complex remains
unclear. Daemonorops fissa complex in West Malesia consists of 9 species
namely D. angustifolia, D. binnendijkii, D. fissa, D.grandis, D. melanochaetes, D.
palembanica, D. sepal, D. stenophylla, and D. trichroa (Beccari 1911). The later
study mentioned only 3 species in Malaya, namely D. angustifolia, D. grandis and
D. melanochaetes (Furtado 1953). However, the last study reported that 5 species
were recognized, D. angustifolia, D. fissa, D. grandis, D. melanochaetes and D.
sepal in Malay Peninsula, Sabah and Sarawak (Dransfield 1979; 1984, 1992).
Rattan classification was mainly based on discontinued of morphological
characters, the basic data for separating among species (Rustiami 2009). However,
morphological characters sometimes have disadvantage where placement of a
taxon in the classification is doubtful (Tellu 2005), therefore another approach is
needed, such as anatomy. Because of uncertain in distinguishing each species
within D. fissa complex, this species complex in this region needs to be reexamined to obtain clear species delimitation, and to describe the relationships
among species within this complex using phenetic approach, based on
morphological and anatomical characters.
Little work has been done on the leaf anatomy of Daemonorops. Anatomy
of Daemonorops can be distinguished from Calamus based on three characters:
guard cells, the presence of longer sklereid cells and commissure, and large cells
contained of tannins (Tomlinson 1961). Another study found that Daemonorops
spp. from Malaya can be distinguished from their sclereid characters (Tomlinson
2006).
The objective of this study was to provide the newest information of
species concept in D. fissa complex, including 1) species diversity, (2)
distribution, (3) morphological variation, (4) anatomical variation and (5) phenetic
relationships.
3
2 MATERIALS AND METHODS
Time and Place
This study was conducted in August 2013 until October 2014 starting with
field exploration in three locations in North Sumatra; Sibolangit, Mount Sinabung
and Deleng Lancuk. Observation of herbarium specimens was done in the
Herbarium Bogoriense (BO) Cibinong Science Centre LIPI, Bogor (West Java).
Plant Materials
Plant materials used to provide morphological data for the present study
were herbarium specimens including preserved material deposited at Herbarium
Bogoriense. All plant materials were represented the species of Daemonorops
fissa complex such as D. angustifolia, D. binnendijkii, D. callicarpa, D. fissa, D.
grandis, D. melanochaetes, D. monticola. D. palembanica, D. trichroa, and D.
stenophylla. A total of 355 herbarium specimens were observed for morphological
examination in this study.
Research Methods
Procedure for observation in this study followed the method described by
Rifai (2012), and de Vogel (1987). The procedure included 14 steps: (1) Selection
of taxon and determination of scope in the study and work to carried out; (2)
Collecting the material as much as possible (3) Reviewing literatures and records
of all the scientific names and data that included in the important taxa; (4)
Selecting the specimens that have been collected in taxonomic units based on
observable characteters; (5) Scrutinizing the available specimens based on
morphological approach; (6) Testing the characters already used by previous
researches, and performing cross check for justifying whether the existing
correlation between various characters could be confirmed or not; (7) Delimiting
the taxa in question according to the study results and resting the plausibility of
newly discovered characters; (8) Searching and determining the relationships
among taxa; (9) Solving the problem concerned to nomenclature; (10)
Constructing a key for identification of acceptable taxa; (11) Labeling on each
specimen examined according to the name of the concluded study; (12) Making
description of each taxa and note for individual specimens analyzed such as
ecology data and distribution; (13) Making draws of organ parts for publication as
needed; (14) Compiling a scientific report to be published.
Field Sampling
Field survey had been done in three areas of North Sumatera, Sibolangit,
Mount Sinabung and Deleng Lancuk, using exploratory method (Rugayah et al.
2004) which explored every location representing each type of ecosystem or
vegetation.
4
Specimen collection followed standard procedures developed by Dransfield
(1986). Data and information recorded from the field included: general data
(location, habitat, height, use of species, and date of collection); habit (cluster/
solitary), stem (total height, with or without leaf sheath diameter, length and
internodes, colour), leaves (number of leaflets, leaflets arrangement, the length
and width of leaves), additional organ (cirrus); inflorescences (length, number,
colour of rachilla), flower (colour, length and widths) ; fruit and seeds. The plants
were then documented, collected and processed for herbarium specimens.
Preparation of herbarium specimens followed standard procedure by
Dransfield (1986). The collection of complete rattan specimens has many
representative characters including:
1- Vegetative parts: Leaf sheath has a knee, ocrea, spine and sometimes a
flagellum (Figure 1A). Leaves of many species are long and large, collected only
a portion of the leaf base, middle section and apex. If the leaf ends with cirrus,
collected the leaf apex with cirrus. If the leaflet is large which does not fit to the
folder, cutting the leaflets off one side (leaving the leaflet bases in place) and
the other side is folded to fit the folder.
A
B
8
2
7
4
1
6
5
3
Figure 1 Part of rattan. [A] Vegetative parts. [B] Generative parts. 1. cirrus, 2.
flagellum, 3. stem bearing a crownshaft, 4. stem bearing open sheaths,
5. knee, 6. ocrea, 7. rachilla, 8. fruit
2. Generative parts: Many rattan species have large and long inflorescence
which could not be collected as a whole specimen (Figure 1B). To collect such
specimens, a part of the inflorescence base was taken, including primary axis,
bract, and the whole partial inflorescence were collected, and then collected the
last partial inflorescence to see the range sizes.
Anatomical Observation
The plant materials used in the present study were mostly dried and pressed
specimens. Samples were taken from the middle leaves located in the middle and
cut along 10cm2. Each species represents of one to two collection numbers.
5
Anatomical observations were prepared from paradermal and transverse leaf
sections. Leaf blade was boiled in HNO3 to manufacture paradermal section then
stained with 1% safranin (Sass 1951). Observed characters included shape and
size epidermal cells and stomata. For preparing leaf transverse section, leaf were
cut using freezing microtome Yamato RV-240 as thin as 20-25 µm. Leaf slices
then were dripped with sodium hypochlorite to remove chlorophyll, then stained
with safranin and then added few drops of glycerin on slide. Observed character
included the number of palisade tissue, palisade thickness, the presence of tannin
cells and the type, number, location and length of bulliform cells. After obtaining
permanent slides, the slide were observed using Nikon Eclipse E100 and
documented using OptiLab Viewer 2.2.
Morphological Observation
Twenty observed morphological characters were described in Table 1.
Morphological character includes several characters from internodes, petiole,
spine, knee, leaflets, inflorescence, fruit and endosperm. (Beccari 1911, Furtado
1953, Dransfield 1979; 1984, 1992, Rustiami 2011).
Table 1 Morphological characters of Daemonorops fissa complex in West Malesia
No
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Characters
Total height (m)
Length of internodes (cm)
Length of petiole (cm)
Spine in leaf sheath scattered
Spine in leaf sheath oblique
circles
Position spine in petiole
Type of knee
State of character (score)
≤ 5 tall (0), 6 – 10 tall (1), ≥10 tall (2)
≤20 (0), 21-30 (1), >30 (2)
4 (0), ≤30 (1), > 30 (2)
absent (0), present (1)
absent (0), present (1)
only one surface (0), on both surface (1)
elongate-folded (0), elongate-flat (1),
elongate inflated (2). bulgy (3)
Number of leaflets (pairs)
30 (0), 50-80 (1), >80 (2)
Width of leaflets (cm)
≤ 1 (0), 1.5-2.5 (1), ≥ 3 (2)
2
Spine density in 2m leaf sheath ≤5 (0), 6-10 (1), ≥ 11 (2)
Spine on leaf sheath mouth
absent (0), present (1)
Type of spine in leaf sheath absent (0), present (1)
mouth net-like
Type of spine in leaf sheath absent (0), present (1)
mouth triangular
Type of inflorescence
pleonanthic (0), hapaxanthic (1)
Length of prophyll (cm)
≤ 30 (0), > 30 (1)
Surface of prophyll
setose (0), velutinose (1)
Length of peduncle (cm)
1 cm (0), > 1 cm (1)
Spine of peduncle
absent (0), present (1)
Surface of fruit
flat (0), concave (1)
Endosperm
slightly ruminate (0), deeply ruminate (1)
6
Data Analysis
A species cluster was constructed based on morphological characters
observed for each species. Coefficient of similarity was analyzed using Simple
Matching (SM). Clustering analysis was analyzed using unweighted pair-group
method with arithmetic average (UPGMA) method (Rohlf 2002). Analysis was
performed using NTSys version 2.02.
3 RESULT AND DISCUSSION
Species Diversity
Species of Daemonorops fissa complex in West Malesia share some
unique characters such as peduncle erect to ± 2 cm, leaves alternate, length of
involucrum 1 cm, fruit globose, and endosperm ruminate. This complex species
consist of eight species namely, D. angustifolia, D. callicarpa, D. fissa, D.
grandis, D. lewisiana, D. melanochaetes, D. monticola and D. stenophylla. Three
species are proposed as new synonym, D. binnendijkii and D. trichroa are
synonym to D. angustifolia, whereas D. palembanica as a synonym to D.
melanochaetes.
Daemonorops binnendijkii was named based on two types, cultivated plant
at Buitenzorg and fruits from the Utrecht Herbarium, but the collection number of
the specimen types were unknown (Beccari 1911). On September, 25 in 1914,
Grashoff collected a specimen named Daemonorops cf. binnendijkii in Blinjoe,
Bangka Island. This spesimen deposits in Herbarium Bogoriense as a specimen
type. Until now, D. binnendijkii was never collected by researches. Because of
lesser data could be collected, D. binnendijkii is rated as Data Deficient (Bachman
2013). Based on morphological observation of specimen herbarium, D.
binnendijkii is similar to D. angustifolia, and later study based on anatomical
characters showed that anatomical characters of D. binnendijkii is same to D.
angustifolia.
Fruit collection by Teysmann no. 3582 collected in Muara Dua,
Palembang is the type of Daemonorops trichroa, and it deposits in Herbarium
Bogoriense. The fruit is extremely similar to fruit of D. angustifolia from Malayan
Peninsula (Beccari 1911). Based on characters armature of the petiole, leaf sheath,
rachis and spathae, Furtado reduce D. trichroa Miq. as a synonym of
Daemonorops angustifolia (Furtado 1937).
Type of Daemonorops palembanica was introduced from Sumatra into
Botanic Garden of Buitenzorg, and planted there under the name of D. lewisiana.
Beccari (1911), considered D. palembanica as a variety of D. melanochaetes.
After critical observation of four collection numbers of D. palembanica from
Lampung and Palembang collected by Gusdorff, it is concluded that D.
palembanica is a synonym of D. melanochaetes based on their similarities in
characters, density spine in leaf sheath mouth, spine in prophyll velutinose and
deeply endosperm.
7
Distribution
The distribution of the Daemonorops fissa complex is given in Figure 2.
D. angustifolia, D. callicarpa, D. fissa, D. grandis, D. lewisiana, D. monticola
and D. melanochaetes are the most wide spread species, extending throughout in
Malay Peninsula, Sumatra, Java and Borneo. D. stenophylla is known only from
the forest of North Sumatra and west Sumatra. D. callicarpa in Sumatra found in
lowland Dipterocarp forest and growed with Calamus castaneus. D. fissa is a new
record from Sumatra. Dransfield (1979; 1984, 1992), has investigated this species
widespread throughout in Borneo and endemic. D. melanochaetes is widespread
in Java. Generally, the forest in Malay Peninsula and Sumatra are comparatively
rich in Daemonorops fissa complex population, each of them having 6 species.
Daemonorops fissa complex grew in lowland rain forest especially along
stream and other wet places or more open areas, swampy area, ridgetop hill
Dipterocarp forest, bamboo forest and river side on altitude from 70 m – 1900 m
sl. All species of Daemonorops fissa complex was found in lowland forest on
altitude 250 – 1900 m asl, but D. monticola was only found above 1000 m asl.
Malay Peninsula
Sumatra
Borneo
Java
Figure 2 Distribution of Daemonorops fissa complex in West Malesia
D.angustifolia ( ), D.grandis ( ), D.calicarpa ( ),D.fissa (
D. lewisiana ( ), D. melanochaetes ( ),D. monticola ( ),
D. stenophylla ( )
8
Morphological Variation
Habit
All species of Daeomonorops fissa complex from West Malesia are climb
clustering rattan. Three species, D. calicarpa, D. lewisiana and D. monticola are
found climbing up to 5 m high or more. Two other species, D. grandis and D.
stenophylla are found climbing to 10 m. Daemonorops angustifolia, D.
binnendijkii, D. fissa and D. melanochaetes are the highest climbers, up to 20 m
or higher.
Leaf sheath
Leaf sheath of Daemonorops fissa complex are varied in colour, spine
position and type. The colour of leaf sheath are dull greenish brown, pale green to
pale brownish, and brownish green. Spine position in leaf sheath are scattered or
oblique circle (Figure 3). Species with spine arranged in oblique circle are D.
calicarpa, D. lewisiana and D. monticola, whereas species with scattered spines
include D. angustifolia, D.fissa, D. grandis, D. melanochaetes and D. stenophylla.
Spine type are net-like spine to triangular spines. All species of D. fissa complex
have triangular spines, but only D. calicarpa has a net-like spine.
A
B
Figure 3 Spine types on the leaf sheath of D. fissa complex. [A] Oblique circle;
[B] scattered
Leaves
The colour of D. fissa complex leaves are dark green. Both upper and
under leaf surfaces are glabrous. The leaflets arrangement is pinnate with narrow
and elliptic shape, but leaflets of D. grandis is broad with lanceolate shape. The
number of leaflets on D. fissa complex is generally up to 50 – 90 pairs on each
side, but D. grandis has up to 30 pairs.
Cirrus
Cirrus is extending from leaf apex. It is varied in length. All species D.
fissa complex have cirrus length up to 100 cm – 125 cm, but D. calicarpa have
short cirrus, 30 cm.
Petiole
Cross section of D. fissa complex petiole are channeled, the petiole adaxial
surface is concave and ridged while the petiole abaxial surface convex.
Daemonorops stenophylla has shorter petiole (4 cm) than that of other species
9
(10-30 cm). Daemonorops angustifolia has the longest petiole length up to 35
cm.
Knee
Type of knee is an important character for identification rattan species.
Baja-lapis (2010) described 5 knee types, including bulgy, elongate-folded,
elongate-flat, elongate inflated and cylindrical with ant holes. Daemonorops
calicarpa, D. grandis and D. stenophylla have elongate-folded knees. D.
melanochaetes and D. monticola has elongate-flat knee. D. angustifolia has
elongate-inflated knee; whereas D. fissa and D. lewisiana have bulgy knee (Figure
4). Knees are covered with spine like the leaf sheath.
A
B
C
D
Figure 4 Type of knee Daemonorops fissa complex. [A] bulgy; [B] elongatefolded; [C] elongate-flat; [D] elongate inflated.
Prophyll
Daemonorops fissa complex has persistent prophyll. The prophylls are
covered many spines, but the density spine is varied. The spine density on
prophyll are rarely to densely spines. Species with rarely spines (setose species)
can be found on D. angustifolia, D. fissa, D. grandis, D. lewisiana, D. monticola,
and D. stenophylla (Figure 5). Two species, D. calicarpa and D. melanochaetes,
have densely spines (velutinose). There are two spine types among Daemonorops,
net-like spine and triangular spine. However, D. calicarpa and D. melanochaetes
have different types of spines, D. calicarpa with net-like spine and D.
melanochaetes with triangular spine. The prophyll of D. fissa complex are tubular
and up to 45 cm long, but D. calicarpa is scarcely longer than 25 cm.
A
B
C
D
E
Figure 5 Density of spine in prophyll of Daemonorops fissa complex.
[A] velutinose with triangular spine; [B] velutinose with netlike spine; [C] setose; [D] net-like spine; [E] triangular spine.
10
Inflorescence
Daemonorops is dioecious plants. There are two inflorescence types,
pleonanthic and hapaxanthic. Stems will die after flowering in hapaxhantic type.
Their inflorescence comprise of numerous lateral inflorescence arising from the
axils of, often markedly, reduced leaves on the upper part of stem. While in
pleonanthic type stem continues to grow even after flowering. Their inflorescence
grow from the lower part of stem which then is continue to grow vegetatively, and
reproduce over a relatively long period throughout its adult life (Baker et. al.
1999). All species of D. fissa complex have pleonanthic inflorescences, but D.
calicarpa and D. lewisiana have hapaxhantic inflorescence.
Flower
Male flowers are up to 6 mm long with corolla splitted to the base into 3
petals. The petals have lanceolate shape and rounded at the base with 4 mm long.
Corolla is twice longer than calyx. Calyx is 2 mm long. Each flower has 6 stamen,
and with anther 2 mm long, female rachilla bears many flowers in dyads consisted
of one female flower and one sterile male flower with empty anthers. Female
flowers are 4 mm long with ovoid, corolla consisted of 3 petals, corolla as long as
calyx. Calyx is truncate or shallowly 3 lobed. Sterile flower (4 mm long) is always
shorter than male flower with empty anthers (Figure 6).
B
A
6
5
2
1
3
C
4
D
7
8
9
11
10
Figure 6 Flower of Daemonorops fissa complex. [A] flower of D.
melanochaetes; [B] Male flower of D. angustifolia; [C] Female
flower of D. melanochaetes; [D] Sterile male flower of D.
melanochaetes. [1] sterile male flower; [2] female flower; [3] male
flower; [4] calyx; [5] corolla; [6] androecium; [7] corolla; [8]
gynoecium; [9] corolla; [10] calyx; [11] empty anther.
Fruit
All species of Daemonorops fissa complex have globose fruit, but their
fruits are differed in fruit scale surface. Only Daemonorops monticola has
11
concave scale surface, the other species have flat scale surface (Figure 7). The
scale of D. fissa complex are arranged in 15 – 18 vertical rows, but the scale of D.
grandis from Aceh is arranged in 21 vertical rows.
A
B
A
Figure 7 Fruit scale surface of Daemonorops fissa complex. [A] concave; [B] flat
Seed
The seed of nearly all rattan has an outer fleshy layer. This fleshy layer is
the outer seed-coat and hence is properly called sacrotesta. sarcotesta varies
greatly in thickness from species to species (Dransfield 1979). All species of
Daemonorops fissa complex have smooth seed surface, but there are two
collections, D. grandis from Mount Kemiri, Aceh and D. angustifolia from
Deleng Lancuk, Sumatra, having reticulate seed surfaces (Figure 8).
A
B
Figure 8 Seed surface of Daemonorops fissa complex. [A] smooth; [B]
reticulate
Endosperm
Sarcotesta is easily separable from the rest of the seed. Endosperm varies
from homogeneous in most genera to shallowly or deeply ruminate in
Daemonorops. A previous study showed that all Daemonorops species have
ruminate endosperm (Dransfield 2008), but endosperms of Daemonorops fissa
complex from West Malesia are varied from slightly ruminate (D. calicarpa, D.
fissa, D. lewisiana and D. monticola) to deeply ruminate endosperm (D.
angustifolia, D. grandis, D. melanochaetes and D. stenophylla) (Figure 9).
A
B
Figure 9 Endosperm of Daemonorops fissa complex. [A] slightly ruminate; [B]
deeply ruminate
12
Table 2. Morphological variation of Daemonorops fissa complex in West Malesia
No
N
Character
Species
1
2
3
4
5
6
7
8
1
Total height (m)
30
6
30
10
7
30
10
8
2
Length of
internodes (cm)
Length of petiole
(cm)
Spine on leaf
sheath scattered
Spine in leaf
sheath
oblique
circle
Position spine in
petiole
Type of knee
35
20
20
35
20
22
30
14
30
15
40
40
15
20
30
4
scattered
oblique
circle
present
scattered
scattered
scattered
absent
oblique
circle
present
scattered
absent
oblique
circle
present
only one
surface
elongatefolded
45
both
surface
bulgy
only one
surface
bulgy
both surface
elongate-flat
only one
surface
elongate-flat
50
both
surface
elongatefolded
30
80
90
65
both
surface
elongatefolded
80
1
1.8
3
1.7
2
1.5
0.8
10
8
5
4
20
10
12
3
4
5
6
7
8
9
10
absent
both surface
elongateinflated
50
Number of
leaflets (pairs)
Widht of leaflets
1.5
(cm)
Spine density in 12
2m2 leaf sheath
absent
absent
11
Spine on leaf present
sheath mouth
present
absent
absent
absent
present
present
absent
12
Type of spine in
leaf sheath mouth
net-like
present
absent
absent
absent
absent
absent
absent
absent
13
Table 2 continued
No
13
14
15
16
17
18
19
20
N
Character
Type of spine in
leaf sheath
mouth triangular
Type of
inflorescence
Length of
prophyll (cm)
Surface of
prophyll
Length of
peduncle (cm)
Spine
of
peduncle
Surface of fruit
Endosperm
Spesies
1
2
3
present
absent
present
pleonanthic
39
hapaxanthi
c
23
setose
5
6
7
8
present
present
present
present
present
pleonanthic
pleonanthic
hapaxanthic
pleonanthic
pleonanthic
pleonanthic
35
40
30
55
40
32
velutinose
setose
setose
setose
velutinose
setose
setose
2
2
1
1
2
2
1.5
1
present
present
present
present
absent
present
present
absent
flat
deeply
ruminate
flat
slightly
ruminate
flat
slightly
ruminate
flat
deeply
ruminate
flat
slightly
ruminate
flat
deeply
ruminate
concave
slightly
ruminate
flat
deeply
ruminate
Labels of individu in the table above are:
1. D. angustifolia
5. D. lewisiana
2. D. callicarpa
6. D. melanochaetes
3. D. fissa
7. D. monticola
4. D. grandis
8. D. stenophylla
4
14
Anatomical Variation
Stomata type of Daemonorops fissa complex on paradermal section have
been studied. The stomata types of all species are tetracytic (Figure 10). The type
has two guard cells surrounded by four subsidiary cells (or cells neighbouring of
the guard cells), two lateral and two polar ones (Metcalfe 1961, Dickison 1999).
Stomata are infrequent in adaxial surface, and are restricted to intercostals region
in abaxia surface. Terminal subsidiary cells are not well differentiated, sometimes
having short size. Almost all stomata are present on abaxial leaf surface, but they
are only fewer in adaxial surface.
A
B
1
100 µm
100 µm
2
Figure 10 Epidermal cell of Daemonorops. [A] abaxial surface; [B] adaxial
surface, 1 stomata, 2 epidermis
Adaxial epidermis is always uniform, having cell rectangular with cell walls
markedly sinuous but they do not have thickening anticlinal walls. Tellu (2006)
showed that the stomata of Daemonorops species have kriptopor type with sunken
position in the epidermis or sub-epidermis. In contrast to stomatal type, stomatal
size shows variation within Daemonorops fissa complex. Daemonorops
stenophylla has the smallest stomata size, 135.25-193.22 µm2, whereas D.
monticola has the largest stomata size, 270.51-318.81 µm2.
1
2
4
3
5
8
6
7
100 µm
Figure 11 Transverse section of leaf Daemonorops. [1] epidermis; [2]
sclerenchym; [3] bulliform cell; [4] palisade; [5] sponge; [6] tannin;
[7] phloem strands; [7] xylem.
15
Bulliform cells are commonly found in Monocotyledon, which have
function for storing water. Bulliform, so called motor cells, are lose turgory under
water deficit conditions, and thus constrict in upon themselves, causing lamina to
fold or roll inward edge to edge (Dickison 1999).
The bulliform cells of Daemonorops fissa complex have thin walls, and
their size are larger than that of the adjacent epidermal cells. There are usually 617 bulliform cells on the adaxial leaf surfaces. From the middle to the margin leaf,
the bulliform cells on the abaxial epidermis are gradually getting smaller, and
becoming the same size to the abaxial epidermal cells. The position of bulliform
cell is varied, in adaxial and abaxial epidermis, between two vascular bundles,
near midvein and mid-vein (Figure 12).
B
A
C
Figure 12 Position of bulliform cell in Daemonorops fissa complex. [A] between
two vascular bundle; [B] near midvein; [C] midvein. Bulliform cell
pointed by arrow.
All eight species of Daemonorops fissa complex have many bulliform cells
on the upper and lower epidermis. The shape of bulliform cells on the upper
epidermis are varied. Bulliform cells were curved downward found in D.
angustifolia, D. binnendijkii, and D. calicarpa. Daemonorops fissa, D. lewisiana,
D. monticola and D. stenophylla have bulliform cells parallel to epidermis, D.
grandis has V shape, whereas the bulliform cells of D. melanochaetes are curved
upward (Figure 13).
The mesophylls of Daemonorops fissa complex are differentiated into
palisade and sponge tissues. Mesophylls usually consist of one layer of palisade
tissue and three to five layers of sponge tissue. Daemonorops melanochaetes and
D. fissa only have one palisade tissue in upper side (dorsiventral), but D.
calicarpa, D. lewisiana, D. grandis, D. melanochaetes and D. stenophylla have
two palisade tissues (isobilateral) (Figure 14).
16
A
B
C
D
Figure 13 Type of bulliform cell in Daemonorops fissa complex. [A] letter V; [B]
curved upward; [C] parallel to epidermis; [D] curved downward
A
B
1
1
2
Figure 14 Palisade tissue in Daemonorops fissa complex. [A] with one layer of
palisade tissue (dorsiventral); [B] with two layer of palisade tissue
(isobilateral); 1. upper palisade, 2. under palisade.
Anatomical key to Daemonorops fissa complex in West Malesia
1. a.
b.
2. a.
b.
3. a.
b.
4. a.
b.
5. a.
b.
6. a.
b.
7. a.
b.
Stomata size ≤ 193.22 µm2 …………...…………………D. stenophylla
Stomata size > 193.22 µm2…………………………………………....2
Number of tannin sac 3 – 5……………………………………...…….3
Number of tannin sac 6 – 8………………………………..D. monticola
Have one type of bulliform cell......................................................4
Have two type of bulliform cell ......………………………...D. grandis
Number of bulliform cell in upper epidermis ≤ 10……………………5
Number of bulliform cell in upper epidermis 12 – 17….....D. lewisiana
Leaves isobilateral type…………………………………….......D. fissa
Leaves dorsiventral type………………………………………............6
Length of upper palisade 20.33 µm2 .................................D. calicarpa
Length of upper palisade > 23.56 µm2.............................................7
Type of bulliform cell curved upward.......................D. melanochaetes
Type of bulliform cell curved downward......................D. angustifolia
17
Phenetic Analysis
In this study, 20 characters, mostly qualitative characters were selected for
phenetic analysis (Table 1). A data matrix of characters was scored according to
the species descriptions.
Cluster analysis separated all samples into two major groups with
similarity coefficient of 0.36 (Figure 15). Group I are composed of 6 species
which are D. angustifolia, D. fissa, D. grandis, D. melanochaetes, D. monticola
and D. stenophylla. Grup II contained, D. lewisiana and D. callicarpa. They were
separated from six characters, total height, number of leaflets, width of leaflets,
type of spine in leafsheath, position spine in petiole and type of inflorescence.
Based on the similarity coefficient, Daemonorops angustifolia, D.
binnendijkii and D. trichroa are identic with similarity coefficient of 100%.
Therefore, it is concluded that D. binnendijkii and D. trichroa are synonym of D.
angustifolia. D. palembanica is a synonym of D. melanochaetes based on three
characters: density spine in leaf sheath mouth, spine in prophyll velutinose and
deeply endosperm. Grup I grouped by spine in leafsheath scattered, position spine
in petiole only one surface and type of inflorescence is pleonanthic. D. lewisiana
and D. callicarpa grouped by total height ≤ 5 m tall, spine in leafsheath oblique
circles and type of inflorescence hapaxanthic.
Strong character in taxonomic research is a character that only belongs to
one group of taxa that can be used to differentiate it to other taxa (Wiley 1981).
In this research, the type of knees, spines and width of leaflets are considered as
strong characters. Type of knee is an important character for identification of
rattan (Baja-lapis 2010). Spine and width of leaflets can be used as a diagnostic
character to species. Based on dendogram, width of leaflets is a diagnostic
characters to separated D. grandis from others. Previous study also showed that
leaflet size is a diagnostic character (Dransfield 1976).
D angustifolia1
D binnendijkii1
D binnendijkii2
D angustifolia2
D angustifolia4
D trichroa
D angustifolia3
D melanochaetes1
D melanochaetes3
D melanochaetes2
D palembanica
D fissa1
D fissa2
D monticola1
D monticola2
D grandis1
D grandis2
D stenophylla
D lewisiana1
D lewisiana3
D lewisiana2
D calicarpa1
D calicarpa2
MW
I
II
0.36
0.52
0.68
0.84
1.00
Coefficient similarity
Figure 15 Dendogram of Daemonorops fissa complex in West Malesia based on
morphological characters
18
Table 3. Anatomical characters of Daemonorops fissa complex in West Malesia
Spesies
Character
D.angustifolia
D.calicarpa
D.fissa
D.grandis
D.lewisiana
D.melanochaetes
D.monticola
D.stenophylla
Shape of epidermis
Rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
Epidermis size (µm2)
20.00-36.75
23.00-39.68
30.00-74.40
24.58-50.00
23.00-39.68
20.00-36.75
24.58-50.00
30.00-74.40
Type of stomata
Tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
Stomata size (µm2)
241.53-265.68
217.37318.81
palisade
217.37241.53
palisade
palisade
270.51318.81
palisade
135.25-193.22
Palisade
217.37265.68
palisade
154.58-241.53
Position sclerenchym
169.07217.37
palisade
Palisade in lower
epidermis
Number of tannin
cell
Arrangement of
bulliform cell in
upper epidermis
Number of bulliform
cell in upper
epidermis
Number of bulliform
cell in lower
epidermis
Bulliform cell size
(µm)
Present
present
absent
present
present
absent
absent
absent
3-5
3-6
5-7
5-7
4-7
3-5
6-8
3-5
curved
downward
curved
downward
parallel
letter v
parallel
curved upward
parallel
parallel
8-9
7
8-10
5-8
12-17
6
10
5-8
8-9
9
8
14
5-7
6
10
8-12
37.20-44.64
24.80
32.24-49.60
24.80-49.60
19.84-49.60
37.20-49.60
29.76-37.20
24.80-32.24
tetracytic
palisade
19
Taxonomic Treatment
Daemonorops
Daemonorops Blume in J.A. & J.H. Schultes, Syst. Veg. 7(2):1333 (1830).
– Type: Daemonorops melanochaetes Blume. West Sumatra, Padang near Ayer
Manchor, August 1878. Beccari 831 (isotype K)
Solitary or clustering rattans, acaulescent to high climbing hapaxanthic (then
always very short stemmed) or pleonanthic, dioecious. Leaf sheaths heavily armed
with spines, spines frequently highly organized. Flagellum absent. Knee
frequently present. Leaves ecirrate in acaulescent species or long cirrate. Leaflets
variously arranged. Inflorescence male and female superficially similar, but within
the genus of two basic types: one with all bracts enclosed within the outermost
bract or prophyll, splitting along their length to expose the flowers (section
Cymbospatha) or the other with bracts splitting along their entire length to leave
no tubular portion and frequently falling (section Piptospatha). Bracts variously
armed. In the section Piptospatha partial inflorescences longer than the
subtending bract; bracteoles and involucres inconspicuous. Male rachilla bearing
male solitary flowers; male flowers with small cup shaped, calyx with three small
lobes; corolla split to the base into 3 petals; stamen 6, slightly epipetalous;
pistillode minute. Sterile male flower found with each female flower, as the fertile
male, but stamens with empty anthers. Female rachilla bearing many flowers in
dyads consists of one female flower and one sterile male flower. Female flower
calyx truncate or shallowly 3 lobed; corolla with 3 petals; gynoecium with 3 stigmas
and with 3 loculs. Sterile flower smaller or at least more slender than the female, with
well formed calyx and corolla, 6 sterile stamens, an abortive ovary. Fruit variously
shaped, tipped with stigmatic remains and covered with reflexed scales. Seed only
one, covered by thin to thick sweet or sour sarcotesta. Endosperm deeply ruminate.
Embryo basal.
Distribution. Geographical distribution of Daemonorops is more restricted than the
genus Calamus
DWI PUTRI RAMADHANI
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015
STATEMENT OF RESEARCH ORIGINALITY AND
INFORMATION SOURCE
This is to verify that my thesis entitled: Daemonorops fissa complex in
West Malesia is my own work and never been submitted to any institution before.
All the incorporated data and information are valid and stated clearly in the text,
and listed in the references.
Bogor, May 2015
Dwi Putri Ramadhani
NIM G353120231
SUMMARY
DWI PUTRI RAMADHANI. Daemonorops fissa complex in West Malesia.
Supervised by TATIK CHIKMAWATI and HIMMAH RUSTIAMI.
Daemonorops fissa complex is highly varied in West Malesia, but
information about this complex is limited. The objective of the research was to
reveal species diversity of Daemonorops fissa complex in West Malesia based on
morphological and anatomical characters.
Evaluation of morphological characters was conducted in Herbarium
Bogoriense (BO) Cibinong Science Centre LIPI, Bogor (West Java). A total of
355 herbarium specimens were observed in this study representing all species of
Daemonorops fissa complex. This study explored Daemonorops fissa complex,
including the following: 1) species diversity, (2) distribution, (3) morphological
variation, (4) anatomical variation and (5) phenetic relationships. Data obtained
then was analized descriptively and phenetically.
Species of Daemonorops fissa complex in West Malesia share some
unique characters such as erect peduncle (to ± 2 cm), alternate leaves, length of
involucrum 1 cm, globose fruits, and ruminate endosperm. This complex species
consists of eight species namely, D. angustifolia, D. calicarpa, D. fissa, D.
grandis, D. lewisiana, D. melanochaetes, D. monticola, D. stenophylla.
Daemonorops binnendijkii, D.palembanica and D. trichroa proposed as new
synonymies. Daemonorops binnendijkii and D. trichroa are new synonyms to D.
angustifolia, and D. palembanica is a new synonym to D. melanochaetes. All
species found in lowland rain forest, especially along stream and other wet places
or more open areas, swampy area, ridgetop hill Dipterocarp forest, bamboo forest,
and river side on altitude from 250 m – 1900 m asl.
Morphological variation of Daemonorops fissa complex in West Malesia
could be differentiated into 20 morphological characters. Cluster analysis
separated all species into two main groups with coefficient similarity of 36%.
Group I consisted of D. angustifolia, D. fissa, D. grandis, D. melanochaetes, D.
monticola and D. stenophylla, while group II consisted of D. calicarpa and D.
lewisiana. They were separated from five characters, total height, number of
leaflets, width of leaflets, type of inflorescence and type of spine in leafsheath.
Leaf anatomical characters of paradermal section could not be used to
separate among species within D. fissa complex, but leaf anatomical characters of
transverse section could be used to identify among species. All examined species
of D. fissa complex have three similar characters, tetracytyc stomata, cryptopor
type with sunken position and cell wall of sinous, but they have different pattern
of bulliform cell and palisade tissues. Both characters are considered as two
valuable new characters for supporting morphological data for species
delimitation.
Keywords: anatomy, Daemonorops fissa complex, morphology, West Malesia
RINGKASAN
DWI PUTRI RAMADHANI. Daemonorops fissa kompleks di Kawasan Malesia
Barat. Dibimbing oleh TATIK CHIKMAWATI dan HIMMAH RUSTIAMI.
Daemonorops fissa kompleks memiliki keanekaragaman yang tinggi di
Malesia Barat, tetapi informasi tentang kompleks ini sangat terbatas. Penelitian ini
mempelajari keanekaragaman jenis Daemonorops fissa kompleks di kawasan
Malesia Barat berdasarkan karakter morfologi dan anatomi.
Evaluasi karakter morfologi dilakukan di Herbarium Bogoriense, Pusat
Penelitian Biologi, LIPI dengan mengamati 355 lembar spesimen herbarium yang
mewakili semua jenis Daemonorops fissa kompleks. Kajian yang dilakukan
meliputi 1) keanekaragaman jenis, (2) persebaran, (3) variasi morfologi, (4)
variasi anatomi dan (5) kekerabatan fenetik. Analisis data meliputi analisis
deskriptif dan analisis fenetik.
Jenis-jenis dari Daemonorops fissa kompleks di kawasan Malesia Barat
memiliki karakter bersama yaitu tanpa tangkai bunga atau memiliki tangkai bunga
sampai ± 2 cm, susunan anak daun berseling, panjang daun pembalut 1 cm,
bentuk buah bulat dan bentuk endosperma termamah. Daemonorops fissa
kompleks di kawasan Malesia Barat terdiri dari delapan jenis yaitu D.
angustifolia, D. calicarpa, D. fissa, D. grandis, D. lewisiana, D. melanochaetes,
D. monticola, D. stenophylla. Daemonorops binnendijkii, D.palembanica dan D.
trichroa diusulkan sebagai sinonim. Daemonorops binnendijkii dan D. trichroa
adalah sinonim dari D. angustifolia, sedangkan D. palembanica adalah sinonim
dari D. melanochaetes. Semua jenis ditemukan di berbagai habitat seperti hutan
hujan tropis, tempat terbuka, hutan Dipterocarpaceae, hutan primer, hutan bambu
dan sepanjang aliran sungai dengan ketinggian sekitar 250 m sampai 1900 mdpl.
Variasi morfologi Daemonorops fissa kompleks dapat dibedakan ke dalam
20 karakter yang digunakan untuk menyusun deskripsi dan kunci identifikasi.
Analisis gugus memisahkan semua jenis ke dalam dua kelompok utama pada
koefisien keserupaan 36 %. Kelompok I terdiri dari D. angustifolia, D. fissa, D.
grandis, D. melanochaetes, D. monticola dan D. stenophylla. Kelompok II terdiri
dari D. calicarpa dan D. lewisiana. Kedua kelompok ini memisahkan berdasarkan
karakter tinggi total, jumlah anak daun, lebar anak daun, tipe perbungaan dan
bentuk duri pada batang.
Karakter anatomi daun D. fissa kompleks dari irisan paradermal tidak dapat
digunakan sebagai pembeda jenis dalam D. fissa kompleks, tetapi irisan melintang
dapat digunakan sebagai pembeda jenis. Semua jenis D. fissa kompleks yang
diamati memiliki tiga karakter yang sama, bentuk stomata tetrasitik, stomata
tenggelam dan tepi sel epidermis bergelombang, tetapi memiliki bentuk sel kipas
dan jaringan palisade yang berbeda. Kedua karakter tersebut dijadikan sebagai
karakter baru untuk menguatkan karakter morfologi sebagai batasan jenis D. fissa
kompleks.
Kata kunci: anatomi, Daemonorops fissa kompleks, morfologi, Malesia Barat
Copyright@ 2015, Bogor Agricultural University
All Right Reserved
It is prohibited to cite all or part of this thesis without referring to and
mentioning the source. Citation only permitted for the sake of education,
research, scientific writing, report writing, critical writing or reviewing
scientific problems. Citation does not inflict the name and honour of Bogor
Agricultural University.
It is prohibited to republish and reproduce all part of this thesis without
copyright permission from Bogor Agricultural University.
DAEMONOROPS FISSA COMPLEX IN WEST MALESIA
DWI PUTRI RAMADHANI
A thesis submitted to fulfill one of the requirements for the
Master Degree in Plant Biology Graduate Program
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2015
Examiner of the examination:
Dr. Rugayah, M.Sc.
Herbarium Bogoriense, Botany Division,
Research Center of Biology – Indonesia Institute of Science,
Cibinong Science Center, Cibinong
Thesis Title : Daemonorops fissa Complex in West Malesia
Name
: Dwi Putri Ramadhani
NIM
: G353120231
Certified by
Supervisor Commitee
Dr Ir Tatik Chikmawati, MSi
Chairman
Dr Himmah Rustiami, SP, MSc
Member
Approved by
The Head of Plant Biology Graduate Program
The Dean of Postgraduate School
Dr Ir Miftahudin, MSi
Dr Ir Dahrul Syah, MScAgr
mExamination date: 6 February 2015
Graduation Date
PREFACE
I am grateful to my supervisors, Dr. Tatik Chikmawati M.Si., and Dr. Himmah
Rustiami, SP, M.Sc from Herbarium Bogoriense (BO) LIPI for their advice, kindness,
patience and valuable discussion throughout this study.
Many people have helped me through good advice, suggestions and supports.
In particular, I would like to thank Mahya, Gilang, Zulfan and Aini Qomariah for
their help during field, Dian and Hariri for their help in GIS Mapping for this study. I
would like to thank Dr. Sunaryo, Drs. Erlin Rachman, Eka Fatmawati T, S.Si., Ujang
Hapid, Widoyanti and Darius for their help and technical supports during my
anatomical study.
I would also like to thank all people in Herbarium Bogoriense for the
information and facilities when I carried out specimen examinations. I sincerely thank
all my friends of Plant Biology Graduate Program for their friendship.
I express deepest appreciation to Indonesian Government through DIKTI for
providing financial support for my research. Finally, my deepest appreciation is
also to my family, Erlis Suriani (mother) and Djulias Muhar (father), Sitha Muriani
(my sister) and Doni Tri Hariansyah (my brother) who has given me love and moral
supports. Undoubtedly, without them it is going to be impossible for me to finally
finish this study.
Bogor, May 2015
Dwi Putri Ramadhani
CONTENTS
LIST OF TABLES
LIST OF FIGURES
LIST OF APPENDIXES
1 INTRODUCTION
2 MATERIALS ANDMETHODS
Time and Place
Plant Materials
Research Methods
Data Analysis
3 RESULT AND DISCUSSION
Species Diversity
Distribution
Morphological Variations
Anatomical Variations
Anatomical Key to Daemonorops fissa complex in West Malesia
Phenetic Analysis
Taxonomic Treatment
Morphological Key to Daemonorops fissa complex in West Malesia
Species Description of Daemonorops fissa complex in West Malesia
4 CONCLUSION
REFERENCES
CURRICULUM VITAE
vi
vi
vii
1
3
3
3
3
6
6
6
7
8
14
16
17
19
19
20
35
35
38
LIST OF TABEL
1
2
3
Morphological characters of Daemonorops fissa complex in West
Malesia
Morphological variation of Daemonorops fissa complex in West
Malesia
Anatomical characters of Daemonorops fissa complex in West
Malesia
5
12
18
LIST OF FIGURES
1
2
3
4
5
6
7
8
9
10
11
12
13
Part of rattan. [A] Vegetative parts; [B] Generative parts. 1. cirrus, 2.
flagellum, 3. stem bearing a crownshaft, 4. stem bearing open
sheaths, 5. knee, 6. ocrea, 7. rachilla, 8. fruit
Distribution of Daemonorops fissa complex in West Malesia
D.angustifolia ( ), D.grandis ( ), D.calicarpa ( ),D.fissa (
D. lewisiana ( ), D. melanochaetes ( ),D. monticola ( ),
D. stenophylla ( )
Spine types on the leaf sheath of D. fissa complex [A] oblique circle;
[B] scattered
Type of knee Daemonorops fissa complex [A] bulgy; [B] elongatefolded; [C] elongate-flat; [D] elongate inflated.
Density of spine in prophyll of Daemonorops fissa complex
[A] velutinose with triangular spine; [B] velutinose with net-like
spine; [C] setose; [D] net-like spine; [E] triangular spine.
Flower of Daemonorops fissa complex. [A] flower of D.
melanochaetes; [B] Male flower of D. angustifolia; [C] Female
flower of D. melanochaetes; [D] Sterile male flower of D.
melanochaetes. [1] sterile male flower; [2] female flower; [3] male
flower; [4] calyx; [5] corolla; [6] androecium; [7] corolla; [8]
gynoecium; [9] corolla; [10] calyx; [11] empty anther.
Fruit scale surface of Daemonorops fissa complex [A] concave;
[B] flat
Seed surface of Daemonorops fissa complex [A] smooth; [B]
reticulate
Endosperm of Daemonorops fissa complex [A] slightly ruminate;
[B] deeply rumninate
Epidermal cell of Daemonorops [A] abaxial surface; [B] adaxial
surface. [1] stomata; [2] epidermis.
Transverse section of leaf, [1] epidermis; [2] sclerenchym; [3]
bulliform cell; [4] palisade; [5] sponge; [6] tannin; [7] phloem
strands; [8] xylem.
Position of bulliform cell Daemonorops fissa complex [A] between
two vascular bundle; [B] near midvein; [C] midvein.
Type of bulliform cell Daemonorops fissa complex [A] letter V; [B]
curved upward; [C] parallel; [D] curved downward.
4
7
8
9
10
11
11
11
11
14
14
15
16
14 Palisade tissue Daemonorops fissa complex [A] with one layer of
palisade tissue (dorsiventral); [B] with two layer of palisade tissue
(isobilateral). [1] upper palisade; [2] under palisade.
15 Dendogram of Daemonorops fissa complex in West Malesia based
on morphological characters
16 Daemonorops angustifolia (Griff.) Mart. [A] herbarium type
collection SING; [B] female and sterile male flower; [C] fruit; [D]
endosperm; [E] knee; [F] latex; [G] cane.
17 Daemonorops calicarpa (Griff.) Mart. [A] herbarium type collection
K; [B] spine oblique circle in leaf sheaths; [C] flower; [D]
endosperm. [1] sterile male flower; [2] female flower; [3] male
flower.
18 Daemonorops fissa Blume [A] herbarium collection BO; [B]
knee; [C] cirrus; [D] fruit; [E] flower; [F] endosperm. [1] sterile male
flower; [2] female flower; [3] male flower.
19 Daemonorops grandis (Griff.) Mart.[A] herbarium type collection
SING; [B] knee; [C] male flower; [D] endosperm. [1] corolla; [2]
stamen; [3] calyx.
20 Daemonorops lewisiana (Griff.) Mart. [A] herbarium type collection
K; [B] spine in leaf sheaths; [C] female and male flower; [D] seed;
[E] endosperm; [F] transversal veinlets.
21 Daemonorops melanochaetes Blume. [A] herbarium type collection
K; [B] knee; [C] prophyll; [D] flower; [E] endosperm. [1] sterile
male flower; [2] female flower; [3] male flower.
22 Daemonorops monticola (Grift) Mart., [A] herbarium type collection
K; [B] prophyll; [C] spine in leaf sheath; [D] male flower; [E]
flower; [F] endosperm. [1] sterile male flower; [2] female flower; [3]
male flower.
23 Daemonorops stenophylla Becc. [A] herbarium type collection K;
[B] knee; [C] prophyll; [D] fruit.
16
17
22
23
25
27
28
31
33
34
1 INTRODUCTION
Daemonorops is a rattan genus included in the subtribe Calamoideae. It is
the second largest rattan genus after Calamus, which was consisted of about 120
species (Rustiami 2011). The genus Daemonorops was first described by Blume
(1849), based on the type specimen Daemonorops melanochaetes. Daemonorops
was distributed from India and South China through the Malay Archipelago to
New Guinea where was represented by one species. Twenty-two species occur in
Southern Asia (Beccari 1911; Dransfield 1979, 2001; Evan et al. 2001; Hodel
1998). Daemonorops species are highly variable, but they are homoplasious in
many morphological characters (Umapathy et al. 2014).
The greatest morphological variation of Daemonorops was found in Malay
Peninsula, Sumatra and Borneo (Dransfield et al. 2008). West Malesia consisted
of Borneo, Sumatra, Malay Peninsula and Java (Raes & van Welzen 2009). High
diversity in West Malesia is supported by geological conditions and climate
history of the complex as well as the fact that this region is an archipelago with a
variety of obstacles. This condition opens up opportunities for the development of
its type in the isolated region. Rising sea levels during the Pleistocene as well as
the isolated islands causes the distribution patterns of plants including rattan is
unique in the West Malesia (Rustiami 2009).
The species of Daemonorops are mostly confined to primary tropical rain
forest on a great various soils, but some species had narrow ecological
requirements. A few species are a rather more weedy nature, found abundant in
forest habitats with high-light intensities such as riverbanks. One species in
Borneo, Daemonorops longispatha, grows on the landward margin of mangrove
forest. Some species are strictly montane, occurring at altitude up to ca. 2500 m
above sea level (m asl). Several species in Borneo are confined to heath forest, or
to limestone or serpentine rock (Dransfield et al. 2008).
Daemonorops spesies have been recognized having many uses.
Daemonorops melanochaetes is used as raw material for the manufacture
of tables, chairs and household crafts, and ripe fruit is edible (Harada et
al. 2005), D. draco for jernang sap seekers and cane can be used to make
household equipments (Sulasmi et al. 2012). In Jambi, umbut (young
shoots) of D. angustifolia is used for ulcer drugs by Suku Anak Dalam
(Jumiati et al. 2012).
The genus is classified into two sections based on its inflorescence
structure, section Daemonorops and Piptospatha (Beccari 1911; Rustiami et al.
2014). Species within section Daemonorops have concave boat-shaped bracts,
which at anthesis are completely enclosed by the prophyll and split longitudinally
to expose their flowers. All bracts are usually persistent. In contrast to section
Daemonorops, species within section Piptospatha have bracts splitting to the base
and only the lower part of inflorescence rachis is enclosed by the prophyll, and the
subsequent rachis bracts are usually deciduous (Dransfield et al. 2008, Rustiami et
al. 2011, Rustiami 2014).
2
Daemonorops fissa complex was classified into the section Daemonorops
with the sheath upright and directly attaching to the stem by peduncle length of ±
2 cm (pers. com 2013). Based on observed specimen in herbarium and compared
to the previous studies, species concept of Daemonorops fissa complex remains
unclear. Daemonorops fissa complex in West Malesia consists of 9 species
namely D. angustifolia, D. binnendijkii, D. fissa, D.grandis, D. melanochaetes, D.
palembanica, D. sepal, D. stenophylla, and D. trichroa (Beccari 1911). The later
study mentioned only 3 species in Malaya, namely D. angustifolia, D. grandis and
D. melanochaetes (Furtado 1953). However, the last study reported that 5 species
were recognized, D. angustifolia, D. fissa, D. grandis, D. melanochaetes and D.
sepal in Malay Peninsula, Sabah and Sarawak (Dransfield 1979; 1984, 1992).
Rattan classification was mainly based on discontinued of morphological
characters, the basic data for separating among species (Rustiami 2009). However,
morphological characters sometimes have disadvantage where placement of a
taxon in the classification is doubtful (Tellu 2005), therefore another approach is
needed, such as anatomy. Because of uncertain in distinguishing each species
within D. fissa complex, this species complex in this region needs to be reexamined to obtain clear species delimitation, and to describe the relationships
among species within this complex using phenetic approach, based on
morphological and anatomical characters.
Little work has been done on the leaf anatomy of Daemonorops. Anatomy
of Daemonorops can be distinguished from Calamus based on three characters:
guard cells, the presence of longer sklereid cells and commissure, and large cells
contained of tannins (Tomlinson 1961). Another study found that Daemonorops
spp. from Malaya can be distinguished from their sclereid characters (Tomlinson
2006).
The objective of this study was to provide the newest information of
species concept in D. fissa complex, including 1) species diversity, (2)
distribution, (3) morphological variation, (4) anatomical variation and (5) phenetic
relationships.
3
2 MATERIALS AND METHODS
Time and Place
This study was conducted in August 2013 until October 2014 starting with
field exploration in three locations in North Sumatra; Sibolangit, Mount Sinabung
and Deleng Lancuk. Observation of herbarium specimens was done in the
Herbarium Bogoriense (BO) Cibinong Science Centre LIPI, Bogor (West Java).
Plant Materials
Plant materials used to provide morphological data for the present study
were herbarium specimens including preserved material deposited at Herbarium
Bogoriense. All plant materials were represented the species of Daemonorops
fissa complex such as D. angustifolia, D. binnendijkii, D. callicarpa, D. fissa, D.
grandis, D. melanochaetes, D. monticola. D. palembanica, D. trichroa, and D.
stenophylla. A total of 355 herbarium specimens were observed for morphological
examination in this study.
Research Methods
Procedure for observation in this study followed the method described by
Rifai (2012), and de Vogel (1987). The procedure included 14 steps: (1) Selection
of taxon and determination of scope in the study and work to carried out; (2)
Collecting the material as much as possible (3) Reviewing literatures and records
of all the scientific names and data that included in the important taxa; (4)
Selecting the specimens that have been collected in taxonomic units based on
observable characteters; (5) Scrutinizing the available specimens based on
morphological approach; (6) Testing the characters already used by previous
researches, and performing cross check for justifying whether the existing
correlation between various characters could be confirmed or not; (7) Delimiting
the taxa in question according to the study results and resting the plausibility of
newly discovered characters; (8) Searching and determining the relationships
among taxa; (9) Solving the problem concerned to nomenclature; (10)
Constructing a key for identification of acceptable taxa; (11) Labeling on each
specimen examined according to the name of the concluded study; (12) Making
description of each taxa and note for individual specimens analyzed such as
ecology data and distribution; (13) Making draws of organ parts for publication as
needed; (14) Compiling a scientific report to be published.
Field Sampling
Field survey had been done in three areas of North Sumatera, Sibolangit,
Mount Sinabung and Deleng Lancuk, using exploratory method (Rugayah et al.
2004) which explored every location representing each type of ecosystem or
vegetation.
4
Specimen collection followed standard procedures developed by Dransfield
(1986). Data and information recorded from the field included: general data
(location, habitat, height, use of species, and date of collection); habit (cluster/
solitary), stem (total height, with or without leaf sheath diameter, length and
internodes, colour), leaves (number of leaflets, leaflets arrangement, the length
and width of leaves), additional organ (cirrus); inflorescences (length, number,
colour of rachilla), flower (colour, length and widths) ; fruit and seeds. The plants
were then documented, collected and processed for herbarium specimens.
Preparation of herbarium specimens followed standard procedure by
Dransfield (1986). The collection of complete rattan specimens has many
representative characters including:
1- Vegetative parts: Leaf sheath has a knee, ocrea, spine and sometimes a
flagellum (Figure 1A). Leaves of many species are long and large, collected only
a portion of the leaf base, middle section and apex. If the leaf ends with cirrus,
collected the leaf apex with cirrus. If the leaflet is large which does not fit to the
folder, cutting the leaflets off one side (leaving the leaflet bases in place) and
the other side is folded to fit the folder.
A
B
8
2
7
4
1
6
5
3
Figure 1 Part of rattan. [A] Vegetative parts. [B] Generative parts. 1. cirrus, 2.
flagellum, 3. stem bearing a crownshaft, 4. stem bearing open sheaths,
5. knee, 6. ocrea, 7. rachilla, 8. fruit
2. Generative parts: Many rattan species have large and long inflorescence
which could not be collected as a whole specimen (Figure 1B). To collect such
specimens, a part of the inflorescence base was taken, including primary axis,
bract, and the whole partial inflorescence were collected, and then collected the
last partial inflorescence to see the range sizes.
Anatomical Observation
The plant materials used in the present study were mostly dried and pressed
specimens. Samples were taken from the middle leaves located in the middle and
cut along 10cm2. Each species represents of one to two collection numbers.
5
Anatomical observations were prepared from paradermal and transverse leaf
sections. Leaf blade was boiled in HNO3 to manufacture paradermal section then
stained with 1% safranin (Sass 1951). Observed characters included shape and
size epidermal cells and stomata. For preparing leaf transverse section, leaf were
cut using freezing microtome Yamato RV-240 as thin as 20-25 µm. Leaf slices
then were dripped with sodium hypochlorite to remove chlorophyll, then stained
with safranin and then added few drops of glycerin on slide. Observed character
included the number of palisade tissue, palisade thickness, the presence of tannin
cells and the type, number, location and length of bulliform cells. After obtaining
permanent slides, the slide were observed using Nikon Eclipse E100 and
documented using OptiLab Viewer 2.2.
Morphological Observation
Twenty observed morphological characters were described in Table 1.
Morphological character includes several characters from internodes, petiole,
spine, knee, leaflets, inflorescence, fruit and endosperm. (Beccari 1911, Furtado
1953, Dransfield 1979; 1984, 1992, Rustiami 2011).
Table 1 Morphological characters of Daemonorops fissa complex in West Malesia
No
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
Characters
Total height (m)
Length of internodes (cm)
Length of petiole (cm)
Spine in leaf sheath scattered
Spine in leaf sheath oblique
circles
Position spine in petiole
Type of knee
State of character (score)
≤ 5 tall (0), 6 – 10 tall (1), ≥10 tall (2)
≤20 (0), 21-30 (1), >30 (2)
4 (0), ≤30 (1), > 30 (2)
absent (0), present (1)
absent (0), present (1)
only one surface (0), on both surface (1)
elongate-folded (0), elongate-flat (1),
elongate inflated (2). bulgy (3)
Number of leaflets (pairs)
30 (0), 50-80 (1), >80 (2)
Width of leaflets (cm)
≤ 1 (0), 1.5-2.5 (1), ≥ 3 (2)
2
Spine density in 2m leaf sheath ≤5 (0), 6-10 (1), ≥ 11 (2)
Spine on leaf sheath mouth
absent (0), present (1)
Type of spine in leaf sheath absent (0), present (1)
mouth net-like
Type of spine in leaf sheath absent (0), present (1)
mouth triangular
Type of inflorescence
pleonanthic (0), hapaxanthic (1)
Length of prophyll (cm)
≤ 30 (0), > 30 (1)
Surface of prophyll
setose (0), velutinose (1)
Length of peduncle (cm)
1 cm (0), > 1 cm (1)
Spine of peduncle
absent (0), present (1)
Surface of fruit
flat (0), concave (1)
Endosperm
slightly ruminate (0), deeply ruminate (1)
6
Data Analysis
A species cluster was constructed based on morphological characters
observed for each species. Coefficient of similarity was analyzed using Simple
Matching (SM). Clustering analysis was analyzed using unweighted pair-group
method with arithmetic average (UPGMA) method (Rohlf 2002). Analysis was
performed using NTSys version 2.02.
3 RESULT AND DISCUSSION
Species Diversity
Species of Daemonorops fissa complex in West Malesia share some
unique characters such as peduncle erect to ± 2 cm, leaves alternate, length of
involucrum 1 cm, fruit globose, and endosperm ruminate. This complex species
consist of eight species namely, D. angustifolia, D. callicarpa, D. fissa, D.
grandis, D. lewisiana, D. melanochaetes, D. monticola and D. stenophylla. Three
species are proposed as new synonym, D. binnendijkii and D. trichroa are
synonym to D. angustifolia, whereas D. palembanica as a synonym to D.
melanochaetes.
Daemonorops binnendijkii was named based on two types, cultivated plant
at Buitenzorg and fruits from the Utrecht Herbarium, but the collection number of
the specimen types were unknown (Beccari 1911). On September, 25 in 1914,
Grashoff collected a specimen named Daemonorops cf. binnendijkii in Blinjoe,
Bangka Island. This spesimen deposits in Herbarium Bogoriense as a specimen
type. Until now, D. binnendijkii was never collected by researches. Because of
lesser data could be collected, D. binnendijkii is rated as Data Deficient (Bachman
2013). Based on morphological observation of specimen herbarium, D.
binnendijkii is similar to D. angustifolia, and later study based on anatomical
characters showed that anatomical characters of D. binnendijkii is same to D.
angustifolia.
Fruit collection by Teysmann no. 3582 collected in Muara Dua,
Palembang is the type of Daemonorops trichroa, and it deposits in Herbarium
Bogoriense. The fruit is extremely similar to fruit of D. angustifolia from Malayan
Peninsula (Beccari 1911). Based on characters armature of the petiole, leaf sheath,
rachis and spathae, Furtado reduce D. trichroa Miq. as a synonym of
Daemonorops angustifolia (Furtado 1937).
Type of Daemonorops palembanica was introduced from Sumatra into
Botanic Garden of Buitenzorg, and planted there under the name of D. lewisiana.
Beccari (1911), considered D. palembanica as a variety of D. melanochaetes.
After critical observation of four collection numbers of D. palembanica from
Lampung and Palembang collected by Gusdorff, it is concluded that D.
palembanica is a synonym of D. melanochaetes based on their similarities in
characters, density spine in leaf sheath mouth, spine in prophyll velutinose and
deeply endosperm.
7
Distribution
The distribution of the Daemonorops fissa complex is given in Figure 2.
D. angustifolia, D. callicarpa, D. fissa, D. grandis, D. lewisiana, D. monticola
and D. melanochaetes are the most wide spread species, extending throughout in
Malay Peninsula, Sumatra, Java and Borneo. D. stenophylla is known only from
the forest of North Sumatra and west Sumatra. D. callicarpa in Sumatra found in
lowland Dipterocarp forest and growed with Calamus castaneus. D. fissa is a new
record from Sumatra. Dransfield (1979; 1984, 1992), has investigated this species
widespread throughout in Borneo and endemic. D. melanochaetes is widespread
in Java. Generally, the forest in Malay Peninsula and Sumatra are comparatively
rich in Daemonorops fissa complex population, each of them having 6 species.
Daemonorops fissa complex grew in lowland rain forest especially along
stream and other wet places or more open areas, swampy area, ridgetop hill
Dipterocarp forest, bamboo forest and river side on altitude from 70 m – 1900 m
sl. All species of Daemonorops fissa complex was found in lowland forest on
altitude 250 – 1900 m asl, but D. monticola was only found above 1000 m asl.
Malay Peninsula
Sumatra
Borneo
Java
Figure 2 Distribution of Daemonorops fissa complex in West Malesia
D.angustifolia ( ), D.grandis ( ), D.calicarpa ( ),D.fissa (
D. lewisiana ( ), D. melanochaetes ( ),D. monticola ( ),
D. stenophylla ( )
8
Morphological Variation
Habit
All species of Daeomonorops fissa complex from West Malesia are climb
clustering rattan. Three species, D. calicarpa, D. lewisiana and D. monticola are
found climbing up to 5 m high or more. Two other species, D. grandis and D.
stenophylla are found climbing to 10 m. Daemonorops angustifolia, D.
binnendijkii, D. fissa and D. melanochaetes are the highest climbers, up to 20 m
or higher.
Leaf sheath
Leaf sheath of Daemonorops fissa complex are varied in colour, spine
position and type. The colour of leaf sheath are dull greenish brown, pale green to
pale brownish, and brownish green. Spine position in leaf sheath are scattered or
oblique circle (Figure 3). Species with spine arranged in oblique circle are D.
calicarpa, D. lewisiana and D. monticola, whereas species with scattered spines
include D. angustifolia, D.fissa, D. grandis, D. melanochaetes and D. stenophylla.
Spine type are net-like spine to triangular spines. All species of D. fissa complex
have triangular spines, but only D. calicarpa has a net-like spine.
A
B
Figure 3 Spine types on the leaf sheath of D. fissa complex. [A] Oblique circle;
[B] scattered
Leaves
The colour of D. fissa complex leaves are dark green. Both upper and
under leaf surfaces are glabrous. The leaflets arrangement is pinnate with narrow
and elliptic shape, but leaflets of D. grandis is broad with lanceolate shape. The
number of leaflets on D. fissa complex is generally up to 50 – 90 pairs on each
side, but D. grandis has up to 30 pairs.
Cirrus
Cirrus is extending from leaf apex. It is varied in length. All species D.
fissa complex have cirrus length up to 100 cm – 125 cm, but D. calicarpa have
short cirrus, 30 cm.
Petiole
Cross section of D. fissa complex petiole are channeled, the petiole adaxial
surface is concave and ridged while the petiole abaxial surface convex.
Daemonorops stenophylla has shorter petiole (4 cm) than that of other species
9
(10-30 cm). Daemonorops angustifolia has the longest petiole length up to 35
cm.
Knee
Type of knee is an important character for identification rattan species.
Baja-lapis (2010) described 5 knee types, including bulgy, elongate-folded,
elongate-flat, elongate inflated and cylindrical with ant holes. Daemonorops
calicarpa, D. grandis and D. stenophylla have elongate-folded knees. D.
melanochaetes and D. monticola has elongate-flat knee. D. angustifolia has
elongate-inflated knee; whereas D. fissa and D. lewisiana have bulgy knee (Figure
4). Knees are covered with spine like the leaf sheath.
A
B
C
D
Figure 4 Type of knee Daemonorops fissa complex. [A] bulgy; [B] elongatefolded; [C] elongate-flat; [D] elongate inflated.
Prophyll
Daemonorops fissa complex has persistent prophyll. The prophylls are
covered many spines, but the density spine is varied. The spine density on
prophyll are rarely to densely spines. Species with rarely spines (setose species)
can be found on D. angustifolia, D. fissa, D. grandis, D. lewisiana, D. monticola,
and D. stenophylla (Figure 5). Two species, D. calicarpa and D. melanochaetes,
have densely spines (velutinose). There are two spine types among Daemonorops,
net-like spine and triangular spine. However, D. calicarpa and D. melanochaetes
have different types of spines, D. calicarpa with net-like spine and D.
melanochaetes with triangular spine. The prophyll of D. fissa complex are tubular
and up to 45 cm long, but D. calicarpa is scarcely longer than 25 cm.
A
B
C
D
E
Figure 5 Density of spine in prophyll of Daemonorops fissa complex.
[A] velutinose with triangular spine; [B] velutinose with netlike spine; [C] setose; [D] net-like spine; [E] triangular spine.
10
Inflorescence
Daemonorops is dioecious plants. There are two inflorescence types,
pleonanthic and hapaxanthic. Stems will die after flowering in hapaxhantic type.
Their inflorescence comprise of numerous lateral inflorescence arising from the
axils of, often markedly, reduced leaves on the upper part of stem. While in
pleonanthic type stem continues to grow even after flowering. Their inflorescence
grow from the lower part of stem which then is continue to grow vegetatively, and
reproduce over a relatively long period throughout its adult life (Baker et. al.
1999). All species of D. fissa complex have pleonanthic inflorescences, but D.
calicarpa and D. lewisiana have hapaxhantic inflorescence.
Flower
Male flowers are up to 6 mm long with corolla splitted to the base into 3
petals. The petals have lanceolate shape and rounded at the base with 4 mm long.
Corolla is twice longer than calyx. Calyx is 2 mm long. Each flower has 6 stamen,
and with anther 2 mm long, female rachilla bears many flowers in dyads consisted
of one female flower and one sterile male flower with empty anthers. Female
flowers are 4 mm long with ovoid, corolla consisted of 3 petals, corolla as long as
calyx. Calyx is truncate or shallowly 3 lobed. Sterile flower (4 mm long) is always
shorter than male flower with empty anthers (Figure 6).
B
A
6
5
2
1
3
C
4
D
7
8
9
11
10
Figure 6 Flower of Daemonorops fissa complex. [A] flower of D.
melanochaetes; [B] Male flower of D. angustifolia; [C] Female
flower of D. melanochaetes; [D] Sterile male flower of D.
melanochaetes. [1] sterile male flower; [2] female flower; [3] male
flower; [4] calyx; [5] corolla; [6] androecium; [7] corolla; [8]
gynoecium; [9] corolla; [10] calyx; [11] empty anther.
Fruit
All species of Daemonorops fissa complex have globose fruit, but their
fruits are differed in fruit scale surface. Only Daemonorops monticola has
11
concave scale surface, the other species have flat scale surface (Figure 7). The
scale of D. fissa complex are arranged in 15 – 18 vertical rows, but the scale of D.
grandis from Aceh is arranged in 21 vertical rows.
A
B
A
Figure 7 Fruit scale surface of Daemonorops fissa complex. [A] concave; [B] flat
Seed
The seed of nearly all rattan has an outer fleshy layer. This fleshy layer is
the outer seed-coat and hence is properly called sacrotesta. sarcotesta varies
greatly in thickness from species to species (Dransfield 1979). All species of
Daemonorops fissa complex have smooth seed surface, but there are two
collections, D. grandis from Mount Kemiri, Aceh and D. angustifolia from
Deleng Lancuk, Sumatra, having reticulate seed surfaces (Figure 8).
A
B
Figure 8 Seed surface of Daemonorops fissa complex. [A] smooth; [B]
reticulate
Endosperm
Sarcotesta is easily separable from the rest of the seed. Endosperm varies
from homogeneous in most genera to shallowly or deeply ruminate in
Daemonorops. A previous study showed that all Daemonorops species have
ruminate endosperm (Dransfield 2008), but endosperms of Daemonorops fissa
complex from West Malesia are varied from slightly ruminate (D. calicarpa, D.
fissa, D. lewisiana and D. monticola) to deeply ruminate endosperm (D.
angustifolia, D. grandis, D. melanochaetes and D. stenophylla) (Figure 9).
A
B
Figure 9 Endosperm of Daemonorops fissa complex. [A] slightly ruminate; [B]
deeply ruminate
12
Table 2. Morphological variation of Daemonorops fissa complex in West Malesia
No
N
Character
Species
1
2
3
4
5
6
7
8
1
Total height (m)
30
6
30
10
7
30
10
8
2
Length of
internodes (cm)
Length of petiole
(cm)
Spine on leaf
sheath scattered
Spine in leaf
sheath
oblique
circle
Position spine in
petiole
Type of knee
35
20
20
35
20
22
30
14
30
15
40
40
15
20
30
4
scattered
oblique
circle
present
scattered
scattered
scattered
absent
oblique
circle
present
scattered
absent
oblique
circle
present
only one
surface
elongatefolded
45
both
surface
bulgy
only one
surface
bulgy
both surface
elongate-flat
only one
surface
elongate-flat
50
both
surface
elongatefolded
30
80
90
65
both
surface
elongatefolded
80
1
1.8
3
1.7
2
1.5
0.8
10
8
5
4
20
10
12
3
4
5
6
7
8
9
10
absent
both surface
elongateinflated
50
Number of
leaflets (pairs)
Widht of leaflets
1.5
(cm)
Spine density in 12
2m2 leaf sheath
absent
absent
11
Spine on leaf present
sheath mouth
present
absent
absent
absent
present
present
absent
12
Type of spine in
leaf sheath mouth
net-like
present
absent
absent
absent
absent
absent
absent
absent
13
Table 2 continued
No
13
14
15
16
17
18
19
20
N
Character
Type of spine in
leaf sheath
mouth triangular
Type of
inflorescence
Length of
prophyll (cm)
Surface of
prophyll
Length of
peduncle (cm)
Spine
of
peduncle
Surface of fruit
Endosperm
Spesies
1
2
3
present
absent
present
pleonanthic
39
hapaxanthi
c
23
setose
5
6
7
8
present
present
present
present
present
pleonanthic
pleonanthic
hapaxanthic
pleonanthic
pleonanthic
pleonanthic
35
40
30
55
40
32
velutinose
setose
setose
setose
velutinose
setose
setose
2
2
1
1
2
2
1.5
1
present
present
present
present
absent
present
present
absent
flat
deeply
ruminate
flat
slightly
ruminate
flat
slightly
ruminate
flat
deeply
ruminate
flat
slightly
ruminate
flat
deeply
ruminate
concave
slightly
ruminate
flat
deeply
ruminate
Labels of individu in the table above are:
1. D. angustifolia
5. D. lewisiana
2. D. callicarpa
6. D. melanochaetes
3. D. fissa
7. D. monticola
4. D. grandis
8. D. stenophylla
4
14
Anatomical Variation
Stomata type of Daemonorops fissa complex on paradermal section have
been studied. The stomata types of all species are tetracytic (Figure 10). The type
has two guard cells surrounded by four subsidiary cells (or cells neighbouring of
the guard cells), two lateral and two polar ones (Metcalfe 1961, Dickison 1999).
Stomata are infrequent in adaxial surface, and are restricted to intercostals region
in abaxia surface. Terminal subsidiary cells are not well differentiated, sometimes
having short size. Almost all stomata are present on abaxial leaf surface, but they
are only fewer in adaxial surface.
A
B
1
100 µm
100 µm
2
Figure 10 Epidermal cell of Daemonorops. [A] abaxial surface; [B] adaxial
surface, 1 stomata, 2 epidermis
Adaxial epidermis is always uniform, having cell rectangular with cell walls
markedly sinuous but they do not have thickening anticlinal walls. Tellu (2006)
showed that the stomata of Daemonorops species have kriptopor type with sunken
position in the epidermis or sub-epidermis. In contrast to stomatal type, stomatal
size shows variation within Daemonorops fissa complex. Daemonorops
stenophylla has the smallest stomata size, 135.25-193.22 µm2, whereas D.
monticola has the largest stomata size, 270.51-318.81 µm2.
1
2
4
3
5
8
6
7
100 µm
Figure 11 Transverse section of leaf Daemonorops. [1] epidermis; [2]
sclerenchym; [3] bulliform cell; [4] palisade; [5] sponge; [6] tannin;
[7] phloem strands; [7] xylem.
15
Bulliform cells are commonly found in Monocotyledon, which have
function for storing water. Bulliform, so called motor cells, are lose turgory under
water deficit conditions, and thus constrict in upon themselves, causing lamina to
fold or roll inward edge to edge (Dickison 1999).
The bulliform cells of Daemonorops fissa complex have thin walls, and
their size are larger than that of the adjacent epidermal cells. There are usually 617 bulliform cells on the adaxial leaf surfaces. From the middle to the margin leaf,
the bulliform cells on the abaxial epidermis are gradually getting smaller, and
becoming the same size to the abaxial epidermal cells. The position of bulliform
cell is varied, in adaxial and abaxial epidermis, between two vascular bundles,
near midvein and mid-vein (Figure 12).
B
A
C
Figure 12 Position of bulliform cell in Daemonorops fissa complex. [A] between
two vascular bundle; [B] near midvein; [C] midvein. Bulliform cell
pointed by arrow.
All eight species of Daemonorops fissa complex have many bulliform cells
on the upper and lower epidermis. The shape of bulliform cells on the upper
epidermis are varied. Bulliform cells were curved downward found in D.
angustifolia, D. binnendijkii, and D. calicarpa. Daemonorops fissa, D. lewisiana,
D. monticola and D. stenophylla have bulliform cells parallel to epidermis, D.
grandis has V shape, whereas the bulliform cells of D. melanochaetes are curved
upward (Figure 13).
The mesophylls of Daemonorops fissa complex are differentiated into
palisade and sponge tissues. Mesophylls usually consist of one layer of palisade
tissue and three to five layers of sponge tissue. Daemonorops melanochaetes and
D. fissa only have one palisade tissue in upper side (dorsiventral), but D.
calicarpa, D. lewisiana, D. grandis, D. melanochaetes and D. stenophylla have
two palisade tissues (isobilateral) (Figure 14).
16
A
B
C
D
Figure 13 Type of bulliform cell in Daemonorops fissa complex. [A] letter V; [B]
curved upward; [C] parallel to epidermis; [D] curved downward
A
B
1
1
2
Figure 14 Palisade tissue in Daemonorops fissa complex. [A] with one layer of
palisade tissue (dorsiventral); [B] with two layer of palisade tissue
(isobilateral); 1. upper palisade, 2. under palisade.
Anatomical key to Daemonorops fissa complex in West Malesia
1. a.
b.
2. a.
b.
3. a.
b.
4. a.
b.
5. a.
b.
6. a.
b.
7. a.
b.
Stomata size ≤ 193.22 µm2 …………...…………………D. stenophylla
Stomata size > 193.22 µm2…………………………………………....2
Number of tannin sac 3 – 5……………………………………...…….3
Number of tannin sac 6 – 8………………………………..D. monticola
Have one type of bulliform cell......................................................4
Have two type of bulliform cell ......………………………...D. grandis
Number of bulliform cell in upper epidermis ≤ 10……………………5
Number of bulliform cell in upper epidermis 12 – 17….....D. lewisiana
Leaves isobilateral type…………………………………….......D. fissa
Leaves dorsiventral type………………………………………............6
Length of upper palisade 20.33 µm2 .................................D. calicarpa
Length of upper palisade > 23.56 µm2.............................................7
Type of bulliform cell curved upward.......................D. melanochaetes
Type of bulliform cell curved downward......................D. angustifolia
17
Phenetic Analysis
In this study, 20 characters, mostly qualitative characters were selected for
phenetic analysis (Table 1). A data matrix of characters was scored according to
the species descriptions.
Cluster analysis separated all samples into two major groups with
similarity coefficient of 0.36 (Figure 15). Group I are composed of 6 species
which are D. angustifolia, D. fissa, D. grandis, D. melanochaetes, D. monticola
and D. stenophylla. Grup II contained, D. lewisiana and D. callicarpa. They were
separated from six characters, total height, number of leaflets, width of leaflets,
type of spine in leafsheath, position spine in petiole and type of inflorescence.
Based on the similarity coefficient, Daemonorops angustifolia, D.
binnendijkii and D. trichroa are identic with similarity coefficient of 100%.
Therefore, it is concluded that D. binnendijkii and D. trichroa are synonym of D.
angustifolia. D. palembanica is a synonym of D. melanochaetes based on three
characters: density spine in leaf sheath mouth, spine in prophyll velutinose and
deeply endosperm. Grup I grouped by spine in leafsheath scattered, position spine
in petiole only one surface and type of inflorescence is pleonanthic. D. lewisiana
and D. callicarpa grouped by total height ≤ 5 m tall, spine in leafsheath oblique
circles and type of inflorescence hapaxanthic.
Strong character in taxonomic research is a character that only belongs to
one group of taxa that can be used to differentiate it to other taxa (Wiley 1981).
In this research, the type of knees, spines and width of leaflets are considered as
strong characters. Type of knee is an important character for identification of
rattan (Baja-lapis 2010). Spine and width of leaflets can be used as a diagnostic
character to species. Based on dendogram, width of leaflets is a diagnostic
characters to separated D. grandis from others. Previous study also showed that
leaflet size is a diagnostic character (Dransfield 1976).
D angustifolia1
D binnendijkii1
D binnendijkii2
D angustifolia2
D angustifolia4
D trichroa
D angustifolia3
D melanochaetes1
D melanochaetes3
D melanochaetes2
D palembanica
D fissa1
D fissa2
D monticola1
D monticola2
D grandis1
D grandis2
D stenophylla
D lewisiana1
D lewisiana3
D lewisiana2
D calicarpa1
D calicarpa2
MW
I
II
0.36
0.52
0.68
0.84
1.00
Coefficient similarity
Figure 15 Dendogram of Daemonorops fissa complex in West Malesia based on
morphological characters
18
Table 3. Anatomical characters of Daemonorops fissa complex in West Malesia
Spesies
Character
D.angustifolia
D.calicarpa
D.fissa
D.grandis
D.lewisiana
D.melanochaetes
D.monticola
D.stenophylla
Shape of epidermis
Rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
rectangular
Epidermis size (µm2)
20.00-36.75
23.00-39.68
30.00-74.40
24.58-50.00
23.00-39.68
20.00-36.75
24.58-50.00
30.00-74.40
Type of stomata
Tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
tetracytic
Stomata size (µm2)
241.53-265.68
217.37318.81
palisade
217.37241.53
palisade
palisade
270.51318.81
palisade
135.25-193.22
Palisade
217.37265.68
palisade
154.58-241.53
Position sclerenchym
169.07217.37
palisade
Palisade in lower
epidermis
Number of tannin
cell
Arrangement of
bulliform cell in
upper epidermis
Number of bulliform
cell in upper
epidermis
Number of bulliform
cell in lower
epidermis
Bulliform cell size
(µm)
Present
present
absent
present
present
absent
absent
absent
3-5
3-6
5-7
5-7
4-7
3-5
6-8
3-5
curved
downward
curved
downward
parallel
letter v
parallel
curved upward
parallel
parallel
8-9
7
8-10
5-8
12-17
6
10
5-8
8-9
9
8
14
5-7
6
10
8-12
37.20-44.64
24.80
32.24-49.60
24.80-49.60
19.84-49.60
37.20-49.60
29.76-37.20
24.80-32.24
tetracytic
palisade
19
Taxonomic Treatment
Daemonorops
Daemonorops Blume in J.A. & J.H. Schultes, Syst. Veg. 7(2):1333 (1830).
– Type: Daemonorops melanochaetes Blume. West Sumatra, Padang near Ayer
Manchor, August 1878. Beccari 831 (isotype K)
Solitary or clustering rattans, acaulescent to high climbing hapaxanthic (then
always very short stemmed) or pleonanthic, dioecious. Leaf sheaths heavily armed
with spines, spines frequently highly organized. Flagellum absent. Knee
frequently present. Leaves ecirrate in acaulescent species or long cirrate. Leaflets
variously arranged. Inflorescence male and female superficially similar, but within
the genus of two basic types: one with all bracts enclosed within the outermost
bract or prophyll, splitting along their length to expose the flowers (section
Cymbospatha) or the other with bracts splitting along their entire length to leave
no tubular portion and frequently falling (section Piptospatha). Bracts variously
armed. In the section Piptospatha partial inflorescences longer than the
subtending bract; bracteoles and involucres inconspicuous. Male rachilla bearing
male solitary flowers; male flowers with small cup shaped, calyx with three small
lobes; corolla split to the base into 3 petals; stamen 6, slightly epipetalous;
pistillode minute. Sterile male flower found with each female flower, as the fertile
male, but stamens with empty anthers. Female rachilla bearing many flowers in
dyads consists of one female flower and one sterile male flower. Female flower
calyx truncate or shallowly 3 lobed; corolla with 3 petals; gynoecium with 3 stigmas
and with 3 loculs. Sterile flower smaller or at least more slender than the female, with
well formed calyx and corolla, 6 sterile stamens, an abortive ovary. Fruit variously
shaped, tipped with stigmatic remains and covered with reflexed scales. Seed only
one, covered by thin to thick sweet or sour sarcotesta. Endosperm deeply ruminate.
Embryo basal.
Distribution. Geographical distribution of Daemonorops is more restricted than the
genus Calamus