Upload
vuongduong
View
252
Download
3
Embed Size (px)
Citation preview
BIOSYSTEMATIC STUDY OF THE FERN GENUSDIPLAZIUM IN WEST MALESIA
TITIEN NGATINEM PRAPTOSUWIRYO
DEPARTMENT OF BIOLOGYTHE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITYBOGOR
2008
STATEMENT OF RESEARCH ORIGINALITY ANDINFORMATION SOURCE
This is to verify that my dissertation entitled: Biosystematic Study of the FernGenus Diplazium in West Malesia is my own work and never been submitted toany institution before. All of the incorporated data and information are valid andstated clearly in the text, and listed in the references.
Bogor, September 2008
Titien Ngatinem PraptosuwiryoNIM P176 00005
ABSTRACT
TITIEN NGATINEM PRAPTOSUWIRYO. Biosystematic Study of the Fern Genus Diplazium inWest Malesia. (Under the supervision of Prof. Dr. Ir. Edi Guhardja, M.Sc., Prof. Dr. Mien A.Rifai, M.Sc., Prof. Dr. Masahiro Kato, M.Sci., and Dr. Dedy Darnaedi, M.Sc. )
Diplazium is a large genus consisting of about 400 species occur mainly in the tropics,sparingly in the subtropic and only locally extending into temperate. It was estimated that 300species of the records were occurred in Malesia. Taxonomically, Diplazium is very difficult andquite insufficiently known. Therefore a comprehensive study on Diplazium in West Malesia wasconducted by using morphological, ecological, geographic distribution, anatomical, palinological,cytological, as well as DNA analysis to understand the diversity and relationship among species.
Based on gross morphological study on 1051 collection number of specimens as well asliving collections, it was concluded that West Malesian Diplazium comprises of 69 species with 14varieties. Thirteen species of them are proposed and described as new species, namely Diplaziumasymmetricum, D. batuayauense, D. crameri, D. densisquamatum, D. halimunense, D. loerzingii,D. megasegmentum, D. megasimplicifolium, D. meijeri, D. parallelivenium, D. profluens, D.subalternisegmentum, and D. subvirescens. Two new varieties are poposed, namely D. accedensvar. spinosum and D. silvaticum var. pinnae-ellipticum. D. pallidum var. montanum and D.accedens var. ridleyi are proposed as new status.
Based on their main habitats, Diplazium can be classified into three major groups, viz.dryland (dominant), riparian and rheophytic species. Species diversity was culminated at 1000-1500 m above sea level. The individuals with different genetic load in the same speciessometimes grow in the different habitat gradients. Based on the range of the geographicaldistribution, West Malesian species can be divided into three types: (1) very wide species (19species), (2) Malesian species (27 species), and (3) locally endemic species (23 species).
Anatomical study on the transversal section of stipe of 27 species showed that thevascular bundle shape is varying among species. Therefore the leaf-trace shapes are importantdiagnostic features which support species delimitation in Diplazium. Spore morphology studyshowed that perine ornamentations support in delimitating species in Diplazium. However thephylogenetic analysis using parsimony revealed that morphological variation of spore isinadequate to depict natural relationship among Diplazium species.
Cytological study on 117 collection number from 54 localities included in 31 speciesfound that West Malesian Diplazium has six ploidy levels with x = 41 (diploid, triploid, tetraploid,pentaploid, hexaploid, and octoploid). New cytological information for science on 19 species arerecorded. They are D. aequibasale (2n = 164), D. angustipinna (2n = 123). D. asymmetricum (2n= 123), D. batuayauense (2n = 164, 205), D. crenatoserratum (2n = 123, 164), D. halimunense (2n= 123), D. hewittii (2n = 123), D. profluens (2n = 164), D. loerzingii (2n = 82, 123), D. pallidum(2n = 82), D. petiolare (2n = 82), D. porphyrorachis (2n = 164), D. riparium (2n = 82, 123), D.spiniferum (2n = 82), D. subserratum (2n = 82, 123, 164), D. subvirescens (2n = 123), D.tomentosum (2n = 82, 205), D. xiphophyllum (2n = 82, 246), and D. wahauense (2n=164).
Phylogenetic analysis on morphological data sets of 69 species using parsimony revealedthat the phylogenetic relationship among species in the genus Diplazium was very difficult toexplain due to the lack of or weak support Bootstrap value. However the lack of or weak supportfor a phylogenetic tree does not strictly indicate that the pattern observed is incorrect but it doeslimit the amount of confidence that can be placed in the relationships between taxa.and theconclusions can be drawn from them. This study showed that some terminal clades formed areconsisting of species that presumed to be closely related by formerly authors.
DNA analysis resulted new gene rbcL sequences data on 25 species. Gene rbcL sequenceis very well in supporting species delimitation and revealing the intraspecific diversity withinspecies of Diplazium. Phylogenetic analysis on 29 species from West Malesia and 9 referencesspecies outside Malesia using parsimony revealed that gene rbcL is more informative thanmorphological data in inferring phylogeny of Diplazium and showed that West MalesianDiplazium is monophyletic. The position of D. porphyrorachis at the basal clade of themorphological tree is supported by the phylogenetic tree generated from molecular data (generbcL sequence). This study also showed the congruence between the clade of riparium groupdrawn by gene rbcL tree and the clade of imparipinnate frond group drawn by morphologicaltree.
RINGKASAN
TITIEN NGATINEM PRAPTOSUWIRYO. Biosystematic Study of the Fern
Genus Diplazium in West Malesia. (dibimbing oleh Prof. Dr. Ir. Edi Guhardja,
M.Sc., Prof. Dr. Mien A. Rifai, M.Sc., Prof. Dr. Masahiro Kato, M.Sci. dan Dr.
Dedy Darnaedi, M.Sc.).
Diplazium merupakan marga besar tumbuhan paku yang beranggotakan
lebih kurang 400 jenis yang sebagian besar ditemukan di daerah tropis, sedikit di
daerah sub tropis dan hanya secara lokal meluas ke daerah beriklim sedang. Dari
jumlah tersebut, diperkirakan 300 jenis terdapat di kawasan Malesia. Marga
tumbuhan paku terestrial ini mempunyai ciri-ciri diagnosa sebagai berikut: Alur
tangkai daun dan tulang daun utama terbuka dan alur ini diteruskan sampai tulang
anak daun berikutnya; alur daun berbentuk U dengan dasar pipih pada sebagian
besar jenis; anak daun basal yang mengarah ke rembang (acroscopic side)
seimbang atau lebih kecil, pinggir lembaran daun tidak menulang; sori menggaris,
ganda (diplazioid) atau tunggal (asplenoid), yang tunggal membuka ke arah urat-
urat daun utama atau urat-urat daun pusat dari cuping utama, yang ganda
membuka dengan arah bertolak belakang.
Secara taksonomi, Diplazium sangat sulit dan kurang dipahami.
Tumbuhan muda mungkin saja subur dan sulit untuk dikenali sebagai suatu jenis.
Banyak takson memiliki variasi morfologi. Adanya poliploidi, apomiksis dan
hibrid dalam marga paku ini menambah sulit dalam membuat pembatasan jenis.
Pengelompokan anak marga dari marga ini secara alami belum pernah dilakukan
walaupun variasi morfologinya sangat luas. Pembatasan marga Diplazium juga
masih meragukan.
Di Malesia pada umumnya dan Malesia Barat pada khususnya, penelitian
sistematika Diplazium dengan menggunakan pendekatan biologi secara
menyeluruh belum pernah dilakukan untuk seluruh kawasan. Oleh karena itu
penelitian biosistematika Diplazium dengan menggunakan pendekatan morfologi,
ekologi, distribusi geografi, anatomi, palinologi, sitologi dan juga analisa DNA
dilakukan untuk memahami keanekaragaman jenis dan hubungan kekerabatannya.
Berdasarkan pengamatan morfologi pada 1051 nomor koleksi specimen
dan juga koleksi hidup, disimpulkan bahwa Diplazium Malesia Barat terdiri dari
69 jenis dan 14 varitas. Tiga belas jenis diantaranya diusulkan sebagai jenis baru,
yaitu Diplazium asymmetricum, D. batuayauense, D. crameri, D.
densisquamatum, D. halimunense, D. loerzingii, D. megasegmentum, D.
megasimplicifolium, D. meijeri, D. parallelivenium, D. profluens, D.
subalternisegmentum dan D. subvirescens. Dua varitas baru diusulkan, yaitu D.
accedens var. spinosum dan D. silvaticum var. pinnae-ellipticum. D. pallidum var.
montanum dan D. accedens var. ridleyi diusulkan sebagai status baru.
Berdasarkan habitat utamanya, Diplazium dapat dikelompokkan dalam
tiga group utama, yaitu jenis lahan kering (dryland species, 64 species), jenis
riparian (5 jenis) dan reofit (2 jenis). Keanekaragaman jenis memuncak pada
ketinggian 1000-1500 m dpl. Individu-individu dengan tingkat ploidi berbeda
pada jenis yang sama seringkali menempati habitat berbeda berdasarkan
ketinggian. Berdasarkan jangkauan distribusi geografinya, jenis Diplazium
Malesia Barat dapat dibagi dalam tiga tipe: 1) jenis tersebar luas (19 jenis), 2)
jenis Malesia (27 jenis) dan 3) jenis endemik setempat (23 jenis).
Penelitian anatomi irisan melintang tangkai daun pada 27 jenis Diplazium
memperlihatkan bahwa bentuk pembuluh vaskular bervariasi diantara jenis dan
penting untuk menyokong pembatasan jenis. Pengamatan morfologi spora pada 46
nomor koleksi yang tercakup dalam 26 jenis memperlihatkan bahwa hiasan perine
menyokong pembatasan jenis Diplazium. Walaupun demikian analisa filogeni
dengan menggunakan parsimoni menunjukkan bahwa variasi perine tidak cukup
untuk menggambarkan hubungan kekerabatan alami diantara jenis Diplazium.
Penelitian sitologi pada 117 nomor koleksi dari 54 lokasi yang mencakup
31 jenis menemukan bahwa Diplazium Malesia Barat mempunyai enam tingkat
ploidi dengan jumlah kromosom dasar x = 41 (diploid, triploid, tetraploid,
pentaploid, heksaploid dan oktoploid). Informasi sitologi baru bagi dunia ilmu
pengetahuan dilaporkan untuk 19 jenis, yaitu D. aequibasale (2n = 164), D.
angustipinna (2n = 123). D. asymmetricum (2n = 123), D. batuayauense (2n =
164, 205), D. crenatoserratum (2n = 123, 164), D. halimunense (2n = 123), D.
hewittii (2n = 123), D. profluens (2n = 164), D. loerzingii (2n = 82, 123), D.
pallidum (2n = 82), D. petiolare (2n = 82), D. porphyrorachis (2n = 164), D.
riparium (2n = 82, 123), D. spiniferum (2n = 82), D. subserratum (2n = 82, 123,
164), D. subvirescens (2n = 123), D. tomentosum (2n = 82, 205), D. xiphophyllum
(2n = 82, 246) dan D. wahauense (2n=164).
Analisa filogeni pada seri data morfologi dari 69 jenis dengan
menggunakan parsimoni menunjukkan bahwa hubungan kekerabatan filogeni
diantara jenis Diplazium sangat sulit dijelaskan karena tidak adanya atau
lemahnya nilai Bootstrap. Bagaimanapun, tidak adanya atau lemahnya
penyokong statistik bagi pohon filogeni tidaklah menandakan bahwa pola-pola
yang diamati tidak benar namun ini hanya membatasi tingkat kepercayaan yang
dapat ditempatkan pada hubungan kekerabatan diantara takson dan dari nilai-
nilai tersebut kesimpulan dapat ditarik. Beberapa cabang ujung pohon morfologi
tersusun dari jenis-jenis yang diduga berkerabat dekat oleh para peneliti
sebelumnya.
Analisa DNA menghasilkan data baru sekuensi gene rbcL dari 25 jenis
Diplazium. Sekuensi gene rbcL sangat bagus untuk menyokong pembatasan jenis,
mengungkap keanekaragaman genetik dalam jenis dan juga lebih banyak
memberikan informasi untuk menduga filogeni Diplazium dibanding data
morfologi. Berdasarkan analisa filogeni pada 29 jenis Malesia Barat dan 9 jenis
referensi dari luar Malesia, Diplazium terbukti monofiletik berdasarkan gene
rbcL. Kedudukan group porphyrorachis pada cabang pangkal pohon filogeni
morfologi disokong oleh pohon filogeni gene rbcL. Cabang group daun menyirip
gasal pada pohon morfologi selaras dengan cabang group riparium pada pohon
gene rbcL.
Penelitian ini tidak mengusulkan suatu kerangka sistematika di dalam
marga Diplazium, sebab: (1) Pohon hipotesa filogeni yang dihasilkan dari data
morfologi tidak didukung oleh alasan-alasan statistik yang obyektif; (2) Pohon
hipotesa filogeni yang dihasilkan dari sekuensi gene rbcL tidak kokoh karena
sebagian besar jenis kunci yang diduga dari pohon filogeni morfologi tidak
dimasukkan dalam analisa disebabkan ketidaktersediaan sampel material segar
pada jenis-jenis tersebut; (3) Analisa filogeni pada seri kombinasi data molekuler
dan non molekuler untuk menduga filogeni Diplazium belum dilakukan.
Copyright @ 2008, Bogor Agricultural University
All Right Reserved
1. It is prohibited to cite all or part of this dissertation without referring to
and mentioning the source.
a. Citation only permitted for the sake of education, research,
scientific writing, report writing, critical writing or reviewing
scientific problems.
b. Citation does not inflict the name and honor of Bogor Agricultural
University.
2. It is prohibited to republish and reproduce all part of this dissertation
without copyright permission from Bogor Agricultural University.
BIOSYSTEMATIC STUDY OF THE FERN GENUSDIPLAZIUM IN WEST MALESIA
TITIEN NGATINEM PRAPTOSUWIRYO
A dissertation submitted to fulfill one of the requirements for theDoctorate Degree at the Study Program of Biology,
Graduate School, Bogor Agricultural University
DEPARTMENT OF BIOLOGYTHE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITYBOGOR
2008
Examiner of first examination:
Dr. Tatik Chikmawati, M.Si.Department of Biology, Faculty ofMathematic and Natural Sciences,Bogor Agricultural University, Bogor
Examiners of second examination:
1. Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc. Department of Biology, Faculty of Mathematics and Natural Sciences, Bogor Agricultural University, Bogor
2. Dr. Rugayah, M.Sc. Herbarium Bogoriense, Botany Division, Research Center for Biology – Indonesian Institute of Sciences, Cibinong Sciences Center, Cibinong
Dissertation Title : Biosystematic Study of the Fern Genus Diplazium in West Malesia
Name of student : Titien Ngatinem Praptosuwiryo
Number of student : P176 00005
Study Program : Biology
Certified by
Supervisor Committee
Prof. Dr. Ir. Edi Guhardja, M.Sc. Prof. Dr. Mien A. Rifai, M.Sc (Chairman) (Member)
Prof. Dr. Masahiro Kato, M.Sci. Dr. Dedy Darnaedi, M.Sc. (Member) (Member)
The Biology Study Program Graduate School
Dr. Dedy Duryadi Solihin, DEA Prof. Dr. Ir. Khairil A. Notodiputro, M.S. (Head) (Dean)
Examination Date: 25 March 2008 Graduation Date:
PREFACE
Research on the West Malesian Diplazium under the tittle Biosystematic
Study of the Fern Genus Diplazium in West Malesia was conducted from 2001-
2007. Field study was carried out at 54 localities in the primary and secondary
forest of Jawa, Sumatra and Kalimantan. Specimens examination was conducted
at Herbarium Bogoriense (BO) and Herbarium of Singapore Botanic Gardens
(SING). Study on the anatomy, cytology, and mode of reproduction type were
conducted at the Anatomical and Cytological Laboratory of Herbarium
Bogoriense. Observation on spore morphology by using Scanning Electron
Microscope and DNA squencing (gene rbcL squencing) were conducted at the
Laboratory of Department of Chemical and Biological Sciences, Faculty of
Science, Japan Women s University and the Laboratory of Molecular Systematic
of Department of Biological Science, University of Tokyo.
A part of this dissertation (Chapter 5) has been published under two
different tittles articles: (1) Cytological Study of Some Species of Ferns Genus
Diplazium in Java I. that published in Floribunda 2 (5): 128-137 (2004) and (2)
Cytological Study of Some Species of Ferns Genus Diplazium in Java II. that
published in Floribunda 2 (8): 209-221 (2005).
I greatly appreciate to my supervisors: Prof. Dr. Ir. Edi Guhardja, Prof.
Dr. Mien A. Rifai, Prof. Dr. Masahiro Kato and Dr. Dedy Darnaedi, for their
advice, guidance and encouragements throughout this study. I sincerely thank
Dr. Irawati and Dr. Eko Baroto Walujo for their advices and technical support.
I am indebted to Prof. Dr. Ryoko Imaichi, Dr. Chie Tsutsumi, Dr. Y. Kita,
Dr. Mami Konomi, Dr. Mamiko Sato, and Mie Hashino for their helpful advices
and technical supports. I would like express my appreciation to Dr. Chin See
Chung and Ms. Serena Lee M. L. for giving me an opportunity to study Diplazium
specimens at the Singapore Botanic Gardens Herbarium. I am grateful to Dr. R.J.
Johns for fruitful discussion and his guidance when I was studying Diplazium
specimens in Singapore Botanic Gardens. I would like to sincerely thank
Dr. Teguh Triono, Dr. Titik Rugayah, and Dr. Joko Ridho Witono for the fruitful
discussion, criticism, and technical supports. I also thank Dr. Sri Sudarmiyati
Tjitrosoedirdjo and Dr. Tatik Chikmawati for their suggestions and corrections.
I sincerely thank Dr. Rudy Lukman for his helpful advices and soft literatures. I
thank Abdulrokhman Kartonegoro, S.Si. and Dr. Nunik S. Ariyanti for taking
some pictures of specimens type and literatures from Leiden. I would like to
thank Prof. Dr. Hiroshi Okada and Dr. Hirokazu Tsukaya for involving me in the
field study in Java and Central Kalimantan. I also thank to Dra. Esti Munawaroh,
Wita Wardani, S.Si., Arief Hidayat, S.Si., Bp. Bambang Purwadi, Bp. Ujang
Hapid and Sri Wahyuni, S.P. for sampling and fieldtrips. I am also grateful to Bp.
Endjum and Ibu Ahati for maintaining the living collections.
I thank the following institute in which I used their facilities: a)
Herbarium Bogoriense, Botany Division, Research Center for Biology
Indonesian Institute of Science (Lembaga Ilmu Pengetahuan Indonesia), b) Center
for Plant Conservation, Bogor Botanic Gardens LIPI, c) Department of
Chemical and Biological Sciences, Faculty of Science, Japan Women s
University, d) Department of Biological Science, University of Tokyo, Japan.
Finally, my grateful goes to my dear father (Bp. Ng. Praptosuwiryo who
has passed away on 25th February 2006) and mother (Ibu Kardiyem) my sisters
and brother (Yu Ni, Dik Tum, and Dimas Bas) for their deep understanding and
moral supports.
This research was partly supported by Proyek Kompetitif Pengembangan
IPTEK LIPI TA 2004 , Prof. Dr. Masahiro Kato (Department of Botany, National
Museum of Nature and Science, Amakubo, Tsukuba, Japan), Prof. Dr. Ryoko
Imaichi (Department of Chemical and Biological Sciences, Faculty of Science,
Japan Women s University, Japan) and The Singapore Botanic Gardens Research
Fellowship 2005.
CURRICULUM VITAE
Titien Ngatinem Praptosuwiryo was born on 27 March 1969 in Boyolali,
Central Java, the second daughter from four children from father the late
Ngadimin Praptosuwiryo and mother Kardiyem. She was graduated from Bogor
Agricultural University (Institut Pertanian Bogor) in 1994. In 1994 to 1996, she
worked at Herbarium Bogoriense, Puslitbang Biologi LIPI, Bogor as an honorary
researcher in ferns. In September 1996, she had an opportunity to continue study
at IPB sponsored by Prof. Dr. Masahiro Kato, the University of Tokyo, Japan and
admitted to the degree of M.Sc. in August 1999. In 1999 to 2000 she continued
working at Herbarium Bogoriense.
In September 2000, she had an opportunity to continue study at IPB for
her PhD. This study was partly supported by MENRISTEK and LIPI. Since 2001
she has been working at the Center for Plant Conservation-Bogor Botanic
Gardens, LIPI, Bogor as a researcher in plant systematic. Her research area is
biosystematic study on ferns.
i
TABLE OF CONTENTS
Page
LIST OF TABLES …………………………………………………….. iv
LIST OF FIGURES ………..……………………………………………. v
LIST OF APPENDIXES ………………………………………………… vi
1 INTRODUCTION
Taxonomical Aspects of Diplazium and Its Systematic Problems …………………………………………………............ 1
The Diversity of Diplazium in Malesia .......................................... 7The Biological Aspects on Systematics Study of Ferns …............ 8
Morphological and Anatomical Evidences in Taxonomy ……………………………………………........ 8
The Constribution of Palynology to Systematics: Spore Morphology Evidence in Pteridophytes ......................... 9 Cytological Evidence in the Revealing Taxonomic
Problems on Diplazium and Its Closely Related Genera ........................................................................ 10
The Utility of Molecular Techniques for phylogeneticstudies of pteridophytes: Gene rbcl Sequences ....................... 11
Objectives ……………………………………….………………. 12
2 DIVERSITY AND ECOLOGY OF DIPLAZIUM
Introduction ……………………………………………………… 13Materials and Methods ……………………………………......... 14Results and Discussion ……………………………………......... 14
Ecology .………………………………………….................. 14Rheophytic Diplazium ……………………...................... 15Riparian Diplazium ………………………....................... 16Diplazium in Dryland ……..……………........................ 16Diversity of Diplazium Based on Elevation …............... 19
Conclusions …………………………………………….............. 22
3 THE DISTRIBUTION OF WEST MALESIAN DIPLAZIUMINSIDE AND OUTSIDE MALESIA
Introduction ……………………………………………………… 23Materials and Methods …………………………………………... 24Results and Discussion ………………………………………….. 25
Very Wide Distribution Species ………………….................. 26Malesian Species …………………………………................. 26
ii
Endemic Species to Island in West Malesia ………................. 27Conclusions ………………………………………………........... 40
4 THE STELAR ANATOMY OF STIPE AND ITS TAXONOMIC SIGNIFICANT IN DIPLAZIUM
Introduction …………………………………………………….. 41Material and Methods …………………………………………... 42Results and Discussion …………………………………………. 42Conclusions …………………………………………………………… 46
5 CYTOLOGICAL AND REPRODUCTIVE STUDIES ON DIPLAZIUM IN WEST MALESIA
Introduction …………………………………………………….. 47Materials and Methods …………………………………………. 49Results and Discussion …………………………………………. 50
Chromosome Number Variations and Mode ReproductionTypes on Diplazium ………………………………………... 50The Relationship between Ploidy Level andMorphological Variation within Species and CloselyRelated Species of Diplazium ............................................... 66Relationship between ploidy level and habitat gradient ....... 71Correlations between reproductive mode and habitat ........... 72
Conclusions ................................................................................ 81
6 PHYLOGENETIC STUDIES OF DIPLAZIUM FROM WEST MALESIA: EVIDENCE FROM MORPHOLOGY
Introduction …………………………………………………...... 83Character Selection and Construction …………………………. 85
Character Selection ………………………………………… 85Character Type …………………………………………….. 85Character Coding …………………………………………… 86
Character Variation within West Malesian Diplazium ……….. 87Materials and Methods ……………………………………….... 100
Taxon Sampling ………………………………………….... 100Character Examination of Diplazium …………………....... 100Phylogenetic Analysis ……………………………………… 101
Results and Discussion ………………………………………… 110Conclusions …………………………………………………..... 117
7 SPECIFIC DELIMITATION AND RELATIONSHIP AMONG SPECIES OF DIPLAZIUM BASED ON SPORE MORPHOLOGY
Introduction .................................................................................. 118Materials and Methods ................................................................. 120Results and Discussion ................................................................. 122
iii
Spore Characters of Diplazium and Its Use in Supporting Species delimitation and Identification …........... 122
Phylogenetic Analysis ……………………………………... 137Conclusions ................................................................................. 143
8 MOLECULAR SYSTEMATIC OF DIPLAZIUM FROM WEST MALESIA
Introduction …………………………………………………….. 146Materials and Methods …………………………………………. 148
DNA Analysis ……………………………………………… 149Phylogenetic analysis ………………………………………. 161
Results and Discussion ………………………………………… 161Infraspecific Genetic Diversity in Diplazium ……………… 161Species Delimitation in Diplazium based on GenerbcL Sequence ……………………………………………... 163
Informative Characters of Gene rbcL Sequencesfor Inferring Phylogenetic Hypothesis of Diplazium ………. 165Phylogenetic Analysis ……………………………………… 166
The Monophyly of Diplazium ………………………….. 166Relationships among species within Diplazium ….…….. 165
Conclusions ……………………………………………………... 175
9 TAXONOMIC STUDY OF THE FERN GENUS DIPLAZIUMIN WEST MALESIA
Introduction …………………………………………………….. 176Materials and Methods …………………………………………. 176Taxonomic Treatment ………………………………………….. 177
10 GENERAL DISCUSSION
Synthesis ……………………………………………………….. 283 General Discussion …………………………………………….. 284 Systematic Implications for the Genus Diplazium ……………... 285
11 CONCLUSIONS …………………………………………………... 187
LITERATURES ……………………………………………………....... 292
LIST OF TABLES
Table 2.1. Classification of Diplazium based on their main habitat …… 18
Table 2.2. Diversity of Diplazium based on elevation …………………. 20
iv
Table 3.1. Distribution of West Malesian Diplazium inside and outside Malesia …………………………………………..…. 28
Table 3.2. Endemic Species of Diplazium in West Malesia …………… 39
Table 3.3. Species diversity and endemism of Diplazium in four mainlands of West Malesia …………………………… 40
Table 5.1. Somatic Chromosome Numbers, Ploidy Level and Mode Reproduction Type of Diplazium from West Malesia ... 52
Table 5.2. Polyploid series of Diplazium in West Malesia Based on Present Study ............................................................................. 73
Table 6.1. Characters, character states, and coding for 88 characters utilized in construction of morphological data set of Diplazium 102
Table 7.1. Spores Description of Diplazium in West Malesia ................ 124
Table 7.2. Characters, character states, and coding for 17 characters utilized in construction of spore morphology dataset of Diplazium ………………………………………… 143
Table 7.3. Coding for 17 characters utilized in construction of spore morphology data set of Diplazium …………………… 144
Table 8.1. List of Taxa Used in This Study …………………………… 152
Table 8.2. Primers Used for Amplifying and Sequencing DNA fromDiplazium (Hasebe et al, 1994) …………………………… 160
Table 8.3. Infraspecific Genetic Variatons of Diplazium based on Gene rbcL Sequences …………………………………. 162
Table 8.4. Interspecific Genetic Variatons of Diplazium based on Gene rbcL Sequences …………………………………. 164
LIST OF FIGURES
Figure 2.1. a-b. Light shade-ferns of Diplazium ………………………. 21
Figure 2.2. Elevational distribution of Diplazium species in West Malesia ……………………………………………………. 21
Figure 4.1. Vascular structure of the leaf axis …………………………. 43
Figure 4.2. Leaf-trace shapes in Diplazium ………………………....... 44
v
Figure 4.3. Leaf-trace shapes in Diplazium. ............................................. 45
Figure 5.1. Somatic chromosome of Diplazium. ……………………… 74
Figure 5.2. Somatic chromosome of Diplazium ………………………... 75
Figure 5.3. Somatic chromosomes of Diplazium ……...……………….. 76
Figure 5.4. Somatic chromosomes of Diplazium ……………………… 77
Figure 5.5. Somatic chromosomes of Diplazium cordifolium …………… 78
Figure 5.6. Somatic chromosomes of Diplazium ...................................... 79
Figue 6.1. Rhizome appeareance of Diplazium …………………………. 89
Figure 6.2. The variation of scale shapes in Diplazium ………………… 91
Figure 6.3. Margin of scales …………………….……………………… 92
Figure 6.4. Stipes appearances of Diplazium ………………….………. 93
Figure 6.5. Frond architectures of Diplazium …………………………… 95
Figure 6.6. Venation types of Diplazium ………………………………. 98
Figure 6.7. Sori variation in Diplazium .………………………...…....... 99
Figure 6.8. Strict consensus of 8 trees of length 1366 from unweighted morphological dataset comprises 88 morphological caharcters. ....................................................... 111
Figure 7. 1. Group I. a and b. D. accedens; c and d. D. bantamense; e. D. lobbianum; f and g. D. pallidum; h-j. D. procumbens; k-l. D. sorzogonense ................................................................. 133
Figure 7.2. Group II. a. D. subserratum; b-c. D. vestitum; d-e. D. vestitum var. borneense; Group III. f-g. D. crenatoserratum; h-i. D. prescottianum; Group IV. j-l. D. silvaticum ...................................................... 134
Figure 7.3. Group V. a-c D. pallidum; Group VI. d-e. D. cordifolium; g-i. D. tomentosum; j.. D. malaccense; k-l. D. megasegmentum; m -o. D. simplicivenium ……………………………………… 135
Figure 7.4. Group VII. a-c. D. profluens; Group VIII. d. D. spiniferum; Group IX. e-f. D. subvirescens ………..... 136
vi
Figure 7.5. Tree number 1 of 100 the most parsimonius trees ………… 141
Figure 7.6. The strict consensus of 100 the most parsimonious trees ………………………………………….. 142
Figure 8.1. Strict consensus of the 200 equally most-parsimonious trees obtained in maximum parsimony analysis of the rbcL sequence data. …………..………………………………….. 169
LIST OF APPENDIXES
Appendix 1. Matrix of 88 Morphological Characters for Maximum Parsimony …………………………………………………. 309
Appendix 2. Gene rbcL Sequence Data ……………………………….. 321
Plate 1. Diplazium asymmetricum Praptosuwiryo ……………………. 351
Plate 2. Diplazium batuayauense Praptosuwiryo ……………………... 352
Plate 3. Diplazium crameri Praptosuwiryo ........................................... 353
Plate 4. Diplazium densisquamatum Praptosuwiryo .............................. 354
Plate 5. Diplazium halimunense Praptosuwiryo ..................................... 355
Plate 6. Diplazium loerzingii Praptosuwiryo ………………………….. 356
Plate 7. Diplazium megasegmentum Praptosuwiryo …………………… 357
Plate 8. Diplazium megasimplicifolium Praptosuwiryo ………………… 358
Plate 9. Diplazium meijerii Praptosuwiryo …………………………….. 359
Plate 10. Diplazium parallelivenium Praptosuwiryo ............................... 360
Plate 11. Diplazum profluens Praptosuwiryo ........................................... 361
Plate 12. Diplazium subalternisegmentum Praptosuwiryo ....................... 362
Plate 13. Diplazium subvirescens Praptosuwiryo ..................................... 363
Plate 14. Diplazium subvirescens Praptosuwiryo ………………………. 364
1
CHAPTER 1
GENERAL INTRODUCTION
1.1. Taxonomical Aspects of Diplazium and Its Systematic Problems
Diplazium was established by Swartz (1801) and typified by Asplenium
plantaginifolium L. (Diplazium plantaginifolium Sw.). Etymologically Diplazium
is formed from the Greek diplazios which means double, because indusia lie on
both sides of the vein.
Diplazium is a large genus consisting of about 400 species occur mainly in
the tropics (Ching 1964a; Copeland 1947; Tagawa & Iwatsuki,1988), sparingly in
the sub tropic and only locally extending into temperate (Kramer et al 1990). This
genus member has diagnostic characters as follow: Groove of frond axis open to
admit the groove of axis of lower order; frond axes U-shaped with a flat base in
most species; acroscopic basal pinnules equal or smaller, laminar margin not
cartilaginous; sori linear, double (diplazioid) or single, the single ones opening
toward the main veins or the central veins of the ultimate segments, the double
ones in opposite directions (van Alderwerelt van Rosenburgh 1908; Holttum
1966; Kato 1977; Tagawa & Iwatsuki 1988; Kramer et al 1990).
a. Species Delimitation.
Taxonomically, Diplazium is very difficult and quite insufficiently known.
It is in great need of monographic study. The young plants may be fertile and
difficult to assign to a species (Kramer et al, 1990). Many taxa are considerably
morphologically diversified. Their morphological variations are continuous
through apparently intermediate forms, which are commonly regarded as putative
hybrids (Takamiya et al 1999).
2
b. Polyploidy and Polymorphic Species Complex
Many cytological complexities including polyploidy from 4x to 8x,
hybridity, apomixis, have been reported in ca. 20% of cytological investigation of
Diplazium taxa (Lovis 1977). Recent studies on Japanese Diplazium revealed that
one of the reason for the taxonomic complexity of the Diplazium species group
with bi- to tripinnate leaves is apomictic reproduction and most of the group’s
members have been found to be triploid apomics (2n=3x=123). Many
polymorphic Diplazium taxa having large evergreen bi-to tripinnate leaves occur
under evergreen broad-leaved forest from the Ryukyus Islands to southwestern
Japan. Several putative hybrids are also known within Diplazium, thus it is
increasing the taxonomical dispute (Takamiya et al 1999). Cytological
observation of Diplazium from Java (Praptosuwiryo & Darnaedi 1994, 2004) and
Lesser Sunda Island (Praptosuwiryo 2003) also showed many polyploid types in
which several species were apomix. Therefore, polyploidy, apomixis and hybrid
in diplazioid ferns creates difficulty in species delimitation.
c. Subdivision of Diplazium.
Natural subdivision of the genus has not been assigned (Kramer et al
1990). Van Alderwerelt van Rosenburgh (1908) divided this genus artificially
into two sections based on its venation, viz. Eudiplazium and Anisogonium. The
first section includes species with free veins such as Diplazium bantamense, D.
crenatoserratum and D. porphyrorachis; while the second section includes
species with anastomosing veins, such as D. cordifolium, D. esculentum, and D.
fraxinifolium. Copeland (1908), in his revision on the Philippine species of
Athyrium, merged Diplazium into Athyrium and recognized 13 non formal group
of Athyrium, namely drynarioid species (A. hyalostegium, A. loheri), A. japonicum
group (A. japonicum, A. grammitoides, A. acrotis), A. filix-femina group (A.
drepanopteron), A. macrocarpum group (A. halconense, A. anisopteron), A.
nigripes group (Diplazium aristulatum, D. aristulatum var. sphanicolum,
Athyrium philippinense, A. brevipinnulum, A. nigripes var. mearnsianum, A.
elmeri, A. stramineum, A. platyphyllum), A. cyatheaefolium group (A.
cyatheaefolium=Diplazium ebenum, A. atratum=D. atratum, A. oligosorum=D.
oligosorum), A. silvaticum group (A. silvaticum, A. blumei= D. polypodioides, A.
3
fructuosum= D. fructuosum, A. dolichosorum= D. dolichosorum, A. maximum, A.
vestitum= D. vestitum, A. davaoense= D. davaoense, A. esculentum= D.
esculentum), A. umbrosum group (A. meyenianum= D. caudatum, A.
sorzogonense= D. sorzogonense= D. woodii, D. brachysoroides= D.
brachysoroides), A. williamsi group (A. deltoideum, A. whitfordii= D. whitfordii,
A. bolsteri= D. bolsteri, D. geophilum, A. williamsi= D. williamsi), A. pinnatum
group (A. pinnatum= D. petiolare, A. cultratum, A. crenato-serratum= D.
crenatoserratum, D. inconspicuum, A. pallidum= D. pallidum), A. accedens group
(A. accedens= D. accedens), A. fraxinifolium group (A. fraxinifolium=D.
bantamense, A. cumingii= D. cumingii, A. tabacinum = D. tabacinum, A. pariens,
A. cordifolium= D. cordifolium), and A. porphyrorachis group (A. merrillii= D.
merrillii). However he did not explain characters delimiting his classification.
Based on the characters such as scales, stipe, lamina and venations, Kato
(1977) recognized genus Diplazium that consisting of five groups, namely: (1)
Diplazium dilatatum group that includes member with groove generally U-shaped
with a flat base, acroscopic basal pinnules or segments equal to or smaller than the
basiscopic or subsequent ones, scales entire or toothed, but not clathrate. The
representative species are D. dilatatum, D. esculentum, D. donianum, D.
mettenianum, D. subsinuatum, D. hachijoense, D nipponicum, D. sibiricum, D.
squamigerum, D. tomitaroanum, D. pullingeri, D. lobatum, D. yaoshanense, and
D. kawakami; (2) Diplazium wichurae group that includes species member with
groove U-shaped, acroscopic base of pinna auricled, adaxial surface of lamina
concave along veins, scales entire, sometimes subclathrate. The representatives
are D. wichurae, D. okudairae, and D. pin-faense; (3) Diplazium mesosorum
group that having characters groove U-shaped with a flat base, acroscopic basal
pinnules or segments equal to or slightly larger than the basiscopic or subsequent
ones, scales entire and subclathrate and only represented by D. mesosorum; (4)
Diplazium javanicum group in which includes species with groove V-shaped,
frond pinnate or imparipinnate, laminar margin entire or undulate, veins sagenoid-
reticulate and scales entire. This group represented by D. heterophlebium, D.
javanicum, D. cavalerianum, and D. marginatum; and (5) Diplazium longicarpun
group. This group includes species with frond pinnate, acroscopic base of pinna
4
truncate, basiscopic cuneate, adaxial surface of lamina not concave along veins,
scales entire. This group represented only by one species, D. longicarpum.
d. The status of Diplazium a separates genus.
The delimitation of Diplazium is still in doubt. Some taxonomist merged
Diplazium into Athyrium, while others maintained them as separate genera.
Beddome (1883) separated Diplazium from Athyrium based on the difference of
sorus shape. Athyrium species have special reniform or round sori, meanwhile
Diplazium has elongated sori. Copeland (1908) transferred Diplazium in
Philippine into Athyrium. In accord with his previous papers, Copeland (1929) has
described a considerable number of diplazioid ferns as species of Athyrium,
merged the whole species of Diplazium. Furthermore Copeland (1947) united
Diplazium into Athyrium because he believed that Athyrium s.str. (Euathyrium)
and Diplazium as a whole is almost surely a phyletic entity. He was unable to find
any line where the genus can be divided to produce distinguishable natural groups.
Ching (1964a, 1964b) splits Diplazium into several genera, e.g.,
Allantoidea, Callipteris, Diplaziopsis, Monomelangium, and Diplazium sensu
stricto. Ching (1964a) delimited Diplazium Swartz as represented by D.
plantaginifolium (Linn) Urban of Tropical America, D. bantamense Blume of
Java and D. donianum (Mett.) Tard.-Blot of the Orient which characterized by
imparipinnate (sometimes simple or trifoliolate) fronds of firm texture with large
similar pinnae (2-6 pairs or rarely more), in similar way the simple frond, the
upper side of pinna-costa has a very shallow or even obsolete longitudinal canal
provided with low rounded edges on each side and not open to the rachis-groove
at point of insertion and by very long, linear and more often double sori extending
from the costa to near the leaf margin. Diplazium sensu stricto is a pantropical
genus of medium-size (Ching 1964a). On the other hand Allantoidia sensu Ching
(1964a) included the bulk of the species of Diplazium of Christensen’s Index
Filicum as represented by Diplazium dilatatum Blume, D. polypodioides Blume,
and their allies. Their short-linear sori and indusia being asplenioid or sometimes
diplazioid but not allantoid. Ching (1964a) delimited Allantoidea R. Brown as
follow: fronds vary from simply to 2-3-pinnate with lateral pinnae gradually
becoming shorter upwards and finally merged into a deltoid, acuminate pinnatifid
5
apical part, with the pinna-rachis or costa or costule of pinnules (in the compound-
leaved species) deeply grooved above and raised (becoming flat upon drying)
knife-edge margins on each side, which are decurrent along the rachis or costa of
pinnae or costule of pinnule, and the grooves of rachis. Pinna-rachis or of costa
and costule are open to each other at the point of insertion, the leaf-texture is
herbaceous or rarely chartaceous and the sori are thick, short-linear or ovoid-
oblong only with the anterior basal one usually diplazioid. It is a large genus of
about 350 species mostly in tropical and subtropical Asia with a few extending to
the temperate region in the Northern Hemisphere.
However, recent molecular phylogenetic study on the ferns species
included in Athyriaceae sensu Ching (1964a) by using evidence from chloroplast
TrnL-F region sequences revealed that Diplaziopsis C.Chr.is included in the
Diplazium Sw. clade (Wang et al 2003). Wang et al (2003) also gave evidence
that Allantoidea R. Br. and Callipteris Bory should be included in Diplazium Sw.
Christensen & Holttum (1934) separated Diplazium from Athyrium
because they thought that merging the whole mass of species of Diplazium in
Athyrium would result too unnatural grouping. Holttum (1940) originally showed
that in Malaya Diplazium and Athyrium are quite distinct, so that he recognized
the genus Diplazium. Holttum pointed out that Diplazium had pinnae and
pinnules of pinnate frond with subequal base, venation never anadromous, sori
elangated along the veins with lateral indusium, the diplazioid sori usually with
two quite separate indusia, near grading into horse-shoe shape, with indusium
continuous all around, while Athyrium had pinnules often with equal bases, an
anadromous venation and division, and sori are always short and broad, the
diplazioid ones often of a horse-shoe shape, the two sides of the sorus being
connected. In contrast to his 1940 treatment, Holttum (1947, 1955) recognized
the close relationship of Diplazium with Athyrium and hence associated the two
genera because the distinction between them were no longer distinct to justify
their separation. Inconsistently, Holttum (1966) without doubt stated that genus
Diplazium should be maintained because cytologically the two groups were
constantly different. According to Holttum (1966) Athyrium has x = 40 and
Diplazium x = 41. Despite Holttum (1966) others taxonomist such as, Ching
6
(1964a), Pichi-Sermolli (1977), Kato (1977; 1994) and Tagawa & Iwatsuki (1988)
maintained Diplazium separated from Athyrium.
Kato (1977) differentiated between the two genera as follows. Athyrium
had stipe bases on ascending to erect rhizomes swollen with pneumatophores,
frond axes V-shaped in transaction, acroscopic basal pinnules larger than others,
laminar margin cartilaginous or not, spines present adaxially at the junction of
costules or not, sori horse-shoe- or J-shaped, or linear, scales entire. Meanwhile
Diplazium had stipe bases neither swollen nor bearing pneumatophores, frond
axes U-shaped with a flat base in most species, acroscopic basal pinnules equal or
smaller, laminar margin not cartilaginous, spines absent, sori linear, scales toothed
or entire. Furthermore, Kato (1977) merged Callipteris Bory, Allantoidea R.
Brown, Hemidictyum Presl, Diplaziopsis C. Christensen, Monomelangium Hayata,
Dictyodroma Ching and Rhachidosorus Ching into Diplazium. In accord with
Kato (1977), recent molecular phylogenetic studies included Monomelangium in
the monophyletic Diplazium clade; Rhacidosorus is separated from monoplyletic
Diplazium clade and not closely related to either Athyrium and Diplazium (Sano et
al 2000a, 2000b).
e. Classification of Diplazium.
Despite the recognition of Diplazium as genus, the higher classification of
Diplazium (family level) is still unclear. Tardieu-Blot (1932) placed this genus in
the tribe Asplenieae of the family Polypodiaceae. Alston (1956), Ching (1964a).
Pichi-Sermilli (1977) and Tagawa & Iwatsuki (1988) placed the genus in
Athyriaceae, the family that proposed for the first time by Ching (1954) and
validated by Alston (1956). Holttum (1966) placed the genus in Denstaedtiaceae,
the largest family of modern pteridophytes sensu Holttum. Tryon & Stolze (1991)
also placed this genus in Dryopteridaceae. Brummit (1992) and Kato (1994)
included Diplazium in Woodsiaceae, a family previously established by Herter
(1949). Nayar (1970) included Diplazium in Athyrioideae, a subfamily of
Dryopteridaceae. Kramer et al (1990) placed Diplazium in Physematieae, a tribe
of Athyrioideae, a subfamily of Dryopteridaceae.
7
Based phylogenetic trees generated from chloroplast TrnL-F region
sequences that inferred using the neighbor-joining and maximum-parsimony
methods, Wang et al (2003) to include Diplazium into subfamily Diplazioideae
and proposed to divide Athyriaceae into five subfamilies : Cystopterioideae,
Athyrioideae, Deparioideae, Diplazioideae and Rachidosorioideae.
1.2. The Diversity of Diplazium in Malesia
Malesia is one of the center megadiversity of tropical plants. With ca.
40,000 species of vascular plants, the Malesian region is among the most species-
rich areas world-wide (Roos 1993).
It is firstly recognized by Swiss botanist, Heinrich Zolliger, in 1858, that
Malesion region is different from the Asia and Australia due to a vast diversity of
vascular plants. It is predicted that not less than 40.000 species of vascular plants
occur in the region (from John 1995). This region extends from Malay Peninsula
to New Guinea and cover more than 3,000,000 km2 at 0-5000 a.s.l. The region
included seven countries, namely Brunai Darrusalam, Indonesia, Malaysia, Timor
Leste, Papua New Guinea, Philippines and Singapore. Phytogegraphically, this
region is divided into three subregion, viz. West Malesia, Central Malesia and
East Malesia. West Malesia, known as Sunda Shelf, cover Malay Peninsula,
Sumatra, Borneo, Palawan, Jawa and Bali. Central Malesia is subregion covering
Philippina, Sulawesi, Moluccas and Lesser Sunda. East Malesia, recognized as
Sahul Shelf or the Papuasia, is a subregion included Irian Jaya and Papua New
Guinea (John 1995).
Malesia is also a species-rich region for Pteridopyta. It is estimated that
4,400 fern species of ca. 12,000 fern species known in the world are distributed in
this region (Roos 1993). New Guunea is the world’s most species-rich island
subcontinent with ca. 2,000 species, Borneo and the Philippines each have 1,000
species, and Java, sumatra and Celebes each have 500 species. Seram Island, a
small island of 18,000 km2 in the Moluccas, is quite rich and is known to have
about 700 species (Kato 1992). Parris et al (1992) reported that more than 600
fern species occur in Mt. Kinabalu (400 km2). Because of the species richness,
Malesia ferns are very suitable for studying their biodiversity and evolution.
8
Malesian region is the center of Diplazium diversity. It is predicted that
75% of species in the world (ca.300 species) are existing in this region (Roos,
1995). However the revision of Diplazium for the Malesian region has not yet
been done. However, some short studies based on local area has been conducted
for Diplazium. Holttum (1940) described 25 species of Diplazium from Malay
Peninsula and recognized 29 species on his monumental book ‘Ferns Flora of
Malaya (Holttum 1966).
Mitsuta (1985) listed 11 species from West Sumatra. In Java, Backer and
Posthumus (1939) described 17 species. Sixty years after that Praptosuwiryo
(1999) recognized 22 species and 4 varieties of Javan Diplazium. Tagawa (1972)
listed 15 species of Diplazium of Borneo based on ferns collected by M. Hirano
and M. Hotta. Iwatsuki & Kato (1984) reported 15 species of diplazioid ferns of
East Kalimantan, moreover Parris et al (1992) listed 31 species from Mt.
Kinabalu. In addition to the records, Kato (1994) reported 32 species of
Diplazium from Ambon and Seram (Moluccas).
1.3. The Biological Aspects on Systematics Study of Ferns
1.3.1 Morphological and Anatomical Evidence in Taxonomy
Morphological characters are very important in systematics, although
many biological approaches are applied in constructing classification system, such
as anatomy, palinology, cytology, and molecular analysis. The morphological
data are used for grouping, identifying, studying the relationship of plants (Davis
& Heywood 1963).
Fern systematists have employed various species concept. Some
systematists employed morphological species concept, using morphology as
primary criterion in recognizing species and formulating initial hyphotheses about
new lineages. In addition, modern floristic surveys are also based primarily on
morphology and provide the first clues to identify certain individuals or
populations as potentially as potentionally unique, and deserving of further
scrutiny (Haufler et al 2000).
The comparative study of plant structure, morphology and anatomy, has
always been the backbone of plant systematics to elucide plant diversity,
9
phylogeny and evolution. The second half of the 20th century has been a
fascinating period in which systematics and structural studies greatly profited
from new techniques and methods. The advancement of new techniques and
methods such as transmission electron microscopy (TEM), scanning electron
microscopy (SEM), sinematografi, cladistics, evolutionary paleobotany, and
molecular systematics and molecular developmental genetics are just exactly
supporting morphological data (Endress et al 2000).
Although morphological data series has revealed more homoplasy than
those from molecular, however combination of morphology and molecular data
might produced more robust phylogeny. Thus, in reality the two data are
mutually complementary. Even, there is an amazing fact that the succes of
molecular phylogenetic studies are predicted based on previous the morphological
data (Endress et al, 2000).
1.3.2. The Role of Palynology in Systematics: Spore Morphology Evidence inPteridopytes
Palynological characters have now been used in plant systematics for
almost two centuries. Studies of a various taxa employing transmission electron
microscopy (TEM) and or SEM (Scanning Electron Microscopy) have elucidated
the complex nature of both exorspore and perispore development (e.g. Mitsui
1986; Tryon 1986). These investigations suggest that fine-scaled comparisons of
spores among taxa may be necessary to distinguish superficial and gross similarity
from true synapomorphy.
Measurement of spores serves as useful probes for establishing
hyphothesis of evolutionary relatioships within polyploid complexes. Spore size
of polyploid is dependent upon two factors: size of cells in diploid progenitors and
ploidy level. Both factors may be used to predict cell size of missing members of
polyploid complexes from cell-size means of the known members, as long as
environmental variation does not compromise the analysis (Barrington et al 1986)
10
1.3.3. Cytological Evidence in revealing taxonomic problems onDiplazium and Its Closely Related Genera
Once a species has been diagnosed, it represents a working hyphothesis
that can be tested through the application of new evidence or analytical methods
(Haufler et al 2000). The first auxiliary tool that had a major impact on species
concept on ferns was the development of techniques for determining
chromosomes numbers and studying meiotic behaviour (Manton 1950). The new
perspectives that were obtained through chromosomal studied led to a revolution
in he recognition of species boundaries in many groups. Polyploids discovery in
taxa that had been considered simply as ‘polymorphic assemblages’, Manton
demonstrated that evolutionary mechanism in ferns were more complex than
previously appreciated and that hybridization between distinct species was an
important component of the history of fern lineages (Manton 1950). This new
approach results a new perception of fern species. Manton (1950) and her
followers analysed meiosis in artificial hybrids to characterize the limits of fern
lineages as reproductively isolated units, and to identify genetic similarities
between genomes. These studies showed that interactions among ‘primary’
species involving hybridization and polyploidy generated ‘secondary’ species
(terminology sensu Grant 1981), and resulted in reticulate species complexes
whose intricacies challenged subsequent systematists.
As reported by Löve et al (1977) cytological research on Diplazium was
firstly conducted by Manton (1953) for Ceylon species. From 1953 until 1977,
only about 15.5% of ca. 400 species were recognized for its chromosome number.
Intensive studies on cytology of Diplazium in Japan (Ohta & Takamiya
1999; Takamiya et al 1999, 2000, 2001; Sano et al 2000) revealed that cytological
approach through chromosomal information, is very helpful in analysing
polymorphic species. However, few cytological investigations on Malesian
Diplazium have been conducted. Manton (1954) initiated cytological examination
on this genus for Malayan species, moreover Holttum & Roy (1965) conducted
similar study for New Guinean species.
Preliminary cytological observation on Diplazium in Java was reported by
Darnaedi (1992) for tetraploid D. cordifolium and diploid D. esculentum. In
11
addition, Praptosuwiryo and Darnaedi (1994) reported cytological research on six
species of Diplazium from Gede-Pangrango National Park, West Java (one species
of its, viz. D. opacum, have been treated as Cornopteris opaca)
Praptosuwiryo (2003) and Praptosuwiryo & Darnaedi (2004) reported
recent cytological infromation of Diplazium from Malesian region. The first
covered two species of Diplazium from Lombok Island, namely D. malaccense
and D. pallidum, while the second report 43 collection numbers from 10 localities
of Java included 10 species.
1.3.4. The Utility of Molecular Techniques for phylogenetic studies ofpteridophytes: Gene rbcl Sequences
The use of morphology in reconstructing phylogeny of ferns is often
complicated due to the lack of phylogenetically informative characters (Haufler &
Rangker 1995). For example, the simplicity of foliar morphology of
Ophioglossaceae has limited number of characters available for reconstructing
classifications and understanding relationships (Hauk et al 2003).
The lack of informative morphological characters in ferns led to the search
for new sources of characters in molecular data, including restriction site and
nucleotide sequence data, to infer phylogenetic relationships (Eastwood et al
2004).
Nucleic acid sequencing is a relatively new approach in plant systematic,
however the power of the techniques and the data generated have made it become
one of the most utilized for inferring phylogenetic history. DNA sequence data
are the most informative tool in molecular systematics. Comparative analysis of
DNA sequences is becoming increasingly important and so valuable in plant
systematics. The major reasons of its valuable area: (1) the characters
(nucleotides) are the basic units of information encoded in organism and (2) the
potential sizes of informative data sets are immense. Systematic informative
variation is essentially inexhaustible and different genes or parts of the genome
might evolve at different rates. Therefore, questions at different taxonomic level
can be addresed using different genes or different region of a gene (Soltis & Soltis
1998)
12
Molecular systematic studies of the pteridophytes have generated robust
phylogenies at all taxonomic scales. In many cases, molecular phylogeny are
highly congruent with morphological-based hyphothesis (Wolf & Conant 1994).
The use of molecular data to infer phylogeny has yielded valuable insight into the
relationships and evolution of ferns, some with taxonomic implications (Hasebe et
al 1995, Gastony & Ungerer 1997, Murakami et al 1999).
1.4. Objectives
The objectives of the research are:
(1) To obtain delimitation of the genus and species concepts
(2) To provide better data on distribution of species in the West Malesia
region
(3) To provide an identification keys to the species and infra species.
(4) To collect data on species diversity.
(5) To understand the anatomical and palinological data in supporting the
species delimitation.
(6) To recognize ecological aspect of species.
(7) To provide cytological map of species in its distribution site.
(8) To unravel species which is having taxonomical problems.
(9) To obtain phylogenetic hypotheses for Diplazium based on
morphological data and gene rbcL sequence.
13
CHAPTER 2
DIVERSITY AND ECOLOGY OF DIPLAZIUM
2.1. Introduction
The number of organisms in a particular area is determined by speciation
and extinction, and by immigration and local loss. The relative importance of
these processes depends upon the scale of investigation. Ecological processes are
crucial to many theories of speciation (Godfray & Lawton 2001).
Delimitation of species is, of course, performed by choice of a certain
combination of critical characters on comparative-morphological basis. Each
Linnaean species is essentially a self-perpetuating population in the genetic sense,
i.e. a mixture of individual which are to a larger or smaller degree genetically
different. The genetically defined characters do not manifest themselves in exactly
the same way in different individuals, as their manifestation in influenced during
the stages of their ontogenetical development by the environmental condition
(Steenis 1957).
Many environmental factors can affect various ferns, including the present
of other plants, the activities of animals (amongst which man now plays a
predominant role in many areas), and a whole range of physical and climatic
factors (Edie 1978). For the sake of simplicity, most authors on fern taxonomy
usually explained the main factors which act on ferns growing under or less
natural conditions. These factors include the substratum where the fern grows,
type of soil, rock, ect., surrounding vegetation, exposure to light and climatic
conditions, microclimate which surrounds the fern itself (temperatures, humidity,
etc.), and availability of water.
In order to grasp a better understanding in species delimitation on
Diplazium and also to recognize the correlation between ecology and species
richness, ecological studies of this genus from Western Malesia are conducted by
doing field studies, specimens examination, and literatures study. The aims of
this study are: (1) to study the ecology of Diplazium species and (2) to recognize
the diversity of Diplazium species in the different habitat gradient.
14
2.2. Materials and Methods
Ecological and distribution data were collected directly in the field in Java,
Sumatra and Borneo and information obtained from the specimens examined.
Specimens vouchers are housed at Herbarium of Bogor Botanic Gardens (BOHB)
and will be distributed to some herbaria, including Herbarium Bogoriense (BO).
In addition, some literatures are cited for comparison.
2.3. Results and Discussion
All species of Diplazium grow terrestrially, except D. lomariaceum. It
grows both terrestrially and epipetric on wet rocks, in disturbed or secondary
forest and primary forest. Most of species are mountain ferns. They distribute
from 10 m – to 3400 m above sea levels. The ecology of West Malesian
Diplazium for each species is summarized in Table 2.1. The ecology and the
distribution of all species are discussed below.
2.3.1. Ecology
Some authors used habit and habitats for classifying ferns. Holttum (1966)
classified ferns into 8 groups based on its habit and habitats, namely: terrestrial
sun-ferns, terrestrial shade-ferns, climbing ferns, epiphytes of sheltered places,
epiphytes of exposed places, rock-ferns and river-bank-ferns, aquatic ferns, and
mountain ferns. Edie (1978) classifies the characteristics of ferns from each type
of habitat in general: terrestrial shade ferns, terrestrial sun ferns, epiphytes, rock
ferns and aquatic ferns. Parris et al (1992) explained the ecological characteristic
of ferns growing in Mt. Kinabalu based on its habit and habitats: roadside and
thicket-ferns at lower elevations, high-elevation thicket-ferns, tree ferns, ferns of
cultivated areas, shade ferns of forest, ferns of mountain ridges, ferns high
elevations, epiphytic ferns, filmy ferns, high-elevation and epiphytes. Chin (1997)
described habitat of tropical ferns and its diversity and then he recognized six
habitats of fern, viz. lowland rainforest, mountain forest, secondary forest,
agricultural areas, swamps and open waters, and urban areas.
15
Based on their main habitat, Diplazium can be included into three major
groups: dryland, riparian and rheophytic. Rheophytes are plant species which
inhabit the beds of swift-running streams and rivers and grows up to flood-level,
but not beyond the reach of regularly occurring flash floods (van Steenis, 1981).
The rheophytic plants are characterized by having particular morphological
characters as follow: a) narrow lanceolate leves or leaflets; b) mattet root systems;
c) short erect, ascending, or creeping rhizomes tightly attached to streambed
substrates; and flexible stems and petioles. Dryland species generally do not occur
in the flooded zone where rheophytes occur, while rheophytes do not occur in
dryland habitat where dryland plants thrive (Imaichi & Kato 1997). As pointed
out by Imaichi and Kato (1997), there is distinct habitat segregation between
rheophyte and dryland species, especially in the humid tropics. Following the
definition described by Lincoln et al (1982), riparian means living or situated on
the banks of rivers and streams, whereas terrestrial is living habitually on the land
or ground surface.
2.3.1.1. Rheophytic of Diplazium
Rheophytic of Diplazium are smallest group of Diplazium in West
Malesia. There are only two species reported in this study, viz. D. aequibasale
and D. wahauense (Table 2.1. and 2.2.). Kato et al (1991) reported three species,
namely D. aequibasale, D. wahaunse and D. subsinuatum. The last species
however has been moved into Deparia and treated as Deparia lancea (Thunb.)
Fraser-Jenk based on morphology, cytology and molecular characters (Sano et al
2000). In this paper D. subsinuatum is excluded. In Sumatra and Borneo, the first
species are usually growing on lowland clayey stream-bank, whereas the second
species, that is only found in Borneo, growing at streambed in flood zone in
lowland.
Van Steenis (1981) classifies rheophytes into obligate and facultative,
according to preference for rheophytic habitats. In Borneo, many plants of D.
esculentum are growing well on river banks in the flood zone, for example at
Sungai Joloi (track to Batikap, Central Kalimantan). This species may be
included in facultative rheophytes. But anatomical observations show that leaves
16
of D. esculentum have poorly developed intercellular spaces of rheophytes (Kato
& Imaichi unpublished), there is no distinct morphological difference between the
reophytic and dryland plants. Kato (1991) regarded such species as evolutionarily
incipient rheophytes.
Kato et al (1991) assumed that most rheophytes are products of primary
speciation from dryland mother species. Diplazium wahauense presumably
derived from D. riparium, Deparia biserialis and De. confluens from D.
petersenii, Phronephrium hosei from P. rhombeum or P. menisciicarpon.
2.3.1.2. Riparian Diplazium
Only few species occur on river banks or stream. They are D.
aequibasale, D. riparium, D. fuliginosum, D. lomariaceum and D. porphyrorachis
(Table 2.2.). However these species do not strictly grow at this habitat. The last
three species are more usual growing in dryland of shady ravine in the forest. In
Borneo, much of individual plants of D. riparium are also growing well in shady
dryland. All species mentioned above, are usually found on lowland mountain
forest.
Environmental condition seems very important component of speciation in
Diplazium although this of course depend upon of how one of species response
the ecological change. Diplazium riparium and its closely related species (D.
aequibasale and D. wahauense) may be a good example for it. Kato et al (1991)
presumed that D. wahauense may have been derived from D. riparium which
occurs in riparian and dryland forest of Borneo. As they explained and supported
by this studies (see Chapter 8), these two species share black scales, somewhat
crisped, entire scales, blackish stipes, dark brown, naked rachises, and
imparipinnate leaves with 4 pairs of entire lateral pinnae. Diplazium wahauense
differs from D. riparium mainly in its narrow pinnae, which are characteristic of
rheophytes.
2.3.1.3. Diplazium in Dryland
Most of Diplazium species are growing terrestrially in dryland forest.
(Table 2.1. and 2.2.). According to the light intensity, the dryland Diplazium can
17
be divided into three groups: a) opened area, b) light shady ferns, and c) deep
shady ferns. The species that adapted to the light intensity are discussed below.
a. Opened area ferns
D. esculentum usually grows on opened areas. This species grows well at
the dump soil of opened areas both in the forest and at the farming areas.
Therefore sometimes this species often form small population at the edges of
rivers or stream.
b. Light shady ferns
There are great number of species growing in the light shady area. Some
species adapted to very humid conditions and living near streams, such as D.
accedens, D. kunstlerii, D. procumbens, D. profluens, D. squarrosum, D.
spiniferum, and D. vestitum. The humidity of the air in primary forest near
streams is always high, even in the places well away from streams its average is
much above that of the open air outsite the forest.
Many species are growing well on dry areas, tolerating drier soil and air.
They are found further from streams on forested hill slopes, viz. D.
asymmetricum; D. atrosquamosum D. bantamense, D. barbatum, D.
batuayauense, D. betimusense, D. cordifolium, D. crenatoserratum, D.
densisquamatum, D. dilatatum, D. dolichosorum, D. donianum , D. fraxinifolium,
D. halimunense, D. hewittii, D. hottae, D. latisquamatum, D. lobbianum, D.
loerzingii, D. malaccense, D. megasegmentum, D . meijerii, D. pallidum, D.
parallelivenium, D. petiolare, D. poiense, D. polypodioides, D. sorzogonense, D.
speciosum, and D. simplicivenium.
Shady ferns usually grow more slowly than sun-ferns. The weak light
prevents plants to make its carbohydrates fast and the high humidity may make
less loss of water and less root activity. However this condition does not prevent
many species of Diplazium from attaining a immense size, such as D. accedens,
D. dilatatum, D. polypodioides, D. subpolypodioides, D. megasegmentum, and D.
sorzogonense.
18
Table 2.1. Classification of Diplazium Based on their main habitat
MajorGroup
Species Number ofSpecies
Rheophytic D. aequibasale, D. wahauense 2Riparian D. aequibasale, D. riparium, D. fuliginosum, D.
lomariaceum, D. porphyrorachis5
Dryland D. accedens, D. albidosquamatum, D. angustipinna, D.asymmetricum, D. atrosquamosum, D. bantamense, D. barbatum,D. batuayauense, D. beamanii, D. betimusense, D. christii , D.cordifolium, D. crameri, D. crenatoserratum, D. crinitum, D.cumingii D. densisquamatum, D. dilatatum, D. dolichosorum, D.donianum, D. esculentum , D. fraxinifolium, D. halimunense, D.hewittii, D. hottae, D. insigne, D. kunstlerii, D. laevipes, D.latisquamatum, D. lobbianum, D. loerzingii, D. malaccense, D.megasegmentum, D. megasimplicifolium, D. melanolepis, D.meijerii, D. moultonii, D. pallidum, D. parallelivenium, D.petiolare, D. poiense, D. polypodioides, D. prescottianum, D.procumbens, D. profluens, D. riparium, D. silvaticum, D.simplicivenium, D. sorzogonense, D. speciosum, D. spiniferum, D.squarrasum, D. subintegrum, D. subalternisegmentum, D.subserratum, D. subvirescens, D. tomentosum, D. tricholepis, D.umbrosum, D. velutinum, D. vestitum, D. xiphophyllum
64
c. Deep shady ferns.
Many species are adapted to deep shady areas, such as D.
albidosquamatum, D. angustipinna, D. beamanii, D. christii, D. cumingii, D.
fuliginosum, D. laevipes, D. lomariaceum, D. moultonii, D. poiense, D.
tomentsum, and D. umbrosum. Therefore they are nearly all have much thinner
fronds than the light shady ferns. The deep shady ferns species do not need to be
tough to avoid being shriveled by the sun’s heat. These species do not need to
store water as much as some light shade ferns do. The deep shady ferns usually
also show more dark green or blue metallic fronds than the light shade ferns
(Figure 2.1.).
This classification is not strict as some species are also grow well in
opened and light shady areas. D. polypodioides are seen in the gap areas of thick
forest and also in the margin forest. There are several species found growing in
both light shady and deep shady areas. D. bantamense, D. batuayauense, D.
cordifolium, D. crinitum, D. crameri, D. dilatatum, D. donianum, D.
fraxinifolium, D. lobbianum, D. loerzingii, D. malaccense, D. melanolepis, D.
riparium, and D. sorzogonense can be found in light shady and deep shady areas.
19
2.3.2. Diversity of Diplazium Based on Elevation
Table 2.3.. and Figure 2.2. show the diversity of Diplazium based on
elevation. The species number of Diplazium culminate at 1000-1500 m. Amount
of 19 species of Diplazium distribute at the elevation. Some studies on diversity
across elevation (Shmida & Wilson 1985; Gentry & Dodson 1987; Colwell &
Hurtt 1994) also reveal that peak diversity occur at intermediate elevations.
Lomolino (2001) predicted that species-density should peak at an intermediate
elevation and the peak should occur at transition zone between the two species-
rich, juxtaposed communities. Because detailed information on climatic gradients,
dispersal, population persistent and anthropogenic disturbance are generally
unavailable, it is difficult to evaluate critically some of the predictions associated
with causal explanation for peaks in diversity at intermediate elevations
(Lomolino 2001).
Some species of Diplazium, such as D. atrosquamosum, D. beamanii, and
D. squarrosum, are only occurring at upper elevation zones (Table 2.3.). Above
3000 m, there was only found D. moultonii. Some scientists (Kikkawa & Wlliams
1971; Gentry & Dodson 1987; McCoy 1990; Rahbek 1995) reported that
speciation and endemicity peak at the intermediate to high elevations. Rahbek
(1995) presumed that upper elevation zones may provide the geographical
isolation required for speciation. If the montane zones are both isolated and large
enough to allow population persistence and divergence over evolutionary time,
they may represent hotspots of speciation and endemicity.
Some species that presumed to be closely related have different in the
range of distribution based on altitude. D. lobbianum and D. bantamense are
presumed closely related. The two species share character combination as follow:
scales toothed; fronds imparipinnate; pinnae oblong-ovate; vein free, forked,
texture chartaceous. Ecologically D. lobbianum is found at 1500-1800 m, whereas
D. bantamense usually below 1500 m.
20
Table 2.2. Diversity of Diplazium based on Elevation
Elevation m Species 20- 500 D. accedens, D. acuminatum, D. aequibasale, D. angustipinna, D. bantamense, D.
batuayauense, D. crenatoserratum, D. crinitum, D. hewittii, D. lomariaceum, D.pallidum, D. petiolare, D. polypodioides, D. porphyrorachys, D. riparium , D.tomentosum, D. wahauense, D. xiphophyllum
500-1000 D. accedens, D. acuminatum, D. bantamense, D.. cordifolium, D. crenatoserratum,D. cumingii, D. hewittii, D. hottae, D. laevipes, D. lomariaceum, D. pallidum, D.petiolare, D. poiense, D. polypodioides, D. simplicivenium, D. sorzogonense, D.tomentosum, D. umbrosum, D. xiphophyllum
1000-1500 D. accedens, D. asymmetricum, D. barbatum, D. cordifolium, D. crenatoserratum,D. cumingii, D. dilatatum, D. halimunense, D. hewittii, D. laevipes, D.lomariaceum, D. latisquamatum, D. moultonii, D. pallidum, D. petiolare, D.poiense, D. polypodioides; D. sorzogonense; D. tomentosum, D. tricholepis, D.umbrosum, D. velutinum, D. xiphophyllum.
1500-2000 D. atrosquamosum, D. barbatum, D. cordifolium, D. fuliginosum, D.latisquamatum, D. lomariaceum, D. moultonii, D. poiense, D. polypodioides, D.sorzogonense, D. speciosum, D. tricholepis, D. umbrosum, D. velutinum
2000-2500 D. atrosquamosum, D. barbatum, D. cordifolium, D. fuliginosum, D.latisquamatum, D. moultonii, D. sorzogonense, D. speciosum, D. tricholepis
2500-3000 D. cordifolium, D. fuliginosum, D. latisquamatum, D. moultonii, D. speciosum
3000-3400 D. moultonii
The individuals with different ploidy level in the same species sometimes
grow in the different altitude (See Chapter 5). Diploid D. pallidum from Java,
Borneo, and Sumatra were found at 200 m, 240 m and 30 to 85 m, respectively.
Whereas the tetraploid ones were found at 1000 – 13000 m.
Some closely related species may occur in overlapped distribution. D.
insigne is evidently closely allied to D. accedens (Holttum 1940). The two species
share in characters: dull brown toothed scales with narrow thickening black
strands, the deltoid deeply lobed apex of fronds and copiously goniopterid
venation. The first species distributes from 600 m to 1200 m and the second
species from 80 m to 1400 m.
21
Figure 2.1. a-b. Light shade-ferns of Diplazium. a. D. hottae; b. D. loerzingii; c-d. Deepshade-ferns. c. D. cordifolium; d. D. tomentosum.
0
5
10
15
20
25
0-500 501-1000 1001-1500 1501-2000 2001-2500 2501-3000 3000-3500
Elevation Range (meter)
Num
ber
of S
peci
es
Fig. 2.2. Elevational distribution of Diplazium species in West Malesia
22
2.4. Conclusions
Based on their main habitat, Diplazium can be divided into three major
groups: dryland, riparian and rheophytic. Most of species are terrestrial dryland
ferns and found at 20 – 3400 m above sea level in the primary and secondary
forest on moist humus-rich soil in light and deep shady places (64 species). There
are only found five of the riparian species, namely D. aequibasale, D. riparium,
D. fuliginosum, D. lomariaceum, and D. porphyrorachis. The rheohytic species
are found in D. aequibasale and D. wahauense.
The most species number of Diplazium were culminated at 1000-1500 m.
The lowest number of species occurred at 2500-3400 m. The individuals with
different ploidy level in the same species sometimes grow in the different gradient
habitats.
23
CHAPTER 3
THE DISTRIBUTION OF WEST MALESIAN DIPLAZIUM INSIDE ANDOUTSIDE MALESIA
3.1. Introduction
The fern genus Diplazium is an important component of tropical rainforest
of the Old World and the New World. It is a terrestrial ferns which is commonly
found in the moist ground at the humus rich soil both in the primary and
secondary forest at 20-3400 m altitude. Most of species grow in shadowed place
and fond of moist humus rich soil. Some species are locally abundant by stream
in the mountains. Only a few species are found in limestone areas and in
rheophitic areas (See Chapter 2).
Since 1801 Diplazium have been the subject of numerous taxonomic
studies conducted, including morphological, anatomical, cytological and
molecular investigations. It is estimated that the genus consisting of about 400
species (Ching 1964a, Copeland 1947). Roos (1995) estimated that 300 species
of its are found in Malesian region. Few species are found in continental Africa
(Kramer et al 1990). In the Neotropics there are nearly 100 species (Pacheco
2004).
Distributional data are important in answering many questions about
polyploidy and speciation (Baack 2004). Usually, increasing in ploidy level is
associated with the origin of novel adaptations (Levin 2002). Polyploid often
occupy different habitats from those of their diploid parents (Soltis & Soltis
2000).
Study of the biogeographical distribution of organisme are also very useful
in inferring the monophyly of a taxon. The elementary questions of historical
biogeography concern areas of endemism and their relationships (Nelson &
Platnick 1981). By analogy to phylogenic systematics, where species or higher
taxa are grouped, in cladistic biogeopgraphy the units grouped are areas of
endemism (Linder 2001). For example, study on molecular phylogenetic and
historical biogeography of Hawaiian Dryopteris (Dryopteridaceae) (Geiger &
Ranker 2005) indicate that Hawaiian Dryopteris is not monophyletic, and there
24
were at least five separate colonizations of the Hawaiian Islands by different
species of dryopterioid ferns, with most of the five groups having closest relatives
in SE Asia.
This chapter presents a preliminary compilation of Western Malesian
Diplazium distribution and their inferred center of diversity in Malesia based on
specimens examined and direct observation in the field and their distribution
outside West Malesia that obtained from literature studies. In the light of current
concerns on the loss biodiversity it is also useful to highlight the areas with hight
species numbers and endemicity. The aims of the research are map the
distribution of Western Malesian Diplazium and determine the endemic species.
3.2. Materials and Methods
The determination of species distribution patterns was studied by
examining specimens deposited at BO and SING, conducting field work in Java,
Sumatra, and Borneo. Beside that many literature were also studied for
determining the range area distribution of each species.
Following those conducted by Parris (2003) in presenting the distribution
of Grammitidaceae in the world, for the purposing of examining the distribution
of Western Malesian Diplazium, it is convenient to divide the world into five
regions: (1) New World (North, Central and South America, the West Indies and
the islands of the Galapagos, Falklands and South Georgia); (2) Africa
(continental Africa, Madagascar and the islands of the Azores, Canaries,
Ascension, St. Helena, Tristan da Cunha group, Seychelles, Comoros, Mauritius,
Reunion, Marion, Crozets and Kerguelen); (3) Asia excluding Malesia (India,Sri
Langka, Nepal, Thailand, Cambodia, Laos, Vitenam, China, Taiwan and Japan);
(4) Malesia (Malaysia, Singapore, Brunei Darussalam, Indonesia, Philippines and
Papua New Guinea), and (5) Pacific (Australia, New Zealand and the islands of
Micronesia, Melanesia and Polynesia).
25
3.3. Results and Discussion
A large number of Diplazium specimens (1051 collections number )
deposited at BO and SING and new collections obtained from the field have been
examined. Sixty nine species of Diplazium are recorded within West Malesian
region. Diplazium of Malay Peninsula, Borneo, Sumatra and Java (including
Bali) comprises of 28, 40, 29, and 30 species, respectively (Table 3.2. ). The
distribution for each species inside and outside West Malesia are presented in
Table 3.1.
Revision on Bornean and Sumatran species has not been conducted since
van Alderwereld van Rosenburgh (1908). For Bornean species, formerly author
who ported the check list of this genus locally were Iwatsuki & Kato (1984),
Tagawa (1972), Kato et al (1991), and Parris et al 1992). For Sumatran species,
the check list provided is only those reported by Mitsuta (1985) of West Sumatra.
Thus study was the first account of Diplazium throughout Borneo and Sumatra.
Beside that three and four species are now recognized as new species for Borneo
and Sumatra, respectively (See Chapter 9).
For Javanese species the recent account after Backer & Posthumus (1939)
was Praptosuwiryo (1999). The first author described 17 species, three species of
them have been included in other genera. Meanwhile second author described 22
species and 4 varieties. Now, eight species were added for Java and seven species
of them are proposed as new species (See Chapter 9).
Holttum (1940, 1966) reported 27 species of Malay Peninsula that include
D. heterophlebium, D. curtisii and D. amplissimum. As stated in Chapter 9, the
first two species are included in the doubtful names. The latest species was
included in Cornopteris as C. atroviridis (v.A.v.R.) M. Kato (Kato 1979). D.
aequibasale , D. christii , and D. fraxinifolium are new record for Malay Peninsula
(See Chapter 9).
Based on the range of its geographical distribution, the West Malesian
species can be divided into three types: (1) very wide distribution species, (2)
Malesian species, and (3) endemic species to island in West Malesia. The three
types of the geographical distribution are discussed below.
26
3.3.1. Very Wide Distribution Species
A conception of very wide species adopted here is referred for species that
is distributed at least two region or more of regional division following those
adopted by Paris (2003). Of the sixty nine species of Diplazium from West
Malesia, 19 of them distribute very wide (Table 3.1.), in the New World. They
are D. accedens, D. bantamense, D. cordifolium, D. crenatoserratum, D.
dilatatum, D. donianum, D. esculentum, D. fuliginosum, D. malaccense, D.
pallidum, D. polypodioides, D. prescottianum, D. riparium, D. silvaticum, D.
simplicivenium, D. sorzogonense, D. subserratum, D. tomentosum and D.
xiphophyllum . Three species of them are distributing across four main region
(Africa, Asia, Malesia, Pacific), viz. D. accedens, D. dilatatum, and D.
esculentum.
Some species that widely distributed are varying in morphology and
genetic (see Chapter 2 and 9). Therefore some varieties are found in these
species. Kato (1995) recorded 2 varieties in D. donianum and D. dilatatum. In
this study (see Chapter 9), D. accedens, and D. cordifolium are rezognized to
have three varieties. Meanwhile in D. pallidum and D. silvaticum are recorded
two varieties.
The very wide species in general are also showing the long distance
gradient distribution. They are commonly growing in lowlands or medium
altitude of mountain forest. For example, D. accedens, D. polypodioides, and D.
pallidum are found at 80-1400 m, 200-1900 m, and 600-1500 m, respectively (See
Chapter 2).
3.3.2. Malesian Species
Malesian species denote for species that having wide distribution range in
Malesia region only. These species may be only found in West Malesia or
distributed throughout Malesia. Most of species are distributed widely in
Malesian region. Some species, such as D. aequibasale, D. angustipinna, D.
kunstlerii, D. latisquamatum, D. procumbens, D. subintegrum and D.
subpolypodioides are only found in West Malesia.
27
3.3.3. Endemic Species to Island in West Malesia
Endemic species concept adopted here refer to a species that restricted to a
particular local or island. List of endemic species of Diplazium are showed in
Table 3.3 Endemic species criteria was determined by inspection of distribution
maps cited from literatures. The total number of local endemic species of West
Malesian species recorded in this study are 23 species. This number are high
enough. Tryon (1970), Smith (1972) and Wagner (1972) have discussed the high
proportion of pteridophytes on oceanic islands. Smith (1972) compiled data from
several islands and showed that ferns have much lower of endemism than
angiosperm at both the genus and species levels. On Hawaii, about 16% of
angiosperms genera are endemic vs. 6.7% of pteridopnytes genera. Ranker et al
(1994) give interpretation for this fact in two fold: first, ferns are more capable of
long-distance dispersal and establishment, second, continued gene flow from
mainland sources may slow speciation rates for ferns, thereby constraining the
evolution of endemic taxa.
Most of endemic species are found mountain forest at 900-1600 m above
sea level. It is evidence that the lower and intermediate elevation of mountain
forest is also the center of endemism areas.
Some endemic species have a relative strict distribution based on
elevation. D. betimusense was found at 300-400 m. D. atrosquamsosum and D.
lobbianum were only found at 1500-2100 and 1600-1800 m, respectively. One
species that has longer distant gradient distribution is D. tricholepis, 100-2100 m.
Some endemic species are found in certain habitats. D. wahauense only
grows at the reophytic areas of Borneo. D. albidosquamatum and D. crinitum
grow in shady places of limestone area. D. betimusense and D. squarrosum were
found growing at shady forest near streams.
This endemic statements, however, can change in accordance with the
science advances. The new exploration and the change of the taxon delimitation
would give new data for a taxon. Consequently taxa that in the past stated as
endemic species would be non endemic species. For example, D. atrosquamoum,
D. barbatum, D. laevipes, and D. vestitum are not endemic to Mount Kinabalu
now (Parris et al 1992).
28
Table 3.1. Distribution of West Malesian Diplazium Inside and Outside Malesia
No. Species Inside MalesiaPM
B S Pa J B Ph C M L S I I J PNGOutsideMalesia
Literatures GeographicalDistributionRange
1. D. accedens Blume V V V V V V V V V V V V North-easternQueensland;Africa,islands ofIndianOcean,Somoa,Thailand.
Present StudyAndrews(1990)Holttum (1940)Tagawa &Iwatsuki(1988)Kato (1992)Mitsuta (1985)
Very wide
2. D. acuminatum Blume V Present study Endemic
3. D. aequibasale (Baker)C.Chr.
V V V V Present studyIwatsuki &Kato (1984)
Malesia
4. D. albidosquamatum Alderw. V Present study Endemic
5. D. angustipinna Holtt.(Holtt.)
Present studyHolttum (1966)
Malesian
6. D. asymmetricumPraptosuwiryo, sp. nov.
V Present study Endemic
7. D. atrosquamosum Copel.)C.Chr. & Holtt.
V Present study Endemic
Note: PM = Peninsular Malaysia, B = Borneo, S = Sumatra, Pa = Palawan, J = Java, B = Bali, Ph = Philippines, C = Celebes, M = Moluccas, LSI = Lesser Sunda Islands, IJ = Irian Jaya, PNG = Papua New Guinea.
29
Table 3.1. Continued
No. Species Inside MalesiaPM
B S Pa J B Ph C M L S I I J PNGOutsideMalesia
Literatures GeographicalDistributionRange
8. D. bantamense Blume V V V V V V Thailand Present studyHolttum (1940)Iwatsuki &Kato (1984)Tagawa &Iwatsuki(1988)Mitsuta (1985)
Very wide
9. D. barbatum C.Chr. inC.Chr. & Holtt.
V Present study Endemic
10. D. batuayauensePraptosuwiryo, sp. nov.
V Present study Endemic
11. D. beamanii M.G. Price V Endemic
12. D.betimusense Alderw. V Present study Endemic13. D. christii C. Chr. V Present study Endemic14. D. cordifolium Blume V V V V V V V V V V V India to
SolomonIsland,Thailand
Present studyHolttum (1940)Tagawa (1972)Tagawa &Iwatsuki(1988)Kato (1992)Mitsuta (1985)
Very wide
30
Table 3.1. Continued
No. Species Inside MalesiaPM
B S Pa J B Ph C M L S I I J PNGOutsideMalesia
Literatures GeographicalDistributionRange
15. D. crameri Praptosuwiryo,sp. nov.
V Present study Endemic
16. D. crenatoserratum (Blume)Moore
V V V V V Thailand Present studyHolttum (1940)Tagawa (1972)Tagawa &Iwatsuki (1988)Kato (1992)Mitsuta (1985)
Very wide
17. D. crinitum (Baker) C..Chr. V Present studyTagawa (1972)Iwatsuki &Kato (1984)
Endemic
18. D. cumingii (Presl) C.Chr. V V Present studyTagawa (1972)Iwatsuki &Kato (1984)
Malesia
19. D.densisquamatumPraptosuwiryo, sp. nov.
V Present study Endemic
31
Table 3.1. Continued
No. Species Inside MalesiaPM
B S Pa J B Ph C M L S I I J PNGOutsideMalesia
Literatures GeographicalDistributionRange
20. D. dilatatum Blume V V V V V V V V V V Thailand,India,Burma, S.China,Taiwan,Ryuku, S.Japan,Indochina,N.Australia.
Present studyHolttum (1940)Tagawa &Iwatsuki(1988)
Very wide
21. D. dolichosorum Copel. V V Present study Malesia
22. D. donianum (Mett.) Tardieu V V V Japan,Taiwan, S.China,Indochina,Thailandand India
Present studyTagawa &Iwatsuki(1988)
Malesia
23. D. esculentum (Retz.)Swartz
V V V V V V V V V V V V Tropic ofAsia toOceania
Present studyHolttum (1940)Tagawa (1972)Tagawa &Iwatsuki(1988)Kato (1992)
Very wide
24. D. fraxinifolium Presl V V V V Present study Malesia
32
Table 3.1. Continued
No. Species Inside MalesiaPM
B S Pa J B Ph C M L S I I J PNGOutsideMalesia
Literatures GeographicalDistributionRange
25. D. fuliginosum (Hook.) M.G.Price
V V V BismarckArch. (New
Ireland)
Present study Malesia
26. D. halimunensePraptosuwiryo, sp. nov.
V Present study Endemic
27. D. hewittii (Copel.) C.Chr V Present studyIwatsuki & Kato(1984)
Endemic
28. D. hottae Tagawa V V Present studyTagawa (1972)Iwatsuki & Kato(1984)
Malesia
29. D. insigne Holtt. V Present studyHolttum (1966)
Endemic
30. D. kunstlerii Holtt. V V V Present studyHolttum (1966)Mitsuta (1985)
Malesia
31. D. laevipes C.Chr. in C.Chr.& Holtt.
V Present studyIwatsuki & Kato(1984)
Endemic
32. D. latisquamatum Holtt. V V V Present studyHolttum (1966)
Malesia
33. D. lobbianum Moore V V V V Present studyVan Alderweldvan Rosenburgh(1908)
Malesia
33
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
34. D. lomariaceum (Christ.)Price
V V V V V V Present studyIwatsuki &Kato (1984)Kato (1992)
Malesia
35. D. loerzingii Praptosuwiryo,sp. nov.
V V Present study Malesia
36. D. malaccense Presl. V V V V Thailand,Indochina
Present studyHolttum (1940)Tagawa &Iwatsuki(1988)Mitsuta (1985)
Malesia
35. D. loerzingii Praptosuwiryo,sp. nov.
V V Present study Malesia
36. D. malaccense Presl. V V V V Thailand,Indochina
Present studyHolttum (1940)Tagawa &Iwatsuki(1988)Mitsuta (1985)
Malesia
37. D.megasegmentumPraptosuwiryo, sp. nov.
V Present study Endemic
38. D.megasimplicifoliumPraptosuwiryo, sp. vov.
V Present study Endemic
39. Diplazium meijeriiPraptosuwiryo
V Present study Endemic
34
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
40. D. melanolepis Alderw. V Present study Endemic41. D. moultonii (Copel.) Tagawa V Present study
Tagawa (1972)Endemic
42. D. pallidum (Blume) Moore V V V V Queensland Present studyHolttum (1966)Kato (1992)Mitsuta (1985)
Very wide
43. D. paralleliveniumPraptosuwiryo, sp. nov.
V Present study Endemic
44. D. petiolare C. Presl. V Present study Endemic45. D. poiense C. Chr. in C.Chr. &
Holtt. V Present study
Iwatsuki & Kato(1984)
Endemic
46. D. polypodioides Blume V V V V V V V V V V V V Thailand,Sri Langka,S. India,Himalaya,Indochina,Taiwan.
Present studyHolttum (1940)Tagawa &Iwatsuki (1988)Mitsuta (1985)
Very wide
47. D. porphyrorachis (Baker)Diels
V Present studyTagawa (1972)
Endemic
48. D. prescottianum (Wall.) Moore V Thailand Present studyHolttum (1940)Tagawa &Iwatsuki (1988)
Endemic
35
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
49. D. procumbens Holtt. V V V Present studyHolttum (1940)
Malesia
50 D. profluens Praptosuwiryo, sp.nov.
V Present study Endemic
51. D. riparium Holtt. V V V V V V Thailand Present studyHolttum (1940)Tagawa (1972)Tagawa &Iwatsuki (1988)Kato (1992)
Malesia
52. D. silvaticum (Bory) Swartz V V V V V Thailand;Mauritus,India,Burma
Present studyHolttum (1940)Iwatsuki & Kato(1984)Tagawa &Iwatsuki (1988)
Malesia
36
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
53. D. simplicivenium Holtt. V V Thailand Present studyHolttum (1940)Tagawa &Iwatsuki (1988)
Malesia
54. D. sorzogonense C. Presl. V V V V Thailand,Vietnam,Indo-China
Present studyHolttum (1940)Iwatsuki & Kato(1984)Tagawa &Iwatsuki (1988)Kato (1992)Mitsuta (1985
Malesia
55. D. speciosum Blume V V V V V Present studyHolttum (1940)
Tagawa (1972)Iwatsuki & Kato(1984)Mitsuta (1985)
Malesia
56. D. spiniferum Alderw. V Present studyTagawa (1972)
Endemic
57. D. squarrasum K. Iwats. & MKato
V Present studyIwatsuki & Kato(1984)
Endemic
37
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
58. D. subintegrum Holtt. V V Thailand Present studyHolttum (1940)Tagawa &Iwatsuki (1988)
Malesia
50. D.subalternisegmentumPraptosuwiryo, sp. nov.
V Present study Endemic
60. D. subpolypodioides Alderw. V V Present study Malesia
61. D. subserratum (Blume) Moore V V V Thailand Present studyHolttum (1940)Tagawa &Iwatsuki (1988)Mitsuta (1985)
Malesia
62. D. subvirescens Praptosuwiryo,sp. nov.
V Present study Endemic
63. D. tomentosum Blume V V V V V Thailand,Burma,Vietnam
Present studyHolttum (1940)Tagawa (1972)TagawaIwatsuki (1988)Kato (1992)Mitsuta (1985)
Very wide
64. D. tricholepis C. Chr. V Present studyTagawa (1972)Iwatsuki & Kato(1984)
Endemic
38
Table 3.1. Continued
No. Species Inside MalesiaPM B S Pa J B Ph C M L S I I J PNG
OutsideMalesia
Literatures GeographicalDistributionRange
65. D. umbrosum (Smith) Beddome V Present study Endemic66. D. velutinum Holtt. V Present study
Holttum (1940)Endemic
67. D. vestitum C. Presl. V V V Present studyKato (1992)
Malesia
68. D. wahauense Kato, Darnaedi etK. Iwatsuki
V Present study Endemic
69. D. xiphophyllum (Baker) C.Chr.
V V V V V Thailand Present StudyHolttum (1940)Tagawa (1972)Tagawa &Iwatsuki (1988)Kato (1992)
Very wide
39
Table 3.2. Endemic Species of Diplazium in West Malesia
No. Species Locality
1. D. albidosquamatum Sumatra: Bengkulu (Lebong Tandai)2. D. asymmetricum
Praptosuwiryo, sp. nov.Java: Mt. Gede, Mt. Halimun (West Java)
3. D. atrosquamosum Borneo: Mt Kinabalu, Tenompok-Humu-humu, Southof Panataran River.
4. D. barbatum Borneo: Mt. Kinabalu (North Borneo), Mt. Besar(South Kalimantan)
5. D. batuayauensePraptosuwiryo, sp. nov.
Borneo: Batu Ayau (Muller Range)
6. D. beamanii Borneo: Mt. Kinabalu7. D. betimusense Sumatra: Betimus River (Sibolangit)8. D. christii Malay Peninsula: G. Muntahak, (Johore)9. D. crameri,
Praptosuwiryo, sp. nov.Sumatra: Kenali (Sukaraja)
10. D. crinitum Borneo: G. Njapa and Long Keluh, Berau (CentralKalimantan)
11. D. densisquamatumPraptosuwiryo, sp. nov.
Sumatra: Bukit Tapan (Jambi)
12. D. halimunensePraptosuwiryo, sp. nov.
Java: Mt. Halimun (West Java)
13. D.megasegmentumPraptosuwiryo, sp. nov.
Java: Mt. Salak (West Java)
14. D.megasimplicifoliumPraptosuwiryo, sp. nov.
Borneo: Bukit Raya
15. D. meijeriiPraptosuwiryo, sp. nov.
Sumatra: Mt. Sago (Payakumbuh)
16. D. melanolepis Sumatra: G. Singgalang17. D. parallelivenium
Praptosuwiryo, sp. nov.Java: Mt. Halimun (West Java)
18. D. profluensPraptosuwiryo, sp. nov.
Java: Mt. Halimun (West Java)
19. D. squarrosum Borneo: Mt. Besar (South Kalimantan)20. D. subalternisegmentum
Praptosuwiryo, sp.nov.Mount Kinabalu, Borneo
21. D. subvirescensPraptosuwiryo, sp. nov.
Java: Mt. Gede and Mt. Halimun (West Java)
22. D. tricholepis Mount Kinabalu, Borneo23. D. velutinum Malay Peninsula: Cameron Highlands24. D. wahauense Borneo: Along Jenta River, Miau Baru, north of
Muara Wahau (Central-East Kalimantan), BelatungRiver, Muller Range (Central Kalimantan)
40
Table 3.3. Species diversity and endemism of Diplazium in four mainlands of West Malesia
Island Account of Species Number Endemics FoundJava 30 6Sumatra 29 6Malay Peninsula 28 2Borneo 40 9
3.4. Conclusions
Sixty nine species of Diplazium are distributed in West Malesia. The total
number of species for each main island are 40, 30, 29, 28 for Borneo, Java,
Sumatra, and Malay Peninsula, respectively. Based on the range of its
geographical distribution, the West Malesian species can be divided into three
types: (1) very wide distribution species, (2) Malesian species, and (3) locally
endemic species.
Most species are only found in Malesia. Nineteen species of West Malesia
distribute very wide. They are D. accedens, D. bantamense, D. cordifolium, D.
crenatoserratum, D. dilatatum, D. donianum, D. esculentum, D. fuliginosum, D.
malaccense, D. pallidum, D. polypodioides, D. prescottianum, D. riparium, D.
silvaticum, D. simplicivenium, D. sorzogonense, D. subserratum, D. tomentosum
and D. xiphophyllum.
Twenty four species are presumed as endemic in the islands of West
Malesia. The highest number of endemic species is occurring in Borneo (9
species): D. atrosquamosum, D. barbatum, D. batuayauense, D. beamanii, D.
crinitum, D. squarrosum, D. subalternisegmentum, D. tricholepis and D.
wahauense. Whereas the lowest number is in Malaya Peninsula (2 species), viz D.
christii and D. velutinum. Six species are endemic to Java and Sumatra. The six
endemic species of Sumatra are D. albidosquamatum, D. betimusense, D. crameri,
D. densisquamatum, D. meijerii, and D. melanolepis, and six species endemic to
Java are D. asymmetricum, D. halimunense, D. megasegmentum, D.
parallelivenium, D. profluens, and D. subvirescens.
41
CHAPTER 4
THE STELAR ANATOMY OF STIPE AND ITS TAXONOMICSIGNIFICANT IN DIPLAZIUM
4.1. Introduction
Anatomical studies of the ferns have had a long and historically
significant place in the professional literatures (e.g. Bower 1912, 1913). These
anatomical studies in particular, played a large part in the conclusions concerning
fern systematic relationships and evolution drawn by Bower. Between the years
of 1960 – 2000, anatomical data have emerged once again as important to an
accurate understanding of relationships and evolution among ferns (e.g. Holttum
& Sen 1961; Bir 1969; Kato 1972; Tryon 1970; Nishida & Nishida 1982; White &
Weidlich 1995; Qiu et al 1995).
Detailed anatomical investigation can serve several useful purposes
including the addition of new knowledge about particular taxa and the useful
application of these kind data on the problems of relationships among fern taxa
(White 1974). Broad and detailed comparative anatomical studies are important
in problems solving of fern systematic and evolution.
One of the kinds of comparative studies usually used in the anatomical
studies on the ferns is vascular (stelar) pattern (White 1974). The importance of
the vascular tissue as a unified tissue system (the stele) was recognized in the late
19th century (e.g. Tiegham & Douliot 1886). Reviewing from the many cases
studies on vascular pattern of ferns, White (1974) came in a concussion that stelar
anatomy is a potentially powerful tool in systematic although the vascular tissue
data alone do not provide the basis on which systematic (or taxonomic)
conclusions are based.
The stelar anatomy of Athyrioid ferns and their relatives has been studied
in detail by some pteridologist (e.g. Tardieu-Blot 1932, Bir 1969, Kato 1972). All
these studies came to the concussion that anatomical evidence is important to
support taxa delimitation. This chapter presents the anatomical data of stipe on 27
species of West Malesian Diplazium. The aim of this study was to collect stelar
anatomical data to support species delimitation in Diplazium.
42
4.2. Materials and Methods
Transection of frond axis, stipe near lamina (upper portion of stipe), were
studied on 27 species collected from West Malesia . A pieces of stipe near blade
of fresh material were hand sectioned with a sharp razor blade. The sections were
embedded in glycerine jelly after staining them with 0.5% methyl green to obtain
semi permanent slides.
4.3. Results and Discussion
As also reported by Praptosuwiryo (1999), this study resulted that all
species under observation have strand of vascular tissue termed amphicribal
vascular bundles. This structure consist of a central strip of xylem completely
surrounded by phloem. Foster & Gifford (1959) termed meristele for this such
concentric strand of vascular tissue. The meristele in the stipe is embedded within
a conjunctive parenchyma which posses large intercellular spaces. Between the
cortical parenchyma and the single layered cuticularized epidermis is a band of
hypodermal schlerenchyma whose fiber are heavily lignified. Meristele structure
in the rhizome and leaf strands is the usual type for Diplazium, Athyrium,
Diplaziopsis, etc. (Tardieu-Blot 1932, Bir 1969). Therefore this structure is not an
important for diagnostic feature which supports the taxonomic separation among
the species of Diplazium.
Leaf-trace are binary. As showed in Figure 4.1. the xylem of a leaf-trace
of most species are same hippocampus-shaped bundle in transaction. The shape
and position of this vascular bundle are varying among species. The similar
anatomy is illustrated by Tardieu-Blot (1932) and Kato (1977). In D.
polypodioides and D. subpolypodioides, the xylem of the leaf-trace develop ridges
and grooves become somewhat W-shaped. This shape is similar with D.
latifolium illustrated by Bir (1969).
The leaf-trace shapes of the stipe, as a whole, are varying among species
and constant among the adult individuals in a species (Figure 4.2. and 4.3.). The
shapes are comprised of five types: (1) uninterrupted V-shaped (Figure 4.2.a-c,
e.), (2) interrupted V-shaped (Figure 4.2.g.), (3) uninterrupted U-shaped (Figure
4.2.i. ), (4) interrupted U-shaped (Figure 4.2.h & k), and (5) W-shaped (Figure
43
4.3.i.). These types similar those illustrated by Tardieu-Blot (1932), Kato (1977)
and Bir (1969).
Figure 4.1. Vascular structure of the leaf axis. a. D. cordifolium; b. D. esculentum;c. D. umbrosum; d. D. procumbens. e. D. polypodioides. Bar = 1 mm for b, c, d, and e. Bar = 0.4mm for a.
The uninterrupted V-shaped leaf-trace is seen in D. tomentosum, D.
angustipinna, and D. cordifolium. Interrupted V-shaped is shown in D.
silvaticum. Interrupted U-shaped leaf-trace is seen in D. accedens var. spinosum,
D. batuayauense and D. D. subserratum.. Most species are possessing
44
uninterrupted U-shaped leaf trace, such D. halimunense, D. procumbens, D.
sorzogonense, D. speciosum, D. subserratum, D. umbrosum and D. vestitum. As
mentioned above, W-shaped leaf trace is found in D. subpolypodioides and D.
polypodioides. Some species may have intermediate shape between two types. For
example, D. riparium and D. crenatoserratum have the intermediate shape between type
one and four.
Figure 4.2. Leaf-trace shapes in Diplazium. Groove U-shaped with flat base. a. D. tomentosum;b. D. cordifolium (simple frond); c. D. angustipinna; d. D. silvaticum var. silvaticum;e. D. riparium; f. D. xiphophyllum; g. D. crenatoserratum h. D. subserratum;i. D. subvirescens; j. D. accedens var. accedens; k. D. accedens var. spinosum. Bar = 2 mm fora, c, d, g, and h. Bar = 1.6 mm for b and e. Bar = mm for f and d. Bar = 2.5 mm for j and k.
45
Figure 4.3. Leaf-trace shapes in Diplazium. a. D. halimunense; b. D. donianum;c. D. simplicivenium; d. D. sorzogonense; e. D. procumbens; f. D. vestitum; g. D. speciosum;h. D. subpolypodioides; i. D. polypodioides; j. D. umbrosum; k. D. spiniferum;l. D. batuayauense. Bar = 1.5 mm for a, b, d and g. Bar = 1 mm for l. Bar = 2 mm for c, e, f, h,i, j, and k.
46
Each type of the leaf traces or the vascular bundle may diversify into the
derivative forms. These character variations can be used to determine a species
among closely related species. D. speciosum and D. sorzogonense are
morphologically very similar. Anatomically, the two species share characters:
vascular bundle form uninterrupted U-shaped, flat base on the two directions,
inward and outward (Figure 4.3.). But, the U-shaped vascular bundle of first
species is with an angle 90º and end slightly ridge, whereas the second species
with an angle 110º and end almost simple. Even, the vascular bundle type of D.
acedens var. accedens (Figure 4.2.j.) is different from D. accedens var. spinosum
(Figure 4.2.k.). Var accedens has an uninterrupted U-shaped vascular bundle
with ridges that formed at the outward of lower base, angles, and ends.
Meanwhile var. spinosum has an interrupted U-shaped with more blunt ridges on
the angle and end. Therefore the leaf-trace shapes are important diagnostic
features that support species delimitation in Diplazium.
4.4. Conclusions
The leaf-trace shape of Diplazium stipe is varying among species and
constant among the adult individuals in a species. Based on the result of this
study and also previous authors (Tardieu-Blot 1932, Bir 1969, Kato 1977) it is
concluded that the leaf-trace shapes of Diplazium can be classified into five main
types: (1) uninterrupted V-shaped, (2) interrupted V-shaped, (3) uninterrupted U-
shaped, (4) interrupted U-shaped, and (5) W-shaped. Each type seems to vary among
the species. Each type may diversify into some different derivative forms that enables to
determine a species among closely related species. Therefore the leaf-trace shapes are
important diagnostic features which support species delimitation in Diplazium.
47
CHAPTER 5
CYTOLOGICAL AND REPRODUCTIVE STUDIES ON DIPLAZIUM INWEST MALESIA
5.1. Introduction
Cytological studies of ferns have given a big influence on the classification
on ferns and ferns allies dating from the publication of Manton (1950), ‘Problems
of Cytology and Evolution in the Pteridophyta’, in which a new simple techniques
was introduced. Her book inspired a great upsurge in chromosomes work on the
ferns and fern allies not only in Europe but also by authors in North America,
India, Japan and New Zealand (Tindale & Roy, 2002). However, by 1977 it was
estimated by Love et. al. that less than 20% of the Pteridophyta had been
examined cytologically, although few genera have escaped attention since that
date. Love et. al. (1977) stated that there are about 3.200 species referred to 446
genera and 65 families on the basis of Pichi Sermolli’s classification (1977).
As reported by Löve et al (1977), cytological observations of Ceylon
Diplazium by Manton (1953) was preliminary cytological research of Diplazium.
From 1953 to 1977 were only about 15.5% of ca. 400 species of Diplazium in the
world known their chromosomal information. In Malesia, it is predicted that
cytological information since Manton (1954) to Praptosuwiryo & Darnaedi (2004)
were only 6% of ca.300 species of Diplazium.
Polyploidization and mode reproduction types has played an important
role in the geographical distribution and evolution of Diplazium. Polyploids,
which contain more than two sets of chromosomes per cell, have important in
plant evolution (Stebbins 1971; Lewis 1980). Up to 95% of pteridophytes may be
polyploid (Grant 1981) and recent polyploids may comprise 45% of extant
homosporous fern (Vida 1976; Haufler & Soltis 1986). Polyploidy may lead to
significant reproductive isolation (Ramsay & Schemske, 1998), and change in
gene expression (Adams et al 2003), and ecological interactions (Segraves et al
1999).
48
Cytological observation have clearly disclosed many apomictic species
within Diplazium (Lovis, 1977). Apomixis is an asexual reproduction via
chromosomal unreduction of spores and gametophytes, and the subsequent
apogamous reproduction (in a strict sense), i.e., asexual reproduction of a
sporophyte from somatic cells of the gametophyte. About 10% of fern species in
world are apomictic (Lovis 1977). About 15% of Japanese ptridophytes are
apomictic (Takamiya 1996, 1997). How many percent of apomictic species of
Malesian pteridophytes are not known because the cytological observations of
peridophytes have not been done completely.
Since the late 20th, cytological and reproductive studies on Japanese
Diplazium have been doing intensively. Takamiya et al (1999) examined 162
plants representing 16 species of Japanese Diplazium, including polymorphic
terrestrial with evergreen bi- to tripinnate leaves. Their studies clearly give
clarification the morphological complexities on some species, moreover would be
a good evident in clarifying its taxomical status. Mitotic and meiotic
chromosomes studies on 80 plants representing seven species with summer-green
bi- to tripinnate leaves were also unraveled many cases of the morphological
complex in Japanese Diplazium (Takamiya et al. 2000; 2001).
Cytological researches on Diplazium in Malesian region are not so many.
Cytological information on Diplazium were firstly reported by Manton (1954) and
Holttum & Roy (1965). The first reported 11 species of Diplazium of Malaya,
and the second observed 4 species of Papua New Guinea. In Java, preliminary
cytological observation of Javanese Diplazium was conducted by Darnaedi (1992)
in which he found tetraploid D. cordifolium and diploid D. esculentum.
Furthermore, Praptosuwiryo and Darnaedi (1994) reported chromosome numbers
of 6 species of Diplazium (one species of its, viz. D. opacum has been changed to
Cornopteris opaca) of Gede-Pangrango National Park. Recent cytological
informations for Malesian region were reported by Praptosuwiryo (2003),
Praptosuwiryo & Darnaedi (2004). The first reported two species tetraploid
Diplazium of Lombok Island, viz. D. malaccense and D. pallidum. While the
second reported cytological research of 10 species of Javanese Diplazium
included 43 collection number of 10 localities.
49
This chapter reports cytological and mode reproduction type studies of
Diplazium from West Malesia included 31 species. This aims of this studies are;
(1) to examine somatic chromosomes number of Diplazium; (2) to determine the
mode reproduction type of Diplazium; (3) to recognize the correlation between
ploidy level and morphological variation in the Diplazium species; 4) to recognize
the correlation between ploidy levels and its habitats.
5.2. Materials and Methods
Living plants of Diplazium included in 33 species and 117 collection
number were collected from 54 localities in West Malesia (Table 5.1.). Living
materials were planted in the green house at Bogor botanic Gardens. Now, the
voucher specimens are deposited in the herbarium of Bogor Botanic Gardens
(BOHB) and its duplicates will be distributed to Herbarium Bogoriense (BO).
Cytological investigations were carried out by observing the somatic
chromosomes at the root tips following a procedure developed by Manton (1950)
and modified by Darnaedi (1991). Root tips pretreated with 0.002 M 8-
hydroxyquinolin for 4-6 hrs at about 18-20°C, and then fixed with 45% acetic acid
for 10 minutes. The fixed roots were macerated with a mixture of 45% acetic acid
and 1 N HCl (1:3) for 3-4 minutes at 60°C, and then stained with 1% aceto-orcein
solution. Chromosomes counting were carried out during observation under a
light microscope with 100x objective lens, and photographs were taken by Nikon
Camera in Herbarium Bogoriense.
Mode reproduction types were determined by counting spores per
sporangium. Sporangium with 64 normal spores treated as sexual reproduction,
while the sporangium with 32 spores treated as apogamous one.
50
5.3. Results and Discussion
Somatic chromosome account and mode reproductive type of Diplazium
from West Malesia included 33 species and 117 collections number collected
from 54 localities are reported (Table 5.1.). All species examined revealed basic
chromosome number x = 41. These are consistent with those reported by many
recently researchers on cytology of Diplazium, such as Ohta & Takamiya (1999)
and Takamiya et al. (1999, 2000, 2001). The 33 species of Diplazium successfully
examined its chromosomes number and mode reproduction type are discussed as
follow.
5.3.1. Chromosome Number Variations and Mode Reproduction Types onDiplazium
Diplazium accedens Blume. Eight individuals of D. accedens revealed
2n = 82 (diploid), three individuals from East Kalimantan and five individuals
from five localities in East Java and West Java.. Somatic chromosomes number of
D. accedens from Kalimantan is firstly report in this study. Eighteen individuals
of plant from four localities in East Java and West Java reported by Praptosuwiryo
& Darnaedi (1994 and 2004) also showed diploid level. Total of 23 plants are
consistently diploid sexual. Manton (1954) in Malaya also reported diploid level
only. Since we understand that ploidy is derived from the diploid one, we
presumed that D. eccedens is a good species or probably native to Java.
Diplazium aequibasale (Baker) C.Chr. Only one individual of D.
aequibasale succeded to be examined and showed tetraploid (2n=164). It is the
first cytological report for this species. There is not a cytological observation
reported yet except this study.
Diplazium angustipinna Holttum. There are two levels ploidy found in D.
angustipinna, viz. triploid (2n=123) and tetraploid (2n=164) from Borneo. The
two cytotypes are the first cytological record for Bornean Diplazium, and the
triploid one is also the first record for science. Manton (1954) reported only n =
82 for Malayan plant. However it is not recognized whether this plant is sexual
tetraploid or apogamous diploid because its somatic chromosome number was not
51
reported. Two specimens types from two different ploidy levels, 2n=123 dan
2n=164, can not differentiated.
Diplazium asymmetricum Prtaptosuwiryo, sp. nov. Four individuals (TNgP
1094, TNgP 1334, and TNgP 1732 ) of this species has been observed
cytologically and showed 2n = 123 (triploid). The small frond of Diplazium
asymmetricum is in a glance similar to those of D. procumbens. However D.
asymmetricum can be differentiated from D. procumbens by the following
characters combination: rhizome short, erect; lamina more incised (up to
tripinnate) basiscopic pinnulae or segments and lobes are larger than the
acroscopic ones (acroscopic and basiscopic pinnuale or segments and lobes
asymmetric); indusia thicker, margin entire. While D. procumbens has rhizome
long creeping; acroscopic and basiscopic pinnulae and lobes almost symmetric;
indusia thin and lacerate at margin.
Diplazium bantamense Blume. Four individuals of D. bantamense
collected from Cangkuang Forest, Mt. Salak, West Java, found to be tetraploid
race. Two individuals are successfully examined their reproduction types. TNgP
1212 showed 32 spores per sporangium presumed as apogamous type and TNgP
1454 showed 64 spores per sporangium as sexual type. The others two plants
have not been recognized their reproduction type. Most of individuals
successfully examined were tetraploid sexual (Praptosuwiryo & Darnaedi, 1994).
As pointed out by Kato (1992), at diploid gametophytes level, sexual species
(tetraploid in sporophyte) are more abundant than apogamosporous ones. Kato
(1992) also explained the hypothetical origins of tetraploid agamosporous type.
Tetraploid agamosporous type may be derived from a crossing between diploid
sexual species and triploid agamosporous. While, triploid agamosporous are
derived from diploid and tetraploid sexual species, as a demonstrated by
Dryopteris sparsa compelx (Darnaedi et al 1989).
Based on this analysis, it is presumed that in Java, diploid sexual and
triploid apogamous of D. bantamense may exist in nature. However up to now
the information concerning the existence of both diploid sexual and triploid
apogamous are not exist. Further cytological survey of this species is urgently
needed to confirm whether the diploid ancestral is still exist or not.
52
Table 5.1. Somatic Chromosomes numbers, Ploidy Level and Mode Reproduction Type of Diplazium from West Malesia
Species ChromosomesNumbers (2n)/ PloidyLevel/ ReproductiveType
VoucherSpecimens
Locality
D. accedens 82/diploid/82/diploid/82/diploid/82/diploid/
82/diploid/
82/diploid/82/diploid/seksual82/diploid/seksual83/diploid/seksual
TNgP1001TNgP1447TNgP1786TNgP1399
TNgP1649
TNgP1211TR 53.1TR 53.2TR 53.3
Gede-Pangrango National ParkG. Salak, West Java. Ca. 1500 s.l.G. Halimun, West Java. Ca. 950 m s.l.Cidaon-Cibunar, Ujung Kulon NationalPark, West Java.Gondang Forest, G. Wilis Utara, Kec.Kare, Kab. Madiun, East JavaG.Salak, West JavaEast KalimantanEast KalimantanEast Kalimantan
D. aequibasale 164/tetraploid/ TNgP2026 Bank of Sungai Abang Ai, TNBD, JambiD. angustipinna 123/triploid/
ca.164/tetraploid/
TNgP1904
TNgP1905b
Kobet Forest, near S. Kobet, trek to BatuAyau, Peg. Muller, KALTENG. 440 m.Kobet Forest, near S. Kobet, trek to BatuAyau, Peg. Muller, KALTENG. 440 m.
D. asymmetricumPraptosuwiryo
123/triploid/apogamous
123/triploid/apogamous
123/triploid/apogamous
TNgP 1094
T.Ng.P 1334
TNgP 1732
Petak 4 Desa Kemutuk Lor Wana WisataBaturraden, G. Slamet, Central Java. 970-1000 m. a.s.l.Cibodas Forest, behind Cibodas BotanicGardens, Mt. Gede, Gede-PangrangoNational Park, West Java. ca. 1450 mTrack Cikaniki – Citalahap, HalimunNational Park, West Java. Ca. 1000 m.
D. bantamense 164/tetraploid/apoga-mous164/tetraploid/
164/tetraploid/Sexual
164/tetraploid/
-/-/Sexual
-/-/Sexual
-/-/Apogamous
TNgP1212
TNgP1321
TNgP1454
TNgP1383
TNgP1516
TNgP1517
TNgP1707
Cangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeG. Slamet, Track to Sang Hyang Ropoh,Central Java. 1060 m s.l.G. Slamet, Track to Sang Hyang Ropoh,Central Java. 1060 m s.l.Cikaniki Forest, Halimun National park,West Java
D. batuayauensePraptosuwiryo
164/tetraploid/-
Ca. 205/pentaploid/-
TNgP1927c
TNgP1909
Above Sungai Talikot Puhung Kucan,Batu Ayau, Muller Range, CentralKalimnatan, Borneo. 450 m.Kobet Forest, near S. Kobet, track to BatuAyau, Muller Range, Central Kalimantan,Borneo. 440 m s.l.
53
Table 5.1. Table continued
Species ChromosomesNumbers (2n)/PloidyLevel/Reproductive Type
VoucherSpecimens
Locality
D. cordifolium“pinnate fronds”“simple fronds”
“pinnate fornds”
“pinnate fronds”
“pinnate fornds”
“simple fronds”
“simple frond”
“pinnate fronds”
“pinnate fronds”
“pinnate fronds”
“pinnate fronds”
“pinnate fronds”
“simple fronds”
‘pinnate frond’
“pinnate frond”
“pinnate frond”
“pinnate frond”“simple fronds”
“simple fronds”
“simple fronds”
D. cordifolium‘pinnate fronds’D. cordifolium‘simple fronds’
164/tetraploid/
Ca.246/hexaploid/
Ca.205/pentaploid/
164/tetraploid/
Ca.205/pentaploid/
Ca.205/pentaploid/
Ca.164/tetraploid/
164/tetraploid/
ca205/pentaploid/
-/-/Sexual
-/-/Sexual
-/-/Sexual
164/tetraploid/
164/tetraploid/
164/tetraploid/
ca.205/pentaploid/
164/tetraploid/205/pentaploid/
164/tetraploid/
164/tetraploid/
ca. 164/tetraploid/ .
328/octoploid/
TNgP1194
TNgP1201
TNgP1307
T NgP1375
TNgP1355
TNgP1204
TNgP1203
TNgP1442
TNgP1441
TNgP1456
TNgP1457
TNgP1460
TNgP1458
TNgP1735
TNgP1774
TNgP1808
TNgP1736TNgP1813
TNgP2284
TNgP2281
TNgP1910
TNgP 1926b
Cangkuang Forest, G. Salak-West Java,Southern SlopeCangkuang Forest, G. Salak-West Java,Southern SlopeCangkuang Forest, G. Salak-West Java,Southern SlopeCangkuang Forest, G. Salak-West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java.Southern SlopeCangkuang Forest, G. Salak-West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak-West Java,Southern SlopeCurug Macan Trail, Cikaniki Forest, G.Halimun, West JavaOwa Trail, Cikaniki Forest, G. Halimun, WestJavaPlot II/C6, Cikaniki Forest, G. Halimun, WestJava. Ca. 950 m.Loop Trail, G. Halimun, West Java.Plot I, near Tea Plantation, Cikaniki ResearchStation, G. Halimun, West Java. Ca. 950 m.Track to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West JavaTrack to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West Java.Near Sungai Anak Kobet, track to Batu Ayau,Batu Ayau. 440 m s.l.Near Sungai Anak Kobet, track to Batu Ayau,Mts. Muller, Desa Tumbang Topus, Kec.Tumbang Konyi, Kab. Murung Raya, CentralKalimantan. ca. 450 m.
54
Table 5.1. Table continued
Species ChromosomesNumbers (2n)/ PloidyLevel/ Reproductive Type
VoucherSpecimens
Locality
D. cordifolium‘pinnate fronds’
164/tetraploid/ ...
ca. 164/tetraploid/ ...
164/tetraploid
TNgP2128
TNgP2129
TNgP2342b
Track S. Sopan-Bukit Batikap, between PonotPenanginan-Camp Lapangan Heliped II, ca.6 msebelah Timur S. Joloi, near Tanjakan NikkiArdilla, Mts. Muller, Central Kalimantan. 240 mTrek S. Sopan-Bukit Batikap, between PonotPenanginan-Camp Lapangan Heliped II, ca.6 mthe eastern of Sungai Joloi, near Tanjakan NikkiArdilla, Peg.Muller, Central Kalimantan. 240 mCikuda Paeh, G. Halimun, West Java
D. crenatoseratum 164/tetraploid/
164/ tetraploid/
164/ tetraploid/
164/tetraploid/
ca. 164/ tetraploid/
ca. 123/triploid/apogami
TNgP2067a
TNgP2026a
TNgP2026c
TNgP2044
TNgP1971
TNgP2075
Bukit Pal, Taman Nasional Bukit Dua Belas,Jambi, Sumatra. ca. 70 m.Bank of Abang Ai, Taman Nasional Bukit DuaBelas, Jambi, Sumatra. 20 m.Bank of Abang Ai, Taman Nasional Bukit DuaBelas, Jambi, Sumatra. 20 m.Bukit Air Keruh, Taman Nasional Bukit DuaBelas, Jambi, Sumatra. 50 m.Air Terjun Tomoroh, S. Belatung, Peg. Muller,KALTENG. 150 m.Bukit Suban Punai Banyak, Taman Nasional BukitDua Belas, Jambi, Sumatra. Ca. 55m.
D. dilatatum 123/triploid/apogamous
123/triploid/apogamous
123/triploid/apogamous
123/triploid/apogamous
123/triploid/apogamous
123/triploid/apogamous
TNgP1073
TNgP1339
TNgP1343
TNgP1011b
TNgP1025
TNgP1526
Petak 55, Desa Karang Mangu, Wana WisataBaturraden, G. Slamet, Central JavaG. Gede, Gede-Pangrango National Park, WestJavaG. Gede, Gede-Pangrango National Park, WestJavaG. Gede, Gede-Pangrango National Park, WestJavaG. Gede, Gede-Pangrango National Park, WestJavaG. Pangrango, 1925 m dpl, Gede-PangrangoNational Park
D. donianum 164/tetraploid/ XIX.C.III.65 Bukit Ubar, Sumatra. Cultivated in Bogor BotanicGardens.
D. esculentum 82/diploid/-
82/diploid/-82/diploid/-
TNgP1291
TNgP1784TNgP2094
Pasir Buntu, Geger BentangG. Pangrango, West JavaCikuda Paeh Trail, Cikaniki Forest, G. HalimunHutan Sungai Air Keruh, TNBD, Jambi
D. profluensPraptosuwiryo
164/tetraploid/ TNgP1798 Canopy Bridge Trail, Cikaniki Forest, G. Halimun
D. halimunensePraptosuwiryo
123/triploid/apogami TNgP 2341b Cikuda Paeh, G. Halimun, West Java
D. hewittii 123/triploid/apogami TNgP1913b Hutan Batu Ayau, Peg.Muller, KALTENGD. loerzingii 82/diploid/-
123/triploid/apogamiTNgP 2339cTNgP 2339d
Cikuda Paeh, G. Halimun, West JavaCikuda Paeh, G. Halimun, West Java
D. megasegmentumPraptosuwiryo
-/-/ Apogamous-/-/ Apogamous
TNgP1451TNgP1452
Cangkuang Forest, G. Salak, West Java, SouthernSlopeCangkuang Forest, G. Salak, West Java, SouthernSlope
55
Table 5.1. Table continued
Species ChromosomesNumbers (2n)/ PloidyLevel/ Reproductive Type
VoucherSpecimens
Locality
D. petiolare 82/diploid/-
82/diploid/-
TT993.2
TT993.3
Cagar Alam Rimbo Panti, Kec. Panti, Kab.Pasaman, West Sumatra. 240-900 m dpl.Cagar Alam Rimbo Panti, Kec. Panti, Kab.Pasaman, West Sumatra. 240-900 m dpl.
D. pallidum 164/tetraplioid
164/tetraploid
164/tetraploid164/tetraploid82/diploid/82/diploid/ -ca.82/diploid82/diploid82/diploid
TNgP1377
TNgP1764
TNgP 1151TNgP1172TNgP1406TNgP2088TNgP2036cTNgP2045TNgP2192b
Cangkuang Forest, G. Salak, Southern slope, WestJava. Ca. 1250 m s.l.Jalur G. Andam, Cikaniki Forest, HalimunNational Park. Ca. 1200 m s.l.Cibodas, G. Gede, JABAR. ca. 1300 m.Cibodas, G. Gede, JABAR. ca. 1300 m.G. Payung, T.N. Ujung Kulon, JABAR. 200 mBukit Lubuk Semak, TNBD, Jambi, 85 mHutan Sukodibeyung, TNBD, Jambi. ±30 m.Hutan Sukodibeyung, TNBD, Jambi. 45 m.S. Joloi, Peg. Muller, KALTENG. ±240 m.
D. polypodioides 82/diploid/82/diploid/
82/diploid/
82/diploid
TNgP1647TNgP1604
TNgP1812
TNgP2285
Gondang Forest, 1000 m s.l., G. Wilis, East JavaBurning Forest of Ngliman, Pace Subdistrict, G.Wilis, East JavaPlot I, near Nirmala Plantation, Cikaniki, G.Halimun, West JavaTrack to Curug Seribu, Taman Wisata AlamGubung Salak Indah, G. Salak, West Java
D. procumbens 123/triploid/Apogamous
123/triploid/Apogamous
-/-Apogamous
-/-/Apogamous
-/-/Apogamous
-/-/Apogamous
TNgP1312
TNgP1348
TNgP1173
TNgP1384
TNgP1453
TNgP1693
G.Gede, Gede-Pangrango National Park, WestJavaCangkuang Forest, Southern Slope of G. Salak,West JavaRawa Denok II, Gede-Pangrango National Park,West JavaCangkuang Forest, Southern Slope of G. Salak,West JavaCangkuang Forest, Southern Slope of G. Salak,West JavaTrack to Wilis Water Fall, Southern Slope of G.Wilis, Dusun Turi, Kec. Geger, Kec. Sendang,Kab. Tulung Agung, East Java
D. porphyrorachis ca. 164/tetraploid TNgP1885 S. Ruhai, G. Pumpung Sapat, Batu Ayau, Peg.Muller, KALTENG. 260 m dpl.
D. riparium ca. 82/diploid
123/triploid
TNgP2147
TNgP 1847
Tract to Sungai Sopan-Bukit Batikap, betweenCamp. Lapangan Heliped II – Sungai Sopan, ca.14 km from Camp. Lapangan Heliped II, ca. 150m to the East of Sungai Joloi, Mt. Muller ProtectedArea. Ca.240 m s.l.Track to Batu Ayau, Muller-Range, CentralKalimantan. 280 m
56
Table 5.1. Continued
Species ChromosomesNumbers (2n)/ PloidyLevel/ Reproductive Type
VoucherSpecimens
Locality
D. simplicivenium 123/triploid/apogamous
123/triploid/apogamous123/triploid/apogamous
123/triploid/apogamous-/-/apogamous
-/-/apogmous
TNgP1229
TNgP1386*TNgP1343
TNgP1371TNgP1179
TNgP 1523
Track Honje Warak-Cibatu Lawang, Gede-Pangrango National Park, West Java. 1545 mCangkuang forest, G. Salak, West JavaCibodas, G. Gede, Gede-Pangrango NationalPark, West Java. 1500 m s.l.Cangkuang Forest, G. Salak, Southern SlopeCibodas Forest, Mt. Gede, Gede-PangrangoNational Park, West JavaCibodas, G. Gede, Gede Pangrango NationalPark, West Java. 1520 m s.l.
D. silvaticum var. silvaticum
ca.164/tetraploid/-ca.164/tetraploid/Sexualca.164/tetraploid/-ca.164/tetraploid/-164/tetraploid/-
164/tetraploid/-
TNgP1300TNgP1301TNgP1302TNgP1303TNgPs.n.21May04TNgPs.n.11May04
Wild Ferns of Bogor Botanic GardensWild Ferns of Bogor Botanic GardensWild Ferns of Bogor Botanic GardensWild Ferns of Bogor Botanic GardensWild Ferns of Bogor Botanic Gardens
Wild Ferns of Bogor Botanic Gardens
D. silvaticumvar.pinnae-ellipticum
123/triploid/apogamous-/-/apogamous123/triploid/apogamous123/triploid/apogamous123/triploid/apogamous-/-/apogamous-/-/apogamous
TNgP2001aTNgP 2007bTNgP2085dTNgP2119cTNgP2019aTNgP 2016aTNgP 2028c
Bukit Lubuk Semak, TNBD, Jambi. 70 m.TNBD JambiBukit Berumbung, TNBD, Jambi. 25 m.Bukit Berumbung, TNBD, Jambi. 25 m.Pematang Berumbung, TNBD, Jambi, 15 m.Pematang Berumbung, TNBD, Jambi, 15 m.Pematang Berumbung, TNBD, Jambi, 15 m.
D. speciosum var.speciosumvar. speciosum
var. speciosum
var. majorvar. speciosumvar. major
82/Diploid/Sexual
82/diploid/Sexual
82/diploid/Sexual
82/diploid/82/diploid/82/diploid/
*TNgP1363
*TNgP1366
TNgP1803
TNgP1727TNgP1745TNgP1758
Cangkuang Forest, G. Salak Nature Reserve,Southern Slope, West Java. Ca. 1600 m s.l.Cangkuang Forest, G. Salak Nature Reserve,Southern Slope, West Java. Ca. 1600 m s.l.Jalur Plot II, Cikaniki Forest, G. Halimun,West JavaCikaniki-Citalahap, G. Halimun, West JavaJalur Owa, Cikaniki Forest, West JavaJalur G. Andam, G. Halimun, West Java
D. procumbens 123/triploid TNgP1344 G.Gede, Gede-Pangrango National Park,West Java
D. speciosum 82/diploid/Sexual82/diploid/Sexual
TNgP1359TNgP1362
Hutan Cangkuang, G. Salak, JABAR. 1600 m.Hutan Cangkuang, G. Salak, JABAR. 1600 m.
D. sorzogonense 82/diploid/Sexual82/diploid/-82/diploid/-82/diploid/-
TNgP1803TNgP1727TNgP1745TNgP1758
Jalur Plot II, Cikaniki , G. Halimun, JABARJalur Cikaniki-Citalahap, G. Halimun,JABARJalur Owa, Cikaniki, JABARJalur G. Andam, G. Halimun, JABAR
57
Table 5.1. Continued
Species ChromosomesNumbers (2n)/ PloidyLevel/ Reproductive Type
VoucherSpecimens
Locality
D. spiniferum(indiv.)(27)
82/diploid/-82/diploid/seksual82/diploid/
TNgP1896aTNgP1896cTNgP1896b
G. Pumpung Sapat, Peg. Muller, KALTENGG. Pumpung Sapat, Peg. Muller, KALTENGG. Pumpung Sapat, Peg. Muller, KALTENG
D. subpolypodioi- des
82/diploid/
-/-/sexual
82/diploid/sexual
-/-/sexual
TNgP1825
TNgP 2285
TNgP2292
TNgP 2303
Track to Perkebunan P.T. Gayatri , G. PasirPacet-G. Galunggung, Lingamulya, Kec.Leuwisari, Kab. Tasikmalaya, West Java.Curug Seribu, Gunung Salak Indah, G.Salak, West Java. 940 mTrack to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, WestJava.940 m.Track to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West Java.940 m.
D. subserratum ca.123/triploid/apogamous
-/-/Sexual
-/-/sexual
-/-/sexual
-/-/sexual
82/diploid/ Sexual
164/tetraploid/
164/tetraploid/
82/diploid/
82/diploid/
123/triploid/
-/-/sexual
-/-/sexual
-/-/sexual
-/-/sexual
-/-/sexual
-/-/sexual
TNgP1072
TNgP1439
TNgP1458
TNgP1459
TNgP1462
TNgP1463
TNgP1379
TNgP1380
TNgP2282
TNgP2283
TNgP2287
TNgP1444
TNgP1705
TNgP1706
TNgP 2282
TNgP 2287
TNgP 2289
Secondary foreest, Petak 55, Desa KarangMangu, Mt. Slamet, Wana WisataBaturaden, Central JavaCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, West Java,Southern SlopeCangkuang Forest, G. Salak, Southern slope,West JavaCangkuang Forest, G. Salak, Southern slope,West Ja va,Cangkuang Forest, G. Salak, Southern slope,West JavaTrack to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West JavaTrack to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West JavaTrack to Curug Seribu, Taman Wisata AlamGunung Salak Indah, G. Salak, West JavaCangkuang Forest, G. Salak, West Java,Southern SlopeCikaniki Forest, Halimun National park,West Java. 1000 m.Cikaniki Forest, Halimun National park,West Java. 1000 m.Curug Seribu, Gunung Salak Indah, G salak,West Java. 940 mCurug Seribu, Gunung Salak Indah, G salak,West Java. 940 mCurug Seribu, Gunung Salak Indah, G salak,West Java. 940 m
58
Table 5.1. continued
Species ChromosomesNumbers (2n)/ PloidyLevel/ Reproductive Type
VoucherSpecimens
Locality
D. subserratum -/-/sexual
-/-/sexual
TNgP 2290
TNgP 2294
Curug Seribu, Gunung Salak Indah, G salak,West Java. 940 mCurug Seribu, Gunung Salak Indah, G salak,West Java. 940 m
D.subvirescensPraptosuwiryo
123/triploid/apogamous
123/ triploid.apogamous
TNgP 1177
TNgP 1013
Cibodas forest, behind Cibodas BotanicGardens, Mt. Gede, Gede-PangrangoNational Park, West Java.Cibodas forest, behind Cibodas BotanicGardens, Mt. Gede, Gede-PangrangoNational Park, West Java.
D. tomentosum 82/diploid/
164/teraploid
205/pentaploid/
TNgP2336b
TNgP2066
TNgP1722
Cikuda-Paeh, Mt. Halimun, West Java. Ca.1300 m.Near Kali Talun – Air Terjun talun, BukitPal, Taman Nasional Bukit Dua Belas.Ca.70 m.Loop Trail, Cikaniki, Mt. Halimun, WestJava. Ca. 1000 m.
D. umbrosum 82/diploid/ TNgP1348 Cibodas forest, behind Cibodas BotanicGardens, Mt. Gede, Gede-PangrangoNational Park, West Java. ±1400 m
D. xiphophyllum 82/diploid/-
±164/tetraploid/-
246/hexaploid/-
164/tetraploid/-
164/teraploid/-
TNgP 1841
TNgP2040b
TNgP1190
RI 867
TD902
Track to Batu Ayau, Peg. Muller, CentralKalimantan. Ca. 280 mRawa Edam, Sukodibeyung SecondaryForest, in TNBD Sumatra.Hutan Cangkuang, G. Salak, West Java. Ca.1200 m.Malampah Nature Reserve, Pasaman Barat,West Sumatra. 950 m dpl.East Kalimantan
D. wahauense 164/tetraploid/ 1972c S. Belitung, Peg. Muller, KALTENG. 150 m.
Diplazium batuayauense Praptosuwiryo, sp. nov. D. batuayauense has
two ploidy levels, tetraploid (2n=164) and pentaploid (2n = ca. 205 ). The two
cytotypes were found on the relatively same habitat. They were growing on
humus rich soil and on rather shady places between the altitudes 440-450 m.
Diplazium cordifolium Blume. There are two kind of morpholodical
types in D. cordifolium, viz. simple frond and pinnate frond. Four ploidy levels
are found in the simple fronds of D. cordifolium. They are tetraploid, pentaploid,
hexaploid (TNgP 1201), and octoploid (TNgP1926b). The tetraploid (TNgP 1203)
was found in Mt. Salak (Ca.1200 m). Two individual pentaploid were found in
Mt. Salak at ca. 1200 m (TNgP 1204) and Mt. Halimun at ca. 950 m (TNgP
1808). The hexaploid and octoploid were growing at Mt. Salak (Ca. 1200 m) and
59
Mts. Muller (Ca.450 m), respectively. While, in the imparipinnate fronds are
found to have tetraploid (TNgP 1194, 1375, 1376) and pentaploid (TNgP 1307,
1355, 1442) from Mount Salak. From Mount Halimun, imparipinnate fronds are
also found to be tetraploid (TNgP 1735, 1736, 1774) and pentaploid (TNgP 1813).
Cytological observation of this species from Malaya reported by Manton (1954)
revealed 2n= >200, might be pentaploid and was not known whether ‘simple’ or
‘imparipinnate’ fronds. Cytological report from Java by Praptosuwiryo &
Darnaedi (2004) gave information concerning existence of the ‘imparipinnate’
tetraploid of this species in G. Patuha and those ‘simple’ pentaploid in Selabintana
G. Gede-Pangrango. The occurance of different morphological characters of
(simple and imparipinnate at the same ploidy level) bring us to the questions
whether they have different ancestor diploid races. Further cytological study is
strongly recommended.
Diplazium crenatoserratum (Blume) Moore. There are two levels ploidy
reported in this study, triploid (2n = 123) and tetraploid (2n=164). The triploid
plant was from Bukit Suban Punai Banyak, Bukit Dua Belas National Park
(Sumatra), minewhile the tetraploid plants were collected from the forest near
Tomoron Water Fall (Muller Range of Central Kalimantan) and Bukit Dua Belas
National Park (Sumatra). The all cytological records of this species are new
information for science .
Diplazium dilatatum Blume. Six individuals of Diplazium dilatatum
collected from West Java and Central Java showed apogamous triploid.
Cytological observations of this species from Gede-Pangrango National Park Wes
Java by Praptosuwiryo and Darnaedi (1994) found some apogamous triploid and
one individual tetraploid. In Japan, two varieties of D. dilatatum has been found,
viz. D. dilatatum Bl var. dilatatum and D. dilatatum Bl var. heterolepis Seriz.
The first variety was reported to have diploid and triploid, while the second
variety is only have triploid race (Takamiya et. al. 1999).
Diplazium donianum (Mett.) Tardieu. Only one plant of D. donianum
successfully examined its chromosome number. This plant is cultivated at Bogor
Botanic Gardens (collected from Bukit Ubar, Sumatra) and showed tetraploid (2n
= 164). Apogamous tetraploid was reported from Japan (Kato 1995). In Japan,
60
Kato (1995) reported D. donianum var. aphanoneuron (Ohwi) Tagawa with
chromosome number 2n = 123 (triploid). This variety is differ from var.
donianum in: lamina thicker and veins invisible beneath. West Malesian D.
donianum has characters: thin lamina and vein visible beneath (Figure 6.6.c.,
Chapter 6). Thus West Malesain plants are D. donianum var. donianum.
Diplazium esculentum (Retz.) Swartz This species usually grows in
opened areas with wet soils and distribute from Tropics of Asia to Oceania.
Mature plants are found to be pinnate and bipinnate. Veins anastomousing with a
few veins from adjacent veins joining into excurrent ones below sinuses between
lobes. Two individuals collected from two localities (Mt. Halimun and Mt. Gede)
showed 2n=82, diploid. Cytological information of this species from eight
localities (Malaya, Ceylon, North and South India) from 1954 to 1970 revealed
diploid without giving type of reproduction information (Love et. al., 1977).
Takamiya et. al. (1999) was also reported sexual diploid D. esculentum of
Shigetomi, Japan. Tetraploid D. esculentum is only found in India (Bhavanandan
& Amal, 1991). It is indicate that diploid D. esculentum is distributed in broad
range.
Diplazium halimunense Praptosuwiryo, sp. nov. There is only one
individual of D. halimunense succesfully examined (TNgP 2341b) and showed 2n
= 123 (triploid). This species is very rare, therefore only two individuals that can
be collected in Mt. Halimun. This species was found growing among the small
populations of D. bantamense and D. donianum. Morphologically, Diplazium
halimunense seems to be intermediate of D. bantamense and D. donianum.
Diplazium hewittii (Copel.) C.Chr. Onle one ploidy level from one plant
succesfully examined, triploid (2n = 123, TNgP 1913b), from Muller Range,
Central Kalimantan. There is no cytological report of this species up to this
present study. This psecies has a wide range of frond morphology, from bipinnate
to tripinnate. Therefore further cytological examination and mode reproduction
type study of this species are needed.
Diplazium profluens Praptosuwiryo, sp. nov. One individual of D.
profluens showed tetraploid (2n=164). The recent survey (in September 2003),
around Cikaniki Station Research, Halimun National Park, West Java, succeed to
61
find this species in shadowed place in the small river bank at track Macan and
track Canopy Bridge.
Diplazium loerzingii Praptosuwiryo, sp. nov. The recent living plants of
this species are found in Mt. Halimun (West Java) and Bukit Tapan (Jambi,
Sumatra). Two individuals from Java showed diploid (TNgP 2339c) and triploid
(TNgP 2339d). This species may have affinity to D. malaccense. This species
differs from D. malaccense in the following characters: lower base of 1-2 pairs
basal pinnae less cut, base of lower pinnae almost equally truncate, texture thicker
or subcoriaceous, upper rachis much gemmiferous, lobes truncate, sori medial on
veinlets or close to margin, attachments sides of indusia darker.
Diplazium pallidum (Blume) Moore. Three individuals of D. pallidum
observed from three different localities from Java showed two ploidy levels. Two
individuals which collected from mountain rainforest above the elevation 1100 m
showed tetraploid level, whereas one individual from lowland rainforest, viz. Mt.
Payung, Ujung Kulon National Park, with the elevation ca.200 m showed diploid
level. The diploid race is very scarcely occurred. Most of individuals of D.
pallidum collected from mountain forest above the elevation 1,200 m from three
localities in Java showed tetraploid level (Praptosuwiryo & Darnaedi, 1994;
Praptosuwiryo & Darnaedi, 2004).
Cytological examination of D. pallidum from Taman Nasional Bukit Dua
Belas (TNBD) in Sumatra and Bukit Batikap, Muller Range Central Kalimantan
has diploid level. These plants grow at 70 – 90 m in TNBD Sumatra and at 200 m
in Muller Range These facts support the hypothesis explained that diploid level of
D. pallidum habits at lowland under 200 m alt.
Diplazium petiolare C. Presl. Two individuals of Diplazium petiolare
collected from Rimbo Panti Nature Reserve, West Sumatra, showed diploid (2n =
82). This cytological account is a new record for Malesian ferns.
Diplazium porphyrorachis (Baker) Diels. Only one individual of D.
porphyrorachis succed its chromosomes counting, viz. tetraploid (2n=164). This
plant (TNgP 1885) was found in Gunung Pumpung Sapat, Muller Range (Central
Kalimantan). This is a new cytological account for Malesia. There is no
cytological report up to this present research.
62
Diplazium procumbens Holttum. Two individuals from two different
localities in, G. Salak and G. Gede, showed triploid only (TNgP 1348 and TNgP
1312). Formerly report by Praptosuwiryo & Darnaedi (2004) from four
localities in West Java and Eas Java also showed 2n=123, triploid. There is no
report of diploid D. procumbens. Cytological observation of this species from
Malaya (Manton 1954) and Ceylon (Manton & Sledge 1954) were also revealed
2n = 123, triploid apogamous.
Diplazium riparium Holttum. Two cytotypes of this species were found,
namely diploid (2n=82) and triploid (2n=123). The two cytotypes were collected
from the same island (Borneo) and in different locality. The diploid one was
collected from tract to Sungai Sopan-Bukit Batikap (Mts. Muller) at ca. 240 m
a.s.l. whereas the triploid from track Batu Ayau (Mts. Muller) at ca. 280 m a.s.l.
Since described by Holttum (1940), there is no chromosome observation for this
species. Thus, the two cytotypes are first record for science.
Diplazium silvaticum (Bory) Swartz. There are two varieties in D.
silvaticum, var. silvaticum that have lanceolate pinnae and var. pinnae-ellipticum
(new variety proposed in Chapter 9) those have elliptical pinnae. The first variety
revealed tetraploid and found in ‘nature habitat’ of the Bogor Botanic Gardens.
Whereas the second variety is triploid and collected from the lowland forest of
Sumatra (ca. 20 m). Manton & Sledge (1954), reported this species from Ceylon
to have ca. 200, while Manton (1953) also reported 2n= ca. 205 from Ceylon.
Abraham et. al. (1962) gave information 2n=205 from South India. Thus,
triploid cytotype was first record for science.
Diplazium simplicivenium Holtt. D. simplicivenium, in a glance, similar to
D. dilalatum. As stated by Holttum (1940) this species differs from D. dilatatum
in the following combination characters: scales wider and longer of coarser
texture; veins all single (a few may be forked on transition pinnae near apex of
fronds but not on the typical larger pinnae, even of large frond); usually not more
than 5 pairs of veins; sori occupying ¾ or longer of the length of veins. Three
individual of D. simplicivenium from three localities proved to be 2n=123,
triploid. Cytological observations of the species outside of Java were also showed
63
triploid, viz. In Malaya (Manton, 1954), Ceylon (Manton & Seldge 1954), and
New Delhi (Bir, 1969).
The rough morphological similarity between D. simplicivenium and D.
dilatatum are sometimes cause difficulties to diagnose the two species fast in the
field. Connecting to the same of chromosome numbers, ploidy level, and the
similarity of morphological appearances, the two species presumable involve in
the complexity of their species relationship.
Diplazium sorzogonense Presl. All plants examined from Mt. Salak and
Mt. Halimun revealed diploid. Two plants from Mt. Salak and Mt. Halimun were
sexual diploid. Morphological examination of both herbarium specimens
deposited at BO and living plants growing in Mt. Salak and Mt. Halimun revealed
that this species is varying in size and in the degree of lobing of its pinnae.
Diplazium speciosum Blume . One of the interested matter of D.
speciosum is its morphological variation of fronds. Therefore, two varieties of this
species have been described, viz. D. speciosum var. speciosum and D. speciosum
var. major. The two varieties can be recognized by the differences of its lobes and
venations. D. speciosum var. speciosum has lobes moderately toothed, end
rounded; veinlets 7-9 pairs, simple, While, D. speciosum var. major has lobes
strongly toothed, end mostly acute; veinlets 10-16 pairs, mostly forked.
However we do not find the differences of the two varieties based on
choromosome numbers. All individuls succesfully examined, include the two
varieties, revealed 2n=82, diploid. Three individuals belonging to D. speciosum
var. speciosum showed sexual type. While two individuals included in D.
speciosum var. major were not recognized its reproduction type.
Diplazium spiniferum Alderw. All three plants collected from G.
Pumpung Sabat (Muller Range, Central Kalimnatan) examined are diploid (2n =
82) and one of them showed sexual. There are no cytological record and mode
type reproduction information for D. spiniferum until this present observation. D.
spiniferum is one species of Diplazium that can grow on limestones area of
mountain forest at altitude 100-1300 m above sea level of Borneo. Stipe dark
green when living with sharply spine and rounded scales are characters that
64
differentiate this species from other bipinnate species of Bornean Diplazium.
The fronds are in varying and usually from bipinnatifid to bipinnate.
Diplazium subserratum (Blume) Moore. Previous cytological reports of
D. subserratum is not known until this research conducted. Four individulas of D.
subserratum from two localities of Mt. Slamet and Mt. Salak showed three ploidy
level: diploid, triploid and tetraploid. Diploid revealed sexual, while those
tetraploid are not known, and those probably sexual of triploid race presumed to
be apogamous.
Diplazium subvirescens Praptosuwiryo, sp.nov. Two indvidual plants
collected from Mt. Gede showed triploid (2n = 123, TNgP 1177 and TNgP 1013).
This species may closely related to D. virescens that distributed in Japan, Korea,
Taiwan, China and Indochina.
Diplazium tomentosum Blume. Three levels ploidy obtained from
cytological observation, viz. Diploid (2n = 82, Mt. Halimun, Java), tetraploid (2n
= 164, Bukit Pal Taman Nasional Bukit Dua Belas, Sumatra) and pentaploid
(2n=205, Mt. Halimun, West Java). Diploid and pentaploid race are new
cytological information for science. Cytological observation of this species from
Ceylon (Manton & Sledge 1954) showed only tetraploid and those from Fraser’s
Hill Malaya (Manton 1954) reported n = 82. The existence of diploid type and
also the higher ploidy level in this species presumed that Malesia, especially Java,
is the center origin for D. tomentosum.
Diplazium umbrosum (Smith) Bedd. Diplazium umbrosum is belonging to
the Diplazium species group with bi-tripinnate leaves. Its segments mostly 3.5-5
mm wide, one basal basiscopic lobes the largest, to 10 by 6 mm, apex blunt or
truncate, margin crenate or lobed 1/3 way to costulet of segments; texture
herbaceous; vein pinnate in each segments 4-6 pairs, mostly forked once in each
crenations, on larger crenation pinnate 2-3 pairs of second veinlets. Sori
elongated from near costulet of segments or on middle veinlets. Cytological
investigation revealed that there is no differences with the former information
from Java by Praptosuwiryo & Darnaedi (2004) collected from Mt. Welirang and
Mt. Patuha), in this recent paper one individual of D. umbrosum from Cibodas,
Mt. Gede, showed 2n=82, diploid. Other cytological observation from Europe by
65
Manton in Jermy (1964) gave 2n=ca.82, diploid also, under the name D.
caudatum (Cav.) Jermy sensu stricto (synonim of D. umbrosum).
Diplazium xiphophyllum (Bak.) C.Chr. Diplazium xiphophyllum is pinnate
species group. Its pinnae 9 pairs, terminal one like the rest; texture thin, drying
light brownish. Pinnae elliptical to 26 cm long, 4 cm wide, narrowed gradually to
slightly unequal cuneate base, ubrubtly to acuminate-caudate apex, margin entire
to irregularly toothed throughout. Veins in small group, at about 55o to costa,
commonly of one basal pairs and one central veins which is forked 1-3 times,
scarcely 4 times, sometimes acroscopic basal veins anastomousing with
basiscopic basal veins of the nearest vein group or with the nearest branch of
central vein near margin.
Three level ploidy for D. xiphophyllum are reported (diploid, tetraploid
and hexaploid). The finding of diploid and hexaploid race from Borneo (Batu
Ayau of Muller Range, Central Kalimantan) and Jawa (Mt. Salak), respectively,
are new record for science. Meanwhile the tetraploid race is new record for Java
and Sumatra. Formerly research was reported tetraploid race from Taiping Hill
(Manton 1954). As stated by Holttum (1966) and this study (Chapter 2 and 9), D.
xiphophyllum commonly grows in lowland forest and valley forest of moderate
mountains. The hexaploid race was found at 1200 m a.s.l. While, in Malaya D.
xiphophyllum was recorded at 914 m s.l. Two individuals tetraploid plant from
Sumatra, TNgP2040b and RI 867, were collected from Jambi at 55 m and from
West Sumatra at 950 m, respectively. It appeared that diploid and tetraploid race
of D. xiphophyllum exist at lower altitude than the hexaploid. This cytological
study on D. xiphophyllum showed a common phenomena that ploidy levels has
any correlation to habitat gradient. .
Diplazium wahauense Kato, Darnaedi et K. Iwatsuki. Diplazium
wahauense was firstly described by Kato et al. (1991) based on specimen
collected from along Jenta River, north of Muara Wahau, East Kalimantan. Since
that time, there is no cytological record for this species up to this present
research.. Due to the difficulties on maintaining the living collections of this
species, of the four collections number collected from Muller Range (Central
66
Kalimantan), there was only one collection that was successfully examined its
chromosome number, viz. TNgP 1972c. It was tetraploid (2n = 164).
5.3.2. The Relationship between Ploidy level and Morphological Variation within Species and Closely Related Species of Diplazium
Summary of the ploidy levels of Diplazium from West Malesia is
presented in Table 5.2. This study showed that intraspecific diversity on West
Malesian Diplazium are high enough. Twelve species of the 31 species
successfully examined are having series ploidy. Thirteen species showed only
polyploid race, from triploid and tetraploid. Whereas nine species revealed only
diploid race. The relationship between ploidy level and morphological variation
within species and closely related species are discussed below.
Diplazium accedens. All species of D. accedens examined are showing
diploid. Nevertheles, morphological variation exist. Three individual of
collections number TR 53 from Sumatra are showing very different appearances:
Roots are not bearing buds, stipes sharply spiny, vascular bundles of stipes
transversal sections near blade showing interupted U shape, rachis not
gemmiferous, basal pinnae much reduced. While collection number TNgP 1001,
1211, 1399, 1447, 1649, and 1786, from Java bearing buds both on roots and
rachise, stipes bearing green protuberances after falling scales, vascular bundles
of stipes transversal sections near blade showing continued U-shape, basal pinnae
slightly reduced.
Diplazium accedens is distributed throughout Malesia and grows well on
moist ground by stream or in humid in evergreen forest, by preferences more or
less shadowed localties, at 60-1550 m altitude. In very large fronds there are extra
areoles between the normal groups adjacent to the main lateral veins; a specimens
having this character formed the basis of Athyrium ridleyi Copeland. Having
opportunity to examine the type specimen of A. ridleyi deposited at SING (Ridley
13970) I agree with Holttum (1966) who also included it in the present species. In
my present field work in 2006 at Bukit Tapan, in Kerinci Seblat National Park
Sumatra, I found the plants similar to the type specimen of A. ridleyi.
67
This species is closely related to D. proliferum (Lam.) Thouars, therefore
the two should perhaps be united (Holttum, 1940). Andrews (1990) placed this
species in the genus Callipteris as Callipteris prolifera (Lam.) Bory and in his
synonymy includes D. proliferum (Lam.) Kaulf., Asplenium decussatum Sw., D.
accedens Bl., and D. proliferum var. accedens (Bl.) v.A.v.R.
As stated by Holttum (1940), the name D. accedens is a little complicated.
The three names of D. accedens, D. repandum, and D. Swartzii, all published in
the same book by Blume (1828), are to be regarded as synonymous. Later authors
have regarded all as synonyms of Lamarck’s earlier name, or of Asplenium
decussatum Sw.; the names Swartzii and accedens have been taken up and used
in the genera Callipteris, Asplenium and Athyrium, but the name D. repandum
appears to have been almost or entirely ignored. Backer & Posthumus (1939),
however, have revived the name D. repandum, apparently on grounds of page
priority (though they do not state this) which is not admitted by Rules. In their
synonymy, however, they include Diplazium proliferum Thouars, an older name;
their use of the name D. repandum is therefore contrary to the rules.
Diplazium angustipinna. Two cytotypes of D. angustipinna (triploid and
tetraploid) do not show any morphological differents. It is presumed that the
mechanism conctrolled is autopoliploidi.
Diplazium bantamense. Diplazium bantamense is usually found in moist
shady forest, chiefly in the hills, sometimes near streams in lowland forest. All
individuals examined here cytologically were collected from highland forest.
Assuming that most of diploid are generally living in tropical area diploid race
probably occur in the lowland. Examinations of many living collection grown in
Bogor Botanic Gardens nursery and dried specimens housed at BO revealed that
fronds of this species have enough variations on both leaf shape and size of
pinnae. Praptosuwiryo & Darnaedi (1994) reported four individuals of the
tetraploid sexual and two octoploid sexual of D. bantamense from Gunung Gede-
Pangrango National Park, West Java. However the features distinguishing the
tetraploid and octoploid type are unclear. Further examination on chromosome
number and its mode reproduction type from all of the range of its habitats,
including from lowland, would clarify their variations.
68
Diplazium cordifolium. Species concept of D. cordifolium stated by
Tagawa dan Iwatsuki (1988) covers individuals having simple frond narrowly
oblong-subdeltoid, cordate at broadest base, narrowing upwards towards
acuminate apex, subentire to undulate at margin and individuals having
imparipinnate fronds with a few pairs of lateral pinnae becoming smaller
upwards, lateral pinnae sessile, bearing gemmae at junction between rachis and
costa. Tagawa & Iwatsuki (1988) treated all morphological variations, the simple
fronds (D. cordifolium Blume) and the pinnate fronds (D. integrifolium Blume),
as one entity, viz. D. cordifolium Blume. The cytological evidence of this
research supports in differentiating D. cordifolium Blume from D. integrifolium
Blume. The simple fronds plants have one serial ploidy from tetraploid,
pentaploid and hexaploid, meanwhile those simply pinnate fronds plants reveal
tetraploid and pentaploid. Observation on living plants revealed that those simple
fronds never change into the simply pinnate fronds. However some characters of
the two groups, such as scales, lamina texture, and venation, are similar.
Mitsuta (1985) recognized two varieties of Sumatran D. cordifolium, var.
integrifolium (Bl.) Mitsuta and var. pariens (Copel.) C.Chr. The two varieties are
differentiated with characters as follow. var. integrifolium has 2-3 pairs of lateral
pinnae, and base of terminal pinnae sessile, while var. pariens with 4-6 pairs of
lateral pinnae, and base of terminal pinna usually wide cuneate.
Cytological observation showed that the simple fronds revealed a series
ploidy (2n=4x, 5x, dan 6x) and those imparipinnate aslo have a series ploidy
(2n=4x dan 5x). Morpholigical observation in the living collections revelaed that
the plant with simple fronds did not change into the pinnate fronds. Further
morphological differences are provided in the identification key in the Chapter 9.
This study showed that genetic variation within D. cordifolium are varying and
has brougth out into the morphological variation. The molecular evidence from
gene rbcL sequence also supported this results (Chapter 8).
Diplazium crenatoserratum. As also reported by Holttum (1940), D.
crenatoserratum is a very common fern of lowland forest. This species shows
much variation in size and in the degree of lobing of its pinnae. Small plants with
blunt entire pinnae may be fertile. The tetraploid type of Tomoroh Water Fall
69
(Muller Range, Central Kalimantan) has small size with blunt entire pinnae while
the plants with larger size and pinnae more incision, margin lobed ¼-1/2 way
towards costa, revealed triploid and tetraploid. Therefore it is presumed that there
are no morphological differences between triploid and tetraploi. However further
cytological observation of this species for all its range distribution are needed to
clarify this preliminary study.
Diplazium dilatatum. In Japan, two variaties of D. dilatatum have been
recognished, viz. D. dilatatum Blume var. dilatatum dan D. dilatatum Blume var.
heterolepis Seriz. (Kato, 1995). The first variety are diploid and triploid, while the
second variety is only triploid (Takamiya et. al. 1999). The triploid race of
Javanese species presumed as var dilatatum, due to morphological similarity with
those of Japanese species. Kato (1995), stated that D. dilatatum var. dilatatum
differs from D. dilatatum var. heterolepis in the scales cahacters. D. dilatatum
var. heterolepis has scales at the stipe base lanceolate, to 20 mm long, black at
margin. While D. dilatatum var. heterolepis has scales at stipe broadly lanceolate,
10-15 mm long, 1-3 mm broad, hardly black at margin. Javanese plants match to
the D. dilatatum var. dilatatum, because its scales characters similar to this
variety, viz. liniery lanceolate, 15 mm long or more, 1 mm wide, yellowish brown
at middle, black and sharply toothed at margin. One individual of the tetraploid
and thirteen individuals of triploid of D. dilatatum from West Java were reported
by Praptosuwiryo & Darnaedi (1994). But, morphological differences between the
two cytotypes are not significant.
Diplazium pallidum. Morphologically, the appearances of the two ploidi
level, diploid and triploid, showed much differences. The tetraploid race showed
following character spots: terminal pinna deltoid and deeply lobed; upper base of
lateral pinnae broadly truncate, lower base narrowly rounded. Whereas the
diploid race has following characters spots: terminal pinna conform to lateral with
one or two lobes; upper base of lateral pinnae rounded, lower base cuneate.
Further examination to see the conssitence of the morphological
differences between the two race of D. pallidum (diploid dan tetraploid) is needed
by examining much more sample from different altitude. In addition, examination
70
of reproduction types, spore mophology, anatomy of the fornds, and isozyme or
DNA analysis would give clarification of the taxonomical status of the two race.
Diplazium silvaticum. There are morphological distinct among the triploid
types and tetraploid types. Tetraploid plants have pinnae 8-13 pairs, lower pinnae
lanceolate, 7-15 x 1.6-3 cm, upper base subtruncate, lower base cuneate.
Meanwhile the triploid ones have pinnae 2-5 pairs, lower pinnae elliptic, 3.3 -5.4
x 2.0-2.7 cm, upper base subrounded, lower base cuneate. Those tetraploid are
sexual, while the triploid are apogamous. It is suggested that the triploid was
originally hybrid. Further studies are needed.
Diplazium subserratum. The differrences of ploidy level is presumed to
be correlated to its morphological variations. The results of the morphological
comparison between diploid (TNgP 1463) and tetraploid (TNgP 1379) is showed
in the following characters: The diploid has stipe 1.5 mm diam., length 15 cm;
lamina 34 cm length, 3.1 cm wide (in the middle), margin entire; veins forked 2-3
times. While those tetraploid has stipe 1.5-2 mm diam., 13.0-16.5 cm length;
lamina 15-29 cm, 2.0-4 cm wide (in the middle); margin 1/6-2/3 portion of base
entire and waved 1/3 – 5/6 portion upper; veins forked 2-4 times. The correlation
between the ploidy level, type of reproduction, and morphological appearance of
this species seem very interesting from the point view of speciation in Diplazium.
Therefore more cytological observations of this species over its range areas of
distribution are needed to verify the correlation between ploidy levels and
morphological differences.
Diplazium tomentosum. 82/diploid/ (TNgP 2336b) ; 164/teraploid (TNgP
2066); 205/pentaploid/ (TNgP1722). There are not any significant qualitative
characters differences among the three cytotypes of D. tomentosum. However
they have several differences in quantitative. The diploid type has lamina up to
21.5 cm by 10 cm, pinnae up to 14 pairs; veinlets on the largest lobe of basal
pinna up to 5 pairs, mostly simple. Tetraploid type has lamina up to 24 cm by 15
cm, pinnae up to 17 pairs; veinlets up to 6 pairs, mostly simple. Whereas the
pentaploid type has lamina up to 32 cm by 17 cm, pinnae to 19 pairs; veinlets on
the largest lobe of basal pinna up to 8 pairs, mostly once forked. These results
indicate that the ploidy seri in this species is autoploid.
71
Diplazium xiphophyllum. Three voucher spesimens of the three cytotypes
(diploid, tetraploid and hexaploid) could not be made comparison because they
are in different stage of growth. However qualitatvely the three cytotypes are not
different. Therefore it is presumed that they are autoploid.
Diplazium aequibasale, D. riparium and D. wahauense. Diplazium
aequibasale (2n=164), D. riparium (2n=82) and D. wahauense (2n=164) almost
have similar morphological appearances. Diplazium aequibasale have allied form
between D. riparium and D. wahauense. D. wahauense is closely related to D.
riparium. Kato et.al. (1991) presumed that D. wahauense derived from D.
riparium which occurs in riparian and dryland forest in Borneo. The two species
share many character, viz. black, somewhat crisped, entire scales, blackish stipes,
dark-brown, naked rachis, and imparipinnate leaves with 3-4 pairs of entire lateral
pinnae. D. wahauense differs from D. riparium mainly in its narrow pinnae,
which are characteristic of rheophytes.
5.3.3. Relationship between ploidy level and habitat gradient
Although data are available for fern floras from different parts of the
wolrd, there is not simple relationship between ploidy level and habitat (Walker
1979). Some studies have revealed that lower-ploids occur in warmer habitats
than higher-ploid, for example Lepisorus thunbergianus (diploid vs. Tetraploid)
(Mitui et al 1987), Woodwardia orientalis (diploid vs. tetraploid) (Mitui 1968),
Dryopteris erythrosora (diploid vs triploid) ( Hyrabayashi 1974), Pteris dispar
(diplaid vs tetraploid) (Nakato 1981), and Diplazium nipponicum (triploid vs
tetraploid) (Takamiya et al 2000). However, many parts of Europe all three
cytotypes (2x, 3x, 4x) of the Polypodium vulgare complex were found in close
proximity (Shivas 1961). Matsumoto (2003) did not find a significant relationship
between latitude and ploidy level in the Cyrtomium falcatum complex. However
there was an association between ploidy level and habitat: diploid grew on sea
cliffs or in dry forests, triploid grew at forest edges, and tetraploid grew in wet
forests. In the study of the Pteris fauriei complex in Taiwan, Huang et al. (2007)
recorded any correlation between habitats, latitude, and elevation of P. fauriei.
72
The three parameters are related to temperatures. In general, triploid plants grow
in low temperature sites than diploid plants.
In many species of Diplazium from West Malesia, however, there are not
any strick correlation between ploidy level and habitat gradient. All species with
only diploid type, viz. D. accedens, D. petiolare, D. polypodioides, D, speciosum,
D. sorzogonense, D. spiniferum and D. umbrosum, are growing from 420 m to
1500 m. Species with two different ploidy levels are growing on almost the same
elevation. The tetraploid D. crenatoserratum were collected from 20 – 150 m sea
level where the triploid one was also found in the range, 55 m sea level. Triploid
and tetraploid D. angustipinna were found at the same altidude, 440 m. In
contrary, pentaploid D. batuayauense was growing at 440 m, whereas the
tetraploid plant was found at 450 m. Overlap distribution was seen in D.
cordifolium. Tetraploid D. cordifolium was growing from 240 m to 1200 m,
whereas pentaploid plants were occuring from 1000 m to 1200 m. As has been
mentioned above, there are only few species that has a correlation between ploidy
levels and habitat gradient, such is D. riparium and D. xiphophyllum.
Some species of West Malesian Diplazium showed a correlation between
ploidy level and altitute gradient. Diploid D. riparium was found at 150 m,
whereas the triploid plant was at 280 m. Tetraploid plants of D. pallidum were
only found above 1000 m, whereas those diploid were growing from 30 m to 250
m. Triploid and tetraploid D. silvaticum are growing at ca. 20 m and ca. 280 m,
respectively. The hexaploid D. xiphophyllum was found at higher altitude than
those of tetraploid and diploid.
5.3.4. Correlations between reproductive mode and habitat
Some studies have found distinct correlation between reproductive mode
and habitat (Holbrook-Walker & Lloyd 1973; Lloyd 1974). Sexual reproduction
is most frequent in moist habitats, whereas apomictic reproduction is more
common in ferns growing in xeric habitats (Tryon 1968). However, some
apomictic species do not live in xeric habitats, for example some Japanese
Diplazium (Takamiya et al 1999) and Cornopteris christenseniana (Park & Kato
2003).
73
Table 5.2. Polyploid series of Diplazium in West Malesia Based on Present Study
Ploidy Level
Species
2X 3X 4X 5X 6X 8X
D. accedensD. aquibasaleD. asymmetricumD. bantamenseD. batuayauenseD. cordifoliumD. crenatoseratumD. dilatatumD. donianumD. esculentumD. halimunenseD. hewittiiD. loerzingiiD. pallidumD. petiolareD. polypodioidesD. procumbensD. profluensD. porphyrorachisD. ripariumD. simpliciveniumD. silvaticumD. speciosumD. procumbensD. sorzogonenseD. subpolypodioidesD. spiniferumD. subserratumD. subvirescensD. tomentosumD. umbrosumD. xiphophyllumD. wahauense
V-------
-V - -VVVV---V--V-VVVV-VVV-
--V---VV
--
V V
V---V--VVV-V---VV----
-V-VVVVV*
V-
- -
-V---VV--V-----V-V-VV
----VV----
- -
-----------------V---
-----V--
--
- -
-------------------V-
---
V*-V--
--
- -
---------------------
TOTAL 15 13 15 3 2 2
Notes:
V Present study
V* Praptosuwiryo & Darnaedi (1994)
74
Figure 5.1. Somatic chromosome of Diplazium. a. D. accedens var. accedens , 2n = 82 ;b. D. accedens var. spinosum, 2n = 82; c. D. angustipinna, 2n = 164 (TNgP 1906b); d. D.angustipinna, 2n = 123; e. D. spiniferum (TNgP 1896a), 2n = 82; f. D. halimuense, 2n=123(TNgP 2341b). Bar = 3 m.
75
Figure 5.2. Somatic chromosome Diplazium. a. D. umbrosum, 2n = 82 (TNgP1348);b. D. petiolare 2n = 82 (TT993-3), c. D. tomentosum, 2n = 82 (TNgP 2336b);d. D. tomentosum, 2n = 205 (TNgP1722); e. D. tomentosum, 2n = 164 (TNgP2066);f. D. asymmetricum, 2n=123 (TNgP 1094). Bar = 3 m.
76
Figure 5.3. Somatic chromosomes of Diplazium. a. D. xiphophyllum (TNgP), 2n = 246; b. D.xiphophyllum (TD902), 2n = 164; c. D. xiphopyllum (TNgP1841 ), 2n = 82; d. D. batuayauense(TNgP 1927), 2n = 123; e. D. porphyrorachis (TNgP1885), 2n = 164. Bar = 3 m.
77
Figure 5.4. Somatic chromosomes of Diplazium. a. D. subserratum, 2n = 82 (TNgP1463),b. 2n = 123 (TNgP2287), c. 2n = 164 (TNgP1379); d. D. subpolypodioides, 2n = 82 (TNgP2292);e. D. procumbens, 2n = 123 (TNgP1348). Bar = 3 m.
78
Figure 5.5. Somatic chromosomes of Diplazium cordifolium. a. 2n = 164 (TNgP1735);b. 2n = 205 (TNgP1204); c. 2n = 246 (TNgP1201); d. 2n = 328 (TNgP 1926b). Bar = 3 m.
79
Figure 5.6. Somatic chromosomes of Diplazium. a. D. pallidum, 2n = 82 (TNgP1406);b. D. pallidum 2n = 164 (TNgP1764); c. D. riparium, 2n = 123 (TNgP 1847);d. D. polypodioides, 2n = 82 (TNgP2285); e. D. dilatatum, 2n = 123 (TNgP1073); f. D. speciosum,2n=82 (TNgP1359). Bar = 3 m.
80
Reproductive studies on Diplazium from West Malesia showed that both
the sexual and apomictic species can occur in the moist habitats. The apogamous
triploid D. procumbens and D. simplicivenium are found in dryland areas of the
slopes on Mt. Gede-Pangrangrango and Mt. Salak in which have the relatively
high amount of rainfall (ca. 200 mm per month in October - May). The
apogamous pentaploid D. batuayauense was also growing at the moist soil in
shady place in the forest. The sexual and apogamous tetraploid of D. bantamense
are found at the relatively same habitats condition. The two race are found at the
moist humus-rich soil of the dryland areas of Cangkuang forest in Mt. Salak.
These results revealed that there are not distinct correlation betweed the
reproductive mode and habitats in West Malesian Diplazium.
In many ferns reproductive barriers are likely to exist between polyploid
and their ancestors. This increases the potential for the development of their
spatial segregation (Vogel et al 1999). Some researchs demonstrate that
microenvironment (habitat) selection is partially responsible for maintaining the
separation of different cytotypes in ferns and flowering plants (Kumaret et al
1987). In West Malesian Diplazium the separation between polyploid and their
ancestors have not been seen clearly. Beacuse most of species with seri ploidy
have not been examined well their mode of reproductive by sampling many
individual plants. Moreover, the ancestors of the many polyploid species
observed have not been found, their ancestors (diploid types) have not been found
yet. The diploid of D. bantamense, D. simplicivenium, D. procumbens, and D.
asymmetricum, for example, have not been recognized their existence. Therefore
further study on cytology and the mode reproductive of West Malesian Diplazium
to understand its speciation mechanism are needed.
81
5.4. Conclussions
All West Malesian species of Diplazium examined showed basic
chromosome number x = 41. New cytological information for science on 19
species are recorded. They are D. aequibasale (tetraploid), D. angustipinna
(triploid), D. asymmetricum (triploid), D. batuayauense (tetraploid and
pentaploid), D. crenatoserratum (triploid and tetraploid), D. halimunense
(triploid), D. hewittii (triploid), D. profluens (tetraploid), D. loerzingii (diploid
and triploid), D. pallidum (diploid), D. petiolare (diploid), D. porphyrorachis
(tetraploid), D. riparium (diploid and triploid), D. spiniferum (diploid), D.
subserratum (diploid, triploid, tetraploi), triploid D. subvirescens, D. tomentosum
(diploid and pentaploid), D. xiphophyllum (diploid, hexaploid), and Diplazium
wahauense (2n=164).
This study showed that intraspecific diversity on West Malesian Diplazium
is high enough. Twelve species of the 31 species successfully examined their
somatic chromosome numbers are having ploidy level series: D. angustipinna
(triploid and tetraploid), D. bantamense (traploid and oktoploid). D. cordifolium
(tetraploid, pentaploid, dan hexaploid, octoploid), D. pallidum (diploid and
tetraploid), D. silvaticum (triploid and tetraploid), D. tomentosum (diploid,
tetraploid, pentaploid), D. subserratum (diploid, triploid, and tetraploid), D.
xiphophyllum (diploid, tetraploid and hexaploid). Thirteen species showed only
polyploid race: D. aequibasale (tetraploid), D. profluens (tetraploid), D.
porphyrorachis (tetraploid), D. procumbens (triploid), D. simplicivenium
(triploid). Whereas nine species revealed only diploid race (D. accedens, D.
esculentum, D. malaccense, D. petiolare, D. polypodioides, D. speciosum, D.
spiniferum, D. sorzogonense, D. subpolypodioides, and D. umbrosum).
Study on the relationships between ploidy level and morphological
variation in some species, such as D. batuayauense, D. crenatoserratum, D.
loerzingii, D. riparium, D. tomentosum, and D. xiphophyllum, suggested that they
are autoploid. Triploid D. silvaticum was presumed alloploid. However further
cytological study are needed to verify the polyploidy mechanisme involved in the
species mentioned above and the others.
82
Study on the relationship between ploidy level and habitat gradient
showed that generally West Malesian Diplazium species are showing no any
strick correlation between ploidy level and altitude, but D. pallidum, D. riparium,
D. silvaticum and D. xiphophyllum. Reproductive studies revealed that many
species are apogamous triploid and both the sexual and apomictic species can
occur in the relatively moist habitats.
83
CHAPTER 6
PHYLOGENETIC STUDIES OF DIPLAZIUM FROM WEST MALESIA:EVIDENCE FROM MORPHOLOGY
6.1. Introduction
Most of pteridologist thought that Diplazium are quite distinct and should
be separated from Athyrium (See Ching 1940; van Alderwerelt van Rosenburgh
1908; Alston 1956; Sledge 1962; Holttum 1940, 1966; Kato 1977, 1995; Edie
1978; Tagawa & Iwatsuki 1988; Andrews 1990; Kramer et al 1990). Strictly
Kato (1977) showed the differences between Diplazium and Athyrium. In
Athyrium, the stipes bases on ascending to erect rhizome swollen with
pneumatophores, frond axes V-shaped in transaction, acroscopic basal pinnules
larger than others, laminar margin cartilaginous or not, spines present adaxially at
the junction of costules or not, sori horseshoe- or J-shaped, or linear, scales entire.
Whereas in Diplazium, the stipes bases neither swollen nor bearing
pneumatophores, frond axes U-shaped with flat base in most species, acroscopic
basal pinnules equal or smaller, laminar margin not cartilaginous, spine absent,
sori linear, scales toothed or entire. Moreover the separation of Diplazium from
Athyrium has been supported by both cytological evidence and molecular data.
The different in the basic chromosome number of Athyrium (x= 40) and
Diplazium (x=41) is useful diagnostic character (Tryon & Tryon 1982).
Preliminary phylogenetic study of Diplazium conducted by Sano et al (2000)
based on chloroplast rbcL gene sequences showed monophyletic of this genus.
By using evidence from chloroplast trnL-F region sequences Wang et al (2003)
also supported the monophyletic of Diplazium clade that include Callipteris
Bory, Allantoidea R. Br. Emend. Ching, and Diplaziopsis C. Chr.
Morphological variation among species of Diplazium are very diverse.
But, a natural subdivision of this genus has not been given yet. Van Alderwereld
van Rosenburgh (1908) tried to divide Malayan Diplazium, include those of the
Malay Peninsula,The Philippines and New Guinea, into two sections based on
only the venation type: (1) Eudiplazium for Diplazium species having free veins
and (2) Anisogonium for Diplazium species that possess anastomousing veins.
84
Based on the characters such as scales, stipe, lamina and venations, Kato (1977)
recognized Japanese Diplazium that consisting of five groups, namely: (1)
Diplazium dilatatum group that includes member with groove generally U-shaped
with a flat base, acroscopic basal pinnules or segments equal to or smaller than the
basiscopic or subsequent ones, scales entire or toothed, but not clathrate; (2)
Diplazium wichurae group that includes species member with groove U-shaped,
acroscopic base of pinna auricled, adaxial surface of lamina concave along veins,
scales entire, sometimes subclathrate; (3) Diplazium mesosorum group that having
characters groove U-shaped with a flat base, acroscopic basal pinnules or
segments equal to or slightly larger than the basiscopic or subsequent ones, scales
entire and subclathrate; (4) Diplazium javanicum group in which includes species
with groove V-shaped, frond pinnate or imparipinnate, laminar margin entire or
undulate, veins sagenoid-reticulate and scales entire; and (5) Diplazium
longicarpun group that includes species with frond pinnate, acroscopic base of
pinna truncate, basiscopic cuneate, adaxial surface of lamina not concave along
veins, scales entire. However the naturalness of the subdivision of Diplazium
circumcribed by van Alderwereld van Rosenburgh (1908) and Kato (1977) have
never been tested and analyzed phylogenetically using morphological datasets.
Therefore phylogenetic studies on Diplazium using morphological datasets
should be introduced.
Most grouping on the tree of life were first inferred using morphological
characters, which have been widely used since Darwin’s time. Most of the
million-plus described species are know only from morphology, and their inferred
phylogenetic positions are based on these characters; with between 90 and 99
percent of life still to be formally recognized, usually initially from morphological
features alone (Lee 2004). Although the pool of morphological characters is
much smaller than molecular characters (See Chapter 8), and often insufficient
for robust phylogentic resolution, is a common view (e.g. Hillis 1987; Givnish &
Sytsma 1997), however morphological datasets often contain as much relevant
phylogenetic signal as typical molecular datasets that have orders of magnitude
more characters (Lee 2004).
85
This chapter presents phylogenetic study of Malesian Diplazium by using
morphological datasets generated from sixty nine species from West Malesia. All
data on the characters analyzed here are cited from this present work (Chapter 9).
This study was conduct to address question of phylogenetic relationships among
species within Diplazium. The objectives of this study were to: (1) to reconstruct
the phylogenetic relationships among the species in the genus Diplazium; (2)
identify monophyletic species groups within Diplazium, (3) establish sister-group
relationships among these monophyletic groups.
6.2. Character Selection and Construction
6.2.1. Character Selection
Cladistic analysis is the most common method currently used to
reconstruct phylogenetic trees, even it is the best method (See Bremer &
Wanntorp 1978, Estabrook 1978, Wiley 1980). For generating a set of trees, it
involves two basis phase-exploration of characters (including selection and
examination), followed by analysis of the data (Thiele 1993). Thiele (1993)
described an ideal morphological character as one in which the character states
vary between terminal units in the analysis but not in other members of the unit
(e.g. conspecific individuals) being represented.
6.2.2. Character Type
In the morphological systematic studies, there are two types of characters
used: qualitative and quantitative. Qualitative characters are mostly obtained by
examination without measurement. These characters can be divided into binary
characters and multistate characters. Binary characters comprises two character
states, such as scales margin (without thickening black strands or with thickening
black strands, in character 14, Table 6.1.). Multistate characters consist of more
than two character states such as lamina division that comprise 7 character states
(simple, pinnatifid, imparipinnate, bipinnatifid, bipinnate, tripinnatified, and
tripinnate, in character 23, Table 6.1.). Generally, qualitative characters are more
86
acceptable and seem to be unambiguous in cladistic analysis because their states
are considered to be clearly defined and no overlapping (Kitching et al 1998).
Quantitative characters are obtained by measurement. Generally
quantitative characters are continuously variable. Continuously variable should
only be exclude if the cladistic analysis cannot handle such data or if it can be
shown empirically that those characters convey no information of phylogenetic
signal relative to other characters in the data matrix (Kitching et al. 1998). In this
study quantitative characters are included in the analysis because they convey
information of phylogenetic signal relative to other characters in data matrix.
Another argument for inclusion of quantitative characters is that some qualitative
characters may be a collection or transformation of quantitative characters. For
example. Leaf shape (a qualitative characters) can be defined by ration of leaf
length to leaf width, which is quantitative caharcters (Thiele 1993.). Characters
number 7, 8, 16, 17, 25, 26, 37, 38, 49, 50, and 56 in the Table 6.1. are
quantitative characters that are important and having tight correlation with the
qualitative characters examined. Excluding these characters would give illogical
relationships among the species. Therefore these characters were included in
analyis.
6.2.3. Character Coding
The crucial point in the phylogenetic analysis using morphological
datasets is how features might be usefully coded so as to reflect accurately our
observations for particular scale problem (Kiching et al 1998). Character coding
is the link between observation and explanation (Strong & Lipscomb 1999).
Therefore, ability, or inability, of a coding method to reflect the evidential
significant of observations should be the primary concern in considering
alternative methods of coding.
Basing on the philosophy above, in cladistic analysis of morphological
characters in Diplazium, a conservative approach of ‘composite characters coding’
(Strong & Lipscomb 1999) was employed. Composite coding refers to the
creation of single multistate character from several potentially dependent
character (method ‘A’ in Kitching et al 1998) that designed to minimize the effect
87
of character linkage. Kitching et al (1998) stated that the more that characters
become linked, i.e. dependent on other characters, the greater is the departure
from independence and consequently the risk that one false homology can obscure
the topologies of true homologies decreased.
Composite coding applied in this study was also designed to reduce the
error that extensive amounts of missing data (a character state scored as ‘?’) can
create in a parsimony analysis (Maddison 1993). The fronds architecture of
Diplazium are varying, from simple to quadripinnate. Consequently taxa with
simple, pinnate, bipinnate fronds do not posses any organ that possessed by those
with pinnate, bipinnate and tripinnate fronds, respectively.
Character 36. (Table 6.1) is an example of composite characters in this
study. It was created from the combination of two characters: 1) Lateral pinnae
exist/not exist and 2) Shape of lateral pinnae oblong / oblong lanceolate /
elliptical / liniery triangular / oblong subtringular / lanceolate / ovate. These two
characters in which the latter is dependent on the former, were combined into a
single character with eight character states (lateral pinna oblong / oblong
lanceolate / elliptical / liniery triangular / oblong subtringular / lanceolate / ovate/
organ not exist). This method reduced the numbers of linked characters and
limited the influence of multiple missing characters states on the analysis for
previously coded characters ‘lateral pinnae oblong / oblong lanceolate / elliptical /
liniery triangular / oblong subtringular / lanceolate / ovate’ in those taxa which
have no lateral pinnae. Maddison (1993) stated that missing character states can
cause difficulties for parsimony analysis if the missing state for the character
occurs in more than one clade.
6.3. Character Variation within West Malesian Diplazium
Previous pteridologist worked on Diplazium have summarized the
principal characters of the genus Diplazium and identified the important
morphological features in identification and classification. Characters those
particularly variable across the taxa and which may be significant in an
evolutionary sense are discussed as follow.
88
a. Rhizome
Most of West Malesian species posses erect rhizome. The appearances of
the erect rhizome are varying. The very short and small rhizome are seen in D.
crenatoserratum and D. tomentosum. The stout and medium rhizome are found in
D. dilatatum, D.kunstlerii, D. megasegmentum, D. polypodioides, and D.
subpolypodioides. The stout long rhizome is present in D. esculentum. Some
species have sreeping rhizome, such as D. donianum, D. procumbens, and D.
subvirescens Mickel (1974) presumed that the main lines of the ferns begun with
probably creeping rhizome. Thus the erect rhizome is regarded as derived. Figure
6.1. showed the rhizome patterns of Diplazium.
b. Scales and Hairs
Scales in Diplazium are varying. As showed by Holttum (1940, 1966),
Kato (1977, 1995), and Tagawa & Iwatsuki (1988), scales are very important
characters in determining species in Diplazium (Figure 6.2.). The scales mainly
attach on upper of the rhizome and stipes. On some species scales are also
present on the rachis or the costa, but in smaller size than those on the rhizome
and stipes. Scales are generally linear lanceolate to obong lanceolate. Rounded or
ovate scales are present on the projections of a muricate stipe such as in D.
megasegmentum and D. profluens (new species proposed, See Chapter 9), D.
kunstlerii, and D. latisquamatum. Rounded or ovate scales which is initially
having a function as an covered organ of spines are seen on the stipe of D.
spiniferum.
Scales are generally attached with a basal point. Some species with
rounded or ovate scales, such as in D. kunstlerii, D. latisquamatum, D.
megasegmentum and D. spiniferum, have scales with a subbasal point attachment
(Figure 6.2.g. in this Chapter; Chapter 9). Scales with a subbasal point attachment
may be derived. Dickason (1946) presumed that a scales with a basal point of
attachment may be more easily derived from a hair than one with broad base or
one peltately attached.
Diplazium scales are entire or toothed, with tooth consisting of two
upturned ends of adjacent marginal cells. The marginal may be with thickening
brown-black strand or not. Toothed scales may show sharp and blunt teeth,
89
regular or irregular teeth construction, and forked or not forked teeth. The
existence of glandular cell at the marginal scales are important in differentiating
among species with entire scales or with blunt toothed scales, such as in D.
tomentosum, D. pallidum and D. spiniferum. Scales with entire margins may be
simpler than those with toothed margins. Because evolution never starts with
complex constructions (Schölch 2000), the toothed margin scales may be derived.
Figure 6.1. Rhizome appearance of Diplazium. a. short-creeping rhizome of D.donianum; b. medium-creeping rhizome of D. subvirescens; c. medium-erect rhizome ofD. accedens; d. short-erect rhizome of D. halimunense; e. stout-erect rhizome of D.megasegmentum.
90
Scale colour are also important in species identification. Scales colour
may be yellowish, pale brown, brown, dark brown to black. For example, black
scales of D. meijerii will be differentiate fast this species from its related species,
D. atrosquamatum, that has brown scales.
The taxonomical evaluation of the articulated hairs (trichomes) in the
Athyriaceae has been conducted by Kato (1973). He summarised that the features
of the articulated hairs are important in the classification of Athyriaceae. In the
genus Diplazium, Kato (1972) showed two kinds of articulated hairs, viz. hairs
with glandular cells and without glandular cells in Monomelangium pullingeri
(that has been proposed to be placed in Diplazium and become D. pullingeri) and
D. tomentosum, respectively. There are only several West Malesian species that
posses hairs. The hairs are without glandular cells. D. squarrosum posseses
stellate hair. D. tomentosum shows simple articulate hairs densely covered stipe,
rachis and costa. D. crenatoserratum posseses simple articulated hair sparsely on
stipes when living and then it will be scarsely found on dried material.
c. Stipe
Diplazium stipe is not swollen and becomes gradually thicker downwards.
The stipe is generally green. The stipe anatomy of some species of Diplazium
has been studied by Tardieu-Blot (1932), Bir (1962, 1969), and Kato (1972). Leaf
traces are binary. The xylem of a leaf-trace is the same hippocampus-shaped
bundle in transaction (See Chapter 4). In this study the anatomical characters of
Diplazium were no used in phylogenetic analysis, because most of species
examined were based on only herbarium specimens.
Stipe characters used in this study were only those from the gross
morphology, such as the size, colour, and the existence of multicellular hairs,
scales, protuberances and spine (Figure 6.4.). Most of Diplazium posses scales
only. Multicellular hairs and scales are seen in D. crenatoserratum, D. silvaticum
D. tomentosum. The multicellular hairs in D. crenatoserratum and D.silvaticum
are usually only seen in living plants, they are fallen when dry. Whereas in D.
tomentosum the multicelullar hairs can be seen both in living plants and dried
materials. The existence of stellate hairs and scales on the stipes and rarchis of
D. squarrosum make this species is very distinct among the West Malesian
91
Diplazium. Green protuberances are seen in D. accedens var accedens and D.
accedens var. swartzii. Spines are present in D. accedens var. spinosum and D.
spiniferum.
Figure 6.2. The variation of scale shapes in Diplazium. a. D. halimunense; b. D.tomentosum; c. D.. silvaticum var. silvaticum; d.. D. esculentum; e. D. petiolare; f. D.batuayauense; g. D. spiniferum; h. D. sorzogonense; i. D. malaccense; j. D. donianum; k.D. silvaticum var. pinnae-ellipticum; l. D. angustipinna.
92
Figure 6.3. Margin of scales. a. entire with glandular cells and irregular thickeningblack strands (D. atrossuamatum); b. margin entire without glandular cells andthickening black strands (D. cumingi); c and d. margin sharply toothed with regularthickening black strands (c. D. subalternisegmentum , d. D. simplicivenium); e. marginentire with glandular cells and thickening dark brown strands (D. sorzogonense); f.margin sharply toothed irregularly with irregularly thickening black strands (D.halimunense); g. Margin with densely glandular cells without thickening black strands(D. batuayauense); h. margin entire without thickening brown or black strands (D.malaccense); i. margine toothed without thickening black strands D. petiolare.
93
When old or in dried specimens, stipes may glabrous, sparsely or densely
scales throughout its length. Chistensen (1911) said that the best and most
constant specific characters is to be found in the dermal appendages, hairs and
scales. A glabrous shoot may be thought ofas more primitive than one having a
potentiality for the developmental of dermal outgrowth (Dickason 2007).
Figure 6.4. Stipes appearances of Diplazium. a. Stipes densely scales (D.simplicivenium); b. stipes dark green, covered sparsely by rounded-ovate scales, scalesfallen when dry (D. spiniferum); c. Stipes covered by green protuberances (D.accedensvar. swartzii) ; d. stipes spiny, scales fallen when old (D. accedens var. spinosum)
94
d. Rachis and costa
As showed by Kato (1977), the rachis and costa groove is usually open to
admit the groove of leaf axis of lower order. D. accedens, D. crenatoserratum, D.
subserratum, D. tomentosum, D. xiphophyllum have rachis that prominently raised
above with shallow grooves and their grooves are U-shaped with flat base
(Chapter 4., Figure 4.2.). The appearance of rachis groove would give more
informative parsimony characters. The cross section of rachis are generally
similar to the cross section of stipes near lamina. Unfortunately, in this study,
anatomical data of the cross section of stipes near lamina were only obtained
from 27 species due to the lack of the living collections in most species.
Therefore grooves characters on rachis or stipes did not used in this analysis,
because they would cause many missing data on many species.
Generally, rachis are glabrous. Some species has fibrillose or sparsely
minutely scales or spiny. Fibrillose rachis seen in D. sorzogonense and D.
loerzingii. Spines are present in D. accedens var spinosum and D. spiniferum.
Because the existence of fibrillose scales or spine are only in few species, in this
study the rachis and costa characters were not used.
e. Fronds architecture
Fronds architectures in Diplazium are varying, they are simple to
quadripinnate. Most of West Malesian species have pinnate to bipinnate fronds.
Simple fronds is seen in D. subserratum, but in very young fronds are pinnate
(Figure 6.5.). Deeply pinnatifid fronds, sometimes with one pair of free to
numerous segments, are seen in D. fuliginosum, D. lomariaceum and D.
porphyrophyllum. D. cordifolium has both simple and imparipinnate fronds.
Imparipinate fronds are in D. aequibasale, D. bantamense, D. lobbianum D.
donianum, D. halimunense (a new species proposed in Chapter 10), D. riparium,
D. xiphophyllum, and D. wahauense. The intermediate fronds between pinnate
and imparipinnate is seen in D. pallidum. Bipinnate-tripinnatifid fronds are
present in D. umbrosum. Tripinnatifid fronds are seen in D. megasegmentum, D.
melanolepis and D. subpolypodioides. In D. subalternisegmentum fronds are
tripinnate. Tripinnatifid to quadripinnate frond is seen in D. moultonii.
95
Figure 6.5. Frond architectures of Diplazium. a. & b. D. subserratum, a. juvenile fronds, b. adultfronds; c. pinnate frond of D. silvaticum var. pinnae-ellipticum; d. imparipinnate frond of D.donianum; e. bipinnatifid frond of D. spiniferum; f. bipinnate frond of D. esculentum.
96
e. Venation types
There are two kind of venation, viz. free venation and anastomosing
venation (Figure 6.6.). Free venation type may be forked or pinnate in the lobes.
Generally forked veins are present in the species with simple to simply pinnate
fronds or in species that have intermediate fronds between simpli pinnate-pinnate
such as D. subserratum, D. aequibasale, D. bantamense, D. lobbianum, D.
riparium and D. pallidum. Veins may forked once or more. Whereas pinnate
veins are seen in the species with pinnate to quadripinnate such as D. petiolare, D.
silvaticum, D. malaccense, D. dilatatum, D. polypodioides and so on.
Reticulate venations are seen in D. accedens, D. angustipinna, D.
esculentum, D. cordifolium, D. cumingi, D. insigne, D. megasimplifolium (a new
species proposed, See Chapter 9), and D.squarrosum. There are some species that
generally have free venation, but occasionally show veins uniting at margin such
as in D. fraxinifolium, D. riparium and D. xiphophyllum.
Kato (1977) showed two types of reticulate venation in Diplazium,
goniopterid venation and sagenoid venation. In West Malesian Diplazium,
goniopterid venation are seen in D. esculentum, D. accedens, D. insigne and D.
squarrosum.. Meanwhile the sagenoid venation are present in D. angustipinna,
D. cordifolium, D. cumingii and D.megasimplifolium.
Anastomosing venation in Ophioglossum is believed advanced, free
venation in other genera as primitive (Kato 1987). However the development of
anastomousing venation from free venation may be parallel, it may evolved in
different phyletic lines.
f. Sori
Diplazium sori are dorsal on the vein and linear, elongated along veins
with lateral indusium (Figure 6.6.). They are either single (Asplenoid) or double
(Diplazioid). Asplenoid sori generally occur along the acroscopic side of a vein
and Diplazioid sori are beared on both sides of the basal acroscopic vein. As
stated by Bower (1928), Holttum (1947), and Kato (1977), Asplenioid sori are
produced by abortion of the shorter arm of J-shaped sori along the basiscopic side
of a vein, in view of all intermediate forms. Bower and Holttum interpreted that
Diplazioid sori are produced by interruption of horseshoe- or J-shaped sori at the
97
distal end. Kato (1977) added that the interpretation of Bower and Holttum may
lead to the possibility of Diplazioid sori on higher veins in addition to the basal
acroscopic vein. Diplazioid sori bifurcate along the bifurcating basal acroscopic
vein, as in Athyrium and Deparia. As pointed out by Kato (1977) Diplazioid sori
are not regarded as a morphological unit, but each one of a pair as a mere single
Asplenioid sorus facing either the costule or costa.
The position of sori in Diplazium is important in species determination.
Mainly, there are two type of sori position, viz. subcostular sori and medial sori.
The subcostular sori can be divided into two group, viz. that running from until
touching midveins and those reaching at proximal end. The medial sori may not
touching midvein (inframedial) or almost perfectly medial. Most of West
Malesian Diplazium posses subcostular sori.
Generally sori are not impressed, but some species show impressed sori.
The impressed are present in D.poiense, D. sorzogonense, and D. subserratum.
The existence of impressed sori on D. sorzogonense becomes one of the
diagnostic characters to differentiate this species from D. speciosum.
In the past, a number of phylogenetic schemes presented for ferns was
based great stress on the position of the sorus (marginal vs. dorsal) with only rare
instances of changing from one to another, as in Bower’s “phyletic slide” of
Bower’s scheme (1923-28). Now, this do not faithfully represent all the diversity
within the ferns, the shift from marginal to dorsal sori occurring several times
(Mickel 1974). In the case of soral position on Diplazium, It is not known
whether those subcostular sori are advance or primitive.
g. Indusia
Most species have firm and persistent indusia. Some species posses very
thin indusia and fragile, such as D. procumbens.
Generally, the indusia is concolour, there is not color difference between
attachment side and margin. But some species posses distinctive indusia, marginal
indusia are paler than those at the attachment side, e.g. in D. cumingii and D.
loerzingii (new species proposed, see Chapter 10).
98
Figure 6.6. Venation types of Diplazium. a-c. Free venation, a. vein pinnate in thelobus, veinlets simple, D. batuayaense; b. vein once forked, D. pallidum var. montanum;c. vein forked 3-4 times, D. donianum; d-e. vein anastomousing, d. goniopteris venation,D.accedens; e. sagenoid venation, D. megasimplifolium.
99
Figure 6.7. Sori variation in Diplazium. a. D. densisquamatum; b. D. cordifolium; c. D.accedens; d. D. xiphophyllum; e. D. megasegmentum; f. D. subvirescens; g. D.asymmetricum; h. D. esculentum.
100
Indusia may be entire or not. It would be one of the important characters
in species determination. Most of Diplazium have entire indusia. Generally,
entire indusia are seen in species with simple to pinnate fronds, such as D.
suberratum, D bantamense, D. cordifolium, D. lobbianum, D. pallidum, D.
riparium, D. silvaticum, and D. wahauense. Whereas non entire indusia (toothed
or fringed or lacerate) are found in species with bipinnate fronds, such as D.
atrosquamosum, D. beamanii, D. betimusense, D. esculentum, D. Kunstlerii, D.
laevipes, and D. meijerii.
6.4. Materials and Methods
6.4.1. Taxon Sampling
All species of Diplazium described in the Chapter 9 (69 species) were
included in this study (Table 6.1.). Athyrium anisopterum was chosen as
outgroup. The description of A. anisopterum taken from Holttum (1966).
Athyrium was chosen as an outgroup because this genus was considered as the
closest relatives of the ingroup (See Kato 1977).
6.4.2. Character Examination of Diplazium
Morphological characters used in this investigation were obtained mainly
from the observations of dried specimens. All characters used by Holttum (1940,
1966), Kato (1995) and Tagawa & Iwatsuki (1988) were assessed. Table 6.1.
showed characters, character states, and coding for 88 character utilized in
construction of morphological dataset of Diplazium. A list of taxa studied and the
scores for characters states are provided in Appendix 1.
The morphological datasets matrix was constructed by scoring character
states of the characters chosen from the species descriptions (Chapter 10). All
qualitative characters were coded as binary and multistate. When characters states
could not be defined due to incomplete specimens examined, it was noted as ‘?’.
Missing character states were marked as ‘-‘.
Quantitative characters are coded by using gap-weigting method (Thiele
1993). The steps are as follows: (1) The raw data are initially ranked as an
101
ordered set of states, arranged according to the values of the means; (2) The data
are then range standardized:
xs = (x- min) / max – min) n
where, x = the raw datum, xs = the standardized datum, n = the maximum
number ordered states allowed by the cladistic computer program, in this study n
= 9; (3) The value is coded as the rounded integer of the standardized values; (4)
The characters are treated as an ordered multistate for analysis.
6.4.3. Phylogenetic Analysis
The data matrix was run using PAUP Version 4.0 beta (Swofford, 1998).
Due to the complexity of the dataset, Heuristic searching was employed. All
characters are of type unordered and have equal weight. Gaps are treated as
"missing”. Multistate taxa interpreted as uncertainty. Starting tree(s) obtained via
stepwise addition. Number of trees held at each step during stepwise addition = 1.
Branch-swapping algorithm used tree-bisection-reconnection (TBR). Initial
'MaxTrees' setting = 100. The number of rearrangements tried were unlimited.
102
Table 6.1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium.
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
1. Rhizome Length Short Long
2.Rhizome habit Erect Suberect Creeping
3.Rhizome appearance
Slender Medium Stout
4. Occurrence ofscales on stipes
Sparsely atbase
Sparselythroughoutits length
Densely atbase
Densely atbase tomiddle
Denselythroughoutits length
5. Attachment side ofscales
Subbasal Basal
6. Scales shape Linear Oblongsubtriangular
Lanceolate Narrowlytriangular
Subulate Oblongovate
Ovate Rounded
7. Scales Length (mm) <2.50 2.50 x<5.75 5.75 x<7.5 7.5 x <9.75 9.75x<11.75
11.75 x<13.75) 13.75 x<16.75 16.75 x<18.75 8.75 x<20.75 75 x
8. Scales Width (mm) <0.6 0.6 x <1.25 1.25 x<2.00
2.00 x<2.75
2.75 x<3.30 3.3 x<3.75 3.75 x <4.75 4.75 x <5.75 5.75 x <6 6
9. Scales colour Yellowish Light brown Palebrown
Brown Darkbrown
Black
10. Scales thickeness Thin Thick11. Existence of teethat scales margin
entire bluntlytoohed
sharplytoodhed
12. Type of teeth onscale marginal
teeth simple(not forked
teeth forked
13. Scales (existence ofmargin glands)
Withoutglandularscells
Withglandulars cell
103
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium.
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
14. Scales marginthickness
Withoutthickeningblack strands
Withthickeningblack strands
15. Stipe appearance Slender Medium Stout
16. Stipe (length, mm) <12.25 12.25 x<28.25
28.25 x<36.75
36.75 x<50.75
50.75 x<64.75
64.75 x<78.25
78.25 x<92.75
92.75 x <106.25
106.25 x <120.75
120.75
17. Stipe thickness (nearbase, mm)
<1.25 1.25 x<2.75
2.75 x < 4 x < 5.25 5.25 x<6.75
6.75 x <7.75
7.75 x < 8.75 8.75 x<10.25
10.25 x11.75
11.75
18. Stipe surface Smooth Protuberance Prickly
19. Hair existence onstipe
Glabrous Glabrescent Tomentose
20. Existence ofscales on upper partof stipe
Glabrescent Scales
21. Existence ofscales on stipe base
Glabrescent Sparsely scales Denselyscales
22. Stramineous darkstramineous
paleolivaceous
yellowish palebrown
darkbrown
.black
23. Lamina (division) Simple Pinnatifid Imparipinnate Pinnate Bipinnatifid Bipinnate Tripinnatified Tripinnate
24. Lamina shape Oblong Oblonglanceolate
Oblongsubtriangular
Oblongovate
Oblongsubdeltoid
Oblongovoid
Elliptic Lanceolate Ovatesubdeltoid
Deltoid
25. Lamina length (cm) 13.50< 13.50 x<30.75
30.75 x<47.25
47.25 x<62.75
62.75 x<77.25
77.25 x<94.75
94.74 x<111.25
111.25 x<128.75
128.75 x<145.25
145.25
26. Lamina width(cm)
<10.75 10.75 x<17.25
17.25 x<27.75
27.75 x<38.25
38.25 x<49.75
49.75 x<57.75
57.75 x<68.25
68.25 x<78.75
78.75 x<88.25
88.25 <
27. Lamina incision Entire Undulate Crenate Divided
104
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
28. Lamina base Cuneate Subcordate Cordate29. Lamina apex Acuminate Attenuate Caudate30. Number of lateralpinnae
Absent less than 5 pair 5-10 pairs 11-16 pairs 17-22 pairs 23 – 28 29 or more
31. Position of mostlateral pinnae to therachise
Absent Patent Ascending
32. Deflection oflowest pinnae
Absent Not deflexed Quitedistinctlydeflexed
33. Reduction oflowest pinnae
not reduced slightly reduced muchreduced
34. Stalk existence oflower lateral pinnae
Absent Sessile Adnate Stalked
35. Stalk existence oflower lateral pinnae
Absent Sessile Adnate Stalked
36. Shape of lateralpinnae
Absent Oblong Oblonglanceolate
Elliptical Linierytriangular
Oblongsubtringular
Lanceolate Ovate
37. Length of the largestlateral pinnae (mm)
(<8) x<12.75 12.75 x<20.25 20.25 x<29.1 29.1x<36.90)
36.90 x<43.1)
43.1 x<50.9)
50.9 x<57.1
57.1 x<64.9
64.9 < x
38. Width of the largestlateral pinnae (cm)
<2.75 2.75 x <6.25 6.25 x <10.25 10.25 x<14.75
14.75 x<18.25
18.25 x<22.75
22.75 x<26.25
26.25 x<30.75
30.75 x<34.25
34.25
39. Upper base oflower lateral pinnae
Absent Truncate Subtruncate Cuneate
40. Lower base oflower lateral pinnae
Absent Truncate Subtruncate Cuneate Subcordate Cordate Rounded Auricled
105
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
41. Margin of lowerlateral pinnae
Absent Entire Subentire Undulate Crenulate Crenate Serrate Toothed Lobed ordivided
42. Margin of upperlateral pinnae
Absent Entire Serrate Toothed Slightlycrenate
Lobed
43. Incision of lateralpinnae
Absent Less than ¼way to costa
1/4 – ½ wayto costa
2/3 - ¾ wayto costa
4/5 – 5/6way to costa
6/7 way tocosta– towithin 1mm ofcosta
divided
44. Apex of lateralpinnae
Absent Rounded Acute Acuminate Caudate Attenuate
45. Pinnulae number Absent less than 10pairs
10 -20 pairs more than 20pairs
46. The existence ofstalk on lowerpinnuae
Absent Shortly stalked Subsessile Sessile
47. The existence ofstalk on upperpinnulae
Absent Shortly stalked Subsessile Sessile
48. Pinnulae shape Absent Lineary Oblong Oblongsubtriangular
Oblonglanceolate
Oblongelliptical
Oblongsubdeltoid
Hastate
49. Length of largestpinnule (cm)
<2.25 2.25 x <3.50 3.50 x <5.75 5.75 x <7.75 7.75 x <9.75
9.75 x<11.75
11.75 x<12.25
12.25 x<15.75
15.75 x<17.25
17.25
50. Width of largestpinnule (cm)
(<0.7 0.7 x < 1.3 1.3 x <1.7 1.7 x<2.4 2.4 x <3.0) 3.0 x <3.7 3.7 x < 4.1 4.1 x < 4.5 4.5 x < 4.9 4.9 )
51. Pinnule base Absent Truncate Subtruncate Broadlycuneate
Subcordate
52. Pinnule margin Absent Entire Serrate Crenulate Lobed
106
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium.
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
53. Pinnule incision Absent less than ¼way to costa
1/4 – ½ wayto costa
2/3 - ¾ wayto costa
4/5 – 5/6way to costa
6/7 way tocosta– towithin 1mm of costa
divided = 7(formingsegments
54. Pinnule (apex) Absent rounded acute acuminate attenuate
55. Shape balanceof pinnulae orsegment lobes
Absent not oblique slightlyoblique
56. The width oflobes
Absent less than 5mm wide
5 mm or morewide
57. Form of lobesapice
Absent Truncate Obtuse Rounded Acute
58. Margine oflobes
Absent Entire Subentire Slightlyserrate
Serrate Slightlytoodhed
Crenate Lobed
59. Form ofterminal pinnae
Absent Similar tolateral ones
Deltoid withlobes at base
Withoutdistinctterminalpinna
60. Texture oflamina or pinnae orpinnulae
Herbaceous Subherbaceous Subcoriaceous Coriaceous Shoftlychartaceous
Chartaceous Papyraceous
61. Surfacedepression
Surface notdepressedbelow sori
Surfacedepressedbelow sori
62. Rachis glabrous minutely hairy minutelyscaly
densely hairy denselyscales
107
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium.
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
63. Rachise(viviparous or not)
Notgemmiferous
Gemmiferous
64. Costae Glabrous Tomentose/hairy
Sparselyminutescales
Denselyscales
65. Vein reticulationtype
Free Anastomousing
66. Vein branchingtype
Pinnate Forked
67. Veinlet number onpinnated vein
less than 4pairs
4- 6 pairs 7 pairs ormore
68. Veinlet branchingon pinnated veins)
Simple Forked 1 – 3times
Forkedmore than3 times
69. Vein (uniting withthe opposite onesforming excurrentveinlets or not)
uniting withoppositeones
not unitingwith oppositeones
70. Vein (forking type) Forked 1-2times
Forked morethen 2 times
108
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
71. Vein (part ofanastomousing)
occasionallyanastomousingnear margim but nevercopiouslyanastomousing
anastomousing¼ from themargin
anastomousing1/3 from themargin
anastomousing1/2 from themargin
anastomousing2/3 from themargin
anastomousingmore than43/4 fromthe margin
72. Veins in small groups at anangle about .... to themidrib or costa
about 45º about 50º about 60ºormore
73. Sori position near basal ofveins
middle ofveins or
from nearbasal of veinsor veinletalmost to reach margin
74. Sori covering on the veins or veinlets
Less than 1/3 ofveinlets length
1/3 of vein orveinlet length
1/2 of vein orveinlet length
2/3 – ¾ Nearly thewhole lengthof veinlets
75. sori existence on veinlets
On part ofveinlets
On all veinlets
76. Sori number on each vein group
1 2-4 5 or more
77. Sori (on basal acroscopic vein)
sometimesdiplazioid
usuallydiplazioid
alwaysdiplazioid
78. Indusia form reniform/hooked crescentic linear79. Indusia Width Narrow Broad
109
Table 1. Characters, character states, and coding for 88 characters utilized in construction of morphological dataset of Diplazium.
Characters (0) (1) (2) (3) (4) (5) (6) (7) (8) (9)
80. Indusia appearance Hardlyevident inmaturesori
Evident inmatrure sori
81. Indusia opening openingbeforemature
openingwhenmature
82. Indusia curling Not rolledback whenold
Rolled backwhen old
83. Indusia colour palebrown
brown darkbrown
84. Colour continuity ofindusia
Concolour Attachmentsside darker
85. Indusia margin entirewhenopening
subentire toothed orfringed orlacerate orcrisped
86. Indusia existence Notpersistent
Persistent
87. Indusiastrengtheness
Fragile Robust
88. Spines at thejunction of costules
Absent Present
110
6.5. Results and Discussion
The eighty eight morphological characters (Table 6.1. and Appendix 1.)
were analysed to determine relationship among the 69 species of West Malesian
Diplazium. The eighty eight morphological characters comprises eighty five
parsimony-informative characters, one constant character, and two parsimony-
uninformative characters. The eight equally most parsimonious trees of 1366 steps
produce a Consistency Index (CI) = 0.24, Retention Index (RI) = 0.48, RC = 0.12,
and Homoplasy Index (HI) = 0.75. Figure 6.8. showed the topology of a strict
consensus tree of the eight equally most parsimonious trees of 1366 steps without
the high level of Bootstrap value.
Due to the low level of Bootstrap support, it is very difficult to state any
statement with confidence about the relationships within West Malesian
Diplazium generated from morphological date derived from unweighted
maximum parsimony analysis. The out group (Athyrium anisopterum) is separated
from the in group without Bootstrap value. The relationships within Diplazium
were also unresolved, without high Bootstrap values but clade that comprised D.
silvaticum and D. petiolare and clade that include D. lomariaceum, D.
megasimplifolium, D. subserratum and D. prophyrorachis with Bootstrap support
65% and 82%, repectively.
The lack of support of the monophyly of the genus Diplazium due to the
high proportion of homoplastic characters (HI = 0.75). Homoplasy – the
independent origin or loss of one or more traits in different organism – can distort
the inference of phylogenetic relationship, tying together similar but unrelated
taxa (Givnish & Sytsma 1997).
Although the phylogenetic tree derived from the maximum parsimony was
lack of or weak support, however, it does not indicate that the pattern observed is
incorrect. Triono (2006) stated that the lack of or weak support for a phylogeny
does limit the amount of confidence that can be placed in the relationships
between taxa and also the conclusions that can be drawn from the inferred
phylogeny. Therefore some clades generated from the phylogenetic analysis of
Diplazium that seems formed from the closely related taxa were discussed below.
111
Figure 6.8. Strict concensus of 8 trees of length 1366 from unweighted morphologicaldataset comprises 88 morphological characters. Bootstrap support values between 50-90 are givenabove line. Only support value above 50% are shown.
112
Two major clades were identified in the morphological parsimony analysis
without well-supported Bootstrap value: Clade I and Clade II (Figure 6.8.).
Clade I consist of D. fuliginosum, D. lomariceum, D. porphyrorachis, D.
megasimplifolium and D. subserratum in which D. megasimplifolium and D.
subserratum are the most closely related. In this clade D. fuliginosum is at the
base clade and diverse into four species, viz. D. lomariceum, D. porphyrorachis,
D. megasimplifolum and D. subserratum, respectively. Clade II included most of
West Malesian species and splited into two clades with very low-supported
Bootstrap value, viz. D. tomentosum alone (Clade II.1.) and all the remaining
species (Clade II.2.) . In the Clade II.2. D. crenatoserratum is positioned at the
basal clade and diverse into two subclade, the small clade in which composed of
D. velutinum, D.silvaticum and D. petiolare (Clade II.2.A.) and the large clade
(Clade II.2.B.)
The affinity of D. fuliginosum, D. lamariceum and D. porphyrorachis. In
the topological tree (Figure 8.6.) D. fuliginosum, D. lomariceeum, D.
megasimplifolium, D. suseratum and D. porphyrorachis form a separated clade
from the other West Malesian species and the wo species, D. subserratum and D.
megasimplifolium (a new species proposed, see Chapter 9), are the most closely
related. The affinity of the two species will be discussed separately below. The
affinity of D. fuliginosum, D. lomariceum and D. porphyrorachis are supported by
the share characters as follow: rhizome short-erect, wiry black roots; fronds
narrowly elliptic, deeply pinnatifid or pinnate, apex coadnate, segments numerous,
usually very dark green when living; scales abundant on stipe and rachis, narrow,
usually dark and shiny; veins free. Price (1983) stated that D. fuliginosum is one
of the most unusual of all diplazia, was not transferred to the genus Diplazium
until recently. This species is strickingly peculiar by the smooth rachis channel
without raised sides, uninterrupted by the insertion of pinna-costa. In small
fronds, and distally on large fronds, the rachis is almost flat above. Other features
unusual to Diplazium are sori informally extending from costa to margin, and thin
translucent pinna margin. D. lomariaceum is very closely related to D.
porphyrorachis and until Price (1983) distinguished it from D. porphyrorachis,
the name seems to have been ignored since Christ himself who described is as
113
Asplenium lamoariaceum, reduced lomariaceum to porphyrorachis in Ann. Jard.
Bot. Buitenz. 15 (1898, p.119). Examining many specimens from K, L, MICH,
NY, P, PNH, UC, and US, Price (1983) showed the signs of apparent gene
interchange between D. porphyrorachis and D. lomariceium. In relation of the
two species with D. fuliginosum, Price (1983) believed that rather than to D.
lomariceum and D. porphyrorachis, the closest relationship of D. fuliginosum is to
D. cumingii (Presl.) C.Chr., with which it agrees in dark frond colour and black
axis aging to greeyish; axes without cartilaginous ridges; scales on stipe abundant,
dark, entire; and indusia black with very narrow pale brown margin, curling back
at maturity; even though D. cumingii is very different in its conform frond apex
and broadly elliptic pinnae. Price (1983) added that almost exactly the same frond
form (and even margin structure) as D. fuliginosum was independently evolved in
the Central American D. harrisonii (Bak.) C. Chr. which otherwise differs
markdedly in scales and the architecture of the axes.
The affinity of D. megasimplifolium and D. subseratum. In the Clade I the
closely related between D. megasimplifolium and D. subseratum is not supported
by well-Bootstrap value. The two species possess similarities on characters as
follow: lamina simple with cuneate bases, margin entire on lower part, apex
acuminate. However the two species are very different. The lamina of D.
megasimplifolium is elliptic, much wider (up to 10.5 cm wide), margin always
entire; veins copiously anastomousing up to 4/5 way of margin. D. subserratum
possess lamina lanceolate, less than 4 cm wide, margin entire to irregularly
crenate; veins free. In juvenile stage lamina of D. subserratum is pinnate.
Holttum (1940) presumed that D.subserratum is probably allied to D. lanceum of
India and China but is larger, and a shorter rootstock which is more or less with
tufted fronds; the scales also appear to be smaller.
The relationship among D. malaccense, D. sorzogonense and D
tricholepis. Of three species, D. malaccense is closest related to D. sorzognense.
Ecologically the two species also grow in similar localities, species of lowland
and mid mountain forest in West Malesia. As also stated by Holttum (1940), D.
malaccense is differing from D. sorzogonense in glabrescent stipe and rachis.less
deeply lobed pinnae, and sori not at all impressed. Whereas, D. sorzogonense
114
seems to have affinity with D. tricholepis, mainly its fibrillose rachis, in deeply
lobed of lanceolate pinnae, oblong lobes and veinlet number in each lobe.
‘Imparipinnate frond group’. ‘Imparipinnate frond group’ refer to the
clade in the Figure 8.6. that comprises D. aequibasale, D. angustipinna, D.
donianum, D. cordifolium, D. halimunense, D. cumingii, D. subintegrum, D.
bantamense, D. lobbianum, D. fraxinifolium, D. xiphophyllum, D. hottae, D.
crameri, D. riparium, and D. wahauense. The affinity of these species seems to
be natural. The affinity among these species are discussed below.
The affinity of Clade D. cumingii. Clade D. cumingii consists of D.
cumingii, D. halimunense, D. cordifolium, D. donianum, D. angustipinna, and D.
aequibasale. In this clade, two pair of species, D. aequibasale and D.
angustipinna, D. cordifolium and D. halimunense, are the closest related, D.
donianum to be the sister clade of D. aequibasale and D. angustipinna, and D.
cumingii at the basal clade. The relationship patterns among the species of Clade
D. cumingii seems unnatural. The similarities on many quantitative characters
among these species that seems not to be correlated with their qualitative resulted
the unnatural patterns. The closest relationship of D. aequibasale is to D. riparium
or D. wahauense. Morphologically, D. aequibasale is intermediate between D.
riparium and D. wahauense. The three species share in characters such as
lanceolate dark brown entire scales and oblong lateral pinnae with cuneate base
and margin entire. D. angustipinna and D. halimunense should to be closely
related to D. cordifolium and D. donianum, respectively. D. angustipinna and D.
codifolium share light brown scales, margin entire with irregularly thickening
black strands, ovate-lanceolate lateral pinna, and copiously anastomousing veins.
While D. donianum and D. bantamense share in irregular sharp toothed scales and
ovate-lanceolate pinnae..
The relationship among D. bantamense, D. lobbinaum, D. subintegrum, D.
fraxinifolium, D. xiphophyllum ,D. hottae. D. crameri. The closest relation of D.
lobbianum and D. bantamense and also D. xiphophyllum and D. fraxinifolium are
very reasonable and natural. The two first species share in the following character
combination: dark brown toothed scales; pinnae ovate-lanceolate with rounded
base, margin entire or serrate near apex; vein free and forked several times.
115
Meanwhile D. fraxinifolum and D. xiphophylum are in the collection often mixed
due to the misidentification. D. fraxinifolium differs from D. xiphophyllum in
lineary lanceolate scales (less that 10 mm long), waved-crenate lateral pinnae,
forking type of veins (5-7 times). D. hottae seems to have close relation to D.
xiphophyllum and D. subintegrum. Tagawa (1972) stated that D. hottae is alled to
D. subitegrum. D. hottae differs in: terminal pinna not lobed at the base and
similar to upper lateral ones, pinnae apparently entire, venation obscure, sori
narrower. D. crameri is more close to D xiphophyllum, and differs from this
species mainly in characters: scales dark brown, pinnae stalked to 1.5 cm long,
ovate-lanceolate, all veins free (D. xiphophyllum is occasinolly showing the
uniting veins near margin) and forked to 2 times.
The affinity of D. riparium and D. wahauense. As explained in Chapter 2,
5 and 9 D. riparium and D. wahauense are morphologically very similar and
closely related (Kato et al 1991). Topological tree showed in Figure 8.6. also
revealed that the two species are closely related. Thus the statement of Kato et
al (1991) has been verified in this study.
The affinity of D. pallidum and D. prescottianum. The morphological
characters between the two species are very similar. Holttum (1940) presumed
that D. prescottianum has close relation to D. pallidum. D. prescottianum differs
from D. pallidum in large brown scales. More lobed edges to pinnae, and many
more soriferous veins in each group.
Relationship among D. velutinum, D. petiolare and D. silvaticum. In the
topological tree inferred from morphological data the relationshop among D.
velutinum, D. petiolare and D. silvaticum are not resolved because the Bootstrap
value is very low (<50%), but the closest related of D. petiolare and D. sivaticum
are supported (Bootstrap value 65%). Morphologically, the two species are very
similar (See Chapter 9). D. petiolare differ from D. silvaticum in characters
combination as follow: D. petiolare has scales linear with distantly teeth and
thickening black strand; upper surface of lamina light green when living, pinnae
lobed ¾ way toward costa, upper base not auricle. Whereas D. silvaticum has
lanceolate scales with closely teeth without thickening black strands; upper
surface of lamina dark green, pinnae lobed ¼-1/2 toward costa
116
This result also showed that this phylogenetic tree generated from
morphological data did not support the taxonomical treatment of van Alderwerelt
van Rosenburgh 1908). Treating Diplazium into two sections based on the
venation type, Eudiplazium for those having free veins and Anisogonium in which
composed of species having anastomousing veins (van Alderwerelt van
Rosenburgh 1908), is not natural. In the topology of the phylogenetic tree
generated from morphological data in this study showed that the species members
of the two sections are mixed and did not form into two distinct lines. D.
megasimplifolium (Anisogonium) is included in Clade I in which also composed
of the other species with free veins (Eudiplazium). In the Clade II the species
with copiously anastomousing veins, D. accedens, D. squarrosum, D.
angustipinna, and D.cordifolium are nested within the free veins species and also
do not fall in one terminal clade. These results suggests that the anastomousing
venation pattern in Diplazium may develop in different line. In evolution this
character occurred parallel. Dickason (1946) stated that free and reticulate
venation are not two fundamentally different things, but rather that reticulate
venation may develop from the open type wherever a diverging meristimatic
center on the margin of the blade primordium meets and merges with an adjacent
center.
Comparison between the phylogenetic tree resulted in this study and the
classification of Kato (1977) could not be done proportionally due to the lack
living collection in many species Nevertheless, all the 27 species of Diplazium
that successfully examined their groove and vascular bundles characters (Chapter
4) could not also be referred to the five Diplazium group of Kato (1977). There
are only two group, D. dilatatum group and D. javanicum group, that could be
reffered to West Malesian species. This topological tree showed that the member
of D. dilatatum group (D. batuayauense, D. dilatatum, D. donianum, D.
polpodioides, D. sorzogonense, D. subpolypodioides, D. speciosum, D.
spiniferum, D. subserratum, D. umbrosum, and D. xiphophyllum) distributed in
many different terminal clades of the main clade (Figure 6.8.). It is indicate that
Kato’s classification could nod be applied on West Malesian Diplazium.
117
6.6. Conclusions
The phylogenetic relationship among species in the genus Diplazium was
very difficult to explain due to the lack of or weak support Bootstrap value.
Therefore the monophyletic species groups in this genus could not be identified
confidently. Consequently, sister-group relationships among these species also
could not be established.
The high homoplastic on morphological characters in Diplazium distort the
inference of phylogenetic relationship among species. Moreover parallel evolution
seems occurring.
Lack of or weak support for a phylogenetic tree does not strictly indicate
that the pattern observed is incorrect but it does limit the amount of confidence
that can be placed in the relationships between taxa.and the conclusions can be
drawn from them. This study showed that some terminal clades formed are
consisting of species that presumed to be closely related by formerly authors and
congruence with the some terminal clades of gene rbcL tree, such as the affinity
of D. porphyrorachis group and ‘imparipinnate frond group’ {((D. subintegrum
(D. bantamense, D. lobbianum)) ((D. fraxinifolium, D. xiphophyllum) D. hottae))
D. crameri) (D. riparium, D. wahauense)} This study also revealed that the
classification of van Alderwereld van Rosenburgh (1908) in dividing Diplazium
into two sections (Eudiplazium and Anisogonium) is not natural. Moreover, this
study gave indication that Kato’s classification (1977) could not be applied on
West Malesian Diplazium. The lack of phylogenetic signal in morphological
datasets shows the need for other more informative data, such as molecular data,
for inferring phylogentic relationships.
Cytological data in the Chapter 5 showed that Diplazium is very
complicated. It indicates that cytological information is very important in
detecting the evolution and speciation in the Diplazium. Therefore in the future
cytological data, including both somatic and gametic chromosome, the
chromosomal behavior in meiosis, and chromosomal karyotype, should be
incorporated in the phylogenetic analysis of Diplazium.
118
CHAPTER 7
SPECIFIC DELIMITATION AND RELATIONSHIP AMONG SPECIESOF DIPLAZIUM BASED ON SPORE MORPHOLOGY
7.1. Introduction
Ferns spores display remarkable variation and have been extremely heplful
in systematic studies. In many cases particular spore morphology are distinctive
for families or genera, or in some instance individual species (Taylor & Mickel
1974). Extensive studies on the spore of Pteridophyta showed that spore
morphology can be used as one of the taxonomic evidence for delimiting taxa.
Ferrarini et al. (1986) reported the illustration, by scanning electron microscope,
of the spores of the Pteridophyta native in Italy amount to 124 specific and
intraspecific taxa. They showed that spore morphology provide supporting
taxonomic evidence for delimiting both generic and species level in the
Pteridophyta. Even they recognized some infraspecific of Asplenium, such. A.
trichomanes, A. ruta-muraria, A. seelosii and A. officinarum, based on its spore
morphological characters. Harris (1955) described spores from 170 taxa of New
Zealand fern based on samples taken from herbarium specimens. Forty six years
after Large & Braggins (1991) revised this work and provided full spores
description of 211 species from New Zealand based on samples taken from fresh
material. These works also reveled that spore morphology among the genera in a
family are diversified so that a tentative general key to the genera of pteridophytes
from New Zealand is given. However they did not discuss dealing the correlation
between palynological characters and its taxomical treatments. Ohta & Takamiya
(1999) used spore morphology characters that observed by using SEM to
distinguish Diplazium griffithii from other species in the Diplazium mettenianum
complex.
Palynological characters may also provide evidence to illustrate the pattern
of relationship. Blackmore (2000) showed the the use of pollen morphology, in
isolation, as a means of explicitly illutrating the relationship pattern of species
included in subtribe Scorzonerinae (Asteraceae). The direction of the
transformation was determined by method of outgroup comparison, because it is
119
very difficult to determine the transformation of characters on the basis of the
evolutionary trend approach. Even it is also difficult to polarize characters on the
basis ontogeny. Therefore one solution to the issue of character polarization is to
analyze the characters as unordered, or unpolarized, unless development evidence
for the transformation is available, and allow parsimony to determine the direction
of change.
In effort to develope new circumscriptions of genera in the Vittariaceae,
with the goal of orgnizing the species into genera that are srictly monophyletic,
Crane (1997) mapped the states for four morphological characters, including
spores type, onto the tree generated from phylogenetic analysis using rbcL gene
sequences data (Crane et al 1995).
The exospore is considered to be extremely important for establishing
evolutionary relationship (van Konijnenberg-van Citteret 1999). Exospore
architecture is very diverse. The detailed exospore structure revealed by light and
electron microscopy was first studied by Pettit (1966). Since then, many
pteridologists have dealt with the subject and its culminated with the work of
Tryon and Lugardon (1991) which describes pteridophytes spores from all over
the world using SEM (Scanning Electron Microscope) and TEM (Transmission
Electron Microscope).
Any determination of what constitutes a primitive (ancestral) or derived
state in the structure of fern spores must be based on homologies. Connecting this
subject, Van Konijnenburg-van Cittert (1999) discussed the evolution of various
characters of spore shape and wall structure based on fossil evidence. The
characters discussed include number of spores per sporangium, spore size, overall
spore shape in connection with laesura, thickness and sculpture of exospore,
absence or presence of a perispore, and its sculpture. The evolutionary trend in
fossil fern spores are summarized as follow: (1) A number of 256 spores per
sporangium is primitive, while 128 or less spores per sporangium is derived.
There is a trend of reduction of the number of spores per sporangium; (2)
Standard spore diameter is between 30 and 60 um. Spore diameter over 60 um is
derived. Spore diameter under 30 um may derived, but is probably the most
primitive stage; (3) Tetrahedral, trilete spores are primitive. Bilateral, monolete
120
spores are derived. The evolution from trilete to monolete spores are seen in
several fern families. Trilete and monolete spores may even occur in one
sporangium; intermediates have been recorded; (4) The most primitive overall
spore shape in trilete spores is probably globose spores, while (rounded) triangular
spores are derived; (5) Exospores between 1 and 2 um are usually primitive, but
thin and smooth exospores correlated with a thick and elaborate parispore may be
derived. Exospores over 3 um and certainly over 6 um, are derived; (6) Smooth or
fine exospore ornamentation is possibly primitive, but in the Schizaeceae the
oldest recorded spores appear only late during evolution; (7) Exospores
sculptures that are certainly derived are: bisculpate spores; strongly raised
laesurae; laesurae bordered by margo; valvate spores; and elaboration of the
equatorial region; (8) Absence of perispore is primitive; presence of perispore is
derived; (9) Thin, smooth or finely ornamented perispores consisting usually of
more than one layer and having an elaborate ornamentation are derived.
This chapter present spore morphology of Diplazium from West Malesia
observed by using both Light Microscope (LM) and Scanning Electron
Microscope (SEM. Due to the available of spores collection, only 27 species
were observed of 69 species recorded in West Malesia. The aims of the research
were to: (1) get supporting data for species delimitation on Diplazium; (2) provide
a tentative key for species based on spore morphology; and (3) infer phylogenetic
relationship between species in the genus Diplazium based on spore characters.
7.2. Materials and Methods
Spores of twenty six species of 46 collection numbers of Diplazium were
observed by using scanning electron microscopy (SEM) for getting detail of its
perine ornamentation. The list of collection number examined are presented on
Table 1. These species were chosen based the spore sample availability.
However the 26 species sampled have represented two main clades and eight
subclade of the topological tree generated from morphological data in Chapter 6.
Spore sample were collected both from fresh material collecting from the
field and herbarium material housed at Herbarium Bogoriense (BO). For
determining its size, both polar (P) and equatorial (E) measurements, and
121
morphological outline, 65 collections number were observed by using light
microcopy (LM).
a. Preparation for light microscopy (LM)
Spore sample, both obtained from fresh spores (air drying without
alcoholic treatments) and from herbarium specimens, were directly transfered on
slides and mounted with glycerine jelly for study with light microscopy. Spore
size, polar (P) and equatorial (E) measurements, were taken for each spores. Each
individual plant was taken 20 spores randomly. Each species was represented by
one to five collection numbers.
b. Preparation for scanning electron microscopy (SEM)
Separation and Attaching Spores on the Holder. Spores sample used were
fresh spores (air drying without alcoholic treatments) and spore of herbarium
specimens housed at BO. Spore sample were separated from the indusium and
sporangium fragment of indusium. Separation was conducted manually. One
spatula of spora sample were spread over on waxpaper 4.5 x 4.5 cm. Waxpaper
with spores were shaked by hand. Spores will separate from the fragments.
Evaporation and Platinum Coating. Spores were spread over and attached
on the holder. Spores samples on the holders were evaporated by using Ion
Sputtering for 5-7 minutes. After that spores sample were coated with Platinum
Carbon by using HITACHI HCP-2 for 10 minutes by using Electron Beam
Coating Method.
Observation and Taking Pictures. Spore sample on the holder coated by
Platinum Carbon were observed by using scanning electron microscope (SEM) S-
800 HITACHI. Spore pictures were taken on the longitudinal ekuator, transversal
ekuator, dan polar views.
c. Description format and taxonomic characters
Description are given in full for each taxon. Terminology used to describe
spores morphology following those used by Harris (1955) and Large & Braggins
(1991).
d. Phylogenetic Analysis
Data matric (Table 5.2.) was analyzed using maximum parsimony, PAUP
Version 4. (Swofford 1998), with the Heuristic search settings. All characters are
122
of type unorder and have equal weight. Gaps are treated as "missing". Starting
tree(s) obtained via stepwise addition. Athyrium filix-femine was used as
reference taxon, out group. Number of trees held at each step during stepwise
addition = 1. Branch-swapping algorithm was runned by using tree-bisection-
reconnection (TBR).
7.3. Results and Discussion
7.3.1. Spore Characters of Diplazium and Its Use in Supporting Species Delimitation and Identification
All Diplazium species have spores morphological characteristic
similarities as follow: golden yellow to brown when released, monolete,
bilaterally symmetrical, and heteropolar; polar outline elliptical, sides convex;
equatorial longitudinal view concave-convex to plano convex; equatorial
transverse view, proximal face planar to concave, distal face convex to
hemispherical. SEM observations revealed that a thick perine is present; laesura
is often concealed by perine. Perine is alate to costate alate, wing-like folds. The
description of each species is presented on Table 7.1.
Character used to describe Diplazium spores are as follow: 1) Pattern: The
general pattern of the perine, may appear, (a) folded, or b) spinulose; 2)
Fenestration: perine may be, (a) fenestrate throughout, (b) fenestrate in the
lacunae of any reticulation, or (c) without fenestrae; 3) Ornamentation: Overall
ornamentation is (a) smooth, (b) veined or wrinckled, or (c) raised ribs within the
lacunae. Large & Braggins (1991) mentioned one more character state, viz. (c)
tuberculate with knob-like elements, however this character state is not found for
Diplazium of Western Part malesian Region. 4) Pattern density: Density classes
are (a) sparse and (b) dense; 5) Perine fold type: Perine folds may vary, general
forms are (a) winged (alate) or (b) ridged (costate); ridges may be rounded or
slightly crested; 6) Fold margins: The edges of the perine folds may (a) extend
into serrate to echinate projections, (b) bear minute teeth, papillae, or (c) be
smooth.
123
Kramer et al (1990) described general morphological chraracteristic for
spores Diplazium, viz. with long, usually prominent, wing-like folds, often with
echinate borders, and sometimes cristate or echinate, but without giving further
explanation for the existence of the grouping based on spores characters. Roubik
(2003) showed alate spores of D. grandifolium of Barro Colorado.
Spores of West Malesian Diplazium fall into nine main pattern groups
based on the perine ornamentation. The nine group are discussed as follows.
(1) Group I. The spore form most common among the diplazioid ferns of
Western Part of Malesian region are costate, non-fenestrate, and ornamented with
a wing-like pattern raised to form a pattern of slightly rounded ridges. D.
accedens, D. bantamense, D. lobbianum, D. polypodioides, D. pallidum, D.
procumbens, D. sorzogonense, D. speciosum, D. subpolypodioides, D. dilatatum,
and D. xiphopyllum are included in this group. The existence of additional
ornaments, however, would differentiate among the species. Reticulation type of
wing-like muri or wing-like costae, terminating margin of wing-like muri, and
surface of perine (both on lacunae and wing-like muri) of these members are in
varying character states. Therefore these characters can be used to support species
delimitation. Spores of D. accedens, D. bantamense, and D. polypodioides are
similar appearance. They are seen incomplete reticulation of wing-like muri or
costae often incomplete with terminating margins entire or smooth. D. accedens
and D. polypodioides differ with D. bantamense in the projection of wing-like
muri or costae. The two first species showed wing-like muri or costae project to 7
m, while the third species has wing-like muri or costae project to 13 m. D.
accedens and D. polypodioides would differ in the size of their irregular polygons
lacunae, the first species is with 10 m or more across while the second species 9
m or less across.
124
Table 7.1. Spore Description of Diplazium in West Malesia
No. Species Spores Description & Specimen Examined
1. D. accedens Blume Monolete, bilaterally symmetrical, heteropolar; polar outlineelliptical, sides convex; equatorial longitudinal view concave-convex to plano-convex; equatorial transverse view, proximal faceplanar to concave, distal face hemispherical; perinate. E:25.34(31.47)36.02±2.54; P: 15.32(19.51)23.59±2.82. Laesure:concealed by perine ridge. Perine: costate-alate, reticulation oftenincomplete; lacunae shallow, project 10-17 m ;wing-like muri orcostae project c.0.5-6 m, terminating margins entire. Exine: oftenvisible through perine, smooth under LM, smooth under SEM.
Specimens Examined: Java: West Java, Mt. Gede, T.Ng.Praptosuwiryo s.n.
2. D. bantamenseBlume
Monolete, bilaterally symmetrical, heteropolar; polar outlineelliptical, sides convex; equatorial longitudinal view concave-convex; equatorial transverse view, proximal face concave, distalface hemispherical; perinate. E: 42.81(53.88)66.05±5.69; P:22.64(31.48)37.47±3.38 Laesure: concealed by perine ridge.Perine: costate-alate, loosely reticulate irregular envelope, costaeform a large reticulation; separated from the spores; reticulationoften incomplete; lacunae large irregular polygons 11-25 macross; costae or alate project 3-13 m, terminating marginsentire; surface of perine smooth. Exine: often visible throughperine, smooth under LM.
Specimens examined: Java: West Java, T.Ng. Praptosuwiryo1707; 1766.
3. D. cordifolium Blume Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex; equatorial longitudinal view (excludingperine) plano-convex; equatorial transverse view, proximal faceplanar, distal face convex; perinate. E: 37.02(42.15)51.62 ±5.09P: 20.23(26.56)31.21±2.97. Laesura: concealed by perine wing.Perine: alat to costate-alate, loose reticulate; irregular envelopeseparated from exine surrounds the spore in continuousanastomosing wings, forming a loose reticulation; lacunae largeirregular polygons 15-20 µm across; thin wing-like muri project2-8 µm, terminating margins are often echinate; surface of perinesparsely echinate or ciliate; echinae project 0.5-0.8 µm. Exine:visible through perine, smooth under SEM.
Specimens examined: Java, West Java, Mt. Salak, T.Ng.Praptosuwiryo 1367, 1457; Mt. Halimun, T.Ng. Praptosuwiryo1708; 1709; 1775.
125
Table 7.1. Continued
No. Species Morphological characteristic of Spores
4. D. crenatoserratum(Blume) Moore
Monolete, bilaterally summetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transversely elliptical, sidesconvex; equatorial longitudinal view (excluding perine) concave-convex;equatorial tranverse view, proximal face concave, distal face convex;perinate. E: 29.86(38.19)45.74±4.95, P: 17.92(23.48)27.36±2.94.Laesura: concealed by thin wing-like muri. Perine: alate, looselyreticulate; irregular envelope, separated from exine, surrounds the spore inanastomosing wings, forming loose reticulation, reticulation irregular andoften incomplete; lacunae irregular polygons, 8-16 µm across, irregularholes and small papillae within; holes caused by the fallen papillae; murithin, wing-like, projected c. 1-8 µm, terminating margin ciliate; surface ofperine fibrous-like and holed, holes irregular and formed by fallen ciliae.
5. D. dilatatumBlume
Monolete, bilaterally symatrical (made asymmetric by prine), heteropolar,polar outline (excluding perine) elliptical, sides convex; equatoriallongitudinal view (excluding perine) concave-convex; equatorialtransverse view, proximal face concave, distal face convex; perinate. E:31.85(45.69)49.56±4.78; P: 19.84(28.55)34.62±3.89. Laesura: concealedby perine. Perine: alate-to costate-alate, irregular envelope separated fromexine surrounds the spore; wing-like muri projected c.6 -15 µm µm,terminating margin are often ciliate; surface of perine smooth. Exine:visible through perine, smooth under SEM. Specimens examined: J.J.Afriastini Bl-170
6. D. esculentum(Retz.) Swartz
Monolete, bilaterally symmetrical, heteropolar; polar outline elliptical,sides convex; equatorial longitudinal view concave-convex to plano-convex; equatorial transverse view, proximal face planar to concave, distalface hemispherical; perinate. E: 31.72(38.39)43.09±3.35P: 22.74(26.37)29.12±1.39. Laesure: not visible, concealed by perine.Perine: smooth under LM, micro rugulate under SEM. Exine: Exine: oftenvisible through perine, granulate under LM.Specimens examined: T.Ng. Praptosuwiryo 637
7. D. lobbianumMoore
Monolete, bilaterally symmetrical (made asymmetrical by perine),heteropolar; polar outline elliptical, sides convex; equatorial longitudinalview concave-convex to plano-convex; equatorial transverse view,proximal face planar to concave, distal face hemispherical; perinate. E:38.20(46.10)51.87±3.22; P: 20.47(26.28)32.81±3.73. Laesure:concealed by perine ridge or wing-like muri. Perine: costate-alate; wing-like muri surrounds the spore without forming reticulation or with formingloose reticulatons, terminating margins ciliate; surface of perine smooth toscatterly ciliate or pappilate. Exine: often visible through perine under LM,smooth under SEM. Specimens examined: T.Ng. Praptosuwiryo 1340
8. D. lomariaceum(Christ) Price
Monolete, bilaterraly symmetrical (made asummetric by perine),heteropolar; polar outline (excludong perine) elliptical; equatoriallongitudinal view (excluding perine) biconvex; equatorial tranverse view,proximal face convex, distal view hemispherical; perinate. E:35.03(39.41)54.55±4.20; P: 19.55(24.57)30.13±3.46. Laesura: concealedby wing-like costae. Perine: alate to costate-alate, alae forming loosereticulation; irregular envelope separated from exine surrounds the spore inwing-like muri reticulation, lacunae large, 13-21µm; wing-like muri thin,project 3-10 µm, terminating margin waved or almost entire. Exine: visiblethrough perine, rough rugulate under SEM. Specimens examined: M.Kato, Gen Murata & JP MogeaB7407
126
Table 7.1. Continued
No. Species Morphological characteristic of Spores
9. D. malaccense Monolete, bilaterally summetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) elliptical; equatoriallongitudinal view planar convex to concave-convex; equatorialtransverse view, proximal face concave, distal view convex.Laesura: concealed by wing-like muri. E:37.73(34.24)27.53±3.03; P: 26.01(20.80)13.33±2.82. Perine:alate to costate-alate, often loosely reticulate; irregular envelopeseparated from exine surrounds the spore in continuousanastomosing wings, forming a loose reticulation; wing-like muriproject 1.5-8 µm, terminating margins sparsely ciliate. Exine:visible though perine, smooth granulate under SEM. Specimensexamined: T.Ng. Praptosuwiryo 831, 944-3
10. D. megasegmentumPraptosuwiryo
Monolete, bilaterally symmetrical (mades asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex; equatorial longitudinal view (excludingperine) concave-convex; equatorial transverse view, proximalface convex, distal face convex; perinate. E:39.61(51.42)57.01±4.28P: 18.51(31.49)35.85±3.59. Laesura: concealed by perine.Perine: alate; alae sometimes loosely reticulate; wing-like muriproject 4-13 m, terminating margins almost entire. Specimens examined : T.Ng. Praptosuwiryo 1451.
11. D. pallidum (Blume)Moore
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical; equatorial longitudinal view (excluding perine) plano-convex; equatorial view tranverse view proximal face convex,distal face hemispherical; perinate. Size: E:32.65(47.96)65.02±8.98; P: 17.08(27.09)34.12±5.12Laesura: concealed by perine. Perine: alate under LM, costateunder SEM, irregular envelope separated from exine surrounds thespore in irregular, wing-like costae, often anastomosing to formloose reticulation, lacunae shallow irregular polygons 15-18 µmwide, muri 0.8 – 5 µm wide, surface of muri smooth and lacunaesmoothly granulate. Exine: visible through perine, granulateunder LM.Specimens examined: T.Ng. Praptosuwiyo 1688, 1759; HolttumSFN 31350; J. v. Borsum 2787.
12. D. polypodioidesBlume
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropalar; polar ouline (excluding perine) transversely elliptical,sides convex; equatorial longitudinal view plano-convex toconcave-convex; equatorial tranverse view, proximal view planarto concave, distal view convex; perinate. Size: E:34.36(39.63)43.17± 2.33, P: 16.31(22.92)27.43±2.97Laesura: concealed by wing- like perine. Perine: costate-alate,loosely reticulate; irregular envelope separated from exinesurround the spore with costae forming loose irregular reticulationor wing-like muri, reticulation often incomplete, lacunae largeirregular polygons, 3-9 µm across; wing-like muri project 1-7µm,terminating margins entire; surface or perine smooth under SEM.Specimens examined: T.Ng. Praptosuwiryo 1582, 1604, 1612,1651.
127
Table 7.1. Continued
No. Species Morphological characteristic of Spores
13. D. porphyrorachis(Baker) Diels
Monolete, bilaterally summetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex; equatorial longitudinal view (excludingperine) plano-convex, ; equatorial tranverse view, proximal,proximal face planar, distal convex; perinate. E:39.89(46.39)50.98±3.53, P: 21.74(27.57)30.49±2.87Laesura: concealed by perine. Perine: alate to costate-alate, alaesometimes anastomosing forming loose reticulation; irregularenvelope, separated from exine, surrounds the spore in looseanastomosing wing, forming a very loose reticulation, reticulationoften incomplete; lacunae very irregular polygons, 10-16 macross; muri thin, wing like muri project 1.5-9.0 m; terminatingmargin ciliate or papillate; surface of perine holed, holes irregularand formed by fallen papillae. Specimens examined: W. Meijer872, T.Ng. Praptosuwiryo 930
14 D. prescottianum(Wall.) Moore
Monolete, bilaterraly symmetrical (made asymmetric by perine),heteropolar, polar outline (excluding perine) transverselyelliptical; equatorial longitudinal view plano-convex; equatorialtransverse view, proximal face planar, distal face concave;perinate.E: 32.72(39.11)51.44±5.16, P: 20.23(24.79)33.56±3.63Laesura: concealed by perine. Perine: alate, costate-alate, looselyreticulate; irregular envelope separated from exine surrounds thespore in continuous anastomosing wings, forming a loosereticulation; lacunae large, irregular polygons 5-8 µm; thin wing-like muri project 4-9 µm , terminating margins ciliate or echinate.Exine: visible through perine, smooth-granulate under SEM.Specimen examined: J.v. Borssum W. (17-6-1953)
15. D. procumbens Holtt. Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical; equatorial longitudinal view (excluding perine) plano-convex; equatorial transverse view, proximal face convex, distalface hemispherical; perinate. Size: E: 43.03(53.78)63.20±4.72,P: 29.52(33.22)37.31±2.39Laesura: concealed by perine. Perine: alate under LM, costateunder SEM, irregular envelope separated from exine surrounds thespore in irregular, wing like costae, often anastomosing to form aloose reticalution, lacunae shallow irregular polygons 8-22 µm,muri 1.6-6.6 µm, surface of muri and lacunae smooth. Specimensexamined: R.E. Holttum 36503; T.Ng. Praptosuwiryo 1455.
16. D. profluensPraptosuwiryo
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline elliptical, sides convex; equatoriallongitudinal view concave-convex to plano-convex; equatorialtransverse view, proximal face planar to concave, distal facehemispherical; perinate. E: 36.44(44.06)63.70±8.92; P:23.89(29.04)43.07±5.73. Laesure: concealed by perine ridge.Perine: micro costate, costae broken, reticulate irregularly, denselyechinate under SEM, echinae single elements irregularly apart andprojecting 1.1-2.1 µm. Exine: often visible through perine,smooth. Specimen examined: T.Ng. Praptosuwiryo 1798
128
Table 7.1. Continued
Species Morphological characteristic of Spores
17. D. simpliciveniumHoltt.
Monolete, bilaterally symmetrical, heteropolar; polar outlineelliptical, sides convex; equatorial longitudinal view plano-convexto concave-convex to plano-convex; equatorial transverse view,proximal face planar to concave, distal face hemispherical;perinate.E: 43.99(50.61)54.60±3.36, P: 23.31(29.42)33.78)±2.79Laesure: concealed by perine ridge. Perine: alate to costae alate,irregular envelope separated from exine surrounds the spore inirregular and in continuous or discontinuous anastomosing wing,forming a loose reticulation; lacunae large irregular polygons,occasionally with granulate deposite within; thin wingilike muriprojected 3-8µm, terminating margins are almost entire. Exine:smooth regulate under SEM. Specimens examined: T.Ng.Praptosuwiryo 1250
18. D. silvaticum(Bory) Sw.
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar, polar outline (excluding perine) transverselyelliptical; equatorial longitudinal view (excluding perine) concave-convex; equatorial transverse view, proximal face concave, distalface hemispherical; perinate. E: 31.59(41.31)49.30±3.82, P:18.64(23.56)26.41±2.46Laesura: concealed by perine. Perine: alate under LM, costate-alate under SEM, irregular envelope separated from exine; alaeform loose reticulation; lacunae irregular polygons c. 6.6-20.0 µmacross, interior of lacunae showing micro irregularreticulate/fenestreta under SEM; muri very thin, wing-like,terminating margin irregularly echinate. Exine: smooth-
19. D. speciosumBlume
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex-straight; equatorial longitudinal view(excluding perine) concave-convex; equatorial tranverse view,proximal face convex, distal face biconvex; perinate. E:29.98(39.29)47.25±5.53; P:17.37(21.69)25.83±2.56. Laesura:concealed by perine ridge. Perine: alate, costate-alate, irregularenvelope separated from exine; alae occasionally form looseretication; lacunae irregular polygons c. 6.6-10.8 µm across,interior of lacunae smooth under SEM; terminating margins ofwing-like muri entire. Exine: rugulate, rugulae large, shallow,irregular anastomousing under SEM after separating from perine.Specimens examined: T.Ng. Praptosuwiryo 1242; 1805
20. D. spiniferumAlderw.
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excludng perine) transversely elliptical,side convex; equatorial longitudinal view concave-convex toplano-convex; equatorial transverse view, proximal face planar toconvex, distal face hemispherical; perinate. E:31.96(39.52)47.77±4.18, P: 14.87(23.92)29.21±4.14. Laesura:concealed by perine. Perine: micro-costate, densely echinateunder SEM, costae broken, densely micro reticulate, coralline;irregular envelope, separated from exine, surrounds the spore indensely coralline with enchinae project 1.5-2.3 µm; lacunae arevery small, irregular polygons less than 0.5-1.5 µm across; heavilyfenestrate throughout under SEM, giving coralline appearance.Exine: visible through perine. Specimens examined: M.Kato, M.Okamoto & EB Walujo B10800
129
Table 7.1. Continued
Species Morphological characteristic of Spores
21. D. sorzogonensePresl
Monolete, bilaterally summetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex; equatorial longitudinal view (excludingperine) plano-convex; equatorial transverse view, proximal faceplanar to convex; distal face hemispherical; perinate. E:27.80(33.03)37.90±3.91, P: 13.95(20.92)26.39±3.82. Laesura:concealetd by perine. Perine: alate, costate-alate, irregularenvelope separated from exine; lacunae irregular polygons c. 8.3-16.6 m accross; terminating margins of costae or wing-like murientire. Exine: smooth under SEM.Specimens examined: T.Ng. Praptosuwiryo 1713.
22. D.subpolypodioidesBlume
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar; polar outline (excluding perine) transverselyelliptical, sides convex-straight; equatorial longitudinal view(excluding perine) concave-convex; equatorial tranverse view,proximal face convex, distal face biconvex; perinate. E:32.49(40.29)47.17±4.08, P: 17.15(21.54)28.28±2.90Laesura: concealed by perine ridge. Perine: alate, costate-alate,irregular envelope, reparated from exine; costae or alae sometimesform loose reticulation; lacunae irregular polygons c. 3.3—12.5µm, interior of lacunae smooth under SEM; muri, wing-like,smooth, terminal margin entire. Exine: smooth under SEM.Specimens examined: T.Ng. Praptosuwiryo 1335, 1605
23. D. subserratum(Blume) Moore
Monolete, bilaterally symmetrical, heteropolar; polar outlineelliptical, sides convex; equatorial longitudinal view concave-convex to plano-convex; equatorial transverse view, proximal faceplanar to concave, distal face hemispherical; perinate. Size:E:27.88(41.19)45.49±5.03, P: 16.54 (27.13)20±5.03. Laesure:concealed by perine ridge. Perine: costate-alate, irregularenvelope, separated from exine; costae form irregular loosereticulation; lacunae irregular polygons c.to 23 µm across; interiorof lacunae and costae smooth. Exine: visible throught perine,smoothly granulate under LM. Specimens examined: T.Ng.Praptosuwiryo 1705, 1704
24. D. subvirescensPraptosuwiryo
Monolete, bilaterally symmetrical (made asymmetric by perine),heteropalar; polar outline (excluding perine) transversely elliptical,sides convex; equatorial longitudinal view (excluding perine)plano convex, equatorial transverse view, proximal face planar,distal view convex; perinate. Laesura: concealed by wing-likealae spinous coralline perine. Perine: alate to costate alate, alae orcostae forming irregular reticulation, reticulation often incomplete;lacunae with echinae and coralline within, wing-like muri project3-11 µm, terminating margins pappilate or echinate; pappilaeproject c.0.7-1.5 µm; surface of perine echinae – coralline withdensely irregular hole, windows-like. Exine: not visible throughperine under SEM. Specimens examined: T. Ng. Praptosuwiryo1178.
130
Table 7.1. Continued
Species Morphological characteristic of Spores
25. D. tomentosumBlume
Monolete, bilaterally summetrical, heteropolar; polar outlineelliptical, sides convex, equatorial longitudinal view plano-concex-concave convex, equatorial transverse view, proximal faceplanar to concave, distal face convex; perinate. E:34.64(41.06)47.19±3.88, P: 23.14(26.77)28.67±2.13. Laesura:concealed by perine. Perine: alate to costate-alate, looselyreticulate; irregular envelope separated from exine surrounds thespore in continuous anastomosing wings, forming a loosereticulation, occasionally wing-like alae only form 3 reticulationsurraound the spore; lacunae large irregular polygons, papillaewithin; thin wing–like muri projected 3-12 µm, terminatingmargin are often ciliate, ciliae projected to ca. 0.5 µm. Exine:visible through perine, granulate under LM. Specimens examined:T.Ng. Praptosuwiryo 827, 829, 911.
26. D. vestitum Presl Monolete, bilaterally symmetrical (made asymmetric by perine),heteropolar, polar outline (excluding perine) transverselyelliptical, sides vonvex; equatorial longitudinal view (excludingperine) concave-convex; equatorial tranverse view, proximal faceconvex, distal face convec to hemispherical; perinate. E:28.21(32.58)35.33±2.26, P: 16.55(19.94)22.73±1.64. Laesura:concelated by perine. Perine: costate-alate, almost no reticulationirregular envelop, separated from exine, surrounds the spore withalae-costae ridge (occasionally) projected 2-5 µm; surface ofperine smooth. Exine: visible through perine, smooth under SEM.Specimens examined: M.Kato & H. Wiriadinata B 6097
Spores of West Malesian Diplazium fall into nine main pattern groups
based on the perine ornamentation. The nine group are discussed as follows.
(1) Group I. The spore form most common among the diplazioid ferns of
Western Part of Malesian region are costate, non-fenestrate, and ornamented with
a wing-like pattern raised to form a pattern of slightly rounded ridges. D.
accedens, D. bantamense, D. lobbianum, D. polypodioides, D. pallidum, D.
procumbens, D. sorzogonense, D. speciosum, D. subpolypodioides, D. dilatatum,
and D. xiphopyllum are included in this group. The existence of additional
ornaments, however, would differentiate among the species. Reticulation type of
wing-like muri or wing-like costae, terminating margin of wing-like muri, and
surface of perine (both on lacunae and wing-like muri) of these members are in
varying character states. Therefore these characters can be used to support species
delimitation. Spores of D. accedens, D. bantamense, and D. polypodioides are
similar appearance. They are seen incomplete reticulation of wing-like muri or
131
costae often incomplete with terminating margins entire or smooth. D. accedens
and D. polypodioides differ with D. bantamense in the projection of wing-like
muri or costae. The two first species showed wing-like muri or costae project to 7
m, while the third species has wing-like muri or costae project to 13 m. D.
accedens and D. polypodioides would differ in the size of their irregular polygons
lacunae, the first species is with 10 m or more across while the second species 9
m or less across.
(2) Group II. Second group included D. subserratum and D. vestitum are
revealed spores with costate-alate and sparsely fenestrate perine, micro holes are
irregularly scattered. The existence and its density of the pappilae on the wing-
like muri would be important in differentiating taxa in this group. D.
subbserratum showed entire terminating margin of wing-like muri, while D.
vestitum have ciliate wing-like of muri.
(3) Group III. The third group included members D. crenatoserratum and
D. prescottianum have alate and sparsely fenestrate perine spores. Additional
decoration of perine on both lacunae and wing-like muri add the variation of spore
sculpture of this group.
(4) Group IV. Spores of D. silvaticum are very distinct among the
diplazioid species of Western Part of Malesian region. Its wings-like muri and
lacunae are coralline and fenestrate, irregular windows or holes large (ca. 0.5-1.5
m across) and decorated with irregular dense echinae on terminating margins of
wing-like muri.
(5) Group V. One species of Diplazium showed a rather simple
ornamentation is D. esculentum. Its perine is micro rugulate without wing-like
muri or costae. Spores with monolete, bilaterally symmetrical, elliptical, sides
convex and regulate perine is also found in other genera, such as Athyrium. Using
LM, Tseng-Chieng (1981) observed spores of this species under the name
Anisogonium esculentum and described that its perine are reticulate with scabrate
processes. As reported by Large & Braggins (1991), spores with micro-rugulate
perine is showed in Athyrium filix-femina. However this species is decorated with
costae sparsely that formed shallow and very large lacunae. A. erythropodum and
132
A. vidalii of Thailand observed by Tseng-Chieng (1981) were also revealed
rugulate spores.
(6) Group VI. Alate non-fenestrate perine sculpture can be seen in D.
cordifolium, D. tomentosum, D. simplicivenium, D. megasegmentum, and D.
malaccense. The variation of wing-like muri ornamentation, such as its
reticulation, lacunae formed, projection, and terminating margins, differentiate
among species. Spores of D. tomentosum and D. malaccense are similar. The
two species would differ in the projection of wing-like muri. The first species is
with wing-like muri project to 14 m, and the second with projection of wing-like
muri to 8 m. D. cordifolium differ from other species of this group in its
incomplete reticulation and large lacunae (to 16 m or more across). Spores of D.
simplicivenium differ from those of D. megasegmentum in the existence of
granulate deposit in lacunae. The first have granulate deposit within lacunae,
while the second is without granulate deposit in the lacunae.
(7) Group VII. Spores that are decorated in echinate perine and non-
fenestrate are found in D. profluens. Echinae is only a single elements and its
projection size is diverse. Density and size projection of echinae would be very
useful for differentiating species in this group. Echinate spores under LM were
also showed in D. subsinuatum and D. virescens of Thailand (Tseng-Chieng
1981). The projection of echinae was also useful to differentiate the two species,
echinae of D. subsinuatum spores were far longer than of those D. virescens.
(8) Group VIII. Diplazium spiniferum, D. lomariaceum, D.
porphyrorhachys are have spores with coralline, echinate and fenestrate
sculptures. Three species can be differentiated by its more detail perine
sculptures. D. spiniferum shows densely corraline with echinae project 1.5-2.3
m with very small lacunae (0.5-1.5 m). D. lomariceum has large lacunae (13-
21 m) with wing-like muri project 3-10 m. The projections of wing-like muri
of D. porphyrorachis (1.5-9.0 m) and its across size of irregular lacunae
polygons (10-16 m) are overlap with those of D. lomariceum. Morphologically,
the two last species are very similar.
133
Fig. 7. 1. Group I. a. and b. D. accedens; c. and d. D. bantamense; e. D. lobbianum; f. and g. D.pallidum; h-j. D. procumbens; k-l. D. sorzogonense. Bar = 15 m for a, b, j, ; Bar = 30 m for c,d, e, g, h, and k. Bar = 3 m for i and j.
134
Figure 7.2. Group II. a. D. subserratum; b-c. D. vestitum; d-e. D. vestitum var.borneense; Group III. f-g. D. crenatoserratum; h-i. D. prescottianum; Group IV. j-l. D.silvaticum. Bar = 30 m for a, b, d, f, h, j, and i. Bar = 3 m for c, e, g, i, and k.
135
Figure 7.3. Group V. a-c D. pallidum; Group VI. d-e. D. cordifolium; g-i. D.tomentosum; j.. D. malaccense; k-l. D. megasegmentum; m -o. D. simplicivenium. Bar =30 m for a, c, d, e, f, g, h, i, k, and m. Bar = 3 m for b, h, l, n, and o.
136
Figure 7.4. Group VII. a-c. D. profluens; Group VIII. d. D. spiniferum; Group IX.e-f. D. subvirescens. Bar = 30 m for a, b, and e. Bar = 3 m for c, d, and f.
(9) Group IX. Spores that have form alate-echinate and fenestrate are
found in D. subvirescens. In a glance, this spore type is similar to those of D.
profluens. However D. subvirescens spores would be differentiated from D.
profluens spores with its micro lacunae sculpture. The lacunae decorated with
coralline and densely irregular windows.
Diplazium spores of Thailand observed by Tseng-Chieng (1981) using LM
revealed alate to costate-alate sculpture in eight species, viz. D. dilatatum, D.
squamigerum, D. wichurae, D. megaphyllum, D. mettenianum, D. petri, D.
phaeolepis and D. doederleinii. More detail of the its perine surface, however,
can not be seen so that the comparison between these costate-alate species and
costate-alate species of Malesia could no be done. Because Tseng-Chieng (1981)
did not provide the spores figures from SEM.
137
This study indicates that perine characteristics are very important in
supporting delimitation of species Diplazium observed. So that a tentative key for
species of Diplazium observed is successfully given.
Key to the Species of Diplazium based on Spore Morphological Characters Observed byUsing SEM
1. Perine alate to costate-alate
2. Perine coralline, fenestrate on wing-like muri ………………………………….......... D. silvaticum
2. Perine not coralline, not fenestrate on wing-like muri
3. Terminating margins of wing-like muri or costae almost entire
4. Reticulation often incomplete
5. The mean of equatorial size over 40 m or more
6. Wing-like costae project up to 13 m; exine smooth under LM D. bantamense
6. Wing-like costae project up to 5 m; exine visible granulate under LM ................................................................... D. pallidum
5. The mean of equatorial size less than 40 m
6. Lacunae (irregular polygons) 10 m or more across …........... D. accedens
6. Lacunae (irregular polygons) 9 m or less across ……............. D. polypodioides
4. Reticulation complete
5. Perine alate
6. Lacunae with granulate deposit within ….................. D. simplicivenium
6. Lacunae without granulate deposit within
7. Exine regulate ……………………………............ D. lomariceum
7. Exine smooth
8. Wing-like muri project up to c. 7 m ........... D. procumbens
8. Wing-like muri project up to c. 13 m .. D. megasegmentum
5. Perine costate-alate
6. Surface of perine (lacunae) smooth granulate . ..... D. subpolypodioides
6. Surface of perine (lacunae) smooth irregular fibrous
7. Wing-like muri project up to c. 4 m ...………… D. speciosum
7. Wing-like muri project up to c.6 m ………….. D. sorzogonense
138
3. Terminating margins of wing-like muri or costae echinate or ciliate
4. Reticulation often incomplete
5. Lacunae to 12 m across
6. Surface of perine smooth or scarcely micro papillate; not coralline ………………………………………………… D. xiphophyllum
6. Surface of perine densely sharp echinae or spine; coralline .… D. subvirescens
5. Lacunae to 16 m or more across
6. Perine not forming holes from fallen ciliae ……………..… D. cordifolium
6. Perine forming holes from fallen ciliae
8. Like-fiber materials on muri not forming continuous lines …………………………… D. porphyrorachis
8. Like-fiber materials on muri forming continuous lines …………………………... D. crenatoserratum
4. Reticulation complete
5. Terminating margins of wing-like muri almost entire or smooth …. D. subserratum
5. Terminating margins of wing-like muri ciliate
6. Costae or alae usually not forming reticulations ……………… D. vestitum
6. Costae or alae forming many reticulations
7. Perine alate
8. Wing-like muri project to 9 m
9. Wing-like muri with irregular holes ………. D. prescotianum
9. Wing-like muri without irregular holes
10. Wing-like muri project up to 8 m …….. D. malaccense
10. Wing-like muri project up to 15 m ….…. D. lobbianum
8. Wing-like muri project to 14 m ………….………. D. tomentosum
7. Perine costate-alate
8. Lacunae without irregular holes within ………...…. D. dilatatum
1. Perine not alate to costate-alate
2. Perine rugulate ………………………………….……………………………… D. esculentum
2. Perine ciliate to coralline-echinate
3. Not fibrous-coralline, not fenestrate . ……………………………………….. D. profluens
3. Thick fibrous-coralline, fenestrate .............................................................. .............. D. spiniferum
139
7.3.2. Phylogenetic Analysis
The evolutionary trends presented by Van Konijnenburg-van Cittert
(1999) can not be applied in plylogenetic relationship in Diplazium because they
appear suitable to be implicated to genus level and above. All derived spore
characters states summed up by Konijnenburg-van Cittert (1999) are exist in
Diplazium (see Table 1.) All species of Diplazium usually reveal 32 and 64 spores
per sporangium for apogamous and sexual, respectively. Therefore the direction
of the characters state transformation was determined by method of outgroup
comparison. Table 5.2. showing the characters and character states utilized in
phylogenetic analysis of Diplazium. The rooting of the inferred phylogenetic
relationship revealed in Figure 1. was conducted by selecting spore description of
Athyrium filix-femina cited from Large & Braggins (1991).
The 17 spore morphological characters (Table 7.2. and 7.3.) were
analysed to determine relationship among the 27 species of West Malesian
Diplazium. The 100 equally most parsimonious trees of 131 steps produce a
consistency index (CI) = 0.53 and retention index (RI) = 0.50.
Figure 7.5. showed tree number 1 of 100 the most parsimonius trees.
However the strict consensus of 100 the most parsimonious trees resulted a
polytomy tree with very low supported Boostrap value (Fig. 7.6.). Separation
between the ingroup and outgroup was also without a high supported Bootsrap
value.
The results showed that there is an incongruence between the tree
generated from gross morphological data and those generated from spore
morphology (See Chapter 6). As showed in Figure 7.6., the concensus tree of
100 most parsimonius tree, relationship among the species of Diplazium based
on spore morphology are polytomy and elusive. Although topological of tree
number 1 of 100 most parsimonious trees showed differentiation among 27
species analyzed (figure 7.5.), however, it did not give any logical relationships
between the two closely related species.
Figure 7.5. showed that both the main clades and sub clades are formed by
the grouping of species that are not closely related species based on the gross
morphological evidences. For example, D. porphyrorachis and D. lomariaceium
140
are closely related (See Chapter 5). In the clade II the two species were separated,
the first species was in the same dichotomous branch with D. cordifolium and the
second species was forming dichotomous branch with D. prescottianum. In the
morphological tree, D. porphyrorachis and D. lomariacium were uniting,
meanwhile D. prescottianum formed a dichotomous branch with D. pallidum. As
explained in the Chapter 6, the reasonable affinity of D. porphyrorachis should
be with D. lomariaceum, and D. pallidum with D. prescottianum.
Incongruence between two independent or more data in phylogenetic
analysis of ferns occasionally occurs. Dubuisson et al (1998) has evaluated the
interaction between three independent data sets (anatomy/morphology, cytology
and molecules) within the ferns genus Trichomanes (Hymenophyllaceae). It was
revealed that the data sets showed high and significant level on incongruence.
The illogical relationship of the spore topological tree result in this study
also indicate that superficial similarity of mature perispores may not be the best
indicator of systematic relationship among species. This indication was also
showed by Rangker (1989) in the study on spore morphology of new aorld
Hemionitis, Gymnopteris, and Bommeria (Adiantaceae). Ranker (1989)
presumed that similar surface patterns of spore may be derived through different
development pathways.
141
Figure 7.5. The cladogram of tree number 1 of 100 the most parsimonius trees
142
Figure 7.6. The strict consensus of 100 the most parsimonious trees. The numberabove line showed supported by the Bootstrap value.
143
Table 7.2. Characters, character states, and coding for 17 characters utilized in construction of spore morphology dataset of Diplazium
(0) (1) (2) (3) (4) (5) (6) (7) (8) (9)1. Equatorial longitudinalview
concave-convex
plano-convex
biconvex {01}
2. Proximal face concave planar convex {01} {12}3. Distal face hemispherical convex concave biconvex4. The Equatorial sizemean
31.47-32.37 32.58-34.24
34.25-38.18 38.19-39.63
40.29-42.15
2. 44.06 45.69-47.39
47.96 50.61-51.42
53.78-53.88
5. The Polar size mean(um)
19.51-19.94 20.80-2154 21.69-22.92 23.48 -24.57
24.88-26.28
26.37-27.57
28.55-29.04
29.42 31.48-33.22
6. The existence ofperine fold
no fold with fold
7. Perine fold type costate costate-alate
alate echinate {12}
8. End of ridge rounded slightlycrested
{01}
9. The existence ofcoralline
not exist exist
10. Type of perine foldreticulation
incompletereticulate
completereticulate
{01}
11. The widest lacunae 1.50 – 2.50 2.60-4.60 4.70-6.70 6.80-10.7 10.8-12.5 10.90-18.5 18.6-20.3 20.4-23.7 23.8-26.112. Fold of CrestMargine
Smooth/entire Waved Papillate Ciliate Echinate {01} {23} {24} {34}
13. The longestprojection of wing likemuri or costae
1-2.1 2.9-4 5 6-7 8 9 10-11 12 13 15
14. Perine surface Smooth Fibrous-like
Ciliate/papilate Holed Echinate Smallgranulate
{05}
15. Existence ofmicroscopic holes onperine
Absent Present
16. Exine appearanceunder SEM
Smooth Granulate Rugulate {01}
17. Exine appearanceunder LM
Smooth Granulate Rugulate {01}
144
Table 7.3. Coding for 17 characters utilized in construction of spore morphology dataset of Diplazium
Species Characters and Characters States
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17Athyrium filix-femina 3 3 0 2 4 1 0 0 0 0 0 0 0 5 0 2 0D. accedens 3 3 0 0 0 1 1 0 0 0 0 0 3 0 0 0 0D. bantamense 0 0 0 9 8 1 1 1 0 0 9 0 8 0 0 0 0D. cordifolium 1 1 1 4 5 1 3 2 0 1 7 4 4 0 0 0 0D. crenatoserratum 0 0 1 3 3 1 2 1 0 0 6 2 4 1 1 0 0D. dilatatum 0 0 1 6 6 1 3 1 0 0 0 3 9 0 0 0 0D. esculentum 3 3 0 3 5 0 - 0 - - 0 - 0 0 0 1 2D. lobbianum 3 3 0 6 4 1 1 1 0 0 0 6 - 2 0 0 0D. lomariaceum 2 1 0 3 3 1 3 1 0 - 0 5 6 0 0 2 2D. malaccense 3 0 1 1 1 1 3 2 0 1 0 3 4 0 0 0 1D. megasegmentum 0 2 1 8 8 1 2 1 0 0 0 0 8 0 0 - -D. pallidum 1 2 0 7 8 1 0 0 0 0 6 0 2 0 0 1 0D. polypodioides 3 3 1 3 2 1 1 2 0 0 3 0 3 0 0 1 0D. porphyrorachis 1 1 1 6 5 1 3 1 0 0 7 0 5 0 0 0 0D. precottianum 1 1 2 3 3 1 3 1 0 1 3 8 4 0 0 0 3D. procumbens 1 2 0 9 9 1 2 2 1 0 8 5 3 0 0 0 0D. profluens 3 3 0 5 6 1 0 1 0 0 0 4 0 4 0 0 0D. simplicivenium 3 3 0 8 7 1 3 1 0 2 0 0 4 6 0 3 0D. silvaticum 0 0 0 4 3 1 2 1 0 0 7 4 - 0 0 3 0D. speciosum 0 2 3 3 1 1 3 1 0 0 4 0 - 0 0 2 0D. spiniferum 3 4 0 3 3 0 1 1 0 0 0 - 0 0 0 0 0D. sorzogonense 1 4 0 1 1 1 3 2 0 0 6 0 - 0 0 0 0D. subpolypodioides 0 2 3 4 1 1 3 2 0 0 4 0 - 0 0 0 0D. subserratum 3 3 0 4 1 1 1 2 0 1 8 0 - 0 0 - 0D. subvirescens 1 1 1 - - 1 3 1 1 0 0 7 - 4 1 - -D. tomentosum 3 3 1 4 5 1 3 2 0 1 0 3 7 0 0 1 0D. vestitum 0 2 3 1 0 1 1 0 0 0 0 0 2 0 0 0 0D. xiphophyllum 3 4 1 4 4 1 3 2 0 0 3 2 6 0 0 0 0
145
7.4. Conclusions
All species Diplazium have monolete, bilateraly symmetrical, heteropolar,
polar outline elliptical, perinates with laesura concealed by its perine. Perine
ornamentation varied from alate to costate alate and rugulate with variation of
additional decoration such as ciliae, coralline, echinae, and fenestrae. Perine
ornamentations support in delimitating Diplazium species. Nevertheless
morphological variation of spore is inadequate to depict natural relationship
among Diplazium species. Superficial similarity of mature perispores would not
indicate on systematic relationship among species as similar surface pattern it
derived through different development pathways.
146
CHAPTER 8
MOLECULAR SYSTEMATIC OF DIPLAZIUM FROM WEST MALESIA
8.1. Introduction
The lack of informative morphological characters in ferns led to the search
for new sources of characters in molecular data, including restriction site and
nucleotide sequence data, to infer phylogenetic relationships (Eastwood et al
2004). Phylogenetic analysis on Diplazium based on morphological characters
(Chapter 6) revealed that Diplazium has only a little informative morphological
characters and a hight homoplasy (HI=0.76). The lack of the informative
morphological characters and the high of homoplasy suggests that additonal
caharcters are required to asses the relationship within this genus.
Advances in molecular biology have provided systematists with a valuable
source of characters. Correlating in inferring phylogeny of the vascular plants,
plant sistematists are depending upon chloroplast genome. Most phylogenetic
reconstructions in plant systematics conducted so far is based on molecular data
from cpDNA genes. The chloroplast genome is well suited for evolutionary and
phylogenetic studies, because: (1) the chloroplast genome is small (typically
between 120 and 200 kb), making it relatively easy to examine the entire genome
via ristriction site analysis; (2) it contains primarily single copy genes; (3) has a
conservative rate of nucleotide subtitution; and (4) extensive background for
molecular information on the chloroplast genome is available (Soltis & Soltis
1998).
The most common gene used to provide sequence data for plant
phylogenetic analysis is the plastid-encoded rbcL gene (Chase et al 1993). It is
located in the large single-copy region of the chloroplast genome and encodes the
large subunit or ribulose 1,5-bisphosphate carboxylase/oxygenase (RUBISCO).
This single copy gene is aproximately 1430 base pairs in length; insertions or
deletions (indels) are extremely rare.
147
The use of gene rbcL sequences for phylogenetic analysis has been
reviewed in Palmer et al (1988). On family level and above, rbcL is preferred for
inferring phylogeny. rbcL are not only widely used in the analyzing the extant
taxa, but also found in the leaves fossil samples from Miocene, ca. 17-20 millions
ago (Golenberg et al. 1990, Soltis et al, 1992; Manen et al, 1995). This gene have
proved useful in elucidating higher-order phylogenetic relationships in
angiosperms (Chase et al, 1993; Duvall et. al. 1993).
Recent use of rbcL sequence data in constructing pteridophyte phylogeny
showed that interspecific as well as higher order relationship could be resolved
(Hasebe et al. 1994; Wolf et al. 1994; Hennequin et al 2006). rbcL also could
circumscribe the genera of the fern family (Crane 1997; Smith et.al. 2006).
Phylogenetic trees based on rbcL sequences support the current classification that
the Matoniaceae and its genera, Matonia and Phanerosorus, each are treated as
monophyletic groups (Kato & Setoguchi 1999). Moreover a large number of rbcL
sequence variation has been reported in various species such as Asplenium nidus
(Yatabe et al 2001), Hymenasplenium obliquissimum (Murakami et al 1998),
Stegnogramma pozoi (Thelypteridaceae) (Yatabe et al 1998), Osmunda
cinnamomea, O. claytonia, and O. regalis (Osmundaceae) (Yatabe et al 1999) and
Cheiropleuria bicupsis (Dipteridaceae) (Kato et al 2001). The substitutions rates
of rbcL in Osmundaceae were estimated to be one nucleotide substitution
between two rbcL sequences occurs only one in 5 million years on average
(Yatabe et al 1999). Considering this slow evolutionary rate for rbcL, the large
amount of rbcL sequence variation within a single morphological species may
suggest that these species contain several cryptic species that are reproductively
isolated from each other.
rbcL gene sequence has been used as a tool in unraveling the taxonomic
problems in ferns. Based on rbcL gene sequence which are also supported by
morphological and cytological characters, Sano et al (2000a) moved D.
subsinuatum and D. tomitaroanum into Deparia lancea and Di. tomitaroana,
respectively. The phylogenetic analysis on nucleotide sequences of the
chloroplast-encoded rbcL gene that included Loxoscaphe thecifera (Aspleniaceae)
and Actiniopteris radiata (Pteridaceae) results robust clades and consequently the
148
two species should be included into Asplenium and Onychium, respectively
(Gastony & Johnson 2001). Moreover characters of rbcL sequences information
are useful in the discovery of cryptic species in ferns, e.g. Asplenium nidus
(Yatabe et al 2001).
rbcL gene sequence has been applied in studying the phylogeny of the
Lady fern group, tribe Physematieae (Dryopteridaceae) (Sano et al 2000a). Based
on the chloroplast rbcL gene sequence included 42 species of tribe Physematieae,
including Diplazium, Sano et al (2000a) showed that: (1) the monophyletic
Diplazium clade included Monomelangium; (2) Athyrium, Cornopteris,
Pseudocystopteris, and Anisochampium form a monophyletic clade and Athyrium
is polyphyletic. However the result of Sano et al (2000a) is preliminary because
they examined only few (10 species) of the ca. 400 species of Diplazium. In their
preliminary inferred phylogeny also showed that Diplazium wichurae from
eastern Asia and D. lonchophyllum from central America form a clade in the rbcL
trees. They suggested that more detailed studies including species from wider
areas should be useful for biogeographic studies of Diplazium.
In this present study, gene rbcL sequences from 54 collections number
representing 29 species of West Malesian Diplazium and 9 collections number of
9 references species outside Malesia were analyzed. The objectives of this were
five-fold: 1) to recognize genetic diversity within species; 2) to obtain supporting
evidences in species delimitation; 3) to obtain more informative characters for
inferring phylogenetic hypothesis of Diplazium; 4) to evaluate the monophyly of
Diplazium, and 5) to investigate species-level relationships within the genus.
8.2. Materials and Methods
DNA analysis was carried out at the Laboratory of Molecular Systematic
of Department of Biological Science, University of Tokyo, from 2002 to 2003
and 2006-2007. Materials were collected from 25 localities of Java, Sumatra, and
Borneo (Table 8.1.).
149
8.2.1. DNA Analysis
a. Plant materials.
Sample used were obtained from fresh dried material (kept in Silica Gell)
collected both from the field and plant growing at green houese of Bogor Botanic
Gardens (Table 8.1.). Voucher specimens were deposited in Herbarium
Bogoriense (BO) and Herbarium of Bogor Botanic Gardens(BOHB).
b. DNA Extraction.
DNA was extracted by a modified CTAB method of Doyle and Doyle
(1978) or using the Dneasy Plant Mini Kit (Qiagen, Inc., Tokyo) following the
manufacturer’s intruction. (1) 500 um CTAB was poured in the 1.5 ml tube and it
was incubated on 60ºC. (2) A piece of lamina (± 4 cm2) was ground on the bowl
by using liquit nitrogen until forming smooth powder . (3) The smooth powder
was poured into the tube containing 500 um CTAB and shaked for 1 minute and
then incubated in 60ºC for 30-60 minutes.
c. Separating DNA from Protein.
(1) 500 um of cloroform and isoamil alcohol mixture was poured into the
tube containing sample and 500 um CTAB and shaked by hand for 10 minutes. (2)
The tube was centrifuged on 20ºC for 20 minutes at 15000 rpm. (3) The top layer
(containing water with DNA) was poured into the 1.5 um tube. Step no.1 – no.3
were repeated once more.
d. DNA Purification.
(1) 400 ul Isoamilalcohol was added into the tube containing DNA liquit
and than kept in the freezer for15 minutes or longer. (2) The tubes were
centrifuged at 0ºC for 15 minutes at 15000rpm. (3) The liquid was disposed of
(the residu was DNA). (4) 300 ul 70% Ethanol was added into the tube and
shaked by using vortex and centrifuged at 20ºC for 10 minutes 15000rpm. (5)
The ethanol was evaporated by using aspirator. (6) Tubes containing DNA was
dried in the desicator for10 minutes. (7) The DNA was dissolved in 100um TE
and than shaked well.
150
e. rbcL Multiplication by Using PCR.
PCR was performed using a DNA thermal cycler (Perkin-Elmer 9700),
Applied Bosystems, CA) with Ex Taq DNA polymerase (TaKaRa Biomedical,
Tokyo). Some samples were amplified with Amp direct (Shimazu, Kyoto), whike
most others were amplified with Ex buffer. (1) 10xPCR Buffer, dNTPmix, 10 pM
primer x 2, MgCl2, DNA template, Taq polymerase and DW (Destilate Water)
were poured into 0.2 ml tubes. Taq must be kept at 0ºC (in the ice box) before
PCR was begun to run. (2) PCR was runned at 94ºC for 3 minutes and turned on
35 cycles at 94ºC for 30 seconds, at 48ºC for 40 second, at 72ºC for 90 seconds,
and at 72ºC for 7 minutes and than the tubes were kept at 4ºC.
f. Checking PCR Product by Using Electrophoretic
(1) making gel agarose LE 1%. (2) 3.5 ul PCR product was mixed with
dye in the parafilm and than poured into sample holes (3) Elektroforesi was
runned (4) When blue colour reaching the forth line electrophoretic was stopped.
Gel was moved and pu t into etidiumbromid for 20 minutes. (5) Gel was
illuminated by using UV beam; if bars pictures with proportion sizes were seen it
was concluded that PCR product have been multiplied well.
g. PCR Product Purification
PCR products were purified with GFX DNA and GEL purification Kit
(Amarsham Pharmacia Biotech, Piscataway, NJ) or with ExoSAP-IT (USB
corporation, Ohio) following the manufacture’s intruction. a. Autocycle. (1) 3
ul premix, 0.5 ul sequence buffer, 0.5 ul MgCl2, 1.6 ul 2pm primer, and 4.4 ul
DNA were put into the 0.2 ml tubes. The premix was containing of dNTP,
ddNTP, buffer and taq polymerase. Taq polymerase must be kept at 0ºC before
autocyle. (2) PCR was begun at 96ºC for 90 second and than runned at 30 cycles
96ºC for 10 second, at 50ºC for 15 second and at 60ºC for 4 minutes and than the
tubes were kept at 4ºC. b. Purifying Autocycle Product. (1) 12.5ul 100 Etanol
and 0.5um 3M sodium asetat were put into 1.5 ml tubes (2) Autocycle product
were added into the tubes and shaked by using vortex. (3) The tubes were kept in
the ice box for 20 minutes. (4) The tubes were centrifuged for 20 minutes at 20ºC
15000rpm and than their liquit were run off. (5) 250ul etanol 70% was added into
151
the tubes and than shaked by using vortex. (6) The tubes were centrifuged at
20ºC for 10 minutes at 15000rpm speed and than ethanol was evaporated by
using aspirator. (7) Autocycle product in the tubes were dried in the desiccator
for 15 minutes.
h. Gel Squencing Preparation
(1) Glass plate for gel was washed off with water and rinsed off with
miliQ and isoprophanol, and than dried at room temperature. (2) preparing gel.
18g urea was mixed with 5 ml 10x TBE buffer and amount of water and than
stirred by using magnetic stirrer for 10 minutes (3) The mixture was added with
water till 48 ml volume. (4) The mixture was evaporated by using aspirator and
than kept in the refrigerator (5) The mixture was evaporated by using dessicator
for 5 minutes. (6) 10 % APS was made in the tube. (7) Glass plate was lay out
on the flat tabel, tweezers were tightened (8) TEMED 30ul and 270ul APS 10%
were poured into the mixture and shaked manually (9) The Gel was poured into
the glass plate and than chomb was installed. (10) After 2 hours, manicure was
smeared for preventing buffer spilled out.
i. Preparing Samples for Squencing
(1) Formamide and blue dextran (5:1) mixtured was made. 3ul DNA
sample for every coll. number were also prpepared. All samples and blue dextran
must alwasy be kept in the ice box and blue dextran. (2) The tubes containing
samples were shaked by using shaker for 5 minutes. (3) The tubes cantaining
were incubated at 98ºC for 2 minutes and than kept in the ice box..
j. Run Sequencer
(1) Chomb was removed from gel and than changed with the chomb with
36 holes. (2) Glass plate gel was washed and rinsed off with isopropanol and
mili-Q till the white line not seen. (3) Glass plate gel was installed on the
chamber sequencer and than cassette recorder was also installed on the
Sequencer. (4) Checking glass plate gell (if the noise voice was heared from the
machine we must clean again the plate. (5) Chamber was installed on the
sequencer and than 1x TBE buffer was poured into the chamber. (6) All of the
bubbles attaching the holes chomb must be removed by using injector.
152
Table 8.1. List of Taxa Used in This Study
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Outgroup
Athyrium distentifolium Unknown AB059577 Yatabe andMurakami(2001)
Athyrium filix-femina (L.) Roth exMeretens
AY818676 Skog et al (2004)
Athyrium niponicum (Mett.) Hance Tokyo, Japan CT1005 TI AB232413 Tsutsumi and Kato(2006)
Athyrium vidalii (Miq.) Koidz. Chiba, Japan Sano 25 CBM D43894 Sano et al (2000b)Athyrium sheareri Aichi, Japan Sano 41 CBM D43892 Sano et al (2000b)Athyrium yokoscense (Fr. Et Sav.)Christ
Chiba, Japan Sano 22 CBM D43893 Sano et al (2000b)
Ingroup
Diplazium angustipinna Holttum (
Near Sungai Kobet, Track to BatuAyau, Muller Range, CentralKalimantan, Borneo. 440 m.
T.Ng. Praptosuwiryo1905
BO Present Study
Diplazium asymmetricum PraptosuwiryoSp. Nov.
Petak 4 Desa Kemutuk Lor, Convertedforest-Natural forest boundary,Wanawisata Baturraden, Mt. Slamet,Cetral Java. 970-1000 m.
T.Ng. praptosuwiryo1094
BO Present Study
Diplazium asymmetricum PraptosuwiryoSp. Nov. (2=1)
Loop Trail, Cikaniki Forest Research,Mt. Halimun, West Java. Ca. 1000 m
T.Ng. praptosuwiryo1728
BO Present Study
Diplazium cordifolium Blume ‘simplefronds’
Cangkuang Forest, Southern Slopes ofMt. Salak, West Java
T.Ng. Praptosuwiryo1202
BO Present Study
153
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium cordifolium Blume‘Pinnate fronds’
Cangkuang Forest, Southern Slopes ofMt. Salak, West Java
T.Ng. Praptosuwiryo1237
BO Present Study
Diplazium cordifolium Blume‘Pinnate fronds’
Cangkuang Forest, Southern Slopes ofMt. Salak, West Java
T.Ng. Praptosuwiryo1238
BO Present Study
Diplazium cf cordifolium Blume Primary forest between S. Anak Kobet-S. Kobet, track to Batu Ayau, CentralKalimantan, Borneo.
T.Ng. Praptosuwiryo1912
BO Present Study
Diplazium accedens Blume Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1001
BO Present Study
Diplazium accedens Blume Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1161
BO Present Study
Diplazium accedens Blume Cangkuang Forest, Southern Slopes ofMt. Salak, West Java.
T.Ng. Praptosuwiryo1211
BO Present Study
Diplazium bantamense Blume Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park,West Java
T.Ng. Praptosuwiryo1160
BO Present Study
Diplazium batuayanense Praptosuwiryo Above Sungai Kobet, Track to BatuAyau, Muller Range, CentralKalimantan, Borneo. 440 m.
T.Ng. Praptosuwiryo1909
BO Present Study
D_cavalerianum (Christ.) C.Chr. Chiba, Japan Sano 11 CBM D43909 Sano et al (2000b)D. chinense (Bak.) C.Chr. Kumamoto, Japan Ohta &Takamiya 739 KUMA AB021718 Sano et al (2000b)
154
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium dilatatum Blume Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1011
BO Present Study
Diplazium simplicivenium Holttum Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1025
BO Present Study
Diplazium simplicivenium Holtum Converted Forest, Petak 55, DesaKarang Mangu, Wana WisataBaturraden, G. Slamet, Baturraden,Central Java. 860 m.
T.Ng. Praptosuwiryo1073
BO Present Study
Diplazium dilatatum Blume Kagoshima, Japan Ohta & Takamiya602
KUMA AB021719 Sano et al (2000b)
Diplazium donianum (Mett.) Tard. Kagoshima, Japan Sano 29 CBM D43911 Sano et al (2000b)Diplazium esculentum (Retz.) Sw. Situ Patengan, Mt. Patuha, Bandung,
West Java.T.Ng. Praptosuwiryo730
BO Present Study
Diplazium esculentum (Retz.) Sw. Gede-Pangrango National Park, WestJava
T.Ng. Praptosuwiryo1227
BO Present Study
Diplazium esculentum (Retz.) Sw. Kagoshima, Japan Hasebe 276001 TI U05619 Hasebe et al (1994)Diplazium hottae Tagawa Near Sungai Anak Kobet, Track to
Batu Ayau, Muller Range, CentralKalimantan, Borneo. 450 m.
T.Ng. Praptosuwiryo1911
BO Present Study
Diplazium lobbianum Moore Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1007
BO Present Study
155
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium lobbianum Moore Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1031
BO Present Study
Diplazium lobbianum Moore Cibodas Forest, Behind CibodasBotanic Gardens, Track I Mt. Gede,Gede-Pangrango National Park, WestJava. 1500 m
T.Ng. Praptosuwiryo1181
BO Present Study
Diplazium lobbianum Moore Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park,West Java
T.Ng. Praptosuwiryo1008
BO Present Study
Diplazium lonchophyllum Kunze Unknown U05920 Wolf et al (1994)Diplazium megasegmentumPraptosuwiryo Sp.Nov.
Cangkuang Forest, Southern Slope, Mt.Salak, West Java.
T.Ng. Praptosuwiryo1450
BO Present Study
Diplazium mesosorum Tochigi, Japan Sano 44 CBM D43910 Sano et al (2000)Diplazium okudairaekamiDiplazium pallidum (Blume) Moore Near Telaga Warna, Gede-Pangrango
National Park, West JavaT.Ng. Praptosuwiryo1172
BO Present Study
Diplazium pallidum (Blume) Moore Mt. Payung, Ujung Kulon NationalPark, West Java
T.Ng. Praptosuwiryo1406
BO Present Study
Diplazium poiense C.Chr. in C.Chr. &Holttum
Unknown TML1
Diplazium polypodioides Blume Cangar Nature Reserve, Mt. Welirang,Batu, East Java. 1370-1400 m.
T.Ng. Praptosuwiryo667
BO Present Study
Diplazium polypodioides Blume Cibodas Forest, Mt. Gede, Gede-Pangrango National Park, West Java
T.Ng. Praptosuwiryo1185
BO Present Study
156
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium porphyrorachis Unknown TML2Diplazium procumbens Holttum Cibodas Forest, Track I of Mt. Gede,
behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1015
BO Present Study
Diplazium procumbens Holttum Cibodas Forest, Track I of Mt. Gede,Behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1047
BO Present Study
Diplazium procumbens Holttum Cibodas Forest, Behind CibodasBotanic Gardens, Track I Mt. Gede,Gede-Pangrango National Park, WestJava
T.Ng. Praptosuwiryo1163
BO Present Study
Diplazium procumbens Holttum Converveted Forest-Natural ForestBoundary, Petak 4 Desa Kemutuk LorWana Wisata Baturraden, Mt. Slamet,Central Java. 970-1000 m.
T.Ng. Praptosuwiryo1094
BO Present Study
Diplazium procumbens Holttum Cimisblung, Mt. Masigit, Mt.Pangrango, Gede-Pangrango NationalPark. 1300 m.
T.Ng. Praptosuwiryo1281
BO Present Study
Diplazium procumbens Holttum Cangkuang Forest, Souther Slopes ofMt. Salak, West Java.
T.Ng. Praptosuwiryo1216
BO Present Study
Diplazium rhachidosorus Unknown TML3Diplazium riparium Holttum Unknown TML4
157
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium sibiricum var. sibiricum Akita, Japan Ohta & Takamiya695
KUMA AB021720 Sano et al (2000b)
Diplazium silvaticum (Bory) Sw. Wild fern of Bogor Botanic Gardens,Java
T.Ng. Praptosuwiryo1300
BO Present Study
Diplazium silvaticum (Bory) Sw. Wild fern of Bogor Botanic Gardens,Java
T.Ng. Praptosuwiryo1302
BO Present Study
Diplazium silvaticum (Bory) Sw. Wild fern of Bogor Botanic Gardens,Java
T.Ng. Praptosuwiryo1303
BO Present Study
Diplazium simplicivenium Holttum Track to Cibeureum, Mt. Gede, Gede-Pangrango National Park, West Java
T.Ng. Praptosuwiryo1141
BO Present Study
Diplazium simplicivenium Holttum Rawa Denok I, Gede-PangrangoNational Park, West Java
T.Ng. Praptosuwiryo1171
BO Present Study
Diplazium simplicivenium Holttum Near Batu Kukusan I, Gede-PangrangoNational Park, West Java
T.Ng. Praptosuwiryo1175
BO Present Study
Diplazium simplicivenium Holttum Cibodas Forest, Mt. Gede, Gede-Pangrango National Park, West Java
T.Ng. Praptosuwiryo1179
BO Present Study
Diplazium sorzogonense C. Presl. Loop Trail , Cikaniki Research Center,Mt. Halimun, West Java. Ca. 1000 m.
T.Ng. Praptosuwiryo1720
BO Present Study
Diplazium sorzogonense C. Presl. Loop Trail , Cikaniki Research Center,Mt. Halimun, Halimun National Park,West Java. Ca. 1000 m.
T.Ng. Praptosuwiryo1725
BO Present Study
Diplazium sorzogonense C. Presl. Loop Trail, Cikaniki Forest ResearchCenter, Mt. Halimun, HalimunNational Park, West Java. Ca. 950 m.
T.Ng. Praptosuwiryo1731
BO Present Study
Diplazium sorzogonense C. Presl. Jalur Owa, Cikaniki Reseacrh Center,Mt. Halimun, West Java. Ca. 975 m.
T.Ng. Praptosuwiryo1744
BO Present Study
158
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium speciosum Blume Geger Bentang, Mt. Pangrango, Gede-Pangrango National Park, West Java.1960 m.
T.Ng. Praptosuwiryo1243
BO Present Study
Diplazium squamigerum (Mett.)Matsum.
Kumamoto, Japan Ohta & Takamiya893
KUMA Sano et al (2000b)
Diplazium subpolypodioides (Alderw.)Alderw.
Cibodas Forest, Mt. Gede, Gede-Pangrango National Park, West Java.
T.Ng. Praptosuwiryo1184
BO Present Study
Diplazium subserratum (Blume) Moore. Converted Forest, Petak 55, DesaKarang Mangu, Wana WisataBaturraden, G. Slamet, Baturraden,Central Java. 860 m.
T.Ng. Praptosuwiryo1070
BO Present Study
Diplazium subserratum (Blume) Moore. Converted Forest, Petak 55, DesaKarang Mangu, Wana WisataBaturraden, G. Slamet, Baturraden,Central Java. 860 m.
T.Ng. Praptosuwiryo1072
BO Present Study
Diplazium subvirescens Praptosuwiryo Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park1500 m.
T.Ng. Praptosuwiryo1178
BO Present Study
Diplazium subvirescens Praptosuwiryo Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National
T.Ng. Praptosuwiryo1012
BO Present Study
Diplazium tomentosum Blume Loop Trail, Cikaniki Forest Research,Mt. Halimun, West Java. Ca. 1000 m
T.Ng. Praptosuwiryo1722
BO Present Study
Diplazium umbrosum (Smith) Bedd. Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1002
BO Present Study
159
Table 8.1. Continued
Species Locality Collectorand Number
HerbariumHoldingVoucher
GenebankAccessionNo.
References
Diplazium umbrosum (Smith) Bedd. Cibodas Forest, Track I of Mt. Gede,behind Cibodas Botanic Gardens GuestHouse, Gede-Pangrango National Park
T.Ng. Praptosuwiryo1050
BO Present Study
Diplazium wichurae (Mett.) Diels Shizuoka, Japan Sano 13 CBM D43915 Sano et al (2000b)Diplazium xiphophyllum (Baker) C.Chr. Eas Kalimantan, Borneo TD 902 Living Coll.
at Nuserryof BBG
Present Study
Diplazium xiphophyllum (Baker) C.Chr. Loop Trail, Cikaniki Forest Research,Mt. Halimun, West Java. Ca. 950 m.
T.Ng. Praptosuwiryo1717
BO Present Study
Diplazium xiphophyllum (Baker) C.Chr. Loop Trail, Cikaniki Forest Research,Mt. Halimun, West Java. Ca. 950 m.
T.Ng. Praptosuwiryo1719
BO Present Study
Diplazium xiphophyllum (Baker) C.Chr. HM 39 Cikuda Paeh, Cikaniki ForestResearch, Mt. Halimun, West Java.
T.Ng. Praptosuwiryo1791
BO Present Study
160
(7) Prerun for 5 minutes. (8) Stopping for a moment and than samples with odd
number collectons were injected into the odd holes. (9) Resume for 5 minutes.
(10) Stop prerun and than the sample with the even collentions number were
injected into the even holes and than the chambercover was installed (11)
Running.
k. Gene rbcL sequence alignment.
The PCR product were sequenced with the Big Dye Terminator Cycle
Squencing Kit Ver. II or Ver. III (Applied Biosystems, CA) following
manufacturer’s intructions with amplification primers of Hasebe et al (Tabel 1.).
l. Homology searching on the data base by internet (using a reprsentative
database of nucleotide sequence, DNA DataBase of Japan (DDBJ,
http://www.ddbj.nig.ac.jp/) to compare predicted gene sequence with the
registered sequence in the database.
Table 8.2. Primers Used for Amplifying and Sequencing DNA from Diplazium(Hasebe et al, 1994)
Primer Sequence (5’ to 3’) Position
rbcL-aF
rbcL-bF
rbcL-sR
rbcL-aR
rbcL-cF
rbcL-sF
rbcL-cR
rbcL-bR
ATGTCACCACAAACAGAGACTAAAGC
TATCCCCTGGATTTATTTGAGGAAGGTTC
GAACCTTCCTCAAATAAATCCAGGGGATA
CTTCTGCTACAAATAAGAATCGATCTCTCCA
TGAAAACGTCGTGAATTCCCAACCGTTTATGCG
ACTGTAGTGGGCAAATTGGAAGGCGAACG
GCAGCAGCTAGTTCCGGGCTCCA
CGTTCGCCTTCCAATTTGCCCACTACAGT
1-26
307-335
335-307
670-640
609-638
988-1016
1373-1351
1016-988
161
8.2.2. Phylogenetic analysis
DNA sequences were edited and aligned by using Clustal W. All
characters were treated as equally weighted and unordered. Maximum
Parsimony (MP) analysis was performed with PAUP 4.0b10 (Swofford 1998)
using the heuristic search option with 100 random replicates of stepwise data
addition with TBR swapping and Multrees set to 200 trees limit. Boostrap analysis
(Felsenstein 1985) was performed with 200 replicates to evaluate internal support.
Athyrium filix-femina, A. distentifolium, A. niponicum, A. sheareri. A.vidalii and
A. yokoscence were selected as outgroup taxa.
8.3. Results and Discussion
Chloroplast rbcL gene sequences of 25 species of Diplazium from West
Malesia were newly generated (Table 1). Length of the sequences of the 29
species are varying, from 640 to 1445 base pair (bp). There are many missing
data on gene between pair number 722 – 1445. Therefore 721-bp region from the
pair number 1 to 721 were used to make comparison between two cytotypes in a
species or among the closely related species and used for phylogenetic analysis.
However the 721-pb region possess many informative characters, viz. 544 (75%).
8.3.1. Infraspecific Genetic Diversity in Diplazium
Intraspecific nucleotides differences in Diplazium are showed in Table
8.3. The differences among individuals in the same species are 0 – 5 nucleotides,
but in D. dilatatum, D. procumbens and D. subvirescens. There are not any
differences on gene rbcL squences in D. esculentum, D. polypodioides, D.
subserratum, D.simplicivenium, D. silvaticum, and D. xiphophyllum (TNgP1717 -
1719). As revealed in Chapter 5, the two first species are diploid. D. subserratum
showed diploid, triploid and tetraploid. D. simplicivenium and D. silvaticum are
triploid and tetraploid, respectively. Whereas the voucher specimen of the last
species was not know its chromosomes number. These results indicate that these
species are autoploid.
162
The diploid and tetraploid of D. pallidum differ from each other by five
nucleotides between pair number 74 – 721. There are missing data on the diploid
(TNgP 1406) on pair number 1-73. Takamiya et al (2001) reported genetic
comparison between triploid and tetraploid of Diplazium doederleinii. The two
cytotypes differ from each other by only two nucleotides in the rbcL sequences,
and there are no variations in cytotype except for a single plant of the triploid.
Tabel 8.3. Infraspecific Genetic Variaton of Diplazium based on Gene rbcL Sequences
Species Number ofNucleotidesDifference
Notes
D. esculentum TNgP 730 – TNgP 1227 0D. esculentum (Kagoshima-Japan) – Java (TNgP1227)
3
D. accedens TNgP 1001 – TNg 1161 4 In the same locality, G. Gede-G.Pangrango National Park
D. accedens TNgP 1161 (G. Gede-Pangrango) –TNgP 1211 (G. Salak)
1 On nucleotide number 649, inwhich TNgP 1161 has Thymineand TNgP 1211 has Cytosine
D. cordifolium ‘simple’ TNgP 1202 – D.cordifolium ‘pinnate’ (TNgP 1237)
1 On nucleotide number 340, inTNgP 1202 Cytosine pyrimidinebase and TNgP 1237 has Adeninpurine base
D. cordifolium TNgP1238 - 1237 10D. cordifolium TNgP 1912 (berbulu) – D.cordifolium TNgP 1237
8
D. xiphophyllum TD902 – TNgP1717 1D. xiphophyllum TNgP1717 - 1719 0D. polypodioides TNgP 677, TNgP 1184, 1185 0D. procumbens 1015 - 1047 5D. procumbens 1015-1216 7D. procumbens 1047-1216 6 Missing data on nucleotide
number 1-4 and 649 to 721 inTNgP 1216
D. pallidum tetraploid ( 1172)– diploid (TNgP1406)
5 Missing data on TNgP 1406 in1-73
D. subserratum TNgP 1070 - 1072 0D. lobbianum TNgP 1031-1181 5 Missing data on TNgP 1181 in
nucleotide number 1-3D. lobbianum TNgP 1181 - 1008 4D. lobbianum TNgP 1031 - 1008 4D. silvaticum TNgP 1300 – 1302 0D.simplicivenium TNgP 1025, 1073, 1141,1171, 1175, 1179
0
D. dilatatum TNgP 1011 (G. Gede Java –ABO21719 (Japan)
16
D. subvirescens TNgP 1012 – D. subvirescensTNgP 1178
22
163
The high number of nucleotides differences of D. dilatatum, D.
procumbens and D. subvirescens are presumed to due they are hybrid. In Java,
the three species mentioned above are triploid apogamous. However their putative
parents have not been found. Further study are needed to prove this suggestion.
8.3.2. Species Delimitation in Diplazium based on Gene rbcL Sequence
These results revealed that rbcL nucleotides variations in Diplazium are
large enough, but between the two pair of the closely related species D.
sorzogonense TNgP 1744 – D. batuayauense and D. xiphophyllum TNgP 1791 -
D. hottae TNgP 1911. Among species are differentiated from each other by 4-37
nucleotides between pair number 1– 721 (Table 8.4.). The more closely related
species the lower the differences number of nucleotides. Morphological character
of D. cordifoliuman is very similar to D. angustipinna. The two species are
sharing in: imparipinnate fronds with copiously anastomosing veins; scales light
brown, lanceolate, margin entire. The pinnate fronds plant of D. cordifolium
differs from D. angustipinna by 11 nucleotides between pair number 103-363.
While the simple fronds plant of D. cordifolium differs from D. angustipinna by 4
nucleotides between pair number 63-717. The distantly related West Malesian
species of D. megasegmentum and D. porphyrorachis are difference each other in
19 nucleotides. The first species is a huge plant with tripinnate fronds whereas the
second species is a small plant with pinnatifid fronds. These results revealed that
gene rbcL sequences are well supporting in delimiting species in Diplazium.
Recently comparison of rbcL sequences among species on ferns showed
that five or more nucleotides differences are enough to differentiate species. In
genus Diplazium (Sano et al. 2000a, 2000b) also showed that rbcL nucleotide
variation are great, more than four nucleotides, between different species, while
there are no or only two sequence changes within species. The intraspecific
variation study at the molecular level (using rbcL sequencing) in the Japanese
Asplenium of sect. Thamnopteris showed that there are as many as 14-23
differences in the rbcL sequences between Asplenium australasicum (J. Sm.) and
A. setoi N. Murak. et Seriz. (Murakami et al. 1999b).
164
Tabel 8.4. Interspecific Genetic Variaton of Diplazium based on Gene rbcL Sequences
Species Number ofNucleotidesDifference
Notes
D. donianum – D. bantamense 18D. bantamense – D. lobbianum TNgP 1031 13D. cordifolium (TNgP 1737) – D. riparium(TML4)
16
D.esculentum – D. cavalerianum 37D. esculentum – D. chinense 16D. wichurae – D. okudairaekami 5D. wichurae – D.lonchophyllum 17D. squamigerum – D. sibiricum 6D. mesososrum – D. rhachidosorus 12D. cavalerianum – D. okudairaekami 36D. pallidum = D. procumbens TNgP 1163D. cordifolium ‘simple’ – D. angustipinnaTNgP 1905
4
D. cordifolium TNgP 1238 – D.angustipinna TNgP 1905
11
D. xiphophyllum TNgP 1791 – D. hottaeTNgP 1911
1
D. speciosum TNgP 1243– D. batuayauenseTNgP 1909
15
D. xiphophyllum TNgP 1791 – D. ripariumTML4
7
D. dilatatum TNgP 1011 – D.simplicivenium TNgP 1025
16 Missing data in 13 sites
D. dilatatum TNgP 1011 – D. procumbensTNgP 1015
12
D. bantamense TNgP 1160 – D. lobbianumTNgP 1181
11
D. sorzogonense TNgP 1720- 1725 -1731 –1744 – D. batuayauense
0
D. simplicivenium TNgP 1179 – D.dilatatum TNgP 1011
18 Missing data on manyplace, mainly in nucleotidenumber 711-721
D.megasegmentum (TNgP 1450) – D.prophyrorachis
19 TNgP 1450 (nucleotidenumber 63-76 is in N)
D. porphyrorachis(missing data onnucleotides number 699 –721)
D. poiense – D. porphyrorachis 14 D. poiense (missing dataon nucleotide number 649-721)
165
The lack or low of nucleotides difference between D. sorzogonense TNgP
1744 – D. batuayauense and D. xiphophyllum TNgP 1791 - D. hottae TNgP 1911
should be re-checked again in the future studies as the length of the gene rbcL
compared in this study are only 721-bp region from the pair number 1 to 721. The
nucleotides differences between the two pair of these species may be existed
between the pair number 722 – 1445. Because morphological studies (See
Chapter 9) showed D. batuayauense and D. sorzogonense are having many
differences. D. batuayauense differs from D. sorzogonense in the following
characters: Scales sharply toothed with thickening dark brown stand irregularly,
fronds are much smaller, never reaching 1 m, lacks the fibrillose scales on stipe
and rachis, pinnae lobed only to ½ way to costa, sori not impressed. Therefore D.
batuayauense is propsed as new species. The morphogical differences between D.
hottae and D. xiphophyllum can be seen in the Chapter 9.
8.3.3. Informative Characters of Gene rbcL Sequences for Inferring Phylogenetic Hypothesis of Diplazium
In the molecular phylogenetic relationships studies on ferns, generally
rbcL shows a high parsimony informative characters. Wolf et al. (1994) reported
465 phylogenetic informative characters of the 1320 nucleotide sites of
Dennstaedtioid ferns. In the molecular phylogenetic relationships study in the
heterosporous fern genus Marsilea Nagalingum et al (2007) obtained 58
parsimony informative characters from 1233-pb. Sano et al. (2000a) reported 342
parsimony informative characters in the phylogenetic analysis of 68 taxa of ferns
in inferring phylogenetic relationship of the Lady fern group, tribe Physematieae,
including Diplazium, by using 1206 bp region between 73-1278. Lu et al. (2007)
showed 144 parsimony informative characters from 1320-necleotide segment of
the rbcL gene in studying the phylogeny of the Polystichoid ferns in Asia. Hauk et
al (2003), in the phylogenetic studies of Ophioglossaceae, the adder’s tongue
ferns that has limited the number of characters available for understanding
relationships, got 341 potentially parsimony-informative characters of the 1321-
bp.
The DNA sequence of West Malesian Diplazium that only included 721
base pair (bp) contained a high parsimony informative characters, 544 (75%) of
166
parsimony-informative characters. This result showed that gene rbcL sequence is
a good enough marker for inferring phylogenetic hiphothesis of Diplazium.
8.3.4. Phylogenetic Analysis
The 721-bp region from the pair number 1 to 721, numbered from the
initial methionin codon of Nicotiana tabacum was used for analysis. As a
references, species outsite of Malesia are included in the analysis: D.
lonchophyllum from Central America and D. cavalerianum, D. mesosorum, D.
rhachidosorus, D. sibiricum, D. squamigerum and D. wichurae from Japan and
(Table 8.1.).
The aligned matrix of the 77- taxon contained 544 (75%) of parsimony-
informative characters, 35 (5%) variable of parsimony-uninformative characters
and 142 (20%) constant characters. The consistency index (CI) and retention
index (RI; Farris 1989) are 0.79 and 0.94, respectively. The topology of Most
Parsimony (MP) of 8 trees and those strict concensus trees are generally similar
(Figure 8.1.).
8.3.4.1. The Monophyly of Diplazium
Recent molecular phylogenetic analysis of Diplazium and closely related
genera revealed that Diplazium is monophyletic. Preliminary phylogenetic study
of Diplazium conducted by Sano et al (2000a) based on chloroplast rbcL gene
sequences showed the monophyletic of Diplazium clade supported by Bootstrap
value 86%. This clade includes Monomelangium pullingeri (Bak.) Tagawa. By
using evidence from chloroplast trnL-F region sequences Wang et al (2003)
showed the monophyletic of Diplazium clade that include Callipteris Bory,
Allantoidea R. Br. Emend. Ching, and Diplaziopsis C. Chr. with supported
Bootstrap value 98%.
All species studied in this study form a monophyletic group. The
monophyly of Diplazium is strongly support in the tree with 100% bootstrap
value. It is revealed that separation of Diplazium from Athyrium are strongly
resolved. Thus, the monophyly of West Malesian Diplazium has been verified
based on this molecular study.
167
Morphologically, Diplazium can be distinguished from other close related
genera (including from Athyrium) by characters combination as follow (Kato,
1977): (1) stipe bases neither swollen nor bearing pneumatophores; (2) Frond
axes U-shaped with flat base in most species; (3) Acroscopic basal pinnules equal
or smaller; lamina margin not cartilaginous, spines absent; (4) Vein free, or
goniopetrid or sagenoid – anstomousing, ending near the margin; (5) Sori dorsal
on the vein and linear, either single (Asplenoid) or double (Diplazioid); generally
Asplenoid sori along the acroscopic side of a vein and Diplazioid sori are confined
to both sides of the basal acroscopic; both parts of double sori of about equal
length; (6) Scales entire or toothed with consisting of two upturned ends of
adjacent marginal cells. This morphological study (see Chapter 9) supported the
previous workers (Ching 1940; van Alderwerelt van Rosenburgh 1908; Alston
1956; Sledge 1962; Holttum 1940, 1966; Kato 1977, 1995; Edie 1978; Tagawa &
Iwatsuki 1988; Andrews 1990; Kramer et al. 1990).) in separating Diplazium from
Athyrium.
8.3.4.2. Relationships among species within Diplazium
In the monophyletic of Diplazium, D. porphyrorachis form a clade alone
and diverges earliest (Clade I), it is the most basally positioned in all studied taxa
(Clade II). But the divergency of D. porphyrorachis from the all taxa studied is
without well-supported Bootstrap value. Subsequently Clade II diverges into two
main clades without well-supported Boostrap value, viz. Clade II.1. that contains
only D. subvirescens (TNgP 1012) and Clade II.2. that diverges again into four
main clades, one main well-supported group, Clade A, with Bootstraping value
100% and three main weakly-supported group, Clade B, C, and D. The
relationships inferred among the species included in these cades are discussed
below. Before discussing the relationship inferred among the species, the
polytomy occurrence in the topological tree generated is discussed first.
In ferns, polytomy relationship patterns among species on the evolutionary
relationship inferred that generated from molecular data are usual. As showed in
the Figure 8.1., Clade A2 diverges into six subclades. Subsequently in the
terminal clade, Diplazium from Japan, Eastern Asia and Central America are also
168
forming a polytomy. The polytomy pattern is also showed in the sister clade of D.
riparium that comprised D. hottae, D. xiphophyllum, D. cordifolium, and D.
angutipinna. Dubuisson et al (1998) showed polytomy relationship pattern of the
sister clade of Trichomanes thysanostomum that consisted of T. minutum, T.
bipunctatum, T. speciosum, and D. diaphanum. In the monophyly of the section
Lepidoglosa (Elaphoglossaceae), Elaphoglossum asterolepis, E. splendens and E.
rufidulum formed a polytomy (Rouhan et al. 2004). Hauk et al (1993) showed the
polytomy pattern in the sister clade of Ophioglossum pusillum that included O.
reticulatum, O. vulgatum, and O. richardsiae. The polytomy patterns indicate that
more data, including more taxa and more informative characters, are needed in
studying the relationships of the taxa in the future.
D. porphyrorachis. D. porphyrorachis is the sister clade of the main
clade (Clade II). Morphologically, D. prophyrorachis is very distict among the
West Malesian Diplazium. D. porphyrorachis posses a suite of morphological
characters distinguishing it from other West Malesian species. As presented in
Chapter 6 and 9 this species is having diagnostic characters as follow: Lamina
pinnatifid lanceolate apex lobed or toothed, the lower 2/3–6/7 deeply pinnatifid
into many or numerous, close, spreading, subfalcate, linear oblong, blunt, slightly
crenate, serrate or toothed segments 8-15 mm broad, which have not seldom the
sides entire and only the apex serrate; lower segments gradually growing smaller,
the 1-4 lowest free and deflexed. The position of D. prophyrorachis at the basal
tree of the rbcL tree is also supported by the topology of the phylogenetic tree
generated from morphological data. In the tree generated from morphological
data, D. porphyrorachis is forming a clade that include its closely related species,
viz. D. fuliginosum and D. lomariaceum and also at the basal clade of the
monophyly of West Malesian Diplazium. As stated by Price (1983) D.
prophyrorachis and its closely related species, D. lomariceum and D. fuliginosum,
constitute unusual Malesian Diplazium that agree in the following characters:
rhizome short-erect, with thick wiry black roots; fronds narrowly elliptic, pinnate
of subpinnate, apex coadunate, segments numerous, usually dark; scales abundant
on stipe and rachis, narrow, usually dark and shiny; veins free.
169
Figure 8.1. Strict consensus of the 200 equally most-parsimonious trees obtained inmaximum parsimony analysis of the rbcL sequence data. Bootstrap values are indicatedabove the branches occurring in more than 50% of 100 bootstrap replicates.
170
Clade C. Clade C without well Bootstrap support comprised D.
procumbens 1047, D. asymmetricum 1094, D. subvirescens 1178, D.
asymmetricum 1728, D. procumbens 1281. Due the very low level of Boostrap
value, it is difficult to state the relationship among them. Therefore each species
is discussed separately below.
D. asymmetricum. This species in a glance is very similar to D.
procumbens as the two species are having affinity in characters combination:
scales on stipes fallen when mature, dark brown, ovate-lanceolate, margin with
blunt teeth; lamina oblong; lobes oblong with blunt or truncate apex; veinlets
forked. Diplazium assymetricum differs from D. procumbens in the following
characters combination: rhizome short, erect; lamina more incised (to tripinnate)
basiscopic pinnulae or segments and lobes are larger than the acroscopic ones;
indusia thicker, margin entire. Therefore it is possible that the two species are
closest related in the phylogenetic tree inferred based on gene rbcL sequence.
Diplazium subvirescens. In the topology of rbcL tree, Diplazium
subvirescens (TNgP 1012) constitute the sister clade of Clade II in which this
speices is also included in unresolved Clade II.2. (TNgP 1178). Cytological
observations on two individuals of this species (Chapter 5) showed only triploid
apogamous and it is presumed hybrid. Suggestion that this species is originally
hybrids may be support by the evidence from the gene rbcl sequence. Two plants
successfully sequenced (TNgP 1012 and TNgP 1178) showed differences each
other in 22 nucleotides. This difference number was high enough although
morphologically the two collection number are very similar and should be treated
as one species. Morphologically, D. subvirescens is closely related to D.
virescens. As mentioned in the Chapter 9., D. subvirescens has affinity to D.
virescens in the characters combination as follows: rhizome long creeping, black
with densely scales on younger part; scales on stipes lineary lanceolate, polish
dark brown with toothed margin; lamina deltoid, deep green, firm herbaceous with
veinlets prominent on both surface; sori oblong to linear, medial to supramedial;
indusia thin-membranaceous, laciniate at margin, irregurarly broken at maturity.
Two collections number that successfully examined their chromosomes number
(TNg 1013 and TNgp 1177) (See Chapter 5) showed triploid and the two
171
collentions number are having similar morphological characters with TNgP 1012
and TNgP 1178. Therefore, based on these cytological and rbcL evidence it is
suggested that this species are hybrids.
Diplazium procumbens. Diplazium procumbens distributes on four
different clades in rbcL Most Parsimonious tree. Firstly, this species is sister
species of D. pallidum that has subcoriaceous pinnate fronds with lineary
lanceolate sharply acuminate toothed pinnae (up to 27 pairs) less than 3 cm wide
with veins forked 1-3 times and soriferous on upper simple branch, with
Bootstrap value 62%. Secondly, D. procumbens form a clade with D. dilatatum
(TNgP 1011) without supporting Bootstrap value. Thirdly, D. procumbes (TNgP
1047; 1281) form a clade with D. asymmetricum and D subvirescens (TNgP 1177)
without supporting Bootstrap value. The latter, D. procumbens (TNgP 1216)
form a clade alone and become the sister clade of well resolved of Clade A and
weakly supported of Clade B dan C.
D. procumbens is bipinnate diplazioid fern with rhizome long creeping,
very narrow slightly toothed scales, veins copiously forked in the oblong lobes
and sori with thin indusia that break down in the middle. Therefore the
relationship between D. pallidum and D. procumbens in the topological tree
generated from gene rbcL is not logical. In the tree generated from morphology,
D. pallidum is closely related to D. prescottianum (See Chapter 6) and this
relation is more logic.
Cytological observations on eight individuals of D. procumbens from five
localities in Java (See Chapter 5., Praptosuwiryo & Darnaedi 2004, 2005) showed
triploid apagamous. DNA rcbL sequence showed that nucleotides differences
within D. procumbens are high enough, viz. from 1 – 10. As mentioned above,
whereas, interspecific nucleotides differences in Diplazium are 4 – 37 (Table 8.4.).
It is indicate that D. procumbens may be polyphyletic and originally hybirids. In
Malesia, D. procumbens is distributed in Malay Peninsula (Holttum 1940),
Sumatra (present study) and Jawa (Praptosuwiryo 1999).
The position of D. procumbens and D. subvirescens in two or more
different clades may be correlated with the hybrid occurrence in these species.
Funk (1985) stated that if hybridization has occurred among the species of a taxon
172
under cladistic analysis the results are varied but always present additional
difficulties. Hybridization results in incongruent intersecting data that obscure the
underlying hierarchy.
Clade B. Clade B consisted of D. donianum, D. tomentosum, D.dilatatum
and D. simplicivenium without Boostrap support. It is difficult to make a
statement about the relationship among these species due to the very low level of
the Bootstrap value, but between D. simplicivenium and D. dilatatum, D.
simplicivenium may has affinity to D. dilatatum. The two species
morphologically very similar (see Chapter 9). Whereas the relationship between
D. donianum and D. tomentosum is difficult to explain and not logic. The two
species are morphologically very different (See Chapter 9). The affinity of D.
donianum shoud be with D. halimunese (not included in rbcL analysis, See
Chapter 9), and D. maoenense Ching. Moore et al (2002) revealed that D.
donianum and D. maoenense are closely related species. D. donianum is different
from D. maoenense in the following characters: D. maonense has lobed terminal
pinna wuth free lobes in lower part, pinnae crenate at margin and serrate near apex
whereas D. donianum has unlobed terminal pinna, pinnae entire at margin and
slightly serrate near apex.
Clade A. Clade A with well supported Bootstrap values level (100%)
included 30 species in which species from Japan (D. chinense, D. rhachidosorus,
D. mesosorum, D. cavalerianum, D. sibiricum, D. squamigerum and D.
okudairaekami), Eastern Asia (D. wichurae) Central America (D. lonchophyllum)
nested. This Clade diverged into two clade, viz A1 that only consisted D.
megasegmentum and Clade A2 that include 29 species. Subsequently A2 diverged
into five branches that are polytomy: (1) A2.1 had one species only (D. poiense);
(2) well supported clade of A2.2. (65%) that comprised D. bantamense, D.
subserratum, D. speciosum, D. lobbianum, and D. silvaticum; (3) Clade A2.3. that
include two individual of D. umbrosum with strong Bootstrap support; (4) Clade
A2.4 that comprised D. accedens, D. polypodioides and D. subpolypodioides
without Bootstrap value support; (5) Clade A2.5. that included D. pallidum, D.
procumbens TNgP 1163, D. esculentum and species from Japan, Eastern Asia and
Central America; and Clade A2.6. Subsequently, Clade A2.6. diverges into two
173
monophyletic group with well-supported Bootstrap value, ‘sorzogonense’ Group
(Clade A2.6.1) and ‘riparium’ Group (Clade A2.6.2). The first group with well-
supported Bootstrap value (96%) includes two species, D. sorzogonense and D.
batuayauense, meanwhile the second group with Bootstrap value 51% includes
D. cordifolium, D. angustipinna, D. xiphophyllum, D. hotta and D riparium. The
relationships among species within these clades are dicussed below.
Clade A.2.6.2 (=‘riparium Group’). This study showed that the closely
related species (D. angustipinna and D. cordifolium; D. xiphophyllum and D.
hottae inferred from phylogenetic analysis using morphology are supported by
gene rbcL sequence data. The monophyletic of ‘riparium Group’ is characterized
by the entire scales and simple pinnate frond with terminal distinct terminal
pinnae, terminal pinnae conform to lateral pinnae. In the monophyletic of
‘riparium Group’ D. riparium is at basal clade and diverge earliest. Of all species
within ‘riparium group’, but D.riparium, are dryland species. D. riparium grows
on both riparian and dryland areas. In this group D. cordifolium and D
angustipinna are the most closely related and become the sister group of D.
xiphophyllum and D. hottae. D. cordifolium and D. angustipinna have similar
morphological characters in glossy light brown lanceolate scales on stipes and
copiously anastomosing veins up to 1/3-1/2 way of margin. In the topology of
rbcL tree, the relationship of D. xiphophyllum and D. hottae are not resolved.
Tagawa (1972) stated that D. hottae whic is occurring on Malay Peninsula and in
Northern Sumatra is allied to D. subintegrum Holtt.
Affinity between D. dilatatum and D. simplicivenium. D. dilatatum and D.
simplicivenium form a clade without supporting Bootstrap value (Clade B2). The
closest related of the two species in rbcL tree is supported by the similarity in
morphological appearance such as the lineary lanceolate of black-margined
toothed scales, the tuft of gigantic bipinnate fronds, and oblong subtriangular
pinnulae. However, in the morphological tree (Sees Chapter 6) these species are
separated into two different terminal clades, but still in the same middle clade.
Relationships among non West Malesian Species. Clade A2.5.2 without
high Bootstraping value, include D. rachidosorus, D. mesosorum, D.
cavalerianum, D. squamigerum, D. wichurae, D. okudairaekami, D.
174
lonchophyllum and D. esculentum. As revealed also by Sano et al (2000), in the
rbcL tree in this study, D. wichurae from eastern Asia and D. lonchophyllum from
Central America consistently form a clade in the rbcL trees and nested among
species from Japan (D. chinense, D. rachidosorus, D. mesosorum, D.
cavalerianum, D. squamigerum, D. okudairaekam ) and West Malesian species.
It is indicate that more detailed molecular phylogenetic studies that include
species from wider areas should be for phylogeographic studies of Diplazium.
Phylogenetic relationships of D. dilatatum group and the others groups of
Kato (1977). The member of D. dilataum group are spread into many different
terminal clades and also positioned at the subbasal clades. Whereas D.
mesosorum dan D. javanicum group are also mixed in the clade in which the
member of D. dilatatum group forming a calde. It is indicate that Diplazium
groups established by Kato (1977) was not monophyletic. Therefore the all of
West Malesian Diplazium can not be referred to the Kato’s Diplazium groups.
Phylogenetic relationships of the section Anisogonium and Eudiplazium
(van Alderwerelt van Rosenburgh 1908). The rbcL tree is congruent with the
phylogenetic tree generated from morphological data in drawing the polyphyletic
of the section Anisogonium and Eudiplazium. Both the member of the section
Anisogonium and Eudiplazium are mixed, the species of Anisogonium nested in
the many places in which the member of Eudiplazium forming terminal clades.
This study represents the first attempt to explore the phylogeny of West
Malesian Diplazium based on gene rbcL sequences. These results showed that
some clades generating from this sequence are congruence with the clades of
phylogenetic tree generated from morphological data. However this result is
preliminary because this study examined only 29 species from West Malesia.
Molecular phylogenetic study that includes all species of Diplazium from West
Malesia and also use more molecular marker would give the more robust or well
resolved phylogenetic hyphothesis tree.
175
8.4. Conclusions
Gene rbcL sequence is very well in supporting species delimitation among
species within the genus Diplazium. Even this study also showed most of West
Malesian species are showing genetic diversity, mainly those having polyploidy
types or seri ploidy. Therefore the division of D. pallidum into two varieties is
supported with this gene rbcL sequences. This study give an indication that
species in which have both morphological difference and seri ploidy, such D.
accedens, D. cordifolium, D. crenatoserratum, D. bantamens, D. subserratum and
D. tomentosum, should be more studied by encompassing much more sample from
their range of distribution.
This study revealed that gene rbcL is more informative in inferring
phylogeny of the genus Diplazium in West Malesia. Therefore the phylogenetic
analysis of gene rbcL sequences found that Diplazium in West Malesia is
monophyletic.
The position of D. porphyrorachis at the basal clade of the philogenetic
tree generated from morphological data and separated from other species is
supported by the phylogenetic tree generated from molecular data (gene rbcL
sequence). This result is in accordance with the statement of Price (1983) that D.
porphyrorachis and close related species are belonging to the different lines
within the genus Diplazium. This study also showed the congruence between the
clade of ‘riparium Group’ drawn by rbcL tree and morphological tree.
The result of this study also showed that the classification of van
Alderwerelt van Rosenburgh (1908) is not monophyletic. The classification of
Kato (1977) in dividing Japanese Diplazium into six group can not be referred to
West Malesian Diplazium. Moreover the Diplazium groups of Kato (1977) are
also not monophyletic.
This study indicates that some uncertain relationships require further
analysis in the future. More data, including both more taxa that include all West
Malesian species and from wider areas and more molecular data, are needed
before a well resolved phylogenetic hypothesis for West Malesian Diplazium can
be offered.
176
CHAPTER 9
TAXONOMIC STUDY OF THE FERN GENUS DIPLAZIUMIN WEST MALESIA
9.1. Introduction
The genus Diplazium was established by Swartz (1801) and typified by
Asplenium plantaginifolium L (Diplazium plantaginifolium Sw.). Since then, a
total of 1401 of binomials have been published by various authors for Diplazium
in the world (IPNI, 2006). Hassler & Swale (2002) listed 474 species and 8
hybrids of Diplazium in the world.
Malesian region is the center of Diplazium diversity as it was predicted
that 75% of species in world (ca.300 species) are existing in this region (Roos,
1995). However, since van Alderwereld van Rosenburgh (1908), there has not
been a comprehenship study, including a revision, on Malesian Diplazium, yet.
Previous workers only studied Diplazium based local area. Holttum (1940, 1966)
in his revision on ferns of Malaya reported 27 species of Malay Peninsula.
Tagawa (1972) listed 15 species of Diplazium from Borneo. Iwatsuki & Kato
(1984) reported 15 species of diplazioid ferns of East Kalimantan. Parris et al
(1992) listed 31 species from Mt. Kinabalu. Mitsuta (1985) listed 11 species from
West Sumatra. In Java, Backer and Posthumus (1939) decribed 17 species. Sixty
years after that Praptosuwiryo (1999) recognized 22 species and 4 varieties of
Java. Kato (1994) reported 32 species of Diplazium from Ambon and Seram
(Moluccas).
This chapter presented the account of West Malesian Diplazium. The aims
of this study are: (1) to revise the genus Diplazium in West Malesia; (2) to
provide the delimitation of the genus and species, and (3) to provide an
identification keys to the species and infra species.
9.2. Materials and Methods
Morphological study on Diplazium covered West Malesian species have
been conducted. This study is based mainly on available specimens housed at the
BO and SING and new collection specimens obtained from field work study in
177
Java, Sumatra and Kalimantan from 1992 to 2006. The supplemented with images
of type specimens from the data bases/websites of the National Herbarium of the
Netherlands, University of Leiden Branch (L), the United States National
Herbarium in the National Museum of Natural History at the Smithsonian
Institution (US), and Herbarium Universitatis Mosquensis (MW) were also used.
The total number of 1051 collections have been examined. Living plants
in the field were studied in Java, Sumatra and Borneo. In addition, the living
plants grown in botanical gardens and cultivated plants in the green-houses of
Bogor Botanic Gardens were also studied. Three species growing naturally in
Bogor Botanic Gardens, viz. D. esculentum, D. silvaticum, and D.
subpolypodioides, were also examined. The process of undertaking this study
follow the steps those described by Davis & Heywood (1963), Rifai (1976), Vogel
(1987) and Maxted (1992).
9.3. Taxonomic Treatment
Diplazium
Diplazium Sw., Schrad. J. Bot. 1800 (2): 61. 1801. –Type: Diplazium
plantaginifolium Sw.; Backer & Posth., Varenfl. Java: 123. 1939; Holttum, Gard.
Bull. S.S. 11: 77. 1940; Tagawa & K. Iwats., Fl. Thailand 3: 449. 1988.
Callipteris Bory, Voy. 1: 282. 1804. –Type: Callipteris prolifera (Lam.)
Bory; Copel., Gen. Fil.: 152. 1947.
Rhizome creeping to erect, scaly. Stipe well-developed, often stout,
occasionally muricate or spiny, rarely persistently scaly, adaxially sulcate; scales
orbicular to linear, margin entire or toothed; teeth composing two adjacent cells.
Lamina simple to pinnately compound; rachis distinctly grooved, occasionally
bearing adventitious buds; adaxial groove of rachis usually with flat bottom,
glabrous; no spine-like appendages at bases of bases of costae or costules; costa
and costules adaxially grooved. Ultimate divisons costate, costae anadromously
pinnate, veins simple or forked, free or anastomousing, without free included
veinlets. Sori elongate along one or both sides of veins; indusium lateral; double
178
sori with two quite separate indusia. Spores monolete, bilaterally symmetrical,
plano-convex or concave-convex, usually prominent, wing-like folds, often with
echinate border, sometimes cristate or echinate, occasionally rugullate.
CHROMOSOME NUMBERS. X=41.
TYPE SPECIES. Diplazium plantaginifolium (L.) Urban.
DISTRIBUTION. Tropical of both hemispheres. Throughout Malesia.
ECOLOGY. Terrestrially, usually on moist humus-rich soil of mountain
slopes of primary and secondary forest of shadowed places. Only a few species
found in the opened areas and grow on the riparian and rheophytic habitats.
Elevation: 20-3400 m.
NOTES. (1) Doubful Species. Seven species of Diplazium published in
some literatures that I doubt and could not found its specimens as follows:
Diplazium allantoideum M.G. Price, D. curtisii Holttum, D. heterophlebium
(Mett.) Diels, D. mesocarpum Alderw., D. falcinellum C.Chr. in C. Chr. &
Holttum, D. megistophyllum (Copel.) Tagawa, D. tabacinum Copel.
(2) Excluded Species. There some species of West Malesian Diplazium
that excluded from Diplazium and then have been included in the other genera,
namely D. chrysocarpum Alderw., D. grammitoides Presl, D. japonicum Bedd.,
D. subsinuatum (Wall. ex Hook ) et Grev.) Tagawa and D. amplissimum. D.
chrysocarpum Alderw was included as Athyrium stramineum Copel.). D.
grammitoides Presl, D. japonicum Bedd., D. subsinuatum (Wall. ex Hook ) et
Grev.) Tagawa were included in Deparia and treated as Deparia confluens
(Kunze) M. Kato, De japonica (Thunb.) M. Kato (Kato 1984), and De lancea
(Thunb.) Fraser-Jenk (1997), respectively. While was D. amplissimum included in
Cornopteris as C. atroviridis (v.A.v.R.) M. Kato (Kato 1979).
(3) Now, West Malesian Diplazium is reported comprising 69 species
with 15 varities. Thirteen new species and two new varieties are described. The
thirteen new species described in this chapter are: D. asymmetricum, D.
batuayauense, D. crameri, D. densisquamatum, D. halimunense, D. loerzingii, D.
megasegmentum, D. megasimplicifolium, D. meijerii, D. parallelivenium, D.
profluens, D. subalternisegmentum, and D. subvirescens. While the two new
varieties descrided are: D. accedens var. spinosum and D. silvaticum var. pinnae-
179
ellipticum. Two new status of Diplazium are proposed: D. pallidum var.
montanum and D. accedens var. ridleyi.
Key to Species of Diplazium in Western Malesia
1. Veins free
2. Lamina simple
3. Rhizome creeping; scales lanceolate, margin entire ………........... D. subsinuatum
3. Rhizome erect or suberect; scales oblong subtriangular, margin shortly toothed. ……………………................................. D. subserratum
2. Lamina simply or compoundly pinnate
3. Lamina simply pinnate to bipinnatifid
4. Terminal pinnae conform to lateral pinnae or deltoid with deeply lobed
5. Scales margin toothed
6. Rhizome short, erect; lateral pinnae commonly 3-18 pairs; terminal pinna larger or deltoid with deeply lobe at base
7. Lateral pinnae commonly 3-5 pairs, 4 cm or more wide; texture chartaceous or thinly papyrceous
8. Rachise non gemmiferae; base of lateral pinnae cuneate …………………………… D. halimunense
8. Rachise gemmiferae; base of lateral pinnae rounded ………………………….. D. bantamense
7. Lateral pinnae commonly 7-18 pairs, 3 cm or less wide; exture thinly coriaceous …………………….. D. lobbianum
6. Rhizome medium, creeping; lateral pinnae commonly to 3 pairs; terminal pinna conform to lateral ones …… D. donianum
5. Scales margin entire
6. Rheophytic or riparian; scales on stipe black; lateral pinnae to 4 pairs
7. Rheophytic; Stipe more than 20 cm long;lateral pinnae shortly stalked 4.5 mm long, lanceolate,
more than 12 cm long, 2 cm broad ……… D. aequibasale
7. Riparian; Stipe less than 20 cm long; lateral pinnae adnate, lineary oblong, usually less than 8 cm long, 1.5 cm broad …. D. wahauense
180
6. Terrestrial; scales on stipe pale to dark brown; lateral pinnae 5 or more pairs
7. Scales pale or light brown to brown;
8. Lateral pinnae ovate-lanceolate, larger ones to 3 wide
9. Scales brown; pinnae to 5 pairs; veins group formingangle 50-60º to costa …………… D. crameri
9. Scales light brown; pinnae to 20 more pairs; veinsgroup forming angle ca. 45º
to costa ….………………………. D. hottae
8. Lateral pinnae elliptical, larger ones 4 cm or more wide ......……. D. xiphophyllum
7. Scales dark brown
8. Lateral pinnae to 10 pairs; texture thinly or softly chartaceous
9. Veins forked to 3 times; pinnae crenate or shallowly lobed,shortly stalked ……. D. prescottianum
9. Veins forked to 4 times; pinnae subentire or toothed near apex, distinctly stalked .... D. subintegrum
8. Lateral pinnae to 27 pairs; texture subcoriaceous …………………………….…. D. pallidum
4. Terrninal pinnae absent, upper pinnae gradually reduced
5. Scales abundant on stipe, rachis, costae, and vein beneath; Lamina nearlyelliptic- lanceolate, to 13 cm broad
6. Lower lamina fully pinnate to 2/3 part; free pinnae to 32 or more pairs; veins forked to 3 times ………..…………………… D. fuliginosum
6. Lower lamina deeply pinnatifid to 5/6; sometimes with one pair of reduced free basal pinnae; veins mostly once forked
7. Lamina elliptic; lobes to 10 mm broad, oblong-lanceolate, narrowing towards apex, subentire ……… D. lomariaceum
7. Lamina lanceolate; lobes to 15 mm broad, liniery oblong, slightly crenate, or toothed and not seldom the sides entire only the apex serrate ………………… D. porphyrorachis
181
5. Scales usually abundant on lower stipe only; lamina oblong subtriangular – oblong lanceolate, to 30 cm broad
6. Scales entire
7. Lamina to about 15 cm wide, subtriangular, costae hairy beneath …………………… D. tomentosum
7. Lamina usually wider, oblong-lanceolate, costae not hairy beneath
8. Margin of pinnae shortly toothed or almost entire; veins forked once to twice, usually soriferous on basal acroscopic side …………………… D. pallidum
8. Margin of pinnae deeply lobed; veins pinnate in the lobes; usually soriferous on all veinlets
9. Rachise fibrillose; margin of pinnaelobed ¾ -11/12 way to costa …. D. sorzogonense
9. Rachise glabrous; margin of pinnae lobed 1/3-2/3 way to costa
10. Scales light brown without thickening blackish strandmargin; lower base of lower pinnae not cut away; soricover veinlets to 2/3 of their lengt …..… D. christii
10. Scales dark brown with thickening blackish strandsmargin; lower base of lower pinnae often much cutaway; sori cover veinlets to ¾ of their length or almostreaching margin
11. Rachise not gemmiferous; lobus rounded; texture papyraceous; sori elongate from near base of veins to near margin of lobus; indusia medium brown, concolour ……….……………… D. malaccense
11. Rachise gemmiferous; lobus truncate; texture subcoriaceous; sori medial or close to margin; indusia dark brown, attachment side darker ....... D. loerzingii
6. Scales toothed
7. Stipe, rachis and costa tomentose ….…………… D. tomentosum
182
7. Stipe, rachis and costa not tomentose
8. Lateral pinnae to 4 cm or more broad, margin lobed to 5/6 way to costa …..………………….. D. speciosum
8. Lateral pinnae less than 4 cm broad, margin lobed to ¾ way to costa
9. Pinnae lobed ¼ - ½ towards costa
10. Scales dark brown or nearly black, without thickening black strand; sori subcostular, touching midveins or nearly so at proximal end
11. Pinnae chartaceous, often auriculate below;scales irregularly andshortly toothed ……… D. crenatoserratum
11. Pinnae thin in texture, not auriculate below;scales strongly toothed …… D. silvaticum
10. Scales brown, with thickening dark brown strand; sori medial or supramedial (not touching midveins) …….... D. batuayauense
9. Pinnae lobed ¾ way towards costa
10. Pinnae lanceolate, texture herbaceous; Veins forming an angle about 60º to costa ……... D. petiolare
10. Pinnae lineary subtriangular, texture thicker; Veins forming an angle 50-55º to costa … D. acuminatum
3. Lamina bipinnate to tripinnate
4. Rhizome erect
5. Lamina bipinnate
6. Scales rounded
7. Pinnulae lobed more than ¾ way to costa
8. Costae bearing scattered small ovate-rounded scales;pinnulae subtriangular, lower stalked to 1 mm long,apex of lobus truncate; sori usually diplazioid onbasal acroscopic veinlets only ………… D. latisquamatum
8. Costae glabrous; pinnulae lanceolate, lower stalkedto 2.5 mm long, apex of lobus rounded-acute;sori diplazioid both on basal and medianacroscopic veinlets ……………………. D. profluens
183
7. Pinnulae lobed ¾ way or less to costa
8. Stipes prickly
9. Stipe muricate, dark brown when dry; lower pinnulaeshortly stalked to 1.5 mm long, margin lobed to1/2 way to costa ………………… D. betimusense
9. Stipe spiny, rather stramineous when dry; lower pinnulaeadnate, margin lobed less than ¼ way to costaor almost entire …………………. D. spiniferum
8. Stipes not prickly
9. Lower pinnulae stalked to 2 mm long; Sori on medialon veins ……………………………………. D. kunstlerii
9. Lower pinnulae stalked less than 1 mm long oralmost adnate; Sori on basal veins ................ D. laevipes
6. Scales ovate-liniery lanceolate
7. Scales toothed
8. Pinnulae lobed 1/3 way or less to costa or almost entire
9. Stipes scales throughout
10. Pinnulae lobed less than 1/4 way to costa (crenulate);veinlets to 9 pairs ….……….……… D. barbatum
10. Pinnulae lobed to 1/3 way to costa; veinlets to 6 pairs
11. Pinnulae to 20 pairs, less than 12 mm broad;veins forked 1-3 time …………. D. melanolepis
11. Pinnulae to 12 pairs, 15 mm or more broad; veinspinnate to 6 pairs of veinlets
12. Pinnulae elliptical, less than 5 cm long, 2 cmbroad; apex of vein whitish-limy(dot like) …………….. D. albido-squamatum
12. Pinnulae oblong, to 6 cm long, 1.5 cm broad,base broadly cuneate; apex of vein not whitish-limy ……………………………. D. crinitum
9. Stipes scales at base only
10. Stipe spiny; pinnulae to 12 pairs, lower stalked to3 mm long; veins to 8 pairs, once forkedon lower lobes …………………..……. D. vestitum
10. Stipe not spiny; pinnulae to 16 pairs, lower stalkedto 1.5 mm long; veins to 5 pairs,commonly 4 pairs, all simple …… D. simplicivenium
184
8. Pinnulae lobed ½ or more to costa
9. Lamina less than 100 cm long; scales light brown, withoutthickening black strand; texture herbaceous … D. umbrosum
9. Lamina more than 150 cm long; scales dark brown, teeth mostlyforked, with thickening black strand; texture papyraceous
10. Stipe prickly; lobes oblong to subquadrangular;sori less than 2.5 mm long …………. D. polypodioides
10. Stipe not prickly; lobes subdeltoid to semiorbicular;sori more than 3 mm long .…………… D. dilatatum
7. Scales entire
8. Stipes not not muricate at base
9. Stipe and rachise not clothed with scattered multicellular brownhairs;
Sori on medial veins, impressed ………….. D. poiense
9. Stipe and rachise clothed with scattered muslticellular hairs; Sori on basal veins, not impressed …………. D. velutinum
8. Stipes muricate at base
9. Stipes densely scales throughout; free pinnulaeto 18 pairs .……………. ………….. D. densisquamatum
9. Stipes densely scales only at base; free pinnulae to 13 pairs
10. Lower pinnulae shortly stalked to 1.5 mm long,lanceolate, more than 6 cm long, lobed to 4/5 way tocosta …………………………… D. atrosquamosum
10. Lower pinnulae adnate – sessile, nearly hastate, less than 5cm long, crenate ….…………...………… D. hewittii
5. Lamina tripinnatifid – tripinnate
6. Pinnulae lobed less than ¾ way to costa
7. Pinnulae to 16 pairs; larger pinnule stalked to 5 mm long,oblong subtriangular, to 18 cm long, 4 cm broad,apex attenuate, margin lobed ½ way to costa ……. D. dilatatum
7. Pinnulae to 20 pairs; larger pinnule adnate, oblong-lanceolate, to 5.5 cm long, 1 cm broad, commonlyless than 4 cm long, 8 mm broad, apex rounded-acute, margin lobed to 1/3 way to costa ……... D. melanolepis
185
6. Pinnulae lobed ¾ way or more to costa
7. Pinnulae lobed to within one mm or less of costule, forming segments
8. Lower costule of larger pinnulae winged; free segment notpresent
9. Scales lineary triangular-lanceolate; Indusia opening whensori mature ………………………. D. subpolypodioides
9. Scales rounded; Indusia opening beforesori mature ………………............ D. megasegmentum
8. Lower costule of larger pinnulae not winged; free segmentspresent
9. Pinnulae 17 pairs or more; larger pinnulae oblong- subtriangular; sori brown when dry
10. Segments crenate or lobed 1/3 way to costulet; sori bearing near basal or on middle veinlets cover ½ of their length; indusia pale brown, attachments side more dark, margin entire ………….. D. umbrosum
10. Segments crenulate, apex rounded; sori bearing on basal costulet cover veinlets 1/3 of their length; indusia brown, concolour, margin fringed ………………….…... D. moultonii
9. Pinnulae less than 16 pairs; larger pinnulae subtriangularlanceolate; sori nearly golden
yellow when dry …………………….. D. chrysocarpum
7. Pinnulae lobed to within more than 2 mm of costa, not formingsegments
8. Scales narrowly linier, margin toothed with thickening blackstrands; Stipes densely scaly near base, surface prickly;
Sori occupying from the base half-way or more to the edge ……............................... D. polypodioides
8. Scales rounded, margin entire; Stipes scales throughout, surfacenot prickly; Sori occupying only the lower half or lessof the veins ………………………… D. latisquamatum
4. Rhizome creeping
5. Margin of pinnae lobed to ¾ towards costa; Sori elongated from basal veinlets …………………………………………………. D. procumbens
5. Margin of pinnae lobed to 2/3 towards costa; Sori elongated on medial or submedial veinlets ……………………………. D. subvirescens
186
1. Veins occasionally uniting at margin or copiously anastomousing
2. Veins rarely anastomousing, occasionally uniting at margin
3. Lateral pinnae 6-9 pairs, elliptical; veins occasionally anatomousing; gemmae present at the junction between rachise and costa .…. D. xiphophyllum
3. Lateral pinnae 2-4 pairs, oblong; veins free or very rarely anastomousing; gemmae absent ………..……………………………………….. D. riparium
2. Veins copiously anastomosing
3. Lamina simply or simply pinnate
4. Lamina simply
5 . Lamina subdeltoid, base cordate, veins anastomousing 1/3-1/2 ………………………... D. cordifolium
5. Lamina elliptical, base subequally cuneate, veinsanastomousing 2/3-4/5 way of margin ….... D. megasimplicifolium
4. Lamina simply pinnate
5. Scales dense on stipe, rachise& costa; lower surface of laminabearing stelate trichome scatterly ………………………. D. squarrasum
5. Scales dense on base of stipes only; lower surface occasionally bearinguniseriat trichome with glandular cells
6. Terminal pinnae differ to lateral ones, deltoid ……. D. accedens
6. Terminal pinnae conform to lateral ones
7. Scales on stipes dark brown nearly black, lateral pinnae oblong, base cuneate; veins anastomousing
1/3 way of margin ……………………………... D. angustipinna
7. Scales on stipes light brown, lateral pinnae lanceolate, lower base rounded to auricled; veins anastomousing¼ way of margin ………………………………. D. cumingii
3. Lamina bipinnate
4. Scales dull brown; stipe spiny toward the base; Pinnulae fully adnate, texturefirmly herbaceous; vein anastomousing like D. accedens …..... D. insigne
4. Scales dark brown; stipe smooth; Pinnulae shortly stalked, texturepapyraceous; Vein in pinnate group in the lobes, 8-10 pairs of side veins, thelower 2-3 pairs of adjacent group anastomousing, forming irregularintermediate excurrent vein leading towards a sinus between
adjacent lobes …………………………………………... … D. esculentum
187
1. Diplazium accedens Blume
Diplazium accedens Blume., En. Pl. Jav.: 192. 1828; Holttum, Gard. Bull.
S.S. 11: 81. 1940; Tagawa & K. Iwatsuki, Fl. Thailand 3: 451. 1988. – Athyrium
accedens (Blume) Milde, Holttum, Rev. fl. Malaya 2: 558. f. 329. 1966.
Diplazium repandum Blume, En. Pl. Jav.: 191. 1828; Backer & Posth.,
Varenfl. Java: 125. 1939. TYPE: Blume s.n. (?, L!).
Diplazium swartzii Blume, En. Pl. Jav.: 191. 1828; Alderw. Ferns. Suppl.
1: 275. 1916.
Callipteris prolifera (Lam.) Bory; Copel., Fern Fl. Philip. 3: 419. 1960;
Andrews, Ferns of Queensland: 72. f. 5.1.C. 1990. TYPE: Callipteris prolifera
(Lam.) Bory.
Asplenium proliferum Lam. in Lam. & Poiret, Encycl. 2: 307. 1786
Athyrium ridleyi Copel., Phil. Journ. Sci. XI (1) : 39. 1908. TYPE:
Ridley 13970 (Isotype, SING!).
Key to the Variety
Rhizome bearing buds. Stipe protuberances, vascular bundles of its transversal sections nearblade continuously U shaped. Rachise not spiny, always gemmiferous.
Stipe sparsely protuberances; protuberance light green. Veinlets pale green when living;extra areola between adjacent normal group of veins few, the line of series extra areola up to1/3 way to costa ................................... ..…………………………………... var. accedens.
Stipe densely protuberances; protuberance deep green. Veinlets light green when living;extra areola between adjacent normal group of veins copious, the line of series extra areola upto 5/6 way to costa ......................................................................................... var. ridleyi.
Rhizome not bearing buds. Stipe sharply spinuous, vascular bundles of its tranversal sections nearblade interrupted U shaped. Rachise bluntly spiny, not gemmiferous ……… var. spinosum.
a. var. accedens
Rhizome stout, short, erect, bearing buds. Stipe stout, green, clothed with
green protuberances toward base, up to 80 cm long, 1-1.2 cm thick, scales at base;
scales linier, 10-13 mm long, 1.2-1.5 mm wide, dull brown, thick in texture with
narrow black strand toothed at margin. Lamina simply pinnate, oblong with
188
acuminate apex in outline, to 110 cm long or more, 40 cm wide, pinnae to 18
pairs; lower pinnae stalked to 7 mm long, sessile in upper ones, basal pinnae or
subbasal pinnae the largest, about 30 cm long, 7 cm wide, oblong with acuminate
apex, base broadly cuneate to subtruncate, margin entire or undulate, or double
crenate; terminal deltoid with 1 or 2 deeply lobed at base; papyraceous, glabrous
above; rachis grooved above, gemmiferous; costa prominent below, grooved
above with distinct ridge; veins pinnate, veinlets 8-10 pairs, veinlets adjacent
groups anastomousing producing a series of parallelogram-shape areolus. Sori
elongate along veinlets, often throughout their length forming areoles; indusia
very thin, fragile, margin entire.
SPORES. Monolete, bilaterally symmetrical, heteropolar; polar outline
elliptical, sides convex; equatorial longitudinal view concave-convex to plano-
convex; equatorial transverse view, proximal face planar to concave, distal face
hemispherical; perinate. E: 25.34(31.47)36.02±2.54; P: 15.32(19.51)23.59±2.82.
Laesure: concealed by perine ridge. Perine: costate-alate, reticulation often
incomplete; lacunae shallow, project 10-17 m ;wing-like muri or costae project
c.0.5-6 m, terminating margins entire. Exine: often visible through perine,
smooth under LM, smooth under SEM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped with an angle about 90º, the outward of the
lower base, angle, and end ridge. Stomata: polocytic and copolocytic.
CHROMOSOMES. 2n = 82 (Cytotype: TNgP1447, BOHB).
DISTRIBUTION. Peninsular Thailand, Malesia, Pacific.
ECOLOGY. Terrestrial, spread throughout Western Part Malesia Region
at 400 – 1550 m sea level in moist soil, by preferences more or less shadowed
localities, in primaryforest, secondary forest in ravine, forest borders, and
meadows.
VERNACULAR NAMES. Paku buwah, paku careham (Sundanese),
pakis angkrik (Javanese).
USE. The yound fronds and bulbis in the axils of leaflets are eaten after
cooking.
189
SPECIMENS EXAMINED. --- JAVA: Arsin s.n.; Backer s.n., 6307,
10891, 16200, 23754; Backer & Posthumus 201; Bakhuizen v/d Brink 2393;
Beguin 83; Brinkman 445; Burk s.n.; Coert 910; Danser 6783; Hallier 201;
Heurn 201, 202, 203; Kooders 373*, 8778, 10891, 21179B, 23356B, 23725B,
41459B, 41508B; Lörzing 933, 1745; Meijer 31, 172; Pleyte 162; Raciborski
109; Sciffner s.n..; Winckel 1279B, 1419B; Zippelius s.n.. ---- MALAY
PENINSULA: Md Nur 32894, s.n. (28-8-1923); M.R. Henderson s.n. (28-8-
1923); Br. G. Allan 3477, s.n. (30-6-1957); Wale 9111. --- SUMATRA: J.A.
Lörzing 5577, 12808, 5562;.Bünnenmejer 4316; D. Darnaedi 107; J.A. Lörzing
5562; C.G.G.J. van Steenis 3470; J.A. Lörzing 12809, 7001; Dr. Cramer no. 25;
J.A. Lörzing 4226; Karta 21; Jacobson 1451, 2498; K. Iwatsuki, Gen Murata, J.
Dransfield & D. Saerudin S- 1542; Bünnenmeyer 4314a; M.Hotta 26131;
W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 12583; H. Surbeck 81; Dr. O.
Posthumus 1105. ---- BALI: O. Posthumus 3722; C.G.G.J. van Steenis 8017; W.
Meijer 10546; W. Meijer 10453.
b. var. spinosum Praptosuwiryo, var. nov.
TYPE: Tahrodin TR53 (holotype, BOHB), East Kalimantan
Stipe sharply spiny, pale green on upper surface when living, black on
lower surface. Pinnae up to 7 pairs; lower pinnae stalked to 3 mm long, basal
pinnae reduce to 6 cm long, 2.3 cm broad, margin undulate; rachise not
gemmiferous.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form interrupted U-shaped with an angle about 90º, angle and end bluntly ridge.
CHROMOSOMES. 2n = 82 (Cytotype: TR 53, BOHB)
SPECIMENS EXAMINED: --- BORNEO: TR 53; YN 04. ---
SUMATRA: H.A.B. Bünnenmeijer 8665.
190
c. var. ridleyi (Copel.) Praptosuwiryo, stat. nov.
Athyrium ridleyi Copel., Phil. Journ. Sci. XI (1) : 39. 1908. TYPE:
Ridley 13970 (Isotype, SING!).
Stipe deep green when living, bearing densely deep green protuberances,
minutely pilosa. Lateral pinnae up to 7 pairs; larger ones up to 10.8 cm broad,
apex caudate, margin entire or crenate. Veinlets very distict, light green. Extra
areola between adjacent normal group of veins copious, the line of series extra
areola up to 5/6 way to costa.
DISTRIBUTION. Malay Peninsula, Suamtra.
SPECIMENS EXAMINED: --- MALAY PENINSULA: Ridley 13970. ---
SUMATRA: T.Ng. Praptosuwiryo 2525a.
2. Diplazium acuminatum Blume
Diplazium acuminatum Blume, En. Pl. Jav.: 193. 1828; Alderw., Mal.
Ferns: 409. 1908.
Rhizome short, erect-suberect. Stipe 34-36 cm long, 2 mm thick when
dry, brown, black and scales at base, glabrous upward; scales lineary triangular, 5-
10 mm long, 0.5-mm broad, dark brown, margin sharply toothed. Lamina
pinnate, subtriangular- lanceolate in outline, 45-50 cm long, 15-23 cm broad, free
pinnae 15-21 pairs below deeply lobed acuminate apex of lamina; lower pinnae
shortly stalked to 6 mm long, upper ones adnate–sessile, lineary subtriangular,
larger pinnae 12-12.5 cm long, 2-2.2 cm broad, base unequal, lower base cuneate,
upper truncate, apex acuminate, margin lobed ½-3/4 way to costa; lobus oblong,
basal acroscopic the largest, 3.5-5.5 mm wide, ends truncate, entire or slightly
toothed; texture firm; rachise gemmiferous on upper part; veins free, forming
angle about 50-55° to costa, pinnate in the lobes, veinlets 5-7 pairs, all simple,
forming angle about 15-20º to main veins. Sori elongate from near costule covers
1/3-2/3 way of their length or almost reaching the margin, basal acroscopic
diplazioid; indusia brown, rolled back, margin entire, firm.
DISTRIBUTION. Sumatra.
ECOLOGY. In the mountain forest at 400-1000 m dpl.
191
SPECIMENS EXAMINED: --- SUMATRA: J.A. Lörzing 14051, 6311; J.
Buumee A. 841.
NOTES. Kato (1994) states that this species is similar and perhaps related
to D. megaphyllum (Baker) Chist from E. Myamma, SW China, Taiwan, Vietnam
and Thailand, in general habit and leaf morphology, but from it in the regularly
anastomosing veins. D. megaphyllum generally has free veins.
3. Diplazium aequibasale (Baker) C.Chr.Diplazium aequibasale (Baker) C.Chr.; Ind. 227. 1905; Alderw., Mal.
Ferns: 404. 1908. J. Fac. Sci. Univ. Tokyo 3 (15): 101. 1991.Rhizome short, erect. Stipe 21-28 cm long, pale brown when dry, black
and scales at base; scales ovate-lanceolate, 3-5 mm long, 1-1.5 mm broad, fragile,
margin entire. Lamina simply pinnate, lateral pinnae 3-4 pairs, terminal pinnae
like the laterals, the largest; lower pinnae shortly stalked to 3-4.5 mm long, upper
adnate, lanceolate, to 14 cm long, 2.5 cm broad, base cuneate, margin entire or
crenate near apex, suddently narrowed near apex, acuminate; texture rather thin,
surface naked; costa rounded beneath, grooved on upper surface; veins group
forming angle about 65-70° to costa, each group of 3 veins, middle vein forked 1-
3 times, occasionally the outer of vein group uniting with the inner close to the
margin. Sori bearing at 1-3 veins in each vein group, those on the outer vein of
the group extending from the costa to margin, rest shorter, acroscopic outer veins
usually diplazioid; indusia brown, narrow, margin entire, persistent, fragile,
opening when mature.
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 2026, BO)
DISTRIBUTION. Java, Sumatra, Malay Peninsula, Borneo.
ECOLOGY. It is rheophytic species and usually grown at lowland clay
stream-bank. 20- 400 m.
SPECIMENS EXAMINED. --- JAVA: Scheffer 16584. --- BORNEO:
A. Kostermans 10448, T.Ng. Praptosuwiryo 2148. --- SUMATRA: O.
Posthumus 695; T.Ng. Praptosuwiryo 2026a, 2026b, 2157. --- MALAY
PENINSULA: R.E. Holttum SFN 24780.
192
4. Diplazium albido-squamatum Alderw.
Diplazium albido-squamatum Alderw., Bull. Buiten. II no. 23. 9. 1916. --
-TYPE: C.J. Brook 272/S. (holotype, BO!), Soelit, Lebong, Bengkulu, Sumatra;
C.J. Brook s.n. (isotype, L!), Lebong Tandai, Bengkulu, Sumatra.
Stipes 38 cm long, 6 mm thick, grooved above, dark brown-blackish,
scales throughout; scales lineary lanceolate or subulate, to 1 cm long, 1 mm wide,
dark brown, margin toothed with thickening black strand. Lamina bipinnate, ca.
70 cm long, (?) cm broad, lanceolate in outline, pinnated pinnae 5 pairs below ca.
5 pairs of pinnatifid pinnae, apical lamina (?); lower pinnae stalked, stalked to 17
mm long, 8-10 cm apart, ovate-lanceolate, 15-24.5 cm long, 8.5-9 cm broad,
pinnulae 5-8 pairs; lower pinnulae shortly stalked to 1 mm long or adnate, 2.5-3
apart, upper pinnulae sessile with subequally broadly cuneate; larger pinnulae
oblong-elliptical, 4.8 cm long, 1.9 cm broad, base subequally subtruncates, apex
acute, margin lobed ¼-1/3 way to costule; lobus 4-6.5 mm wide, ends rounded-
subtruncate; texture subherbaceous, upper surface glabrous, to apex vein
whitish-limy, dot-like; rachise obscurely dark brown, minutely scales, at length
base persistent scales fallen away rugged; veins free, pinnate, mid-veins forming
angle ca. 60º to costule; veinlets 4-6 pairs, length base persistent scales fallen
away rugged all simple, forming angle about 20-25° to costule. Sori elongate
along veinlets, cover ½-2/3 of its length, acroscopic basal diplazioid, others
asplenoid; ndusia dark brown, stiff, margin entire, opening when mature.
DISTRIBUTION. Sumatra.
ECOLOGY. Grown in limestone.
SPECIMENS EXAMINED. --- SUMATRA: C.J. Brooks 272/S.
NOTES. young plants simply pinnate, with the fronds similars to the
largest pinnae of the adult ones but less copiously soriferous.
193
5. Diplazium angustipinna (Holttum) Holttum
Diplazium angustipinna (Holttum) Holttum, Rev. Fl. Malaya 2. Appendix
II: 637. 1966. Athyrium angustipinna Holttum, Gard. Bull. S.S. 14: 8. 1953;
Rev. Fl. Malaya 2: 549. 1966. --- TYPE: R.E. Holttum SFN 39416 (holotype,
SING!), Frasser’s Hill, Pahang, Malay Peninsula.
Rhizome short, erect. Stipe to 36 cm long, 3 mm thick, scales at base;
scales narrow, to 8 mm long, 1 mm broad, medium margin entire, some larger
ones with black firm edges. Lamina simply pinnate, fertile ones to 41 cm long,
lateral pinnae 4-6 pairs below a terminal leaflet of similar shape; largest fertile
pinnae on different fronds 10-18 cm long, 1.6-2.8 cm broad, widest 1/3 from the
base, tapered evenly to the narrow apex and to the base which is truncate on a
stalk 1 mm. long, margin slightly and irregularly sinuous; frond and rachis
glabrous except for very short hairs in groove of upper surface of rachis and small
scales on bases of costae; rachise not proliferous; veins anastomousing freely in
outer 1/3 of each half of the lamina, forking 2 or 3 times before anastomousing.
Sori long, diplazioid on acroscopic branch of each first forking, reaching from
costa 2/3 towards the margin, also 1-3 short sori on distal anastomousing vein-
branches; indusia narrow, firm, perisistent, margin entire.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form uninterrupted V-shaped, with an angle 70º, ends simple, not forming a ridge.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 1904, BO),
164 (T.Ng. Praptosuwiryo 1905b, BO).
DISTRIBUTION --- Malay Peninsula, Borneo.
ECOLOGY --- Terrestrially on humus rich soil or rock soil in dense
jungle. 400-900 m.
SPECIMENS EXAMINED --- BORNEO: T.Ng. Praptosuwiryo 1904,
1905b. --- MALAY PENINSULA: R.E. Holttum SFN 3946; King’s 8026.
194
6. Diplazium asymmetricum Praptosuwiryo, sp. nov. Plate 1.TYPE. Java. West Java, Mt. Halimun, track Cikaniki-Citalahap, ca. 1000
m., 27 September 2003, T.Ng. Praptosuwiryo 1728 (holotype, BO).
Rhizoma breve erectum. Stipites ad 40 cm longi, basi squamis subulatis
vel oblongis ovatis ad 4.5 mm longis 1.5 mm latis margine dentatis vestitae.
Lamina deltoideus, 46-50 longae, 46-50 latae, bipinnata-tripinnatifida, pinnate
pinnae 3-4 jugatae infra pinnatifis pinnae, apice deltodeus peniti-lobus. Pinnae
infimus maximae, stipipatae ad 4 cm longae, ubtriangulare, libere pinnulae 6-10-
jugatae infra apice deltoideus pinnatifidus; pinnulae infimus basiscopicus
maximae, stipitae ad 1.5 mm longae, subtrianglare, asymmetricus, base
subcordatus vel truncates, apice acuminatae, cetera margine lobatae ad 4/5 costam
versus; lobi angulum plus minusve 90° cm costa formantes, ad 8 mm latis supra
basi, margine leviter denticulatus, apice subtruncatus vel obliquely subrotundatus.
Vennae in uno lobo ad 6-jugatae, plerumque furcatae. Sori basaliter, aliquando
medianus, in 1/3-1/2 longitudine venularum sedentes, in basi venulae
acroscopicus diplazioideus. Indusia badius, fragilis, maturis aperiens.
Rhizome short, erect. Stipe pale green when living, scales at black base,
glabrous upwards, 30-101 cm long, 5-8 mm diam. near base; scales dark brown,
subulate-oblong ovate, to 4.5 mm long, 1.5 mm wide, margin toothed. Lamina
bipinnate-tripinnate, deltoid in outline, 46-50 cm long, 46-50 cm broad, pinnate
pinnae 3-4 pairs below pinnatifid pinnae, apex deltoid with deeply lobed. Basal
pinnae the largest, stalked to 7 cm long, subtriangular in outline, pinnule 6-10
pairs below acuminate deeply lobed deltoid apex of pinnae, basiscopic pinnules
the larger than acroscopic ones; pinnulae shortly stalked to 4 mm long,
subtriangular, asimetrical, base subcordate-truncate, apex sharply toothed
attenuate or caudate, margin lobed 4/5-7/8 way to costule (on bipinnate lamina),
or forming segments (on tripinnate lamina); basiscopic segments larger than
acroscopic ones, to 4 cm long, 1.8 cm broad, base truncate or broadly cuneate,
apex acute, margin lobed to 1/3 way to costulet; lobus almost at right angle, to 4
mm broad (on tripinnate lamina) or to 8 mm broad above base (on bipinnate
leaves), margin slightly toothed, ends subtruncate-obliquelly subrounded. Veins
pinnate in the lobes, veinlets to 6 pairs, usually once forked. Sori from near
195
costule, sometimes medial, cover 1/3-1/2 of veinlets, diplaziod on basal
acroscopic veinlets. Indusia pale brown, fragile, margin entire, opening when
maturity, rolled back.
PARATYPE. JAVA. West Java: Mt.Gede, Cibodas Forest, behind
Cibodas Botanic Gardens, ca. 1450 m, 19 August 2002, T.Ng. Praptosuwiryo
1334 (BO); Mt. Salak, Southern Slope, Cangkuang Forest, 18 December 2002,
T.Ng. Praptosuwiryo1365 (BOHB ); Mt. Halimun, Track Cikaniki-Cikuda Paeh,
ca. 1250 m, 29 September 2003 T.Ng. Praptosuwiryo 1780 (BO).
CHROMOSOMES. 2n = 123.
DISTRIBUTION. Only found from Java, from Mt. Gede, Mt. Salak and
Mt. Halimun.
ECOLOGY. Growing on moist humus-rich soil, shadowed places of
mountain forest at altitude 1000-1500 above sea level.
ETYMOLOGY. This species is named in relation to the specific
characters of its pinnae and pinnulae. Pinnae and pinnulae are usually
asymmetric, the acroscopic pinnulae and segments or lobes are larger than the
basiscopic.
SPECIMENS EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 1728, 1334,
1365, 1780.
NOTES. Diplazium asymmetricum is similar to D. procumbens The two
species share in scales on stipes fallen when mature, ovate-lanceolate dark brown
scales with blunt teeth, deltoid lamina, oblong lobes with blunt or truncate apex,
and forked veinlets. Diplazium assymetricum differs from D. procumbens in the
following characters combination: rhizome short, erect; lamina more incised (to
tripinnate) basiscopic pinnulae or segments and lobes are larger than the
acroscopic ones; indusia thicker, margin entire.
196
7. Diplazium atrosquamosum (Copel.) C.Chr. & Holtt.
Diplazium atrosquamosum (Copel.) C.Chr. & Holttum, Gard. Bull. S.S. 7:
274. 1934. --- Athyrium atrosquamosum Copel., Phillip. J. Sci. 12c: 59. 1917.
--- TYPE: Clemens 11051 (holotype PNH?†; isotype MICH, s.n.; photo of isotype
at UC, K, s.n.).
Rhizome short, erect. Stipe light brown, 65 cm long, 7 mm thick.,
muricate and scales fallen. Lamina bipinnate, subdeltoid, 56 cm long, 40 cm
broad, pinnated pinnae to 4 pairs; pinnae stalked to 5.5 cm long, lanceolate, 32 cm
long, 14 cm broad, free pinnulae 12 pairs below pinnnatifid apex of pinnae, basal
acroscopic slightly reduced; lower pinnulae shortly stalked to 2 mm long, upper
andate or sessile, lanceolate, to 9.2 cm long, 2 cm broad, apex sharply acuminate,
margin lobed ¾- 4/5 way to the costule; lobes oblong, 2-5-3.5 mm wide above
base, ends truncate, slightly toothed; rachis gemmiferous at the adjacent to the
costae at the apex of lamina, glabrous; costae and costule sometimes scales,
mainly at the adjacent costae and costule or costule and costulets; veins black,
mainvein forming angle about 70º to costa, free, pinnate in the lobes; veinlets 5-6
pairs, simple, forming an angle about 35º to the costulet. Sori on basal veinlets,
elongate from near costulet cover 1/3-1/2 way to margin, indusia pale brown, thin,
broad, persistent, margin lacerate when opening.
DISTRIBUTION. Borneo.
ECOLOGY. Terrestrial in mountain forest. Elevation: 1500-2700 m.
SPECIMENS EXAMINED. --- BORNEO: R.E. Holttum SFN 25429; J.
M.S. Clemens 29716, 28103, 32558.
8. Diplazium bantamense Blume
Diplazium bantamense Blume, En. Pl. Jav.: 191. 1828; Copel., Polypod.
Philipp.: 72. 1905; Backer & Posth., Varenfl. Java: 128. 1939; Holttum, Gard.
Bull. S.S. 11: 82. 1940; Tagawa & K. Iwats., Fl. Thailand 3: 455. 1988. –
Athyrium bantamense (Blume) Milde, Bot. Zeit.: 353. 1870: Holttum, Rev. Fl.
Malaya 2: 558. f. 330. 1966.
Rhizome short, erect, suberect. Stipe up to 90 cm long, 4-6 mm thick near
base, glabrescent, clothed with scales at base; scales narrow, to 15 mm long, 2
197
mm wide at base, dark brown, margin toothed. Lamina simple pinnate, oblong in
outline, up to 50 cm long, 26 cm wide; lateral pinnae 2-5 pairs, ascending, shortly
stalked, upper ones sessile, oblong, rounded at base, widest at a little above the
base, gradually narrowing towards acuminate apex, margin subentire or serrate
near apex, up to 25 by 5.5 cm; terminal pinnae similar to lateral ones or
occasipnally with large lobe at base; texture chartaceous, glabrescent; rachis
grooved above, often gemmiferous at the junction with costa of upper pinnae;
costa raised below, grooved above; veins free, 2-3 times forked. Sori elongate
along veinlets, usually diplazioid. Indusia brown, fragile, margin entire.
SPORES. Monolete, bilaterally symmetrical, heteropolar; polar outline
elliptical, sides convex; equatorial longitudinal view concave-convex; equatorial
transverse view, proximal face concave, distal face hemispherical; perinate. E:
42.81(53.88)66.05±5.69; P: 22.64(31.48)37.47±3.38. Laesure: concealed by
perine ridge. Perine: costate-alate, loosely reticulate irregular envelope, costae
form a large reticulation; separated from the spores; reticulation often incomplete;
lacunae large irregular polygons 11-25 m across; costae or alate project 3-13
m, terminating margins entire; surface of perine smooth. Exine: often visible
through perine, smooth under LM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form uninterrupted U-shaped with an angle about 120º, base almost flat, end
ridges on both dirrection, outward and inward, bend outward to form an angle
150º. Stomata: polocytic and copolocytic.
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 1454,
BOHB), 328 (Cytotype: Tin 1-3, BOHB).
DISTRIBUTION. Malesia throughout.
ECOLOGY. Terrestrial in lowland and lower montane forest. Occurs in
shady places in hills and occasionally by streams in the lowlands, but not
common. 20-1600 m.
SPECIMENS EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 1366, 1369,
1373, 1383, 1212, 1454, 1510, 1511, 1516, 1517, 1752. ---BORNEO: M. Kato,
M.Okamoto, K.Ueda, D. Darnaedi & E.B. Walujo B-8355; M. Kato, M. Okamoto
& E.B. Walujo B-10609; M. Kato, G. Murata, Y.P. Mogea B-3773; A.A. Church,
198
A.C. 319. --- SUMATRA: Dr. Cramer 40; W.J.J.O. de Wilde and B.E.E. de
Wilde-Dyfjes 12793; O. Posthumus s.n. (27 July 1925); J.A. Lörzing 14730; J.A.
Lörzing 5342a; J. v. Borssum W. 2767; J.J. Afriastini 613; J.A. Lörzing 6894. ---
MALAY PENINSULA: G.H. Addison SFN 37159.
9. Diplazium barbatum Christensen
Diplazium barbatum C.Chr. in C.Chr. & Holttum, Gard. Bull. S.S. 7: 272.
pl. 59. 1934. --- TYPE: Borneo, Mt. Kinabalu, Holttum SFN 25386 (Holotype,
BM n.v.; isotype, US!; K n.v., SING n.v.).
Rhizome short, erect. Stipe ca. to 30 cm long, light brown throughout,
densely scales toward base; scales light brown, linear lanceolate, 3-1.4 mm long,
0,25-1 mm broad, concolours, without thickening black strands, margin toothed,
teeth distinctly forked. Lamina broad lanceolate, ca. to 65 cm long, 19 cm broad,
pinnae 10 pairs; lower shortly stalked to 3 mm long, pinnae pinnate, ovate
lanceolate, to about 12 cm long, 5.3 cm broad, free pinnulae 2-4 pairs below
pinnatifid apex of pinnae; pinnulae sessile, almost at right angle to costa, oblong,
margin entire, ends acute-rounded, slightly toothed; texture papyraceous, rachis
and costa densely minute scales; veins free, pinnate in the pinnulae; veinlets to 9
pairs, once forked. Sori elongate from near costule continuing to acroscopic
branch of veinlets covers 2/3-3/4 of their length, indusia pale brown, margin
entire, persistent.
DISTRIBUTION. Borneo.
ECOLOGY. Terrestrial in mountain forest. Elevation 1400-2100 m.
SPECIMENS EXAMINED. --- BORNEO: J. & M.S. Clemens 32607;
R.E. Holttum 25386; M. Kato, G. Murata & Y.P.Mogea.
NOTES. As notified by Christensen & Holttum (1934), D. barbatum is
somewhat resembling D. speciosum in general habit, colour and texture, but very
distinct by its densely squamose stipe, rachis and costae..
199
10. Diplazium batuayauense Praptosuwiryo, sp. nov.
TYPE: Borneo, Central Kalimantan, Mts. Muller, Batau Ayau, Above S.
Talikot Puhung Kucan, ca. 450 m, 14 June 2004, T.Ng. Praptosuwiryo 1927
(holotype, BO).
Rhizoma breve erectum. Stipites gracilis, 19-5-23 cm longae, 3 mm
crassa, squamis caducus brunneus, linear-lanceolatus, 4-10 mm longae, 0.5 mm
latis, margine irregularis dentibus furcatis cum filum niger spissescens. Lamina
pinnata, lanceolatis, 41 cm longis, 20 cm latis, pinnae 15-jugatae infra apice
pinnatifidus; pinnae inferiora stipitae ad 3.5 mm longis, superiora adnatus,
lanceolatae, 11.2 cm longis, 2 cm latis, basi inaequaliter truncatae ad cuneatae, e
margine 1/3-1/2 costam lobatae; lobi 4-5.5 mm latae supra basim, truncatae,
leviter crenati. Rachides non gemmiferae, glabrae. Venulae distinctus in
superficiebus ambabus, liberis, pinnatus in uno lobo, venae principalis angulum
55º cum costa formantes, venulae 4-5-jugatae, simpliciter. Sori medial or close to
margin of lobes, cover 1/3-1/2 of veinlets length (3 mm length). Indusiis
brunneis, concolour, persistens, ineteger.
Rhizome short, erect. Stipe slender, 19.5 -23 cm long, 3 mm thick, fallen
scales; scales brown, lineary lanceolate, 4-10 mm long, 0.5 mm broad, margin
toothed, teeth apart, with thickening dark brown strand irregularly, glandular cells
present,. Lamina pinnate, lanceolate in outline, 41 cm long, 20 cm broad, pinnae
15 pairs below pinnatifid apex of lamina; lower pinnae stalked 3.5 mm long,
upper pinnae adnate, one pair basal pinnae bending downward with cut away
base; pinnae lanceolate, widest at 1/3 part from basal, 11..2 cm long, 2 cm broad,
base unequally truncate on lower pinnae, cuneate at upper ones, apex acuminate,
margin lobed to within 4 mm of costa (or 1/3-1/2 way to costa); lobus widest at
base, 4-5.5 mm wide above base, ends truncate, slightly crenate. Rachise not
gemmiferous, glabrous. Texture thin. Veins distinct on both surface, free, pinnate
in the lobus, main vein forming angle 55° to costa, veinlets 4-5 pairs, all simple,
reaching margin. Sori medial or close to margin of lobes, cover 1/3-1/2 of
veinlets length (3 mm length). Indusia brown, persistent, margin entire.
200
PARATYPE. BORNEO: Central Kalimantan, Mts. Muller, Above S.
Talikot Puhung Kucan, track to Batu Ayau, 450 m, 14 June 2004, T.Ng.
Praptosuwiryo 1927a, 1927b, 1927c, 1927d, 1927e.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form interrupted U-shaped, with an angle 115º, angle slightly ridges, end almost
simple.
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 1927c,
BOHB), 205 (Cytotype: T.Ng. Praptosuwiryo 1909, BOHB).
DISTRIBUTION. Borneo
ECOLOGY. Terrestrial in lower montane forest. It is Grows on humus
rich soil in shady places.
NOTES. This species closely related to D. sorzogonense. Diplazium
batuayauense differs from D. sorzogonense in the following characters: Scales
sharply toothed with thickening dark brown stand irregularly, fronds are much
smaller, never reaching 1 m, lacks the fibrillose scales on stipe and rachis, pinnae
lobed only to ½ way to costa, sori not impressed. D. batuayauense is also similar
to D. malaccense.
ETYMOLOGY. The specific epithet is from the locality where this
species found, Batu Ayau, the mountain forest of Mts. Muller in Central
Kalimantan, Borneo.
11. Diplazium beamanii M.G. Price
Diplazium beamanii M.G. Price, Contr. Univ. Mich. Herb. 16: 193, f. 1-3.
1987. Type: Borneo: Pinosuk Plateau, 1400 m, Beaman 10724 (Holotype,
MICH!; Isotype , K!., MSC n.v.).
Rhizome erect. Stipe stramineus, 66 cm long, 7 mm thick, densely scales
at base; scales pale brown, ovate, 10 mm long, 4.5 mm broad, margin entire.
Lamina ovate, 60 cm long, bipinnate to deeply tripinnatifid at base, pinnae (?)
pairs; pinnae stalked to 2.5 cm long, subbasal the largest, to 37 cm long, ca. 17 cm
broad, pinnulae to 13 pairs, basal acroscopic slightly reduced; pinnulae stalked to
ca. 1 mm long, 2 cm distance, subtriangular-lanceolate, to 8 cm long, 2 cm broad,
base truncate, apex acuminate, margin lobed to 5/6 way to costa (within 2 mm of
201
costa); segments to ca. 5 mm wide, ends subtruncate, margin slightly denticulate-
crenulate; veins free, pinnate, veinlets to 5 pairs, usually simple. Sori from near
costule cover 1/3 way of veinlets (1.0-2.5 mm long), basal acroscopic often
diplazioid; indusia broad, margin irregularly fringed, persistent.
DISTRIBUTION. Borneo.
ECOLOGY. Terrestrial. Lower montane forest by stream. Elevation:
1400 m. Endemic to Mount Kinabalu.
SPECIMEN EXAMINED. --- BORNEO: Beaman 10724.
12. Diplazium betimusense Alderw.
Diplazium betimusense Alderw., Bull. Buit. III. 2. 142. 1920. ---TYPE:
J.A. Lörzing 5718 (Holotype, BO!; Isotype, L!), Sumatra, Sibolangit, Betimus
River.
Rhizome short, erect, 17 mm thick. Stipe dark brown, 48 cm long,
sparingly muricate, scales towards the base, scales on stipes dark brown, rounded-
ovate, margin entire, deciduous, leaving the stipes roughish by their persistent
bases, 3.5 mm long, 3 mm broad. Lamina bipinnate, (?) cm long, (?) cm broad,
pinnae (?) pairs; lower pinnae stalked to 5.3 cm long, oblong subtringular, 40 cm
long, 18 cm broad, pinnulae 6-9 pairs below pinnatifid deltoid apex of pinnae;
lower pinnulae shortly stalked to 1.5 mm long, upper ones adnate-sessile,
lanceolate, 8-10.5 cm long, 2-3 cm broad, base subequally truncate-cuneate,apex
acuminate-caudate, sharply toothed, margin lobed to within 7 mm of costa (or 1/3-
1/2 to cosule; lobus more or less widest at base, 5-6.5 mm broad, ends truncate,
slightly toothed; rachise dark brow, glabrous; texture firm; veins free, pinnate in
the lobus, main veins forming angle 50-55° to costule, veinlets 4-5 pairs, forming
angle about 10-15 º to main veins, distinct on both surface, simple, all reaching
the margin. Sori subbasal or medial, covers to 1/3 of their length, sometimes
diplazioid on basal acroscopic veinlets; indusia narrow.
DISTRIBUTION. Sumatra.
ECOLOGY. Growing on shade part of forest, near river at ca. 400 m sea
level.
202
SPECIMENS EXAMINED. --- SUMATRA: J.A. Lörzing 5718
13. Diplazium christii C.Chr.
Diplazium christii C.Chr., Index Fil.: 229. 1905; C.Chr. & Holttum,
Gard. Bull. S.S. 7: 270. 1934.
Rhizome (?). Stipe 35 cm long, 4 mm diam., scales at base , dark brown;
scales on stipes light brown, concolours, shining, 3-7 mm long, 0.5-1.5 mm diam.
near base, margin entire, without thickening black strands. Lamina pinnate,
lanceolate, 65 cm long, 18 cm broad, pinnae 19 pairs below deeply deltoid apex of
lamina (?); pinnae shortly stalked to 3 mm long, lanceolate 15 cm long, 1.8 cm
broad, base cuneate, apex acuminate, margin lobed ½ way to costa; lobus oblique,
widest at base, to 6 mm above base, ends subtruncate, slightly toothed; veins free,
pinnate in the lobes; veinlets usually 5-6 pairs, simple. Sori elongate at middle
veinlets covers 2/3 of their length; indusia light brown, broad, margin entire,
opening when mature.
DISTRIBUTION. Malay peninsula.
ECOLOGY. In dense forest, near summit 600 m.
SPECIMENS EXAMINED. --- MALAY PENINSULA: R.E. Holttum
SFN 19912.
14. Diplazium cordifolium Blume
Diplazium cordifolium Blume, En. Pl. Jav.: 190. 1828; Holttum, Gard.
Bull. S.S. 11: 83. 1940. – Anisogonium cordifolium (Blume) Bedd., Ferns Br.
Ind.: t. 331. 1870; Hand. Ferns Br. Ind.: 191. f. 92. 1883. -- Athyrium cordifolium
(Bl.) Copel., Holttum, Rev. Fl. Malaya 2: 548. f. 322. 1966. TYPE: C.J.A. van
Hasselt s.n. (?, L!), Kapala Tjibarran, Java.
Diplazium integrifolium Blume, En. Pl. Jav.: 190. 1828. TYPE: Blume
s.n. (?, L!), Bogor, Java.
Callipteris cordifolia (Blume) J. Sm., Copel., Polypod. Philipp.: 70.
1905.
Athyrium pariens Copel., Phil. Journ. Sci. IIIc : 299. 1908.
203
Key to the varieties
1. Lamina simple, oblong subdeltoid, base cordate, apex acuminate; veins forked to 5 times ………………………………………………... var. cordifolium
1. Lamina simply pinnate
2. Terminal pinnae usually the largest, oblong subdeltoid; lateral pinnae oblong, base cordate, apex acuminate; texture coriaceous …………………... var. integrifolium
2. Terminal pinnae usually similar shape to lateral ones, lanceolate; lateral pinnae lanceolate, base rounded to moderately auricled at basiscopic and truncate to cunetae at acroscopic, apex attenuate; texture subcoriaceous ……. var. pariens
Rhizome erect, suberect, scales on younger part. Stipe pale brown, 3-4
mm diam., 30-55 cm long, grooved above, brown, nearly black and scales toward
base; scales oblong ovate with acuminate apex, 6-9 mm long, 1-2 mm wide near
base, shining dark brown, margin entire with black thickening above base to tip
when old, occasionally with glandular cells. Lamina of two kinds, simple and
imparipinnate; simple ones oblong subdeltoid, 23-30 cm long, 8-13 cm wide
above base, widest 1/3 from base, base cordate, apex acuminate, margin entire;
imparipinnate ones nearly oblong deltoid in outline, 28-45 cm long, 24 cm wide;
lateral pinnae 1-6 pairs, upper smaller, usually terminal ones largest, oblong
subdeltoid, to 13-18 by 4-8 cm; upper pinnae sessile, broadly cuneate at base,
lower pinnae shortly stalked, the largest 16 by 5.5 cm, cordate at base, apex
acuminate, margin entire, rachis distincty beneath, gemmiferous at the junction
with costa; texture coriaceous; veins at about 45-70°to costa on pinnate ones, 50-
80º to midrib on simple ones, forked close to midrib, lower branch forked again 2-
5 times, anastomousing irregularly about 1/3-1/2 from margin. Sori elongate
along veinlets on both side or on acroscopic ones; indusia thin, persistent, margin
entire.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) plano-convex; equatorial
transverse view, proximal face planar, distal face convex; perinate. E:
37.02(42.15)51.62 ±5.09 P: 20.23(26.56)31.21±2.97. Laesura: concealed by
perine wing. Perine: alat to costate-alate, loose reticulate; irregular envelope
separated from exine surrounds the spore in continuous anastomosing wings,
204
forming a loose reticulation; lacunae large irregular polygons 15-20 µm across;
thin wing-like muri project 2-8 µm, terminating margins are often echinate;
surface of perine sparsely echinate or ciliate; echinae project 0.5-0.8 µm. Exine:
visible through perine, smooth under SEM.
SPECIMENS EXAMINED. MALAY PENINSULA. --- SINGAPORE:
H.N. Ridley 5867. --- BORNEO: Turaya 1894.
DISTRIBUTION. Malesia throughout, eastward to Solomon Islands.
a. var cordifolium
Diplazium cordifolium Bl., En. Pl. Jav.: 190. 1828. ---TYPE: J.C.A. van
Hasselt s.n. (L!, No.Reg. L 0051534), Kapala Cibaran, Java.
Lamina simple, about 27 cm long, 13 cm broad; veins branching to 8
times, anastomousing 1/3 from margin to the costa; veins group forming an angle
about 70ºto costa (at the middle), soriferous on outer veins, inner also, diplazioid
on basal acroscopic.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted V-shaped, with an angle 65º, end simple, not forming a
ridge.
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 1203, BO),
205 (Cytotype: T.Ng. Praptosuwiryo 1204, BO), 246 (T.Ng. Praptosuwiryo 1201,
BO), 328 (Cytotype: T.Ng. Praptosuwiryo 1926b, BO).
DISTRIBUTION. Sumatra, Malay Peninsula, Java, Borneo.
ECOLOGY. Occurs on moist sandy soil or humus rich soil of mountain
slopes in dense forest at low or medium altitudes. 400 – 2000 m.
SPECIMENS EXAMINED. ---- JAVA. West Java: A. Hidayat & H.
Wiriadinata AH 501; T.Ng.Praptosuwiryo 1201, 1202, 1203, 1204, 1206, 1207,
1208, 1456, 1460, 1461, 1737, 1768, 1807, 1813, 1710, 1768, 1807, 1808. ---
BORNEO: T.Ng. Praptosuwiryo 1910, 1926a, 1926b, 2128a, 2128c, 2128e,
2194a, 2194c, 2128b, 2194d, 2194e; Amdjah 271; J. & M.S. Clemens 26898;
Amdjah 273; J. & M.S. Clemens 33810; Amdjah 718; Kunio Iwatsuki, M. Kato,
Gen Murata & Y.P Mogea B-779; K. Iwatsuki, M. Kato, G. Murata & Y.P.
Mogea B-1935; A.Kostermans 8130; Amdjah 718; Kostermans 9051; Teysman
s.n.; M.Kato, M. Okamoto & E.B. Walujo B-10079; M. Kato & H. Wiriadinata B-
205
6362; M. Kato & H. Wiriadinata B-4727; M. Kato & H. Wiriadinata B-7098; M.
Kato & H. Wiriadinata B-6900; Kunio Iwatsuki, M. Kato, Gen Murata & Y.P.
Mogea B-2961; K. Iwatsuki, M. Kato, Gen Murata & Y.P. Mogea B-2271. ---
MALAY PENINSULA: King 6274; M.A. Donk no B.T. 10; R.E. Holttum 9898;
R.E. Holttum 31219; R.E. Holttum 23340; R.E. Holttum 10799; Md Nur SFN
19773. --- SUMATRA: S. Prawiroatmodjo 2576; W.J.J.O. de Wilde & B.E.E. de
Wilde-DUyfjes 19946; v. Steenis 3792; D. Darnaedi 81; L. Ajoeb 90; L. Ajoeb
96; J. v. Borssum W. 2788.
b. var. integrifolum
Diplazium integrifolium Blume, En. Pl. Jav.: 190. 1828. ---TYPE: Blumes.n. (L!, No.Reg. L 0051533 ), Java. Diplazium cordifolium Blume var.integrifolium (Blume) Mitsuta, Acta Phytotax Geobot. 36 (1-3): 78. 1985.
Lamina imparipinnate. Terminal pinnae usually the largest, oblong
subdeltoid; lateral pinnae 1-6 pairs, the largest 16 by 5.5 cm, oblong, base cordate,
apex acuminate; texture coriaceous; veins 1-3 forked, anastomousing in the
marginal 1/3-1/2.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) plano-convex; equatorial
transverse view, proximal face planar, distal face convex; perinate. E:
37.02(42.15)51.62 ±5.09 P: 20.23(26.56)31.21±2.97. Laesura: concealed by
perine wing. Perine: alat to costate-alate, loose reticulate; irregular envelope
separated from exine surrounds the spore in continuous anastomosing wings,
forming a loose reticulation; lacunae large irregular polygons 15-20 µm across;
thin wing-like muri project 2-8 µm, terminating margins are often echinate;
surface of perine sparsely echinate or ciliate; echinae project 0.5-0.8 µm. Exine:
visible through perine, smooth under SEM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped with an angle about 110º, base flat on inward and
outward direction, end ridges equally on both both directions.
CHROMOSOMES. 2n = 164 (T.Ng. Praptosuwiryo 2128, BO).
DISTRIBUTION. Sumatra, Java, Malay Peninsula, Borneo.
206
SPECIMENS EXAMINED. --- JAVA: TNgPraptosuwiryo 1192, 1205,
1206, 1207, 1208, 1305, 1367, 1368, 1375, 1374, 1457, 1706, 1708, 1709, 1729,
1735, 1736, 1742, 1774, 1775, 2280, 2281, 2284, 2288, 2291, 2293, 2297, 2298,
2300, . --- SUMATRA: de Vogel & Vermeulen 7504.; Kerling s.n., T.Ng.
Praptosuwiryo 2524a, 2524b, 2539. --- BORNEO: M. Kato, M. Okamoto, K. Ueda
& E.B. Walujo B-7397; M. Kato, M. Okamoto, K. Ueda & E.B. Walujo B-7386;
Masahiro Kato & H. Wiriadinata B-4984; M. Kato & H.Wiriadinata B-5243;
Halllier 1727; M. Kato, M. Okamoto & E.B. Walujo B-10236; M. Kato, M.
Okamoto & E.B. Walujo B- 10580; M. Kato, M. Okamoto & E.B. Walujo B-
9622; M. Kato, M. Okamoto, E.B. Walujo B-10847.
c. var. pariens (Copel.) C. Chr., Gard. Bull. S.S. 7 (3): 274. 1934; Ind.
Fil. Suppl. III: 75. 1934; Mitsuta, Acta Phytotax. Geobot. 36 (1-3): 78. 1985.
Lamina pinnate, pinnae 4-8 pairs; lower lateral pinnae shortly stalked to 3
mm, upper ones sessile, lanceolate, gradually reduce in size upwards, larger ones
12 cm long, 2.8 cm wide, attenuate at apex, rounded to moderately auricled at
basiscopic and truncate to cunetae at acroscopic bases, margin entire, terminal
pinnae similar to the lateral ones; texture subcoriaceous, rachis gemmiferous at
the adjacent to costa; veins 1-3 forked, anastomousing in the marginal 1/3-1/2.
SPECIMENS EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 1192, 1205,
1305, 1367, 1368, 1369, 1374, 1375, 1457, 1708.
NOTE: Mitsuta (1985) also recognized two varieties of D. cordifolium
of Sumatra, var. integrifolium (Blume) Mitsuta and D. pariens. The two varieties
are differentiated with characters as follow: Var. integrifolium has 2-3 pairs of
lateral pinnae and base of terminal pinna sessile, while var. pariens with 4-6 pairs
of lateral pinnae and base of terminal pinna usually wide cuneate.
15. Diplazium crameri Praptosuwiryo, sp. nov.
TYPE: Sumatra, Sukaraja, Kenali, 27 August 1915, DR. Cramer 41
(Holotype, BO).
Rhizoma erect (?). Stipites dilutus brunneus nitidus, 38 cm longis, 4 mm
crassis, basi piceus squamis caducus lanceolatus, c.7 mm longis, 2 mm latis,
integris, brunneis. Lamina simpliciter imparipinnatae, oblongae, 25.5 cm longae,
207
21 cm latae; pinnae 5-jugatae, stipitae ad 1.5 cm longis, 3.5-4 cm seorsum, ovate-
lanceolate, gradatin decrescente in statura, pinnae infernus ad 12.5 cm longis, 3
cm latis, basi cuneatae, marginae subintegrae, versus apice leviter serratus, apice
acuminatae, glabrae, in sicco superne atrobrunneae, inferne brunneae, pinnae
terminalis ad lateralis conformes; rhachides non gemmiferae, glabrae, costae
inferne prominens; vennae distingubilis, libere, angulum 50-60° cum costa
formantes, propinquus costae furcated, ramus superis simpliciter et soriferus,
ramus infernis simpliciter vel furcatis, plerumque furcatis. Sori e propinquus
costae elongati ad 7/9 venulae occupants, soris infimus acroscopicus
diplazioideus. Indusiis latis, brunneis, margine leviter crispatus, persistens.
Rhizome erect (?). Stipe pale brown, glossy, 38 cm long, 4 mm thick,
black at base, fallen scales; scales lanceolate, ca. 7 mm long, 2 mm broad, margin
entire, dark brown. Lamina simply pinnate, oblong, 25.5 cm long, 21 cm broad,
pinnae 5 pairs, terminal pinnae conform the lateral ones; lateral pinnae stalked to
1.5 cm long, 3.5-4 cm apart, below apical pinnae adnate, ovate-lanceolate,
gradually decrising in size, lower pinnae to 12.5 cm long, 3 cm broad, margin
entire, slightly serrate towards apex, base equally cuneate; rachise not
gemmiferous, costa raised below, glabrous; texture very firm. Veins free, distinct
on both surface, forming angle 50-60 to costa, forked near costa, upper branch
simple and soriferous, lower branch simple-once forked, commonly once forked.
Sori elongate from near basal veinlets (0.5-3.5 mm distance from costa) cover to
7/9 of lengt veinlets (irregularly), acroscopic basal diplazioid, other asplenoid.
Indusia broad, brown, margin slightly crisped, persistent.
DISTRIBUTION. Sumatra.
ECOLOGY. This is a dry land terrestrial fern that grows in shady places
of primary forest .
NOTES. Diplazium crameri may closely related to D. xiphophyllum.
The two species share stramineous stipe, lanceolate entire concolour scales,
simply pinnate oblong lamina, base of pinnae cuneate and texture subcoriaceous.
Diplazium crameri differs from D. xiphophyllum in its scales dark brown, pinnae
stalked to 1.5 cm long, ovate-lanceolate, all veins free and forked to 2 times.
Meanwhile D. xiphophyllum has pinnae elliptical, lower pinnae shortly stalked,
208
veins often anastomousing near margin and forked to 5 times. There is one
specimen only found among assemblage of specimens at BO that collected from
Sumatra. (22 Sept. 2007).
ETYMOLOGY. This species is named after DR. Cramer, the first
collector of this species.
SPECIMENS EXAMINED. --- SUMATRA: DR. Cramer 41.
16. Diplazium crenatoserratum (Blume) Moore
Diplazium crenatoserratum (Blume) Moore, Ind. Fil.: 121. 1859; Bedd.,
Hend. Ferns Br. Ind.: 177. 1883; Backer & Posth., Varenfl. Java: 130. 1939;
Holttum, Gard. Bull. S.S. 11: 99. 1940. –Asplenium crenatoserratum Blume,
Enum. Pl. Jav.: 177. 1828. –Athyrium crenatoserratum (Blume) Milde, Bot. Zeit.:
353. 1870; Holttum, Rev. Fl. Malaya 2: 561. f. 322. 1966.
Rhizome short, erect. Stipe 30-50 cm long, glabrescent, pale brown, black
and scales at base, distinctly grooved above; scales narrowly oblong subtriangular,
about 5 by 1 mm, concolour, dark brown, margin toothed irregularly, teeth not
forked. Lamina pinnate without distinct terminal pinnae of the fornd, oblong
triangular in outline with attenuate apex, widest at base, 33-70 cm long, 14-20 cm
or more wide; rachis grooved above; pinnae 9-22 pairs, stalked 3.5-5 mm long,
liniery oblong, 7-11 cm long, 1.4-2.7 cm wide, upper ones sessile with cuneate
lower base, lower ones stalked, upper base strongly auricled, less lower base or
slightly rounded; margin lobed ¼- ½ to costa or almost entire, lobes irregularly in
size; texture softly chartaceous; veins pinnate in lobes to 3 pairs or once-twice
forked in subentire pinnae. Sori alongate from nearly costa almost reach margin
of pinna, single or occasionally diplazioid on acroscopic veinlets of forked veins,
usually diplazioid on acroscopic of basal pinnate veins; indusia thin brown,
fragile, persistent, margin entire.
SPORES. Monolete, bilaterally summetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) concave-convex;
equatorial tranverse view, proximal face concave, distal face convex; perinate. E:
29.86(38.19)45.74±4.95, P: 17.92(23.48)27.36±2.94. Laesura: concealed by thin
209
wing-like muri. Perine: alate, loosely reticulate; irregular envelope, separated
from exine, surrounds the spore in anastomosing wings, forming loose
reticulation, reticulation irregular and often incomplete; lacunae irregular
polygons, 8-16 µm across, irregular holes and small papillae within; holes caused
by the fallen papillae; muri thin, wing-like, projected c. 1-8 µm, terminating
margin ciliate; surface of perine fibrous-like and holed, holes irregular and formed
by fallen ciliae.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an interrupted V-shaped formed from two long oval leaf traces, end simple
without ridges at both inward and outward directions.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 2075, BO),
164 (Cytotype: T.Ng. Praptosuwiryo 2067, BO).
DISTRIBUTION. Peninsular, Malaysia, Sumatra, Java, Borneo,
Moluccas, Thailand.
ECOLOGY. Terrestrial in primary and secondary forest, dry land, shaded
in slope and rigde of lowlands and montane forest. It can grow on various soil
from yellow-red loamy, red clay, and on humus rich soil. 20 – 1000 m.
SPECIMENS EXAMINED. --- JAVA: Backer 10644, 9927, 9999,
25555; Bakhuizen v/d Brink 3330, 3416, 6403, 7282. --- MALAY PENINSULA:
H.N. Ridley 3068, S.Z. Lin S2 2002-65; Sidek & Rozali Jaman RJ 1783; V.L.
Gurung 2. --- BORNEO: A. Kostermans 9747, 4145, 4139; K. Iwatsuki, M.
Kato, G. Murata & YP. Mogea, B-2273; B-2274; K. Iwatsuki, M.Kato, G. Murata
& Y.P.Mogea B-2958; K Iwatsuki, M. Kato, G. Murata, Y.P. Mogea B-717; B-
736; F.H. Endert 2796; Amdjah 723. BANGKA: Anta 728; Huguenin s.n., A.C.
Church, U.W. Mahyar, A.Ruskandi, Nurdin -192; A.C. Church, U.W. Mahyar,
A.Ruskandi, Nurdin -442; Sibatak Luang S 24733. --- MENTAWAI ISLANDS:
Iboet 557; Amdjah 186. --- SUMATRA: W.J.J.O de Wilde & B.E.E. de Wilde-
Duyfjes 19733; W. Meijer 15818; O. Posthumus 664; T.Ng. Praptosuwiryo
2011a, 2011b, 2011c, 2142a, 2142c, 2044, 2067a, 2067b, 2073a, 2073b, 2074a,
2074b, 2074c, 2074d, 2075, 2075c, 2075d, 2084a, 2085a, 2085b; 2208, 2210.
210
17. Diplazium crinitum (Baker) C.Chr.
Diplazium crinitum (Baker) C.Chr.; Ind. Fil. 230. 1905. ---Athyrium
crinitum (Bak.) Copel. Brittonia 1: 73. 1931. ---Athyrium vestitum of Copel.
Sarawak Mus. Journ. 2: 377. 1917.
Rhizome (?). Stipe 25-45 cm long, 2.5-7 mm diam. near base, light
brown, densely scales throughout; scales on stipes yellowish, shining, lanceolate,
4-7 mm long, 0.5-1 mm broad, margin toothed with thickening black strands
irregularly, fragile; teeth forked irregularly. Lamina bippinate, ovate in outline,
(?) cm long, 24 cm broad, pinnae (?) pairs. Pinnae shortly stalked to 6 mm long,
oblong in outline, 25 cm long, 10 cm broad, pinnulae to 11 pairs; lower pinnulae
adnate, upper sessile below deeply deltoid apex of pinnae; pinnulae oblong, larger
ones to 6 cm long, 1.5 cm broad, base broadly cuneate, apex acuminate, margin
lobed 1/3 way to costule; lobes oblique, widest at base, ends truncate or slightly
rounded, subentire; rachise dark brown, grooved on upper surface, densely minute
toothed scales; costa and costule also densely scales as like as rachise; texture
herbaceous-memranaceos; veins free, pinnate in lobes, veinlets 4-6 pairs, simple,
soriferous 1-4 pairs from basal. Sori elongate from near cosulet covers 1/4-2/3 of
their length, diplazioid on basal acroscopic; indusia light brown, attachment side
darker, margin entire, fragile.
DISTRIBUTION. Borneo.
ECOLOGY. Limestone areas. Terrestrial on mountain slope in deep
shade. 50-100 m.
SPECIMENS EXAMINED. --- BORNEO: M. Kato & H. Wiriadinata B-
5953; B-5955; B-5956; M. Kato, H. Wiriadinata B-5414; M. Kato & H.
Wiriadinata B-5953; B-5935.
18. Diplazum cumingii (Presl) C. Chr.
Diplazum cumingii (Presl.) C. Chr. Ind. Fil. 230. 1905. --- Athyrium
cumingii (Presl) Milde. --- Ochlogramma cumingii Presl. --- Calliptiteris
alismifolia J. Smith (nomina nudum). --- TYPE: Cuming 116 (Isotype, MW!).
Rhizome short, erect. Stipe dark brown or nearly black, 25.5-39 cm long,
3-4 mm diam. near base, blackish, fallen scales at base. Scales on stipe dark
211
brown, blackish, concolour, subulate, 4-7 mm long, 0.5-1.0 mm broad, margin
entire. Lamina imparipinate, oblong, 37-45 cm long, 19-32 cm broad, lateral
pinnae 2-3 pairs, terminal pinnae like the rest, the largest. Pinnae subopposite,
shortly stalked 2.5-3 mm long, oblong, base cuneate, apexcaudate, margin entire,
to 19 cm long, 4.7 cm broad; terminal pinnae 21.5-26 cm long, 6.8-7.0 cm broad.
Coatae distinctly raised below, dark brown when dry, minutely scales on lower
surface. Veins in small group of 2-4 veins, forming an angle about … ºto costa,
veilets forked 2-3 times again, outer and inner veins uniting with the nearest veins
forming areoles 1 mm wide, or nastomousing 1/7-¼ way from margin to costa.
Sori elongate from costa continuously to near margin, forming areoles near
margin, outer acroscopic diplazioid, inner also. Indusia black, margin entire, firm,
persistent, opening when old, margin of indusia leaving line scar between veins
like false veins.
DISTRIBUTION. Borneo
ECOLOGY. Terrestrial on stream bank in deep shade, lowlands ever
green rain forest, on humus rich slope. 270 – 1500 m.
SPECIMENS EXAMINED. ---BORNEO: Kato & H. Wiriadinata B-
4729; B-5060; M Okamoto & E.B. Walujo B-11491; M.Kato, M. Okamoto &
E.B. Walujo B-11227; M. Kato, M. Okamoto, K. ueda, D. Darnaedi & E.B.
Walujo B-8353; M. Kato, M. Okamoto & E.B. Walujo B-9882; M. Kato, Gen
Murata, & Y.P. Mogea B-3775.
NOTE: The sori and the indusia of this species are very distinctive.
Indusia are rolled back when old, always leaving a distinct ridge indicating the
original position of their outer edges.
DISTRIBUTION. Borneo, Philippines.
19. Diplazium densisquamatum Praptosuwiryo, sp. nov. Plate 4.
Type: SUMATRA. Jambi, Kerinci Seblat National Park, Sungai Penuh,
Bukit Tapan, secondary forest, 1290-1300 m, 5 September 2006, T.Ng.
Praptosuwiryo 2491 (holotype, BO).
Rhizoma breve erecta. Stipites ad 47.5 cm longae, 0.6 cm cassi fere basi,
squamis brunneis nitidus integris ovatis-lanceolatis 7-17 mm longae 0.5-2.5 mm
latae densus penitus vestitae. Lamina ad 95 cm longae, 65 latae, bipinnata
212
deltoideus; pinnate pinnae ad 6 jugatae, pinnatida pinnae ad 7 jugatae infra apice
pinnatifidus. Infimus pinnae stipes ad 1 cm longae, 38-41 cm longae 10.5-13.0 cm
latae, lanceolate, libere pinnulae 13-18-jugatae; pinnulae infimus acroscopicus
leviter dimidia ad 2.8 cm longis, 1 cm latis; pinnulae maximae stipitae ad 1 mm
longis, superiora adnatus-sessilis, oblong subtriangulare, ad 6.8 cm longiae, 1.9
cm latae, basi truncatae, apice acuminatae, e margine ¾ costam versus lobatae;
lobi oblongis, basi basiscopicus maximae, ad 4 mm latae, plerumque 3 mm latae,
truncati, leviter dentatis. Venulae libere, pinnatae in lobo, 5-6-jugatae,
plerumque 5-jugatae, simplices vel in lobo infimus basiscopicus furcatis Rachise
et costae, minute squamis, non gemmiferae. Sori e basi venarum ½, ad 1.5 mm
longae, soris infimus acroscopicus diplazioideus. Indusiis latis, brunneus,
concolorus, marginem lacerus, persistens.
Rhizome short, stout, erect, scales densely on younger part. Stipes dark
brown, black toward base, 47.5 cm long, 0.6 cm cm thick, scales densely
throughout; scales dark brown, ovate-lanceolate, 7-17 mm long, 0.5-2.5 mm
broad, margin entire, without thickening black strands, glandular cell present.
Lamina bipinnate, deltoid, to 95 cm long. 65 cm broad; pinnate pinnae 6 pairs,
pinnatifid pinnae 7 pairs below pinnatifid apex of lamina; lower pinnae stalked to
1 cm long, lanceolate, 38-41 cm long, 10.5-13.0 cm broad, free pinnulae 13-18
pairs; basal acroscopic pinnulae a little reduced to 2.8 long, 1 cm broad; larger
pinnae short stalked to 0.5-1 mm long, upper adnate-sessile, oblong subtriangular,
5.6-6.8 cm long, 1.7-1.9 cm broad, base truncate, apex acuminate, margin lobed ¾
way to costa; lobes oblong, basal basiscopic the widest, 2-4 mm wide, commonly
3 mm wide, ends truncate, slight toothed. Veins free, pinnate in the lobes, veinlets
5-6 pairs, commonly 5 pairs, commonly simple, once forked on basal basiscopic
lobes. Rachise and costa minutely scales, not gemmiferous. Texture
subpapyraceous. Sori elongate from basal covers ½ of its length, 1.5 mm long,
basal acroscopic diplazioid. Indusia broad, dark brown, concolour, margin
lacerate, persistent.
DISTRIBUTION. This species has hitherto only been found in shady
forest of Bukit Tapan, Kerinci Seblat National Park, Sumatra.
ECOLOGY. In shady place of dry land in secondary forest. 1000-1200 m.
213
SPECIMENS EXAMINED. SUMATRA: T.Ng. Praptosuwiryo 2491.
NOTES. Diplazium desisquamatum is in a glance similar to the small
plants of D. polypodioides. However the dense scales throughout the stipes with
ovate –lanceolate and entire margin will differentiate from D. polypodioides
fastly. The stipes of D. polypodioides are densely covered by lineary lanceolate
sharply toothed scales at base only. Moreover D. desisquamatum is differ from D.
.polypodioides in characters combination as follow: larger pinnulae lobed to ¾
way to costa and veinlets 6 pair or less, while D. polypodiodes has larger pinnulae
lobed more deep close to costules and veinlets to 11 pairs.
ETYMOLOGY. The species epithet is from the Latin densus and
squamatus meaning scales are dense in illustrating the densely scales throughout
the stipes and rachis.
20. Diplazium dilatatum Blume
Diplazium dilatatum Blume, En. Pl. jav.: 194. 1828; C.Chr. Holttum,
Gard. Bull. 7: 273. 1934. Backer & Posth., Varenfl. Java: 132. 1939; Holttum,
Gard. Bull. S.S. 11: 85. 1940.
Rhizome stout, short, erect. Stipe to 80 cm or larger, 8 mm or more thick
near base, dark green when living, black and very densely scales at base; scales
narrowly linier, to 15 mm long, 1 mm wide, yellowish brown at middle, blackish
brown and sharply toothed at margin. Lamina bipinnate-tripinnatifid, about 90 cm
long, 70 cm wide, variable in size; pinnae oblong, narrowing toward acuminate
apex, to 58 cm long, 70 cm wide, pinnules to about 16 pairs below deltoid lobed
apex of pinna; basal pinnules usually a little reduce; larger pinnules stalked to 5
mm long, to 18 long, 4 cm broad, oblong subtriangular with attenuate apex, basal
lobes a little reduce, base truncate-cordate on stalked ones, lobed ½ way to costa
or a little more; lobes slightly subdeltoid-semiorbicular, apex rounded, margin
subentire or serrate, 15 by 9 mm, commonly less; rachis glabrescent, costa with
scattered narrow brown scales to 20 by 3 mm; veins in lobes pinnate with 5-7
pairs of simple or forked veinlets. Sori elongate along veinlets from near base to
2/3 length
214
SPORE. Monolete, bilaterally symatrical (made asymmetric by prine),
heteropolar, polar outline (excluding perine) elliptical, sides convex; equatorial
longitudinal view (excluding perine) concave-convex; equatorial transverse view,
proximal face concave, distal face convex; perinate. E: 31.85(45.69)49.56±4.78;
P: 19.84(28.55)34.62±3.89. Laesura: concealed by perine. Perine: alate-to
costate-alate, irregular envelope separated from exine surrounds the spore; wing-
like muri projected c.6 -15 µm µm, terminating margin are often ciliate; surface of
perine smooth. Exine: visible through perine, smooth under SEM.
ANATOMY. Stomata: Polocytic, copolocytic and seppolocytic.
CHROMOSOMES. 2n= 123 (Cytotype: T.Ng. Praptosuwiryo 1073)
DISTRIBUTION. India, Burma, S. China, Taiwan, Ryuku, S. Japan,
Indochina, Malesia throughout to N. Australia.
ECOLOGY. Terrestrial. On humus rich soil of mountain slopes primary
forest or mixed forest at 100-1800 m sea level.
VERNACULAR NAMES. Pakis layung (Javanese), paku beunyeur
(Sundanese).
USE. The young fronds can be eaten as vegetable after cooking.
SPECIMENS EXAMINED. --- JAVA: Adelbert 142; Alston 12771;
Dillewgn 706; Donk s.n., 603; Lefebu 113; Matthew 610; Mogea 2344, 2345;
Mousset 50, 760; Popta 225; Posthumus 3584, 3768, 3939; Sapiin 2660, 2713,
2717; Zippelius 239. --- BORNEO: M. Kato, G. Murata & Y.P. Mogea B-3873;
R.E. Holttum SFN 25555; M.Kato, G.Murata & Y.P. Mogea B-3738; M. Kato,
M. Okamoto, & E.B. Walujo B-10036.
NOTES. Kato (1995) recognized two varieties D.dilatatum in Japan, viz.
var. dilatatum and var. heterolepis. The first variety has scales on stipe base
lanceolate, to 20 mm long, black at margin, while the second variety has scales on
stipe broadly lanceolate, 10-15 mm long, 1-3 mm broad, hardly black at margin.
215
21. Diplazium dolichosorum Copel.
Diplazium dolichosorum Copel., Philip. J. Sci. 1 Suppl.: 151. 1906; M.G.
Price, Contr. Univ. Mich. Herb. 16: 195. 1987. --- TYPE: Philippines,
Mindanau, Zamboanga, San Ramon, 800 m, Copeland 1716 (Holotype, s.n.)
Rhizome stout; scales lineary, 1 cm long, black-dark brown. Stipe
crowded, ca. 35 cm long, 8 mm thick, base densely scales, muricate, upper;
subglabrous; scales dark, lineary, 4-8 mm long; lamina ovate, 1-1.5 mm long,
bipinnate, apex pinnatifid; larger pinnae 45 cm long, 20 cm broad, lower smaller;
pinnulae shortly stalked, subfalcate, to ca. 10 cm long, 2 cm broad, base truncate,
apex serrate, margin lobed to 1/3 way to costa; lobus truncate, toothed; rhachis
glabrescent, grooved on upper surface; texture papyraceous, glabrous, upper
surface shiny, lower surface pale; veins free, pinnate in the lobus, veinlet to 6
pairs, simple, lower curved. Sori lineary, to 8 mm long; indusia broad, brown.
DISTRIBUTION. Borneo, Philippines
ECOLOGY. Terrestrial. Hill dipterocarp forest. Elevation: 1000 m.
SPECIMEN EXAMINED: Borneo: Beaman 10641 (K).
NOTES. This species is described based on photograph of specimen
deposited at Rijks Herbarium, Kew.
22. Diplazium donianum (Mett.) Tardieu
Diplazium donianum (Mett.) Tardieu, Asplén. Tonkin: 58, t 5 1 & 2.
1932; Tagawa & K. Iwats., Fl. Thailand 3 (3): 455. 1988. Asplenium donianum
Mett., Farngatt. 6: 177. 1859.
Rhizome creeping, 4-5 mm thick, blackish, with leaves 1-2 cm apart,
scales on younger part; scales brown shining, ovate-narrowly lanceolate, 2.5-9
mm long, 1-3 mm broad, apex acuminate, brown, margin toohed irregularly with
thickening black strand. Stipe longer than lamina, 28-57 cm long, 3-4.5 mm
thick, dark brown and scales at base, upper pale green when living. Lamina
simply pinnate, oblong in outline, 30-42 cm long, 15-19 cm broad, lateral pinnae
subalternate, to 4 pairs, terminal pinnae conform to lateral ones; pinnae stalked 3-
216
6 mm long, ovate-lanceolate to narrowly oblong, to 16.2 cm long, 3-4.5 cm broad,
base cuneate, apex acuminate, margin entire, slightly toothed; rachise glabrous;
texture firm-papyraceous, upper surface rather light green, glossy, lower pale
green; slightly serrate near apex; veins group forming angle 55-60º to costa, free,
forked near costa, upper branch simple or one forked and soriferous, lower forked
1-4 times again, outer usually also soriferous but shorter. Sori bearing on outer
veins, sometimes also on inner ones, basal acroscopic usually the longest,
diplazioid, elongate from near costa almost reaching the margin; indusia
concolour, brown, rolled back, margin subentire, persistent, opening when mature.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped, with an angle 110, flat base both inward and
outward, angles not forming ridges, ends slightly ridges outward.
CHROMOSOMES. 2n = 164 (Cytotype: XIX.C.III.65, BOHB, living
plant is cultivated at Bogor Botanic Gardens).
DISTRIBUTION. Japan, Taiwan, S. China, Indochina, Thailand, India,
Sumatra, Java, New Guinea.
ECOLOGY. On mountain slopes in light shade or in dense primary forest,
at low elevations lower than 1300 m.
SPECIMENS EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 2340. ---
SUMATRA: XIX.C.III. (Cultivated at Bogor Botanic Gardens).
NOTES. This species is new record for Java and Sumatra. Kato (1995)
also reported chromosomes number ’n’= 164 (apogamous).
23. Diplazium esculentum (Retz.) Sw.
Diplazium esculentum (Retz.) Sw., Scrad. Jour. Bot. 1801 92): 312. 1803;
Syn. Fil. : 92. 1806; Holttum, Gard. Bull. S.S. 11: 86. 1940.
Rhizome erect. Stipe 5-10 mm diam., 14-80 cm long, brown, glabrescent,
black and scales toward base; scales narrowly linier, 7-13 mm long, 1-1.5 mm
wide, concolour, dark brown, margin toothed with thickening black strand.
Lamina pinnate-bipinnate, large, various in size; lower one-two pairs of pinnae
usually reduced; larger pinnae 27-47 cm long, 12-25 cm wide, bearing numerous
217
of pinnules; pinnules oblong, narrowing towards acuminate or attenuate apex,
varying much in size, larger ones 8-13 cm long, 1.5-2.8 cm wide, lower shortly
stalked about 1 mm long, the rest sessile truncate-subcordate or broadly cuneate,
auricled one or both side at base, margin crenate or lobed about 2 mm or less from
margin to ¼ way to costa; lobes or crenation truncate or rounded, and serrate at
apex; texture papyraceous, sometimes fibrillose hairy beneath; rachis glabrescent
or occasionally bearing fibrillose hairs beneath; costa or costae grooved with
distinct ridge above, bearing scattered minutely scales; vein in pinnate group in
the lobes, 8-10 pairs of side veins, the lower 2-3 pairs of adjacent group
anastomousing, forming irregular intermediate eccurrent vein leading towards a
sinus between adjacent lobes. Sori occupying almost the whole length of the
veins, often also on part of the joint excurent vein, 1-4 pairs from basal sometimes
diplazioid; indusia not so thin, dark brown, persistent, margin toothed when
opening.
SPORES. Monolete, bilaterally symmetrical, heteropolar; polar outline
elliptical, sides convex; equatorial longitudinal view concave-convex to plano-
convex; equatorial transverse view, proximal face planar to concave, distal face
hemispherical; perinate. E: 31.72(38.39)43.09±3.35; P: 22.74(26.37)
29.12±1.39. Laesure: not visible, concealed by perine. Perine: smooth under
LM, micro rugulate under SEM. Exine: Exine: often visible through perine,
granulate under LM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an interrupted U-shaped with angle about 100º, base flat, end ridges outward
to form an angle about 140º. Stomata: polocytic and seppolocytic.
CHROMOSOMES. 2n= 82 (Cytotype: T.Ng. Praptosuwiryo 1784, BO)
DISTRIBUTION. India to Fiji, throughout Malesia, north to Japan.
ECOLOGY. On moist soil in light shade, ravine, swampy ground,
grassland (not no dry), hedges, dith banks, roadside, riverside in the forest at
elevation 5-1600 m sea level.
USE. Young fronds of this species is sold in the local marked as
vegetables. It is eaten after cooking.
218
SPECIMENS EXAMINED. SPECIMENS EXAMINED --- JAVA:
Adelbert 435; Arsin 19455; Backer 4479, 7855, 12004, 12574, 17745, 18638;
Backer & Posthumus 4, 108, 162, 308, 400, 419, 601; Bakhuizen v/d Brink 1440,
2401, 3715, 5508; Beguin 76; Dorgello 1938; Hallier 678b; v. Heurn s.n.
(Pujon); Kooders 22754B; Lörzing 698, 699; Oosten a.17; Polak s.n. (Rawa
Lakbok); Posthumus 1455a; T.Ng. Praptosuwiryo 630, 635, 637, 638, 1784,
1785; Raciborski s.n. (Kota Batu); Rumka 3; Rutten 323; Zippelius 178. ---
MALAY PENINSULA: Newton, Z. Teruya 2343. --- BORNEO: K. Iwatsuki M.
Kato, Gen Murata & Y.P.Mogea B-194; M. Kato & Y.P.Mogea B-224;
K.Iwatsuki, M. Kato, M. Okamoto, K. Ueda & E.B. Walujo B-7190; M. Kato &
H. Wiriadinata B-6868; O. Posthumus 2065. ---SUMATRA: CH Lamourex
5652. --- BORNEO: T.Ng. Praptosuwiryo 2017, 2080, 2094b, 2094c, 2161b,
2161c, 2248a, 2248b.
24. Diplazium fraxinifolium Presl
Diplazium fraxinifolium Presl, Rel. Haenk 1: 49. 1825.
Athyrium fraxinifolium (Presl) Milde, Bot. Zeit. 28: 353. 1870.
Rhizome erect, 12 mm thick. Stipe 44.5-67 vm long, 4-7 mm thick, fallen
or sparsely scales at base; scales on stipe 2.5-8 mm long, 1-1.5 mm broad, margin
entire, brown, concolour. Lamina simply pinnate, 37.5-46 cm long 22-44 cm
broad, lateral pinnae 4-5 pairs, terminal pinna conform to the rest; pinnae 4-8 cm
distance, lower stalked 6-7 mm long, upper adnate, elliptical, 21.5-26 cm long,
4.5-6 cm broad, base cuneate, margin waved- crenate entirely or only toward
apex, or lobed to 1/6 way to costa, apex acuminate or caudate; veins group
forming angle 50-65º, each crenation for one vein group; vein forked 5-7 times,
outer veinlets in each veinlets group uniting with outer adjacent veinlets group at
1/5-1/4 way from margin or less. Sori almost on each veinlet, cover ½-5/6 of
veinlet length (2.5-17 mm long), diplazioid on basal acroscopic, others asplenoid
and opening toward acroscopic, except subbasal acroscopic ones opening towards
basiscopic; indusia pale brown, margin entire and darker, persistent, opening
when mature.
DISTRIBUTION. Malaya, Borneo, New Guinea, Philippines.
219
ECOLOGY. Terrestrial on slope in light-deep shade, lowland rain forest
at 20-750 m s.l.
SPECIMEN EXAMINED. --- Borneo: C. Boden Kloss 19028 (BO,
SING); M. Kato & H. Wiriadinata B-5568; B-4728; M.Kato, M. Okamoto & K.
Ueda B-11615, B-11659.
25. Diplazium fuliginosum (Hook.) M.G. Price
Diplazium fuliginosum (Hook.) M.G. Price, Brit. Fern Gaz. 10: 260. 1973;
Gard. Bull. S.S. 36 (1): 26. 1983. Asplenium fuliginosum Hook. Sp. Fil. 3: 120.
1859; C.Chr. Gard. Bull. S.S. 7: 280. 1934; Athyrium fuliginosum (Hook.) Copel.
Philip. J. Sci. 56: 476. 1935; Asplenium lugubre Hook. Second Cent. Ferns
(1861) t.3 (non Liebm. 1849). Type: North Norneo, Mt. Kinabalu, H. Low (K).
Athyrium longissimum Copel. Philip. J. Sci. 38: 139. 1929; Fern Fl.
Philip. 3: 411. 1961; Diplazium longissimum (Copel.) C.Chr. Ind. Fil. Suppl. 3:
74. 1934. Type: Philippines, Leyte, Dagami, Ramos BS 15269, Aug. 1912
(MICH).
Rhizome short, erect. Scales on stipes blackish, shining, margin entire, 6
mm cm long, 1.5 mm broad. Stipe 2.5-7 cm long, 3-3.5 mm diam, blackish when
dry, densely scales. Lamina very narrowly elliptic, 31-79 cm long, 9-13 cm broad
at about 1/3 of upper part, lower 2/3 part fully pinnate, upper 1/3 part pinnatifid,
free pinnae 15-32 pairs, lower pinnae 17 pairs gradually reduced downward;
pinnae subfacate, sessile, lanceolate, to 4.8 -6.7 cm long, 1.2-1.7 cm broad, base
truncate, apex sharply acute, margin subentire; texture very thin; rachis densely
scales beneath; veins free, 1-3 forked, soriferous on acroscopic branch. Sori
elongate along veinlets from basal almost reaching the margin; indusia dark
brown, margin entire, opening when mature.
DISTRIBUTION. Bismarck Arch. (New Ireland), New Guinea
(widespread), North Borneo (Mt. Kinabalu, common), Philippines (Leyte, one
collection).
ECOLOGY. Occurring in shaded moist ravines in montane forests, c.
1000-3000 m.
220
SPECIMENS EXAMINED --- BORNEO: R.E. Holttum SFN 25529;
J.M.S. Clemens 33723; J. & M.S. Clemens 31797.
26. Diplazium halimunense Praptosuwiryo, sp. Nov. Plate 5.
TYPE: Java: G. Halimun, Cikuda Paeh-Cikaniki, ca. 1300 m, 25 February
2006, T.Ng. Praptosuwiryo 2341 (holotype, BO).
Rhizoma breve erectum. Stipites 31-43 cm longae, in sicco stramineus,
basi brunneus squamatuss deciduis; squama linearis lanceolatis, ad 7 mm longae,
1 mm latae, margine irregularis dentatus cum filum niger spissescens. Lamina
pinnata, oblongus, pinnae laterales ad 3-6-jugatae, pinna terminalis ceteris similis,
rachis non prolifera; pinnae stipitatae ad 6 mm longae vel adnatus, ovate-
lanceolate, 15-21 cm longae, 3.5-5.5 cm latae, basi subequaliter cuneate, margine
fere integrae, apicem acuminatus leviter serratus. Venae libere, angulum 60-70°
cum costa formantes, furcatis Sori basalis vel medius, 1/8-4/5 venulae
occupantes. Indusia brunneus, marginem subintegris, persistens.
Rhizome short, erect, scales on younger part. Stipe 31.5-43 cm long,
stramineus when dry, dark brown and fallen scales at base; scales lineary
lanceolate, 5-7 mm long, 1 mm broad, margin with thickening black strand,
irregularly toothed. Lamina simply pinnate, oblong in outline, lateral pinnae 3-6
pairs, terminal pinnae conform to lateral ones; pinnae ovate-lanceolate, 15-21 cm
long, 3.5-5.5 cm broad, base cuneate, margin entire, slightly serrate near apex,
apex acuminate; rachis non gemmiferae, texture thinly papyraceous, light green
when living. Veins free, veins group forming angle 65-70º, forked near costa,
upper branch simple and soriferous, lower branch 2-3 times forked again, outer
basiscopic also soriferous but shorther and asplenoid. Sori linear from near costa
or submedial, cover 1/8 – 4/5 of their length, basal acroscopic the longer and
diplazioid. Indusia brown, margin subentire, opening when matur.
PARATYPE. JAVA, West Java, Bandung, Mt. Patuha, Situ Patengan, ca.
1300-1400 m, 26 Dec. 1997, T.Ng. Praptosuwiryo 749 (BO).
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an interrupted U-shaped with angle about 110º, flat base on both inward and
outward directions, end bluntly ridges to form an angle 140º.
221
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 2341, BO)
DISTRIBUTION. Java.
ECOLOGY. This species is growing on moist humus-rich soil of shady
places in mountain forest. In the Mt. Halimun, D. halimunense was found
growing among D. donianum and D. bantamense. Ca. 1300 m.
SPECIMEN EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 749, 2341.
NOTES. D. halimunense may be closely related to D. bantamense, but the
last two species appears to differ from the first species. Diplazium halimunese
has irregularly sharp toothed scales with thickening black strands, while the
margin scales of D. bantamense are minutely and regularly toothed; Pinnae of D.
halimunense are cuneate at base with margin entire, while pinnae of D.
bantamense are round at base with subentire or serrate at posterior portion. It also
resembles D. donianum, but D. halimunense has erect rhizome, while D.
donianum with creeping rhizome. The two species have similarities in irregular
sharp toothed scales and ovate-lanceolate pinnae.
Cytological observation of one individual of D. halimunense (TNgP
2341b) showed 2n = 123 (triploid). Whereas most indivual of D. bantamense
from Java are tetraploid and only some individual oktoploid. One collection
number of D. donianum planted in Bogor Botanic Gardens (from Sumatra) is
tetraploid (See Chapter 5).
ETYMOLOGY. The specific epithet is formed from name of the locality
where this species is found, Mt. Halimun, Halimun National Park, West Java.
27. Diplazium hewittii (Copel.) C.Chr
Diplazium hewittii (Copel.) C.Chr., Index Fil. Suppl. 1: 26. 1913.; C.Chr.
& Holttum, Gard. Bull. S.S. 7: 273. 1934.
Athyrium paripinnatum Copel., Philip. J. 11 (2): 147. 1915.
Diplazium paripinnatum Alderw., Handb. Suppl. 1 : 266. 1916.
Rhizome short, erect. Scales on stipes concolour, dark brown-blackish,
shiny, ovate, to 2.5 mm long, 2 mm broad, margin entire, glandular cell on tip
when young. Stipe 25-67 cm long, 2.5-3.5 mm diam. near base, light brown,
slightly muricate, black at base. Lamina pinnate – bipinnate, subtrangular in
222
ouline, 30 – 70 cm long, 20-27 cm broad. Pinnae opposite-subopposite; lower
pinnae stalked to 1.5 cm long, lanceolate, 22 cm long, 5.7 cm broad, pinnate, apex
acumimate; lower pinnulae adnate – subsessile, upper pinnule sessile below
deeply lobed apex of pinnae; larger pinnulae nearly hastate, 1.8-3.3 cm long, 0.4 –
1.0 cm broad, base broadly cuneate, upper base distinctly auricle, lower base less,
apex acute-acuminate, margin crenate; middle pinnae with pinnulae sessile; upper
pinnae gradually less pinnate-pinnatifid; texture subcoriaceous; surface naked,
lower pale, upper dark green when living; rhacise naked; veins free, forked in
each crenatiron, oblique. Sori elongated along veinlet from near costule covers
5/6 way to margin, acroscopic branch diplazioid; indusia brown, fragile,
persistent, margin entire.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 1913b,
BO).
DISTRIBUTION. Borneo
ECOLOGY. Growing on humus rich soil, low sandy ridges, on shaded
places in the forest at 400-450 m.
SPECIMENS EXAMINED. --- BORNEO: M. Kato, M. Okamoto, K.
Ueda & E.B. Walujo B-7383; M. Kato, M. Okamoto, K. Ueda & E.B. Walojo B-
7902; B-7903. K. Ueda & D. Darnaedi B-8835; T. Ng. Praptosuwiryo 1913a;
1913b; 1913c; 1908a; 1908b; 1908c; 1942; 2171; 2172a; 2172b, 2174; 2177;
2178; 2179; 2180.
NOTES. Christensen & Holttum (1935) stated that a series of specimens
collected by Mjoberd in Sarawak has shown that Athyrium Hewittii, A.
sarawakense, and A. paripinnatum of Copeland probably all forms of the same
species, different in size but otherwise scarcely distinguishable.
My collections from Muller Range, Borneo, showed that this species has
variation in morphology from a young to adult. Young plants in which spores
have not been produced are showing pinnate fronds while adult plants having
fronds with pinnate-bipinnate.
223
28. Diplazium hottae Tagawa
Diplazium hottae Tagawa, Acta Phytotax. Geobot. 25 (2-3): 65. 1972. ---
TYPE: Hotta 15185 (Holotype, KYO, n.v.), Gunung Mulu, Sarawak.; Kokawa &
Hotta 2364, 2415 (Paratype, KYO, n.v.).
Rhizome shorth, stout, erect. Scales on stipes light brown, shining,
concolours, 9-15 mm long, 1-2 mm broad.margin entire, Stipe 56-67.5 cm long, 3-
6 mm diam. at base stramineous, deeply grooved upper surface, base dark brown,
scales, upward glabrous. Lamina oblong, lanceolate, ca. 85 cm long, 36 cm broad,
pinnae 11-21 pairs; lower pinnae stalked to 5 mm long, lanceolate, larger pinnae
28 cm long, 2.5 cm broad, base narrowly cuneate, apex acuminate and slightly
crenate, margin entire; rachis glabrous, gemmiferous at the adjacent between
rachise and costa near terminal pinna; veins group forming an angle about 45º to
costa; veins forked near costa, upper branch simple, lower branch forked again
once-twice; Sori elongate along acroscopic branch, mostly not so close to costa,
almost reaching the margin; indusia light brown, shining, margin entire, rolling
back, not so broad.
DISTRIBUTION. Sumatra, Borneo,
ECOLOGY. Terestrial on rather dry mountain in light shade. 300 –
1350 m.
VERNACULAR NAMES. Paku rahang (Dayak).
USE. Dayak use this plant for medicine. The very young fronds are
crushed for sore poultice.
SPECIMENS EXAMINED. --- BORNEO: M. Kato, M. Okamoto, K.
Ueda 7 E.B. Walujo B-7428; M. Kato, M. Okamoto, K. Ueda & E.B. Walujo B-
7675; K. Ueda & D. Darnaedi B-11571; M. Kato, M. Okamoto, K. Ueda & E.B.
Walujo B-7879; T.Ng.Praptosuwiryo 1911. --- SUMATRA: C.J. Brooks 330 S.,
T.Ng. Praptosuwiryo 2536a.
NOTES. Tagawa (1972) stated that D. hottae is allied to D. subintegrum
Holtt. Occurring on Malay Peninsula and in Northern Sumatra. This species
differs with D. subintegrum constantly in: (1) terminal not lobed at base and
similar to apper laterals ones in shape and size, (2) uppermost 1 or 2 lateral pinnae
gemmiferous at the base on the rachis, (3) pinnae apparently entire, but in reality
224
provided with minute and remote incision on margin, (4) venation obscure,
acroscopic veinlet of group simple and soriferous, basiscopic one once forked and
sterile, (5) sori all asplenioid, narrower, very unequal in length, their anterior ends
arranged in an intramarginal uneven line, (6) scales at the base of stipe much
longer.
29. Diplazium insigne HolttumDiplazium insigne Holttum, Gard. Bull., S.S. 9: 123. 1937; Holttum,
Gard. Bull. S.S. 11: 87. 1940. --- TYPE: R.E. Holttum SFN 21635 (holotype,
SING!), Trans Valley, Frasser’s Hill, Pahang, Malay Peninsula.
Rhizome stout, short, erect. Stipe stout, c.1 m long, spiny toward the base,
the spines 2 mm long, each at first bearing a scale; scales dull brown 1.5 cm long
by 1.5 cm wide at base, with a narrow black toothed edge, deciduous. Lamina to
1.5 m long, bipinnate; lowest pinnae about 28 cm long and 6.5 cm wide, narrowed
and stalked at the base, the margins lobed half-way to the costa, the apex
acuminate; middle pinnae largest, to 60 cm long and 16 cm wide, pinnate;
pinnules slightly oblique, adnate to the rachis (the lowest ones narrowly, the upper
fully adnate and grading into the lobed apical lamina of the pinna), to 9 cm long
and 2.3 cm wide, cunete at the base at an angle of about 45o on each side,
narrowed gradually from the base and then suddenly at 1.5 – 2 cm, from the apex,
margins slightly serrate; texture firmly herbaceous; veins anastomousing as in D.
accedens. Sori few or copious, the lowest acroscopic sorus in each vein-group
usually diplazioid, and occasionally a few others also.
ECOLOGY. In moist shady valley forest. 600-1200 m
DISTRIBUTION. Malay Peninsula.
SPECIMENS EXAMINED. --- MALAY PENINSULA: R.E. Holttum
SFN 21635.
NOTES. As stated by Holttum (1940) this species is evidently closely
allied to D. accedens and the apex of afrond might pass for that species, but its
copious bipinnate form marks it as a quite distinct species. Holttum (1940) added
that this species is also similar to D. Smithianum from Ceylon, but appears to be
225
much larger; probably both D. Smithianum and D. insigne are local derivatives of
D. accedens (or D. proliferum).
30. Diplazium kunstlerii Holttum
Diplazium Kunstlerii Holttum, Gard. Bull. S.S. 11: 88. f. 2. 1940. –
Athyrium kunstlerii (Holtt.) Holttum, Rev. Fl. Malaya 2: 564. f. 335. 1966. ---
TYPE: R.E. Holttum SFN 31194 (holotype, US!).
Rhizome suberect, about 2 cm diam., scales on younger part. Stipe 7 mm
diam at base, 70 cm long, glabrescent, dark brown, nearly black and scales at
base; scales rounded. Lamina bipinnate, about 107 cm long, 60 cm wide; pinnae
about 11 pairs; larger pinnae 52 cm long, about 16 cm wide, on stalk to 11 cm
long, upper pinnae with shorter stalk; lowest pinnules to about 5 cm apart, largest
ones 9.2 cm long, 2.7 cm broad, on stalk to 2 mm long, base broadly cuneate to
truncate, margin lobed to 2/3 way toward costa; lobes slightly oblique, 5-7 mm
wide, base dilated, margin slightly toothed, apex rounded; texture thicker than in
allied species; rachis gemmiferous at upper portion; veins in each lobe 6-9 pairs,
oblique, mostly simple, distinct but hardly prominent on both surface. Sori at
middle part of the veins or nearly close the costules, basal acroscopic veinlets
usually diplazioid; indusia narrow, hardly evident in mature sori, very dark brown.
DISTRIBUTION. Malaya, Java.
ECOLOGY. Locally abundant near streams in shady forest in the
foothills. Ca. 900-1000 m.
SPECIMENS EXAMINED --- MALAY PENINSULA: R.E. Holttum
SFN 31194. --- JAVA: Winckel 1556B.
31. Diplazium laevipes C.Chr. in C.Chr. & Holttum
Diplazium laevipes C.Chr. in C.Chr. & Holttum, Gard. Bull. S.S. 7: 271.
Pl. 58. 1934. TYPE: Borneo, Mt. Kinabalu, H. 25259 (Holotype, SING!).
Stipe 55-75 cm long, ca. 3-8 cm diam.near base, glabrescent, dark brown
and scales towards base. Scales on stipes rounded, to 3.5 mm diam., dark brown,
margin entire, more thick toward central. Lamina subdeltoid, bipinnatifid-
226
bipinnate, pinnae numerous; pinnae shortly stalked 5-15 mm long, oblong
subtriangular-ovate outline, basal pinnae smallest, oblong lanceolate, base
subcordate-subttruncate, subbasal pinnae 32-37 cm long, 8-19 cm wide, pinnulae
to 10 pairs below pinnatifid or deltoid lobed apex of pinnae, acroscopic basal
basal pinnulae a little reduced; pinnulae almost at right angle to costa, lower
shortly stalked less than 1 mm long, oblong subtriangular, 10 cm long, 3.5 cm
broad, base subtruncate, apex acuminate with slightly toothed, margin lobed 1/3-
2/3 way to costule; lobus oblong, slightly oblique, to 4 mm above base, acroscopic
basal a little reduce, ends truncate, entire-slightly toothed; texture stiff , blackish
on upper surface and dark brown beneath when dry; rachis and costa deeply
grooved and rather tomentose above, gemmiferous, gemmae bearing at adjacent
between upper part of rachis and costa; veins pinnate in each lobe, free, not so
distinct on both surface, forming an angle about 65-80º to costule; veinlets simple,
to 9 pairs, occasionally once forked on one-two pair basal lobus, forming an angle
about 25-35º to costulet. Sori elongate from basal veinlet cover ¼-2/3 their
length, diplazioid on basal acroscopic; indusia dark brown, fragile.
DISTRIBUTION. Borneo.
ECOLOGY. Occurring on forest floor in deep shade near river or ravine
at 900-1300 m.
SPECIMENS EXAMINED. --- BORNEO. Kalimantan Timur: M. Kato,
M. Okamoto, E.B. Walujo B-11261, B-9596, B-10748, B-11486, B-9625, B-
7399; M. Kato, M. Okamoto, K. Ueda, D. Darnaedi & E.B. Walujo B-8345; M.
Kato, M. Okamoto, K. Ueda & E.B. Walujo B-7399.
NOTES. As pointed out by Christensen & Holttum (1934), D. laevipes is
closely related to D. spiniferum. The two species are quite similar in size, colour
and division, but the step of D. spiniferum is prckly, the segments all entire, the
sori far from the costule, and the frond coriaceous.
227
32. Diplazium latisquamatum Holttum
Diplazium latisquamatum Holttum, Gard. Bull. S.S. 9 : 124. 1937. Fig. 3.
Gard. Bull. S.S. 11: 90. 1940. Fig. 3; --- TYPE: R.E. Holttum SFN 31311
(holotype, SING!), S. Terla, Ulu Telom, Pahang, Malay Peninsula. –Athyrium
latisquamatum (Holtt.) Holttum, Rev. Fl. Malaya 2: 563. f. 334. 1966.
Rhizome short, erect. Stipe 52-100 cm long, 5-9 mm thick, dark brown,
nearly black at base, scales throughout, more dense at base; scales rounded-ovate,
dark brown, center thicker, margin entire, 4-8 mm long, 2.5-4 mm broad,
thickening black strand irregularly near apex. Lamina bipinnate-tripinnatifid, 50-
125 cm long, about 40-80 cm wide; pinnae stalked 2.5-5.5 cm long, lanceolate,
larger pinnae 26-54 cm long, 15-18 cm wide, pinnulae 10-13 pairs below
pinnnatifid apex of pinnae, basal pinnules a little reduce, smaller on acroscopic
than on basiscopic side; pinnulae sessile on upper ones, shortly staked on lower
ones, the larger 6.5-11 cm long, 1.8-3 cm wide, base broadly cuneate to
subtruncate, gradually narrowed to acuminate apex, margin lobed to within 1.5-2
mm of the costa (3/4 way to costa), lobes slightly oblique, 2.5-6 mm wide,
commonly 5 mm wide, apex truncate, margin entire or toothed near apex; texture
thin, firm; rachis occasionally gemmiferous at the adjacent to the costae at the
apex of lamina, glabrous; costae bearing scattered small rounded-ovate scales
beneath; veins pinnate in lobes, veinlets 5-7 pairs, simple or forked. Sori covering
¼-1/2 way of veinlet, acroscopic basal veinlets usually diplazioid. Indusia thin,
brown, broad, torn from margin to the base when opening, persistent.
DISTRIBUTION. Malay Peninsula, Java, Borneo.
ECOLOGY. In moist shady valley of montane forest. These plants
often grow in very wet ground. Elevation: 1200-2700 m.
SPECIMENS EXAMINED --- JAVA: Matthew 609; Meijer 1453, 1818;
Popta 203; Raciborski 108. --- MALAY PENINSULA: R.E. Holttum SFN 31311
(holotype, SING!). --- BORNEO: J. & M.S. Clemens 27122, 27951, 28391,
29716, 32516, 32952.
228
33. Diplazium lobbianum Moore
Diplazium lobbianum Moore, Ind. Fil.: 331. 1861; Alderw., Malay. Ferns
Hand.: 408. Asplenium lobbianum Hook., 2nd Cent. of Ferns, t. 17. 1861.
Rhizome short, erect. Stipes up to 70 cm or larger. 7 mm diam. when
dry, glabrescent, dark brown, black and scales toward base, distinctly grooved on
upper surface; scales subulate, to 15 mm long, 5 mm wide at base, brown, toothed
with thickening black strand at margin. Lamina simple pinnate, oblong-ovate in
outline, up to 53 cm long, 27 cm wide near subbasal, pinnae to 18 pairs below
deltoid acuminate with deeply lobed of terminal frond; pinnae subhorizontal,
ascending, oblong lanceolate to linier-oblong, lower ones on stalk to 3 mm long,
basal pinnae up to about 10-14 cm long, 2-2.8 cm wide or larger, subbasal pinnae
14-15.5 cm long, 2.3 cm wide, upper base subtruncate, lower base slightly
rounded, upper pinnae sessile with obliquely rounded-cuneate at base; margin
entire on lower part, toothed near apex, apex acuminate, subfalcate; texture thinly
coriaceous, vein-group at an angle 50º to costa, forked near costa, upper branch
simple, soriferous, lower branch forked 1-2 times again, scarcely 3 times,
basiscopic and also one pair subbasal soriferous. Sori on acroscopic veinlets
elongate nearly from the costa almost reaching the margin or more far (covers ½-
3/4 way to wargin), usually diplazioid, basiscopic veinlets soriferous also, shorter,
mostly simple; ndusia frim, fragile, brown, margin entire, persistent.
SPORES. Monolete, bilaterally symmetrical (made asymmetrical by
perine), heteropolar; polar outline elliptical, sides convex; equatorial longitudinal
view concave-convex to plano-convex; equatorial transverse view, proximal face
planar to concave, distal face hemispherical; perinate. E: 38.20(46.10) 51.87
±3.22; P: 20.47(26.28)32.81±3.73. Laesure: concealed by perine ridge or wing-
like muri. Perine: costate-alate; wing-like muri surrounds the spore without
forming reticulation or with forming loose reticulatons, terminating margins
ciliate; surface of perine smooth to scatterly ciliate or pappilate. Exine: often
visible through perine under LM, smooth under SEM.
ANATOMY. Tranverse section of stipe near lamina: vascular bundle
form an anterrupted U-shaped with angle about 115º, base flat on inward and
229
outward directions, angle and end not forming ridges. Stomata: polocytic and
seppolocytic.
CHROMOSOMES. 2n=164 (Cytotype: T.Ng. Praptosuwiryo 720, BO).
DISTRIBUTION. Java, Philippines, New Guinea.
ECOLOGY. On moist ground or humus-rich soil in mountain slopes
secondary and primary forest in altitude beween 1500-1800 m sea level.
SPECIMENS EXAMINED. --- JAVA: Backer 14716, 15910; Donk 612,
632, 771; Popta 204; Posthumus 204; T.Ng. Praptosuwiryo 720, 722, 723, 1190,
1240, 1239, 1245, 1246, 1251, 1346, 1487 (G. Slamet), 1488 (G. Slamet);
Raciborski 52.
34. Diplazium lomariaceum (Christ) Price
Diplazium lomariaceum (Christ) Price, Gard. Bull. S.S. 36 (1): 27. 1983.
Asplenium lomariaceum, Verh. Naturf. Ges. Basel 11: 229. 1895. Type: central
Celebes, Takalekadjo, F.& P. Sarasin 994, 8 Feb. 1895 (P, lectotype, n.v.)
Diplazium merrillii Copel. Philip. J. Sci. 2C: 128., t.2A. 1907; Hieron.
Bot. Jahrb. 56: 134. 1920; Athyrium merrillii (Copel.) Copel. Philip. J. Sci. 3C:
300. 1908; 56: 476. 1935; Fern Fl. Philip. 3: 411. 1961. Type: Philippines,
Mindoro, Mt. Halcon., Merrill 5914, Nov. 1906 (Lectotype MICH; US, n.v.)
Diplazium porphyrolepium v.A.v.R. , Bull. Jard. Bot. Buitenz. II, 20: 11.
1915. Type: Celebes, Soemalilah, Capt. Van Vuuren’s Explor. Comm., Rachmat
418 (BO?; L).
Diplazium porphyrophyllum v.A.v.R., Bull. Jard. Buitenz. II, 28: 18.
1918. Lectotype: Ceram, Wai Lantabi, L. Rutten’s Explor. Comm., Kornassi
1240, 4 May 1918 (L!).
Athyrium altum Copel. Philip. J. Sci. 38: 138. 1929; Fern Fl. Philip. 3:
411. 1961; Diplazium altum (Copel.) C.Chr. Ind. Fil. Suppl. 3: 72. 1934. Type:
Philippines, Mindanau, agusan, Mt. Urdenta, Elmer 14081, Oct. 1912 (MICH; L,
NY, US, n.v.) .
230
Scales blackish, shiny, entire, to 9 by 1 mm, gradually narrowed to a hair-
tip, abundant and persistent on stipe, rachis, costae, and vein beneath, Stipe of
fertile fronds to 15 cm long, of sterile tp 9 cm. Lamina narrowly elliptic, to 50 by
9.5 cm, deeply pinnatifid, one pair of reduced basal pinnae sometimes free and
sessile; lobes to 4.5 by 1 cm, oblong-lanceolate, narrowing towards apex,
subentire, blunt. Colour dark greenish-brown, more or less shiny below when dry,
dark bluish-green when living; rachises above with a channel formed by raised
cartilaginous sides, either continuous or interrupted at each junction with a middle
of a lobe, and paleate at that point whether or notinterrupted; indusia brown,
margin erose. Spore brown, with irregular short wing.
SPORES. Monolete, bilaterraly symmetrical (made asummetric by
perine), heteropolar; polar outline (excludong perine) elliptical; equatorial
longitudinal view (excluding perine) biconvex; equatorial tranverse view,
proximal face convex, distal view hemispherical; perinate. E:
35.03(39.41)54.55±4.20; P: 19.55(24.57)30.13±3.46. Laesura: concealed by
wing-like costae. Perine: alate to costate-alate, alae forming loose reticulation;
irregular envelope separated from exine surrounds the spore in wing-like muri
reticulation, lacunae large, 13-21µm; wing-like muri thin, project 3-10 µm,
terminating margin waved or almost entire. Exine: visible through perine, rough
rugulate under SEM.
DISTRIBUTION. Philippines, Borneo, Sumatra, Celebes, Ceram, New
Guinea.
ECOLOGY. This species usually grows in moist montane forest, 400-
2000 m.
SPECIMENS EXAMINED --- BORNEO: Garry Shea SHEA 23160;
Maskuri 845; Veldkamp 7874. -- SUMATRA: CJ Brooks 322S.
NOTE: Price (1983) state that the holotype was destroyed in 1945 at the
PNH so he designated the MICH specimen as lectotype. D. porphyrolepium and
D. porphyrophyllum are not excep not exceptional in any way. The latter had a
syntype purportedly from Sumatra, Brooks 322S. Price (1983) designated the
specimen from Ceram at L as lectotype. Athyrium altum is a form with narrow
fronds not otherwise distinguishable, Three specimens from eastern Kalimantan
231
(Kortermans 9089, Meijer 577, 872 – all L) differ by having fewer and brownish
paleae but agree in distribution of paleae, and in frond form and colour.
Diplazium lomariaceum is very closely related to D. pophyrorachis and until now
the name sees to have been ignored since Christ himself reduced lomariaceum to
porphyrorachis in Ann. Jard. Bot. Buitenz. 15 (1989, p. 119).
35. Diplazium loerzingii Praptosuwiryo, sp. nov.
TYPE: SUMATRA. North Sumatra, E. Mount Sibajak, Upper Petani
Valley, primary forest, 1250-1350 m, 5 Feb. 1929, J.A. Lörzing 15124 (holotype,
BO).
Rhizoma breve erectum. Stipites ad 45 cm longi, in sicco brunnei, basin
versus aquamis brunneis nitidis lanceolatis, ad 13 mm longis 1.5 mm latis integris
acuminatis vestiti. Lamina pinnata lanceolatis, ad 75 cm longa et 32 cm supra
basin lata, pinnae ad 24-jugatae; rachis supra sulcata, infra acem gemmifera.
Pinnae subbasales maximae (1-3 jugatae deflexae), petiolulatae ad 5 mm longae,
lanceolatae, ad 17.5 cm longae, 2.8 cm latae, basi truncatae, apice attenuatae,
margine ½-2/3 costam lobatae; lobi maximi ad basim, 5-9 mm lati, apice truncati,
margine apices leviter dentati. Rachis supra sulcata, glabrae, infra apicem
gemmifera. Textura subcoriacea vel plus solid; in vivo colorae supra atroviridis,
subtus pallidus. Venae perspicuus in uno lobo 3-5 jugatae, plerumque 5 jugatae,
omnes simplices. Sori medius in 1/3-3/4 longitudine venularum. Indusia
brunneus, in affixus atro brunneus, marginem integrum, non revolutus, persistens.
Rhizome short, erect, scales densely on younger part; scales lanceolate, 7-
13 mm long, 0.5-1.5 cm broad, dark brown-blackish, with thickening black
strands, shining, margin entire. Stipe 33-45 cm long, 4-5 mm thick, pale brown
when dry, black at base, fallen scales. Lamina lanceolate, pinnate, 61-75 cm long,
24-32 cm broad, pinnae 22-24 pairs below pinnatifid triangular apex of lamina.
Lower pinnae stalked to 3-5 mm long, 2.3-3.6 cm apart, oblong-lanceolate, larger
ones 10.3-21.3 cm long, 1.8-3.2 cm broad, base of 1-3 pairs basal pinnae cut away
lower, base truncate, margin lobed to ½-2/3 way to costa, apex attenuate, toothed;
upper ones adnate-sessile; lobus widest at base, larger ones 5-9 mm broad, ends
truncate, slightly toothed. Rachise gemmiferous, bearing buds at the adjacent
232
between rachis and costa, glabrous, costa glabrous. Texture subcoriaceous or
firmer, dark green upper surface, pale green lower when living. Veins free,
pinnate in the lobus, mid-veins of lower lobus forming angle 70-75 ° to costa,
veinlets 3-5 pairs, commonly 5 pairs, distinct, simple, all reaching margin,
forming angle 25-30º to midveins. Sori medial, (leaving sterile part of veinlets
1.5-2.5 mm) or close to margin of lobus, elongate cover 1/3-3/4 of veinlet length,
basal acroscopic diplazioid. Indusia broad, brown, attachment side darker, margin
entire, opening when mature, not rolled back, persistent.
PARATYPE. JAVA. West Java, Mt. Halimun, track Cikuda Paeh-
Cikaniki, ca. 1300 m, 25 February 2006, T.Ng. Praptosuwiryo 2339 (BO).
SUMATRA. Sibolangit Nature Reserve, 400-500 m., 10 December-1928, J.
Buumee A. 841; Jambi, Kerinci Seblat National Park, Sungai Penuh, Bukit
Tapan, secondary forest Pal 823-829, 1360 m, 9 Sept. 2006, T.Ng. Praptosuwiryo
2519, 2520, 2521 (BO).
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 2339c , 123
(T.Ng. Praptosuwiryo 2339d).
DISTRIBUTION. Java and Sumatra.
ECOLOGY. Secondary and primay forest, light-deep shady places, on
moist and humus rich soil. 1250-1400 m.
NOTES. Diplazium loerzingii is closely related to Diplazium malaccense.
This species differs from D. malaccense in the following characters: lower base
of 1-2 pairs basal pinnae less cut, base of lower pinnae almost equally truncate,
texture thicker or subcoriaceous, upper rachis much gemmiferous, lobes truncate,
sori medial on veinlets or close to margin, attachments sides of indusia darker.
ETYMOLOGY. This species is named after J.A. Lörzing who collected
this plant for the first time. He collected D. loerzingii in 1920 at Mt.Sibajak
Sumatra. Nine year after that he collected this plant again in the same place. C.J.
Brook also found this species at Mt. Dempo (Sumatra) in 1923. I found this plant
at Mt. Halimun (Java) and Bukit Tapan (Kerinci Seblat National Park, Sumatra)
respectively in Februari and September 2006.
233
36. Diplazium malaccense Presl.,
Diplazium malaccense Presl, Epim.: 86. 1849; Tard. & C.Chr. in Fl. Gén.
I.-C. 7(2): 258. 1940; Tagawa & K. Iwats., Southeast As. St. 5 : 104. 1967; Acta
Phytotax. Geobot. 23 : 56. 1968. -- Athyrium malaccense (Presl.) Hottum, rev. Fl.
Malaya 2 : 525. 1955.
Rhizome short, ca. 1.5-1.8 cm diam, erect, densely scales on young part.
Stipe 39- 48.5 cm long, 0.3-0.5 cm diam near base, densely scales at base; scales
10-12 mm long, 1 mm broad, lineary triangular, brown. Lamina pinnate, oblong
elliptical –subtriangular in outline, 33-64.5 cm long, 15-30 cm broad, pinnae 16-
19 pairs below lobed apex of lamina. Lower pinnae stalked 2-5 mm long, 1-2
pairs basal pinnae bending downward, lineary- subtriangular, 9-19.5 cm long, 1.1-
2.8 cm broad, base very unequal, lower base cut away, upper truncate, upper
pinnae with base subequally truncate or cuneate, Pinnae thin in texture, drying
rather light green, margin lobed 1/3-2/3 way to the costa, apex acuminate; lobes
about 4-7 mm wide at base, oblique, apex rounded to subtruncate, slightly
toothed. Veins 4-7 pairs in each lobe, all simple. Sori from near base of veins to
near margin of lamina, acroscopic basal sorus usually diplazioid. Indusia medium
brown, thin but firm, narrow, margin entire.
SPORES. Monolete, bilaterally summetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) elliptical; equatorial
longitudinal view planar convex to concave-convex; equatorial transverse view,
proximal face concave, distal view convex. Laesura: concealed by wing-like
muri. E: 37.73(34.24)27.53±3.03; P: 26.01(20.80)13.33±2.82. Perine: alate to
costate-alate, often loosely reticulate; irregular envelope separated from exine
surrounds the spore in continuous anastomosing wings, forming a loose
reticulation; wing-like muri project 1.5-8 µm, terminating margins sparsely ciliate.
Exine: visible though perine, smooth granulate under SEM.
ECOLOGY. In evergreen forest, terrestrial on rather moderate slope, in
light shade, on humus-rich soil ground. 20 – 1000 m.
SPECIMENS EXAMINED. --- MALAY PENINSULA. Pahang: M.R.
Henderson 11235; Md Nur 10508; R.E. Holttum 20788; A.B. Murdok s.n. (12
June 1913); H.N. Ridley s.n. (1891). Negeri Sembilan: R.E. Holttum 9565; Md
234
Nur s.n. (29 Nov. 1923). Melaka: A. Derry 14. Penang: R.E. Holttum SFN
31196. --- BORNEO. Kalimantan Timur: K. Ueda & D. Darnaedi B-8982; M.
Kato, M. Okamoto & E.B. Walujo B-9070; M. Kato, M. Okamoto, K. Ueda &
E.B. Walujo B-7398; M. Kato, M . Okamoto & E.B. Walujo B-10909; M. Kato,
M. Okamoto, K. Ueda, E.B. Walujo B-7871; K. Iwatsuki, M. Kato, G. Murata &
Y.P. Mogea B-2472. Kalimantan Selatan: M. Kato, G. Murata & Y.P. Mogea B-
4155; 4175. --- SUMATRA: W.J.J.O. de Wilde & B.E.E. de Wilde – Duyfjes
12560; W.J.J.O. de Wilde & B.E.E. de Wilde – Duyfjes 12920; T.Ng.
Praptosuwiryo 2218, 2220, 2209, 2217, T.Ng. Praptosuwiryo 2223, 2224, 2225,
2226, 2229, 2230, 2233, 2235, 2236, 2237, 2238.
37. Diplazium megasegmentum Praptosuwiryo, sp. nov. Plate 7.
TYPE. JAVA. West Java, Mt. Salak, Southern Slope, Cangkuang Forest,
19 December 2002, T.Ng. Praptosuwiryo 1382 (foto) (holotype, BO).
Rhizoma breve erectum, crassum. Stipites ad 122 cm longis, prope basin
10 mm crassi, in sicco brunneis, basin versus squamis deciduis nigris rotundis vel
ovatis peltatis integris imbricates vestitae. Lamina ad 100 cm longae, 80 cm latae,
subdeltoideus, tripinnatifidus, pinnae ad 14-jugatae; pinnae inferus bipinnatidus,
stipites ad 3.5 cm longae, lanceolatus, 50 cm longae 26 cm latae, libere pinnulae
ad 16-jugatae, apice pinnatifidus; pinnulae inferus subalternatus, 2.5-3.5 cm
seorsum, stitipatae ad 3 mm longae, oblongus-lanceolatus, ad 13 cm longae 3.5
cm latae, basi truncatae, apice acumitae, margine lobatae segmenta formantes,
segmenta inferiora plus minusve opposite, 2.5-5.0 mm seorsum, sessilis;
segmenta superus subalternate; segmenta maximae 2.4 cm longae et 8 mm latae,
apice rotundatus-acutus, margine 1/3-1/2 versus venae principalis; pinnae medius
pinnatus, pinnae superus pinnatifidus. Venae liberis, in segmento pinnatus,
venulae ad 6-8-jugatae, in lobo furcatis vel bifurcate. Sori e basi venulae ramis
acroscopicus 1/3-1/2 extensi, plerumque in basi venulae acroscopicus
diplazioideus. Indusiis brunneus, margine laciniatus, ante sporangiis maturis
aperiens, persistens.
235
Rhizome erect, stout. Stipes dark brown, 122 cm long, 10 mm diam. near
base, fallen scales; scales rounded, ca. 6 mm long and broad. Lamina bipinnate-
tripinnatifid, subdeltoid in outline, 100 cm long, 80 cm broad, pinnae to 14 pairs,
lower pinnae bipinnatifid, middle pinnae pinnate, upper pinnae pinnatifid. Larger
pinnae stalked to 3.5 cm long, lanceolate, 50 cm long, 26 cm broad, free pinnulae
16 pairs below pinnatifid apex of pinnae. Lower pinnulae subalternate, 2.5-3.5
cm apart, shortly stalked to 3 mm long, oblong-lanceolate, 13 cm long, 3.5 cm
broad, base truncate, apex acuminate, margin lobed deeply to whitin 1 mm of
costa forming segments; lower segments opposite, 2.5-5.0 mm distance, sessile;
upper segments subalternate below deeply lobed acuminate apex of pinnulae;
larger segments oblong, 2.4 cm long, 8 mm broad, apex rounded-acute, margin
lobed 1/3-1/2 way to main vein. Veins free, pinnate in the segments, to 6-8 pairs
Veinlets once-twice forked in lobes. Sori bearing on basal acroscopic branch of
veinlets, elongate from basal covers 1/3 -1/2 of their length, diplazioid on basal
acroscopic veinlets. Indusia brown, persistent, opening when sori young, margin
laciniate.
PARATYPE. Java: West Java, Mt. Salak, Southern Slope, Cangkuang
Forest, 28 December 2002, T.Ng. Praptosuwiryo 1443, 1450, 1451, 1452.
SPORES. Monolete, bilaterally symmetrical (mades asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) concave-convex;
equatorial transverse view, proximal face convex, distal face convex; perinate. E:
39.61(51.42)57.01±4.28; P: 18.51(31.49)35.85±3.59. Laesura: concealed by
perine. Perine: alate; alae sometimes loosely reticulate; wing-like muri project 4-
13 m, terminating margins almost entire.
DISTRIBUTION. Java.
ECOLOGY. Diplazium megasegmentum is growing well on moist humus-
rich soil in shady places.
NOTES. Diplazium megasegmentum is very distinct among Javanese
species Diplazium in the combination of the following characters: pinnules cut to
within 1 mm of costa to form a big oblong segments to 8 mm broad that lobed
again to ½ way to main vein; basalis sori which not more than half away to the
236
main vein. It is grows on moist humus-rich soil in shady forest in southern slope
of Mt. Salak West Java.
ECOLOGY. The species epithet is from the Latin mega and segmentum
meaning the large segment in illustrating the large size of its large lobed
segments.
38. Diplazium megasimplicifolium Praptosuwiryo, sp. nov. Plate 8.
TYPE: Borneo: Bukit Raya, Tumbang Tubus, Veldkamp 7998, 150 m, 5
May 1983 (holotype, BO).
Rhizoma breve erectum, gracilis. Stipites ad 15 cm longae, in sicco
brunneus dilutus, glabratus, basi deciduis squamatus; squamis (?). Lamina
simpliter, ellipticus, ad 40 cm longae, 7.5 – 10.6 cm latae, angustatus versus
apicem acuminatus, gradatin decrescens versus basin qui subito contractus e
latitude 1 cm vel minus, basi cuneatus, margine integrae; subcartaceus; rhachides
prominens; venae in parvus caterva in angulum 65-70º cum costa formantes, ad
5-7 mm seorsum, copiosus anastomantes (e margine 2/3-4/5 versus costae). Sori
in venae lateralis elongate, venae interiora et venae exteriora areolae c.1.5 mm
latis formantes, ramis basalis acrocopicus et venulae anastomosis diplazioideus.
Indusiis dilutis brunneis, integris, persistens.
Rhizome short, erect, slender. Stipe 14-15 cm long, pale brown, glabrous,
fallen scales at base. Scales on stipes (?). Lamina simple, elliptical, to about 40
cm long, 7.5 – 10.6 cm wide, narrowed toward acuminate apex, narrowed
gradually to the base which is suddenly contracted from a wide of 1 cm or less,
base subequally cuneate, margin entire. Texture thin, subcartaceous pale green on
upper surface when dry, paler beneath. Rachis distinct on both surface; veins in
small group at an angle about 65-70º to the costa, the group about 5-7 mm apart,
each vein group forked at the costa, lower branch forked 5-7 times, lateral
members of the vein group copiously anastomousing (2/3-4/5 way from margin to
costa). Sori elongate along lateral veins, both outer and inner vein group forming
areoles about 1.5 mm wide, basal acroscopic branch and also those anastomousing
veinlets diplazioid. Indusia thin, pale brown, margin entire, persistent.
DISTRIBUTION. Borneo.
237
ECOLOGY. This species grows terrestrially on lowland mountain forest
at 150 m. Hitherto it has only been found in Bukit Raya.
NOTES. This species is very distinct among Malesian Diplazium in
characters combination as follow: Lamina simple elliptical with base subequaly
cuneate, veins copiously anastomousing to 4/5 way of margin to form areoles
about 1.5 mm wide on both ounter and inner veins group. This species similar to
simple frond of D. cordifolium in its copiously anasomousing veins so that the
specimen was formerly identified as D. cordifolium by de Joncheere.
Unfortunately, scales that is the important characteristic for identifying Diplazium
is fallen, not found on the specimens.
ETYMOLOGY. The species epithet is chosen in illustrating the large
simple frond.
39. Diplazium meijerii Praptosuwiryo, sp. nov. Plate 9.
TYPE: Sumatra: Central Sumatra, Payakumbuh, Northern slope of Mt.
Sago, 900-1200 m, 21 July 1955, W. Meijer 3772 (Holotype, BO).
Rhizome subprocumbent. Stipites ad 62 cm longis, prope basin 4 mm
crassi, in sicco dilutus brunneus, basi nigris squamis; squama nitidus nigris,
linearis lanceolatis, ad 9 mm longis 1 mm latis, margine regularis dentatus cum
filum niger spissescens, dentatis furcatis. Lamina ad 50 cm longis et latis,
bipinnata deltoideus; pinnae inferiora stipitae ad 6.5 mm latis, ad 9 cm seorsum,
ascendens, oblongis subtriangularis, ad 28 cm longis 12 cm latis, punnulae ad 13
jugatae; pinnulae inferiora stipitae at 2 mm longis, superiora adnatae vel sessile,
lanceolatis, ad 7.5 cm longis, 2 cm latis, basi truncates, apice acuminatus, margine
2/3-3/4 costam lobatae; lobus basalis acroscopicus multus deminutus ad 2 latis,
lobi magniora ad 6 mm latae, obiquus, basin latissimus, apice truncates vel to
roundrotundatus, integer; venae liberis, in lobo pinnatus, venulae ad 6-8-jugatae,
simplices vel furcatis. Textura tenuis papyraceous. Sori medius in 1/4-3/4
longitudine venularum.. Indusiis tenuis, paleis brunneis, persistens, margine
laciniatus.
238
Rhizome subcreeping (?), densely scales on younger part. Stipe to 62 cm
long, 4 mm thick, light brown when dry, black and scales at base; scales lineary
lanceolate, to ca. 9 mm long, 1 mm broad, black, shiny, margin toothed, with
regularly thickening black strands, teeth forked at tip. Lamina bipinate, deltoid,
ca. 50 cm long and broad; lower pinnae stalked to 6.5 cm long, to 9 cm apart,
ascending, oblong subtringular, to 28 cm long, 12 cm broad, pinnulae to 13 pairs;
lower pinnulae stalked to 2 mm long, upper adnate to sessile, lanceolate, to 7.5 cm
long, 2 cm broad, base truncate, apex sharply acuminate, margin lobed 2/3-3/4
way to costa; basal acroscopic lobes much reduce to 2 mm broad, larger lobes to 6
mm broad, oblique, broadest at base, apex truncate to rounded, entire; rachise not
gemmiferouse; veins free, pinnate in the lobes, veinlets 6-7 pairs, commonly 7
pairs, simple or once forked. Texture thinly papyraceous. Sori medial, cover ¼-
3/4 of veinlets length; indusia thin, pale brown, persistent, margin laciniate.
DISTRIBUTION. Sumatra.
ECOLOGY. Terrestrially, in primary forest at 900-1200 m.
NOTES. Diplazium meijeri may closely related to D. latisquamatum. D.
meijeri differs from D. latisquamatum in its black lineary toothed scales with
regular thickening black strands; sori medial. D. latisquamatum has dark brown
round-ovate entire scales with irregularly thickening black strand; sori basalis.
ETYMOLOGY. This species is named after W. Meijer, the collector of
this species.
40. Diplazium melanolepis Alderw.Diplazium melanolepis Alderw., Bull. Jard. Bot. Buit. II nr. XI. 8. 1913. -
--TYPE: Sumatra, G. Singgalang, C.G. Matthew 507 (holotype, BO!).
Rhizome (?). Scales on stipes dark brown, concolour, lanceolate, 5-12 mm
long, 1-2 mm broad, margin toothed, Stipe 56 cm long, 7 mm thick,blackish when
dry, scales throughout, dense toward base. Lamina bipinnate-bipinnatifid, pinnae
(?) pairs; lower pinnae stalked to 13 mm long, 7.5 cm apart, almost at right angle,
lanceolate, widest one third from the base, 34.5 cm long, 8 cm broad, pinnulae 20
pairs below pinnatifid apex of pinna; pinnulae adnate-sessile, oblong-lanceolate,
239
one pair basal reduced to 12 mm long, 3.5 mm broad, larger pinnulae to 5.2 cm
long, 0.9 cm broad, commonly less than 4 cm long, 0.8 cm broad, apex rounded-
acute, margin lobed to 1/3 way to costule; lobus 2-2.5 mm broad, oblique, ends
subtruncate, slightly toothed. Rachise and costae scales. Texture firm. Veins
forming angle 40-50º to costule, free, forked at costule, upper branch simplend
soriferous, usually diplazioid, lower brach simple or forked once-twice again,
basiscopic branch usually also soriferous, asplenoid. Sori from basal cover 2/3-3/4
of their length. Indusia dark brown, usually rolled back, persistent, margin entire.
DISTRIBUTION. Sumatra.
ECOLOGY. Occurs in light shade, on mountain forest at ca. 1900 m
above sea level.
SPECIMENS EXAMINED. --- SUMATRA: C.G. Matthew 507.
41. Diplazium moultonii (Copel.) Tagawa
Diplazium moultonii (Copel.) Tagawa, Acta Phyotax. Geobot. 25: 67.
1972. --- Athyrium moultonii Copel. Journ. Str. Br. Asiatic Soc. No. 63. 71.
1912; C.Chr. Gard. Bull. S.S. 7: 267. 1934.
Lamina tripinnatifid, (?) cm long, 25 cm broad; pinnae numerous, (?)
pairs. Pinnae alternate, about 3-4 cm apart, costa forming an angle about 70-80º
to rachis, stalked to about 8 mm long, oblong subtriangular, to about 28 cm long,
8 cm broad, pinnulae to 24 pairs below pinnatifid apex of pinna, gradually
decrease in size towards apex of pinna; pinnulae almost at right angle to costa,
shortly stalked to 33 mm long; lower pinnule oblong subtriangular, to about 6 cm
long, 1.6 cm wide, base subequally truncate, lower segmented, segments 1-4 pairs,
upper deeply lobed to 4/5 – 6/7 way to costulet (deeply lobed to within 1 mm of
costule), apex acuminate; segments adnate to sessile, forming an angle about 60º
to costule, oblong, to about 9 mm by 4 mm, apex rounded, crenulate. Veins
pinnate in each segment, free, to 4 pairs, veinlet simple in each crenation, distinct,
black when dry. Sori from basal costulet cover veinlets 1/3 of their length.
Indusia brown, thin, broad, persistent, margin tearing of.
DISTRIBUTION. Borneo.
240
ECOLOGY. Growing on shade parts of ridge mountain forest at 1100-
1850 m sea level.
SPECIEMNS EXAMINED: BORNEO: Kalimantan Timur: M Kato, M.
Okamoto & E.B. Walujo B-9806; M. Kato, M. Okamoto & E.B. Walujo B10018;
M. Kato, M. Okamoto & E.B. Walujo B10925; M. Kato, M. Okamoto & E.B.
Walujo B-9566.
42. Diplazium pallidum (Blume) Moore
Diplazium pallidum (Blume) Moore, Ind.: 333. 1861; Bedd., Hand. Ferns
Br. Ind: 175. 1883; Copel., Polypod. Philipp.: 73. 1905. Backer & Posth.,
Varenfl. Java: 129. 1939; Holttum, Rev. Fl. Malaya Appendix II : 637. 1966. –
Asplenium pallidum Blume, En. Pl. Jav.: 177. 1828. -- Diplazium montanum
Alderw., Bull. Jard. Bot. Buitenz. II Ser., 28: 19. 1918. – Athyrium montanum
(Alderw.) Holttum, Rev. Fl. Malaya 2: 555. 1966.
Key to the varieties
Terminal pinna deltoid and deeply lobed; upper base of lateral pinnae broadly truncate, lower basenarrowly rounded ……………………….…………………………………… var. pallidum
Terminal pinna conform to lateral or with one or two lobes; upper base of lateral pinnae rounded,lower base cuneate …………………………………………………………… var. montanum
a. var. pallidum
Rhizome erect, suberect. Stipe 17-57 cm long, black and scales densely at
base; scales dark brown, nearly black, concolours, 7-13 mm long, 1-1.8 mm wide.
Lamina nearly oblong, pinnae 11-27 pairs; rachis grooved on upper surface;
pinnae 7.8-23.5 cm long, 1.2-2.6 cm wide; lower pinnae on stalk up to 5 mm long,
upper pinnae sessile, apical lamina of the frond deltoid and deeply lobed; upper
pinnae with cuneate base, lower pinnae with base broadly truncate, lower base
narrowly rounded; margin of pinnae toothed, apex toothed acuminate;texture
subcoriaceous; veins free, forked near the costa, upper branch simple and
soriferous, lower branch forked again 1-3 times. Sori on acroscopic veinlet
241
simple, sometimes double, reaching from the costa almost to margin of pinnae.
Indusia firm, persistent, dark brown, margin entire.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical;
equatorial longitudinal view (excluding perine) plano-convex; equatorial view
tranverse view proximal face convex, distal face hemispherical; perinate. Size: E:
32.65(47.96)65.02±8.98; P: 17.08(27.09)34.12±5.12. Laesura: concealed by
perine. Perine: alate under LM, costate under SEM, irregular envelope separated
from exine surrounds the spore in irregular, wing-like costae, often anastomosing
to form loose reticulation, lacunae shallow irregular polygons 15-18 µm wide,
muri 0.8 – 5 µm wide, surface of muri smooth and lacunae smoothly granulate.
Exine: visible through perine, granulate under LM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped with an angle 105º, base flat, ends bluntly ridge
both inward and outward.
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 1151,
BO).
DISTRIBUTION. Malay Peninsula, Sumatra, Java.
ECOLOGY. Grows in shady forest in the hills and mountains, 1000 –1600 m.
SPECIMENS EXAMINED. --- JAVA: T.Ng. Praptosuwiryo 1151, 1237,
1335, 1377, 1484, 1505, 1630, 1739, 1759, 1764; W.S. Hoover, J.M. Hunter, H.
Wiriadinata, D.Girmansyah & A. Ruskandi ARs 29; A. Hidayat & H. Wiriadinata
AH 558. SUMATRA: T.Ng. Praptosuwiryo 2246.
b. var. montanum (Alderw.) Praprosuwiryo, com. nov.Diplazium montanum (Alderw.), Bull. Buitenz. II Ser. 28:19. 1918; Holtt.,
Gard. Bull. S.S. 11: 92. 1940. –Athyrium montanum (Alderw.) Holttum, Rev. Fl.
Malaya 2: 555. f. 327. 1966.
Pinnae to 16 pairs, 8.5-21 cm long, 1.3-2.3 cm wide; lower pinnae on stalk
2-4 mm long, upper base rounded, lower base cuneate; upper pinnae sessile with
242
broadly cuneate base; terminal pinane like others, occasionally with one-two
rounded lobes; margin of pinnae shortly toothed or almost entire.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng.Praptosuwiryo 1406, BO).
DISTRIBUTION. Sumatra, Java.
ECOLOGY. Terrestrial. Dense jungle, rich soil on rock, wet ground.
Elevation: 20 – 400 m.
SPECIMENS EXAMINED. --- SUMATRA: T.Ng. Praptosuwiryo 2244,
2537; J. v. Borssum W. 2787; J. v. Borssum W. 2334; J. v. Borssum W. 2338, ---
MALAY PENINSULA. Pahang: L.B. Molesworth – Allen 4247; M.R.
Henderson 18583; Haniff 4047; L. Wray 3617; King 10959; H.N. Ridley 14209;
Hullet s.n. (Aug. 1880) Selangor: H.N. Ridley s.n. (Dec. 1896). --- JAVA:
T.Ng.Praptosuwiryo 1406. --- BORNEO: T.Ng. Praptosuwiryo 2163, 2163b,
2163c.
NOTES. All specimens examined here are treated as D. montanum
v.A.v.R. by Holttum (1940). He stated that this species differs from D. pallidum
Bl. In having the apical pinna like others (in D. pallidum the apical lamina of the
frond is deltoid and deeply lobed), and in having the lower pinnae with much
narrower bases (in D. pallidum the lower pinnae have s broadly truncate upper
base). He added, based his field experience, that D. pallidum has fertile fronds
from young stage of development, whereas D. montanum does not produce fertile
fronds untuil the palants are full f\grown. D. pallidum occours in Sumatra on
higher parts of the mountains, matching Java specimens exactly; D. montanum
appears to occur at lower levels in Sumatra, thus falsifying its name. In Peninsula
D. pallidum does not occur. D. montanum occurs in forest at low and medium
elevations.
243
43. Diplazium parallelivenium Praptosuwiryo, sp. nov. Plate 10.
TYPE: Java, West Java, G. Halimun, Track Cikuda Paeh – Cikaniki, ca.
1300 m s.l., 25 February 2006, T.Ng. Praptosuwiryo 2338 (holotype, BO).
Rhizoma breve erectum. Stipitus 62 cm longis, 8 mm crassis in sicco
niger in basi, squamis sparsim penitus; squama bruneis nitidis linearis lanceolatis
ad 6-12 mm latae 0.5-1 latae, margine dentatis cum filum niger spissescens.
Lamina ad 94 cm longis et latis, bipinnata sudbeltoideus, pinnae 9-jugatae; pinnae
inferiora stipitae ad 3.2 cm longis, oblongis subtriangularis, ad 48 cm longis 29
latis, pinnulae 9-11-jugatae; pinnulae inferiora stipitae ad 3.5 mm longis, pinnulae
superiora adnatae latisissimis cuneatis; pinnulae magniore ad 14.7 cm longis 3.2
cm latis, lanceolatis, basin truncatus, margine 1/5-1/3 costam lobatae, plerumque
1/5-1/4 costam lobatae, apice attenuatis; lobi ad 9 mm latis truncates dentatis, lobi
basalis basiscopicus magissimathe; venae liberis, in lobo pinnatus, venae
principalis angulum fere 60º cum costa formantes, venulae 6-7-jugatae, simplices
vel furcatis in lobi basalis acroscopicus, venulae basasimus acroscopicus
nondilatatus. Sori costulis, in ½ vel minus longitudine venularum (2.5-7.5 mm
latis), basalis acroscopicus diplazioideus. Indusiis tenuis angustatus dilutus
brunneus, persistens, revolutus, laceratus.
Rhizome stout, short, erect, scales on younger part. Stipe 62 cm long, 8
mm thick when dry, black at base, sparsely scales throughout; scales dark brow,
lineary lanceolate, 6-12 mm long, 0.5-1 mm broad, margin toothed with
thickening black strand regularly. Lamina subdeltoid, ca. 94 cm long and broad,
bipinnate, pinnae 9 pairs; lower pinnae stalked to 3.2 cm long, oblong
subtriangular, to 48 cm long, 29 cm broad, pinnulae 9-11 pairs; lower pinnulae
stalked to 3.5 mm long, upper pinnulae adnate with broadly cuneate; larger
pinnulae to 14.7 cm long, 3.2 cm broad, lanceolate, base truncate, margin lobed
1/5-1/3 way to costa, commonly 1/5-1/4 way, apex attenuate; lobes to 9 mm wide,
basal basiscopic the largest, ends truncate, toothed; veins free, pinnate in the
lobes, midveins commonly forming angle about 60º to costa, veinlets 6-7 pairs,
simple or once forked on bosal acroscopic lobes, basal acroscopic ones not
dilated. Sori costular, cover ½ or less of veinlets length (2.5-7.5 mm), basal
244
acroscopic diplazioid.. Indusia very thin, narrow, light brown, persistent, rolled
back, margin lacerate.
DISTRIBUTION. Java.
ECOLOGY. This psecies is found in moist soil, red clay mixed with
humus rich soil, in shady place at about 1300 m.
NOTES. Diplazium parallelivenium closely related to D. vestitum. This
species differ from D. vestitum on the following characters combinations: Stipe
smooth, not muricate or spiny; pinnulae lanceolate; midvein commonly forming
angle 60º to costa; sori is not more than ½ of veinlets length; indusia rolled back,
lacerate. D. parallelivenium is also closely related to D. dolichosorum . The two
species share lineary black scales, pinnulae with truncate base and shallow
incision margin to form truncate lobes, and free simple veinlets. D.
parallelivenium differ from D. dolichosorum on the following character
combinations: lamina subdeltoid, pinnulae lanceolate with sharply acuminate or
attenuate apex, basal acroscopic veinlets parallel to the other veinlets; while D.
dolichosorum have lamina ovete, pinnulae oblong with acute or acuminate apex,
basal acroscopic veinlets curved.
ETYMOLOGY. The species epithet parallelifolium is used because basal
acroscopic veinlets are parallel to the others. It is a distinct character that
differentiate from other bipinnate West Malesian Diplazium species.
44. Diplazium petiolare C. PreslDiplazium petiolare C. Presl, Epimel. Bot.: 86. 1849.
Rhizome short, erect, densely scales on younger part.. Stipe slender, 41.5-
51 cm long, 2-2.5 mm diam. near base, light brown, black at scales base, upper
glabrescent; scales lineary triangular, concolour, dark brown, margin sharply
toothed. Lamina pinnate, subtriangular in outline, 34-45 cm long, 17.5-26 cm
broad, pinnae 15-17 pairs below pinnatifid apex of lamina; lower pinnae stalked
2-4 mm long, lanceolate, to 8.2-13 cm long, 2.1-3 cm broad above base, base
subequally cuneate, apex sharply acuminate, margin lobed to ¾ way to costa;
lobes about 5 mm wide above base, slightly oblique; rachise glabrescent; texture
herbaceous; light green above when dry, paler beneath, surface naked; veins free,
245
pinate, forming an angle about 60º to costa, veinlets 4- 7 pairs, simple, forming an
angle about 25º to costulets. Sori almost on all veinlet, sometimes the last pair no
sori, elongate from near base cover their length, basal veinlets acroscopic,
sometimes subbasal also; indusia brown, persistent, margin entire, opening when
mature.
CHROMOSOMES. 2n = 82 (Cytotype: TT993.3, BOHB).
DISTRIBUTION. Sumatra, Borneo.
ECOLOGY. Occurring on shade places in the forest. Elevation: ca. 450 –
-1000 m sea level.
SPECIMENS EXAMINED. --- SUMATRA. Sumatra Utara: K. Iwatsuki,
G. Murata, J. Dransfield & D. Saerudin S-77. Aceh: S-772; K. Iwatsuki,
G.Muarata, J.Dransfield & D. Saerudin S-688.
45. Diplazium poiense C.Chr. in C.Chr. & HolttumDiplazium poiense C.Chr. in C.Chr. & Holttum, Gard. Bull. S.S. 7: 269.
1934. --- TYPE: R.E. Holltum 25380 (Holotype, SING!), Borneo, Tenompok,
Mt. Kinabalu, 1430 m.
Rhizome erect, suberect. Stipe dark brown, 35-40 cm long, ca. 4 mm
thick, scales at base; scales brown, shining, lineary lanceolate, 6-8 mm long, 0.25
- 1 mm broad, concolours, margin entire, without thickening black strand.
Lamina pinnate-bipinnate, oblong lanceolate, 38.5 cm long, 17 cm broad, pinnae
27 pairs below pinnatifid apex of lamina; lower painnae stalked to 4 mm long,
subfalcate, to 8.5 cm long, 1.8 cm broad, lower base narrowly cuneate, upper base
truncate, apex acuminate, margin lobed ½-5/6 way to costa; lobus oblong,
oblique, basal acroscopic the largest, 3-5 mm wide, subtruncate-acute, slightly
toothed; veins free, pinnate in the lobus, rather hardly observed, veinlets to 10
pairs, simple. Sori impressed, elongate ½-2/3 from margin; indusia broad,
persistent, brown, concolour, margin entire, opening when mature.
DISTRIBUTION. Borneo.
ECOLOGY. Growing on shade part of mountain forest at ca. 500-1700 m
sea level.
246
SPECIMENS EXAMINED. --- BORNEO: Kalimantan Timur: M Kato,
M. Okamoto, K.Ueda, & E.B. Walujo B-7408; M Kato, M. Okamoto, K. Ueda &
E.B. Walujo B-7403; J. & M.S. Clemens 28412; M. Kato, M. Okamoto & E.
B.Walujo B-10908; M. Kato, M. Okamoto & E. B.Walujo B-10922; M. Kato, M.
Okamoto & E. B.Walujo B-9804; M. Kato & H. Wiriadinata B-4811.
46. Diplazium polypodioides BlumeDiplazium polypodioides Blume, En. Pl. Jav.: 194. 1828; Holtt., Gard.
Bull. S.S. 11: 93. 1940. Diplazium asperum Blume, En. Pl. Jav.: 195. 1828.
Diplazium ophiodontum (Copel.) C.Chr., Index Filicum Suppl. III, 75.
1934. Athyrium ophiodontum Copel.; Phil. Journ. Sci., 46: 214. 1931. --- TYPE:
M. Ramos Bur. Sci. No. 77188 (isotype, SING!), Penablanca Cagayan Province,
Luzon.
Diplazium polypodioides Bl., Pl. Jav.: 194. 1928; Backer & Posth.,
Varenfl. Java: 130. 1939; Holtt., Gard. Bull. S.S. 11: 93. 1940.
D. asperum Bl., Pl. Jav. 195: 1828. –Athyrium asperum (Bl.) Milde, Bot.
Zeit.: 353. 1870; Holtt., Rev. Fl. Malaya 2: 571. f. 320. 1966.
Athyrium blumei (Bergsmann) Copel., Fern Fl. Philip. 3: 396. 1960.
Rhizome stout, erect, to about 10 cm diam., 15 cm cm hight, scales like on
stipes. Stipe stout, about 9 mm dian near base, 36=110 cm long, densely scaly
near base, surface prickly; scales narrowly linier, 20-50 by 1-2 mm, thick, dark
brown, black and toothed at margin. Lamina bipinnate-tripinnatifid, ovate-
subdeltoid in outline, 50-156 cm long, to about 70 cm wide; pinnae 21-60 cm
long, 11-17 cm wide, narrowed oblong subtriangular, shortly stalked, with 13-26
pairs of pinnules below deltoid lobed apex of pinna; largest pinnuler 6.5-16.5 by
1.2-2.1 cm, oblong with acuminate apex, base subtruncate, sessile or lowest
shortly stalked, margin deeply lobed near costules; lobes oblong to
subquadrangular, oblique, rounded to obtuse at apex, sharply serrate, 6-15 mm
long, 2-5 mm wide; texture papyraceous, dark green, paler beneath; veins pinnate
in each lobe, veinlets 5-11 pairs, usually simple, sometimes once forked. Sori
247
usually on almost all veinlets, from base half-way or more close to margin, basal
veinlet sometimes diplazioid; indusia thin, persistent.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropalar; polar ouline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view plano-convex to concave-convex; equatorial
tranverse view, proximal view planar to concave, distal view convex; perinate.
Size: E: 34.36(39.63)43.17± 2.33, P: 16.31(22.92)27.43±2.97. Laesura: concealed
by wing- like perine. Perine: costate-alate, loosely reticulate; irregular envelope
separated from exine surround the spore with costae forming loose irregular
reticulation or wing-like muri, reticulation often incomplete, lacunae large
irregular polygons, 3-9 µm across; wing-like muri project 1-7µm, terminating
margins entire; surface or perine smooth under SEM.
ANATOMY. Vascular bundle form uninterrupted U-shaped with an angle
105º, base and angle ridges on both inward and outward directions, end
develops large ridges and grooves becomes somewhat W-shape.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 1647, BO).
DISTRIBUTION. Assam, Sri Langka, SW China, Taiwan, throughout
Malesia.
ECOLOGY. On moist ground, humus rich soil in shadowed places in
mountain slopes forest elevation 200-1900 m sea level.
VERNACULAR NAMES. Pakis benik (Bengkulu).
USES. Young fronds of this species are sold in the local marked of
Sumatra as vegetables.
SPECIMENS EXAMINED --- JAVA: Alston 12671; Backer 8898;
9185, 9812 bit, 9847, 10242, 10412, 10494, 12024, 13204, 12906, 14158, 16043,
16231, 18447, 18621; Backer & Posthumus 44, 91, 122, 207, 258, 541;
Bakhuizen v/d Brink 88, 2011, 2530, 3252, 5677; Beguin 12, s.n. (5-1927);
Clason s.n. (27-5-1931), F101, F97, K173; Blume s.n., Clason-Laarman K142,
K162; Dillerogn 707; Donk 355, 357, 602; Groenhart s.n. (Malang); Hallier s.n.
(23-11-1894), 682, 685; Heurn s.n. (7-1935); Kern 8459; Koorders 15442B,
22881B, 41113B, 41164B, 43738B; Laarman K169; Posthumus 1460a, 1666;
Popta 84, 94; D.R. Pleyte 32; T.Ng. Praptosuwiryo 533, 536, 549, 605, 609, 703,
248
711, 714, 755, 763, 1391, 1581, 1582, 1583, 1599, 1605, 1606, 1612, 1613, 1617,
1623, 1640, 1651, 1339; Raciborski s.n. (Telaga Warna), s.n. (Kota Batu), s.n.
(Mt. Guntur); Rumka s.n. (18-11-1829); Sapiin 2677; Scheffer s.n. (3-9-1870);
Schiffner s.n. (27-2-1894), s.n. (5-1-1894); Ciets 12; Vriese s.n. (5-5-1926);
Winckel 1248B, 1281B, 1413B, 1422B; Wisse 627. --- SUMATRA: P. Buwalda
6340; Dr. P. Buwalda 6340. --- MALAY PENINSULA: Md Nur SFN 32937,
King F90, King 10849; Ridley 14206; A.G. Piggott 26; Md Nur SFN 11515; R.E.
Holttum SFN 9770. SUMATRA: T.Ng. Praptosuwiryo 2242.
NOTES. I justify Athyrium ophiodontum Copel. as new synonym of D.
polypodioides Bl. The size of the middle pinnae, 11 x 3 cm, is slightly larger than
that recorded for D. polypodioides in Holttum (1940). Otherwise there is no
differences between the Philippine specimen and the materials from BO and
SING.
47. Diplazium porphyrorachis (Baker) Diels
Diplazium porphyrorachis (Baker) Diels, Nat. Pfl. 1 (4): 225. 1899; Price,
Gard. Bull. S.S. 36 (1): 28. 1983. Asplenium porphyrorachis Baker, J. Bot. 17:
40. 1879; Icon. Pl. 17, t.1650. 1886; C.Chr., Gard. Bull. S.S. 7: 279, p.p. 1934;
Athyrium porphyrorachis (Baker) Copel. Philip. J. Sci. 3C: 300. 1908;
Polypodium subserratum Hook. Sp. Fil. 4: 202. 1863 (non Diplazium subserratum
(Blume) Moore, 1861). Type: Borneo, prob. W. Sarawak, A.R. Wallace s.n. 1855
(K).
Rhizome erect or short-creeping, bearing weary roots, clothed with
lanceolate-acuminate, dark-brown scales at the extremity. Stipes ca. tufted, firm,
slender, 5-50 cm long, scaly at the base, black-hairy or scaly above, the hairs or
scales deciduous; scales 2 mm long, 0.25 mm broad, dark brow, stiff, deciduous.
Lamina pinnatifid, 16-25 cm long, 3.5-5.5 cm broad, lanceolate, apex lobed or
toothed, the lower 2/3–6/7 deeply pinnatifid into many or numerous, close,
spreading, subfalcate, linear oblong, blunt, slightly crenate, serrate or toothed
segments 8-15 mm broad, which have not seldom the sides entire and only the
apex serrate; lower segments gradually growing smaller, the 1-4 lowest free and
deflexed; texture subcoriaceous; rachise and under surface slightly covered with
249
scattered scales; veins distinct, mostly once forked. Sori oblique, parallel,
reaching ca. the edge; indusia dark brown, persistent, margin entire, opening when
mature.
SPORES. Monolete, bilaterally summetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) plano-convex, ; equatorial
tranverse view, proximal, proximal face planar, distal convex; perinate. E:
39.89(46.39)50.98±3.53, P: 21.74(27.57)30.49±2.87. Laesura: concealed by
perine. Perine: alate to costate-alate, alae sometimes anastomosing forming loose
reticulation; irregular envelope, separated from exine, surrounds the spore in loose
anastomosing wing, forming a very loose reticulation, reticulation often
incomplete; lacunae very irregular polygons, 10-16 m across; muri thin, wing
like muri project 1.5-9.0 m; terminating margin ciliate or papillate; surface of
perine holed, holes irregular and formed by fallen papillae
CHROMOSOMES. 2n = 164 (Cytotype: T.Ng. Praptosuwiryo 1885,
BO).
DISTRIBUTION. Malay Peninsula, Borneo, Celebes (v.A.v.R., 1908)
ECOLOGY. Terrestrial on on small river banks in the forest shade at 200-
1270 m sea level.
VERNACULAR MANES. Paku tuot (Dayak)
SPECIMENS EXAMINED. --- BORNEO. East Kalimantan: Tabang (32
km point), West Kutai, M. Kato & H. Wiriadianata B-6159; G. Kongkat-G.
Kongbotak, Alt. 350-200, Masahiro Kato & Harry Wiriadianata B-5176;
Masahiro Kato & Harry Wiriadianata B-5259; Masahiro Kato & Harry
Wiriadianata B-6348; Masahiro Kato & Harry Wiriadianata B-6879; Masahiro
Kato & Harry Wiriadianata B-4733; Masahiro Kato & Harry Wiriadianata B-
6214; T.Ng. Praptosuwiryo 1845, 1883, 1884, 1885, 1890, 1925, 1936.
NOTES. As noted by Price (1983), D. porphyrorachis is very closely
related with D. lomariaceum and some specimens shown sign of apparent gene
interchange. He shows the different of the two species in the key with characters
combination as follow. D. lomariaceum has scales blackish that numerously bear
on stipe, rachise, costae and vein beneath, and also present above where costae
250
and meet rachis; frond narrowing gradually towards base, drying dark greenish
brown. While D. porphyrorachis reveals scales brown that present on stipe,
rachis, and costae beneath; frond not or only moderately narrowed downwards,
drying dull grayish brown.
48. Diplazium prescottianum (Wall.) Moore
Diplazium prescottianum (Wall. ex Hook) Moore, Ind. Fil.: 156, 334.
1859; Holttum, Gard. Bull. S.S. 11: 94. 1940.; Tagawa & K. Iwats., Southeast As.
St. 5: 103. 1967; Acta Phytotax. Geobot. 23: 56. 1968. –Asplenium
prescottianum Wall. ex Hook., Sp. Fil. 3: 251. 1860. –Diplazium sylvaticum var.
prescottianum (Wall. ex Hook.) Bedd., Handb.: 178. 1883. –Athyrium
prescottianum (Wall. ex Hook.) Holttum, Rev. Fl. Malaya 2: 557. 1955.
Rhizome short, erect, bearing thick black roots; scales dark brown, about
15 by 1 mm, entire. Stipe about 80 cm long, stramineous with dark brownish
base, glabrous. Frond oblong, about 50 by 20 cm, pinnate; lateral pinnae stalked,
ascending, falcate, caudateiattenuate at apex, rounded to moderately auricled at
acroscopic and cuneate at basiscopic bases, up to 15 by 1.7 cm, shallowly crenate,
terminal pinna not distinct with deeply lobed base, the lower ones like the upper
lateral pinnae; thin chartaceous, dark green with paler lower surface, glabrous;
costa raised below, veins twice or thrice forked, all free, glabrous. Sori elongate
along basal anterior veinlets or rarely along basal posterior ones; indusia thin but
firm, persistent.
SPORES. Monolete, bilaterraly symmetrical (made asymmetric by
perine), heteropolar, polar outline (excluding perine) transversely elliptical;
equatorial longitudinal view plano-convex; equatorial transverse view, proximal
face planar, distal face concave; perinate. E: 32.72(39.11)51.44±5.16, P:
20.23(24.79)33.56±3.63. Laesura: concealed by perine. Perine: alate, costate-
alate, loosely reticulate; irregular envelope separated from exine surrounds the
spore in continuous anastomosing wings, forming a loose reticulation; lacunae
large, irregular polygons 5-8 µm; thin wing-like muri project 4-9 µm , terminating
margins ciliate or echinate. Exine: visible through perine, smooth-granulate under
SEM.
DISTRIBUTION. Malaya Peninsula.
251
ECOLOGY. On humus-rich mountain slopes in dense forest at 1500 –
2000 m sea level.
SPECIMENS EXAMINED. MALAY PENINSULA. Perak: Kinta s.n
(January 1885), C. Curtis 3369; Larut, C.G. Mathew s.n. (1905). Singapore:
Ridley s.n. (1892); J.Sinchlair 9376. Selangor: H.L. Hume 8997; H.L. Hume
9347.
NOTES. No. Coll. J. Sinclair 9376 (12 March 1958) showed a few
differences to the description of Holttum (1940), basal veinlets of some pinnate
vein group area sometimes uniting. But J. Sinclair stated in his notes that this
specimen matches the type specimen of D. Prescottianum (Aspidium
Prescottianum Wall.). For clarification, therefore, the type specimen should be
seen and compared to the Sinclair’s specimen.
49. Diplazium procumbens Holttum
Diplazium procumbens Holttum, Gard. Bull. S.S. 11: 95. 1940. Fig. 4. --
- TYPE: Malay Peninsula, Pahang, Frasser’s, R.E. Holttum SFN 36503 (holotype,
SING!).
Rhizome creeping, 8-13 mm diam. Stipe 30-47 cm long, glabrescent, pale
brown, black at base. Lamina deltoid, bipinnate, 41.5 – 73 cm long, (?) wide;
largest pinnae 25-27 cm long, 12-23 wide; pinnules sessile or the shortly stalked
to 2 mm long, larger ones to 7.5 by 2-2.7 cm, base truncate, apex acuminate,
margin lobed ¾ way or more to costae; large lobes 6-7 mm wide, oblong, almost
at right angle to costae, apex rounded, slightly toothed toward apex; veins 6-7
pairs, all forked in larger lobes, usually simple in smaller ones; costae bearing
scattered small narrow brown toothed scales beneath. Sori spreading from cortule
1/3-1/2 of the length of the veins; indusia very thin, pale brown.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical;
equatorial longitudinal view (excluding perine) plano-convex; equatorial
transverse view, proximal face convex, distal face hemispherical; perinate. Size:
E: 43.03(53.78)63.20±4.72, P: 29.52(33.22)37.31±2.39. Laesura: concealed by
perine. Perine: alate under LM, costate under SEM, irregular envelope separated
252
from exine surrounds the spore in irregular, wing like costae, often anastomosing
to form a loose reticalution, lacunae shallow irregular polygons 8-22 µm, muri
1.6-6.6 µm, surface of muri and lacunae smooth.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 1173,
BO).
ANATOMY. Transverse section of stipe near lamina: Vascular strand
form an uninterrupted V-shaped or U-shaped with an angle 100º, end ridge in both
inward and outward directions do form an angle 120º.
DISTRIBUTION. Malay Peninsula, Sumatra, Java.
ECOLOGY. Terrestrial. Rock soil, on open wet ground by stream.
SPECIMENS EXAMINED. --- MALAY PENINSULA. Pahang: R.E.
Holttum 11478; 3800 ft., R.E. Holttum 21645; R.E. Holttum SFN 36503, C.H.
Addison 37201; I.H. Burkill & R.E. Holttum 8844; E.J.H. Corner s.n. (12 Agust.
1937); R.E. Holttum 21645. Perak: King 7894. --- JAVA: T.Ng. Praptosuwiryo
1142, 1238, 1455, 1344, 1348, 1388, 1392, 1520, 1522, 1565, 1693, 1699, 1816,
1714, 1715, 1734, 1772, 1795, 1821. --- SUMATRA: T.Ng. Praptosuwiryo ...
50. Diplazium profluens Praptosuwiryo, sp. nov. Plate 11.
TYPE: Java, West Java, G. Halimun, Jalur Macan, ca. 950 m, 1 Oct.
2003, T.Ng. Praptosuwiryo 1820 (holotype, BO).
Rhizoma breve erectum, Stipites 68 cm longae, 7.5 mm crassi fere basi, in
sicco nigellus penitus, basi squamus caducus; squamis rotundatis vel ovatis, ad ca.
5 mm longae, 4 mm latis, brunneis nitidis, integris. Lamina bipinnatae; pinnae
inferiora stipitae ad 3.7 cm longae, lanceolatae, 51 cm longae, 23 cm latae,
pinnulae ad 14-jugatae infra apice pinnatifidus; pinnulae inferiora stipites ad 2.5
mm longae, 2.5-4 cm spatium, superiora adnatus vel sessilis; pinnulae
lanceolatae, basiscopicae plerumque maximae, 11-13.5 cm longae, 3.5-3.9 cm
latae, basi subaeque truncatae, apice acuminate, e margine 7/8-8/9 costam lobatae;
lobi oblongi, basiscopicae plerumque longiora, oblique, apice rotundatae vel
acutae, margine apicem leviter dentate. Venulae libere, pinnatae in lobo, venulae
principalis inferiora angulum 70º vel patentisimus cum costa formantes, venulae
7-9 –jugatae, prerumque 7–jugatae, plerumque furcatis, 1-2 –jugatae superiora
253
simpliciter. Sori e basi profluens versus venulis acroscopicae ramis 1/3-1/2
longitudine venularum sedentes, sorus infimus acroscopicus diplazioideus, sorus
medius acroscopicus intermidum diplazioideus. Indusiis brunneus, concolorous,
persistens, integer.
Rhizome stout, erect. Stipe 68 cm long, 7.5 mm thick at base, blackish
throughout when dry, fallen scales at base; scales on stipes fallen without leaving
spines or protuberances on stipes surface, rounded, blackish, margin entire.
Lamina bipinnate; lower pinnae stalked to 3.7 cm long, lanceolate, widest at 1/3
part from base, 51 cm long, 23 cm broad, pinnulae 14 pairs below deeply
pinnatifid apex of pinnae; lower pinnulae stalked to 2.5 mm long, 2.5-4 cm apart,
upper adnate to subsessile with broadly cuneate base; pinnulae lanceolate,
basiscopic ones usually larger, 11.1-13.5 cm long, 3.6-3.9 cm broad, base
subequally truncate, apex acuminate, margin lobed to within 2-3 of costa (or 7/8-
8/9 way to costa); lobus oblong, basiscopic ones usually longer, end oblique,
rounded-acute, slightly toothed; veins free, pinnate in the lobes, lower main vein
forming angle 70° to almost at right angle, veinlet 7-9 pairs, commonly 7 pairs,
veinlets usually once forked, except those 1-2 pairs upper part (simple). Sori from
basal running to acroscopic branch of veinlets, covers 1/3-1/2 way of veinlets
length, diplazoid on basal acroscopic veinlets, sometimes others middle
acroscopic veinlets also diplazioid. Indusia dark brown, concolours, persistent,
margin entire.
PARATYPES. JAVA. West Java: Mt. Halimun, Cikaniki Forest, Jalan
Macan, 1 October 2003, T.Ng. Praptosuwiryo 1818; track Cikaniki-Cikuda Paeh,
near Sungai Cikuda Paeh, ca. 950 m, 30 September 2003, T.Ng. Praptosuwiryo
1798.
CHROMOSOMES. 2n = 164 (T.Ng. Praptosuwiryo 1798, BO).
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline elliptical, sides convex; equatorial longitudinal
view concave-convex to plano-convex; equatorial transverse view, proximal face
planar to concave, distal face hemispherical; perinate. E: 36.44(44.06)63.70±8.92;
P: 23.89(29.04)43.07±5.73. Laesure: concealed by perine ridge. Perine: micro
costate, costae broken, reticulate irregularly, densely echinate under SEM, echinae
254
single elements irregularly apart and projecting 1.1-2.1 µm. Exine: often visible
through perine, smooth.
DISTRIBUTION. Java.
ECOLOGY. Diplazium profluens usually grows in moist humus-rich soil
at shady places of small river banks.
NOTES. D. profluens is closely related to D. latisquamatum and shared
characters as follow, scales on stipes round or ovate, entire, and nearly black;
sori basalis and occupying only lower half or less of the veins, margin of pinnulae
lobed to within 2-3 of costa. It is different from D. latisquamatum on veinlets 7-9
pairs, middle acroscopic veinlets also diplazioid, margin of pinnulae lobed to
within 2-3 mm of costae (7/8-8/9) way to costa).
This species is also similar to D. kunstlerii, but the two species is
differentiated on their pinnulae incision. Pinnulae D. kunstlerii is usually lobed
2/3 way to costa, while pinnulae of D. profluens lobed 7/8-8/9 way to costa.
51. Diplazium riparium Holttum
Diplazium riparium Holttum, Gard. Bull. S.S. 11: 97. f.5. 1940; Tagawa
& K. Iwatsuki., Fl. Thailand 3: 454. 1988. --- Athyrium riparium (Holttum)
Holttum, Rev. Fl. Malaya 2: 554. f. 326. 1966. --- TYPE: Malay Peninsula,
Selangor, Semenyih, H.L. Hume 8186 (holotype, SING!).
Rhizome short, erect. Stipe 25-52 long, brown when dry, black and scales
at base; scales 4-8.5 mm long, 0.5-1 mm wide, dark brown or nearly black, margin
entire. Occasionally with glandular cells. Lamina pinnate, pinnae 2-3 pairs,
terminal one like the others; argest pinnae oblong, 25-5 cm long, 6 cm wide,
shortly stalked to 4 mm long or sessile, base cuneate, margin entire, suddenly
narrowed near apex, apex caudate; texture chartaceous; rachis without buds; costa
rounded beneath, glabrous, grooved on upper surface; veins at angle about 60º to
costa, in small group, each group of 3 veins, middle vein forked 1-3 times, the
outer of vein group uniting each other near margin of pinnae or 1/5 or less length
from margin. Sori elongate along 2-4 to each vein-group, those on the outer veins
of the group extending almost from the costa to margin, rest shorter, acroscopic
255
outer veins usually diplazioid; indusia not so thin, pale brown, fragile, margin
entire.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
continued V-shaped with an angle 65º, ends simple, not forming a ridge.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 2147, BO),
123 (Cytotype: T.Ng. Praptosuwiryo 1847, BO).
DISTRIBUTION. Java, Sumatra, Malay Peninsula, Borneo.
ECOLOGY. Occurring on in jungle, rock soil, usually in wet places and
often on stream banks at elevation 60– 1250 m sea level.
USE. Dayak people use the young frond as vegetable after cooking.
SPECIMENS EXAMINED. --- SUMATRA: T.Ng. Praptosuwiryo 2003a,
2003b, 2007, 2038, 2089, --- BORNEO. Kalimantan Timur: M. Kato, M.
Okamoto, K. Ueda, D. Darnaedi & E.B. Walujo B-8126; B-8235; T.Ng.
Praptosuwiryo 1838, 1847, 1894, 1916, 1940, 1952a, 2108, 2123a, 2123c, 2145,
2147b, 2191c, 2193b, 2193c, 2200. --- BORNEO: Kostermans 10448; M. Kato,
M. Okamoto, K. Ueda, D. Darnaedi & E.B. Walujo B-8126, B-8235; M. Kato, M.
Okamoto & E.B. Walujo B-10070. MALAY PENINSULA. Perak: King 7894;
L. Wray, 3590, 3650. Johor: R. E. Holttum S.F.N. 18301. Kelantan: Md Nur s.n.
(14 February 1924). Melaka: H.N. Ridley 1589; H.N. Ridley s.n. (Dec.1899).
Trengganu: R.E. Holttum SFN 15323.
52. Diplazium silvaticum (Bory) Sw.
Diplazium silvaticum (Bory) Sw. Syn. Fil. : 92. 1806; Blume, En. Pl. Jav.:
1828; Bedd., Hand. Ferns Br. Ind.: 177. 1883; Copel., Polypod. Philipp.: 73.
1905; Backer & Posth., Varenfl. Java: 130. 1939; Holttum, Gard. Bull. S.S. 11:
99. 1940. -- Callipteris silvatica Bory in Belanger, Voy. Bot. 1: 282. 1804 (n.v.).
Key to the varieties
Pinnae up to 13 pairs; lower pinnae lanceolate, 7-15 cm long,
upper base subruncate .............................................................................. var. silvaticum
Pinnae up to 6 pairs; lower pinnae elliptical, 3 -5.5 cm long,
upper base subrounded ................................................................... var. pinnae-ellipticum
256
a. Var. silvaticum (Bory) Sw.
Rhizome erect, sub erect, scales. Stipe to about 40 cm long, scales; scales
very dark, nearly black, to about 10 mm long, 1.5 mm wide, margin toothed, teeth
mostly forked. Lamina pinnate, 18.5-63 cm long, 14.5-38 cm wide near base;
pinnae numerous, 8-13 pairs, upper ones sessile, lower ones shortly stalked to 3
mm long; rachis gemmiferous; pinnae lanceolate, 6.7-15 by 1.6-3 cm, lower ones
with upper base subtruncate, lower base cuneate, upper pinnae with broadly
cuneate base to truncate, more or less auricle above but not below, apex
acuminate, margin lobed to ¼ way to costa, texture thin, lobes oblique, apex
truncate, toothed, veins in pinnate group with 3-4 pairs of lateral veinlets. Sori
elongate from near base of vein diplazioid, other simple, basiscopic veins of
basiscopic basal lobes sometimes also diplazioid; indusia brown, broad, persistent,
margin entire, opening when mature.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar, polar outline (excluding perine) transversely elliptical;
equatorial longitudinal view (excluding perine) concave-convex; equatorial
transverse view, proximal face concave, distal face hemispherical; perinate. E:
31.59(41.31)49.30±3.82, P: 18.64(23.56)26.41±2.46. Laesura: concealed by
perine. Perine: alate under LM, costate-alate under SEM, irregular envelope
separated from exine; alae form loose reticulation; lacunae irregular polygons c.
6.6-20.0 µm across, interior of lacunae showing micro irregular
reticulate/fenestreta under SEM; muri very thin, wing-like, terminating margin
irregularly echinate. Exine: smooth-granulate under LM.
ANATOMY. Vascular bundle form an uninterrupted U-shaped with and
angle about 120º, base flat inward and outward directions, angle not forming
ridge, end almost simple or slightly ridges.
CHROMOSOMES. 2n=164 (Cytotype: T.Ng. Praptosuwiryo 1300,
BOHB).
DISTRIBUTION. Africa to Samoa, throughout Malesia.
ECOLOGY. Occurs in forest in the lowlands and hills, and to moderate
altitudes in the mountains. 15 – 1350 m.
257
SPECIMEN EXAMINED. --- JAVA: Backer 2241, 18023, 18445;
Backer & Posthumus 256, 651; Donk s.n.; Pleyte 29; Posthumus 1736, 1498,
4027, 3542; Raciborski s.n.; v.A.v.R. s.n.; T.Ng. Praptosuwiryo 1300, 1301. ---
MALAY PENINSULA: M.R. Henderson 22422; Ridley 5808; R.E. Holttum
24704; H.N. Ridley s.n. (Dec. 1891); Curtis s.n. (September 1890); M.R.
Henderson 19498. --- SUMATRA: C.H. Lamoureux 5580.
b. Var. pinnae-ellipticum Praptosuwiryo, var. nov.
Stipe 7.5 – 16.0 cm long, 3 mm diam. at base, pale green, densely shortly
hairy throughout, scales toward base; scales subulate to linearly triangular, to 7
mm long, 1 mm wide, brown, margin toothed, teeth mostly forked. Lamina
pinnate, ovate-subtriangular in outline, light green, 16.5-25 cm long, 8.5-12.5 cm
broad, 2-6 pairs below pinnatifid deltoid apex of lamina; lower pinnae stalked to
2.5 mm long, elliptical, 3.3 -5.4 cm long, 2.0-2.7 cm wide, upper base
subrounded, lower base cuneate; upper pinnae upper adnate-sessile, oblong
sobtriangular, upper base truncate, lower base broadly cuneate; margin lobed 1/6-
1/4 way to costa, apex acute; lobus 5-10 mm wide at base, ends truncate to
subemarginate, almost entire; veins free, distinct on both surface, pinnate in each
lobe, 3-4 pairs, simple.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 2001a,
BOHB).
DISTRIBUTION. Sumatra
ECOLOGY. On secondary forest, rather wet humus rich soil in shadowed
places at 15-70 m sea level.
SPECIMENS EXAMINED. --- SUMATRA: Jambi: T.Ng. Praptosuwiryo
2001a, 2001b, 2001d, 2001i, 2016a, 2019a, 2019b, 2019c, 2019d, 2020a, 2020c,
2028c., 2028e, 2083d.
258
53. Diplazium simplicivenium Holttum
Diplazium simplicivenium Holttum, Gard. Bull. S.S. 11: 100, 101. f.6.
1940. –Athyrium simplicivenium (Holtt.) Holttum, Fl. Malaya 2: 573. f. 340.
1966. ---TYPE: Malay Peninsula, Pahang, Frasser’s, R.E. Holttum 36507
(holotype, SING!).
Rhizome short, erect. Stipe 100 cm or more long, 15 mm thick, rough
near base, fallen scales at base; scales 2 cm long or more, 1-2 mm wide near
base, dark brown, margin toothed, teeth simple (not forked). Lamina bipinnate,
55-150 cm long, 50-100 cm wide, or larger; pinnae c.15 cm apart, oblique; 45-66
cm long, 17-24 cm wide, free pinnules 15-20 pairs below the lobed apex of pinna;
largest pinnule stalked to 1.5 mm, 7.4-12 by 1.8-2.1 cm, upper pinnules sessile,
base truncate, apex acuminate, margin lobed 1/5-¼ way towards costa; lobes 6-7
mm wide, slightly oblique; texture thin; veins distinct on both surface, upper
surface hairy, pinnate, veinlets 4-5 pairs, all simple. Sori covered the basal 1/3-¾
or more of veinlets; indusia thin, firm, persistent.
SPORE. Monolete, bilaterally symmetrical, heteropolar; polar outline
elliptical, sides convex; equatorial longitudinal view plano-convex to concave-
convex to plano-convex; equatorial transverse view, proximal face planar to
concave, distal face hemispherical; perinate. E: 43.99(50.61)54.60±3.36, P:
23.31(29.42) 33.78±2.79. Laesure: concealed by perine ridge. Perine: alate to
costae alate, irregular envelope separated from exine surrounds the spore in
irregular and in continuous or discontinuous anastomosing wing, forming a loose
reticulation; lacunae large irregular polygons, occasionally with granulate deposite
within; thin wingilike muri projected 3-8µm, terminating margins are almost
entire. Exine: smooth regulate under SEM.
ANATOMY. Vascular bundle form an uninterrupted U-shaped with an
angle about 90º; base flat inward, slightly ridges outward, angle ridges, end
distinctly ridges to form an angle 100º.
CHROMOSOMES. 2n = 123 (Cytotype: T.Ng. Praptosuwiryo 1386,
BO).
DISTRIBUTION. Malay Peninsula, Sumatra, Java, Thailand.
259
ECOLOGY. Found in shady mountain valleys and occasionally in
exposed places by roadside drains. 400 – 1600 m.
SPECIMENS EXAMINED. --- JAVA: Koorders 23538B; T.Ng.
Praptosuwiryo 1136, 1139, 1351, 1341, 1342, 1348, 1349, 1351, 1371, 1449,
1519, 1521, 1523, 1526, 1530, 1761, . --- SUMATRA: T.Ng. Praptosuwiryo
2512; 2535. --- MALAY PENINSULA. Pahang: C.E. Carrr 22388; R.E.
Holttum 36507 (27-12-1939) (HOLOTYPE), R.E. Holtum 21636. Perak: King
2214; King 2346. Selangor: H.L. Hume 9110.
54. Diplazium sorzogonense C. Presl.
Diplazium sorzogenense (C. Presl.) C. Presl., Tent. Pterid.: 114. 1836;
Copel., Polypod. Philipp.: 74. 1905; Bedd. Hand. Ferns Br. Ind.: 181. 1883. ---
Asplenium sorzogonense C. Presl., Rel. Haenk. 1: 45. 1825 (n.v.). --Athyrium
sorzogenense (C. Presl.) Milde, Bot. Zeit. 1870: 354; Holttum, Rev. Fl. Malaya 2:
552. f. 325. 1966.
Rhizome erect, suberect. Stipe 20-65 cm long, dark brown, black at base,
scales throughout; scales narrowly linier, 20 by 1-3 mm, concolour, dark brown,
margin entire with black strand. Lamina pinnate, oblong lanceolate in outline
with nacute apex, 40-85 cm long, 20-45 wide; rachis grooved shallowly above,
fibrillose; pinnae commonly 20 pairs below pinatified apex of frond, basal ones or
more pairs, reflexed, upper ones adnate or decurrent to form distinct apical
portion, middle pinnae shortly stalked, patent to ascending, narrowly lanceolate,
13-22 cm long, 2-2.4 cm wide above base, apex acuminate, subtruncate to cordate
at base, deeply lobed to ¾-6/7 way to costa; fibrillose beneath, shallowly grooved
above; lobes almost at right angle to costa, oblong, oblique, rounded at apex,
toothed; texture papyraceous; veins pinnate, black, distinct beneath, veinlets
commonly 6-8 pairs, mostly simple. Sori elongate along veinlets, nearly from
main veins ½-2/3 length of veinlets, sometimes almost marginal, diplazioid only
on acroscopic basal veinlets; indusia thin, brown, persistent, crescentic.
SPORES. Monolete, bilaterally summetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) plano-convex; equatorial
260
transverse view, proximal face planar to convex; distal face hemispherical;
perinate. E: 27.80(33.03)37.90±3.91, P: 13.95(20.92)26.39±3.82. Laesura:
concealetd by perine. Perine: alate, costate-alate, irregular envelope separated
from exine; lacunae irregular polygons c. 8.3-16.6 m accross; terminating
margins of costae or wing-like muri entire. Exine: smooth under SEM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped, with an angle 110º, base flat both inward and
outward, angles without ridge, ends almost simple.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 1803, BO).
DISTRIBUTION. Peninsular Thailand, throughout Malesia.
ECOLOGY. On slopes in lowland and lower montane , hill forest.
Locally common on shady ridges between streams in forest in the hills and to
moderate altitudes in the mountains. 10-2150 m.
SPECIMENS EXAMINED. --- MALAY PENINSULA: R.E. Holttum
s.n. (31-12-1939); R.E. Holttum s.n. (20-3-1929); Md. Nur 11198; B.M. Allen
2036; Ridley 13432; M.R.Henderson 18612; Kiah SFN 35012; R.E. Holttum
SFN 20747. --- BORNEO: M. Kato, G. Murata & Y.P. Mogea B-3810. ---
SUMATRA: W.J.J.O. de Wilde and B.E.E. de Wilde-Duyfjes 19408; Dransfield
3274; Harry Wiriadinata 1540; H. Surbeck 151; Dr. J. Winkler 2121; C.G.G.J.
van Steenis 710; T.Ng. Praptosuwiryo 2092a, 2092c, 2092d, 2092f, 2092h, 2092i.
--- JAVA: T.Ng. Praptosuwiryo 1713, 1720, 1725, 1737, 1743, 1744, 1745, 1746,
1805, 1753, 1758, 1760, 1803, 1802. BORNEO: T.Ng. Praptosuwiryo 2092a,
2092b, 2092c, 2092d, 2092h, 2092i.
261
55. Diplazium speciosum Blume
Diplazium speciosum Blume, En. Pl. Jav.: 193. 1828; Copel., Polypod.
Philipp.: 74. 1905; Backer & Posth., Varenfl. Java: 127. 1939; Holtt., Gard.
Bull. S.S. 11: 103 1940. –Athyrium speciosum (Blume) Milde, Bot. Zeit.: 354.
1870; Holttum, Rev. Fl. Malaya 2: 559. 1966.
Key to the varieties
Lobes moderately toothed, end rounded. Veinlets 7-9 pairs, simple ….... var. speciosum
Lobes strongly toothed, end mostly acute. Veinlets 10-16 pairs, mostly forked ... var. major
a. var. speciosumRhizome (?). Stipe (?) cm long, distinctly grooved above, scales towards
base, dark brown, black at base. Scales on stipe shining brown, narrowly linier,
about 17 by 1-1.5 mm, without thickening black strand at margin, margin toothed.
Lamina about 85 cm long, 36 cm wide, pinnate, oblong lanceolate, pinnae 15-24
pairs; rachis grooved; pinnae spreading, ascending oblong lanceolate, upper one
adnate to decurrent to form indistinct apex of frond, middle pinnae shortly stalked,
to 22 cm long, 4 cm wide above base, baee truncate, apex acuminate, margin
deeply lobed 5/6 way to costa; costa distinctly grooved above with distinct ridge;
lobes oblong, to 21 mm long, 6 mm wide, end rounded. Margin toothed; texture
herbaceous, but firm; veins in each lobes pinnate, distinct beneath, veinlets 7-9
pairs, simple. Sori not impressed, elongate nearly from the costule cover 2/3
length of veinlets, acroscopic basal ones diplazioid. Indusia not so thin, brown,
persistent, margin entire.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex-straight; equatorial longitudinal view (excluding perine) concave-convex;
equatorial tranverse view, proximal face convex, distal face biconvex; perinate.
E: 29.98(39.29)47.25±5.53; P:17.37(21.69)25.83±2.56. Laesura: concealed by
perine ridge. Perine: alate, costate-alate, irregular envelope separated from exine;
alae occasionally form loose retication; lacunae irregular polygons c. 6.6-10.8 µm
262
across, interior of lacunae smooth under SEM; terminating margins of wing-like
muri entire. Exine: rugulate, rugulae large, shallow, irregular anastomousing
under SEM after separating from perine.
CHROMOSOMES. 2n = 82 (T.Ng. Praptosuwiryo 1359, BO).
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped with an angle about 90º, base flat both inward
and outward, ends slightly ridge.
DISTRIBUTION. Malay Peninsula, Sumatra, Java.
ECOLOGY. Shady jungle, especially in sheltered, moist depressions,
there typical and covering a good deal of the ground. Found in light or partial
shade on a few high mountain peaks. Elevation: 1500-2600 m.
SPECIMENS EXAMINED --- JAVA: Backer 14723, 15847; Lörzing
2595; Posthumus s.n. (Tangkuban Prau), 109; Raciborski s.n. (G. Salak); v.
Slooten 184. SUMATRA: T.Ng. Praptosuwiryo 2221, 2227, 2228, 2229.
b. var major (Bedd.) HolttumDiplazium speciosum var. major (Bedd.) Holttum, Gard. Bull. S.S. 11:
103. 1940. – D. sorzogenense v. major Bedd., Supl. 40. 1892.Lamina to 138 cm long, 30 cm wide; middle pinnae to 20-25.5 by 4-5.5
cm, base truncate, apex acuminate, margin deeply lobed to 9/10 way to costa;
costa bearing scattered brown small linier subtriangular scales with margin
toothed benath; lobes various in length, to 7 mm wide above base, end mostly
acute, strongly toothed; veinlets 10-16 pairs, mostly once forked, occasionally
twice.
CHROMOSOMES. 2n = 82.
DISTRIBUTION. Malay Peninsula, Borneo, Sumatra, Java
ECOLOGY.
SPECIMENS EXAMINED. MALAY PENINSULA: A.G. Piggott 1087;
Scortechini s.n. --- BORNEO: M Kato, M Okamoto & E.B. Walujo B-9801; B-
9797; B-9795; R.E. Holttum SFN 25444. SUMATRA: J.A. Lörzing 14901;
15540; J.A. Lörzing 16255; C.J. Brooks 350 S; J.A. Lörzing 15934; J.A. Lörzing
5987; H.A.B. Bünnenmeijer 2708; 5426; 5427a.
DISTRIBUTION. Java, Sumatra, Malaya, Borneo, Seram.
263
NOTES. This species is similar to D. sorzogonenense. As stated by
Holttum (1940) and Kato (1994), it differs from D. sorzogonense in the toothed
scales, non-fibrillose scales on the rachise, and non-impressed sori.
56. Diplazium spiniferum Alderw.
Diplazium spiniferum Alderw., Mal. Ferns Suppl.: 265. 1917; D.
muricatum (Copel.) C. Chr., Ind.Suppl. 27. 1913; Athyrium muricatum Copel.,
Philipp. Journ, VIIc, 61.
Rhizome short, erect. Stipe 65 cm long, 8 m diam, near base, scales,
spinuous, pale brown or rather stramineous, shining, black at base; scales rounded,
ca. to 4 mm long and broad, entire. Lamina bipinate, 95 cm long, 58 cm broad,
pinnated pinnae 5 pairs, pinnatifid pinnae 3 pairs, simple pinnae 8 pairs below
pinnatifid apex of lamina; lower pinnae oblong lanceolate, to 32 cm long, 9 cm
broad, free pinnulae to 9 pairs, commonly 8 pairs; pinnulae almost at right angle,
adnate-sessile, oblong, base broadly cuneate, apex acute-acuminate, margin
usually lobed less than 1/4 way to costule or almost entire; basal lobus the largest,
to 3 mm wide; rachis veins free, forming an angle about 45º to costule, veinlets to
4 pairs, commonly 3 pairs, simple, forming an angle about 10-15° to main veins.
Sori at middle veinlest covers ¼-1/3 of theirs length, basal acroscopic diplazioid;
indusia dark brown, narrow, margin torn, persistent.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excludng perine) transversely elliptical, side
convex; equatorial longitudinal view concave-convex to plano-convex; equatorial
transverse view, proximal face planar to convex, distal face hemispherical;
perinate. E: 31.96(39.52)47.77±4.18, P: 14.87(23.92)29.21±4.14. Laesura:
concealed by perine. Perine: micro-costate, densely echinate under SEM, costae
broken, densely micro reticulate, coralline; irregular envelope, separated from
exine, surrounds the spore in densely coralline with enchinae project 1.5-2.3 µm;
lacunae are very small, irregular polygons less than 0.5-1.5 µm across; heavily
fenestrate throughout under SEM, giving coralline appearance. Exine: visible
through perine.
264
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped, with an angle 90º, base flat inward and ridges
outward, angles distinctly ridges, ends ridges outward and forming an angle 130º.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 1896c,
BO).
DISTRIBUTION. Borneo.
ECOLOGY. Growing on humus rich soil or limestones area of mountain
forest at altitude 100-1300 m sea level.
SPECIMENS EXAMINED. --- BORNEO. East Kalimantan: M. Kato &
H. Wiriadinata No. B 4868; M. Kato & H. Wiriadinata No. B 5845; M. Kato &
H. Wiriadinata No. B 5847; M. Kato & H. Wiriadinata No. B 6428; M. Kato &
H. Wiriadinata No. B 4818; M. Kato, M. Okamoto, K. Ueda, D. Darnaedi &
E.B.Walujo No. B 8277; M. Kato, M. Okamoto & E.B. Walujo B-10735; M.
Kato, M. Okamoto & E.B. Walujo B-10848; M. Kato, M. Okamoto & E.B.
Walujo B-10800; Kunio Iwatsuki, M. Kato, Gen Murata & Y.P. Mogea B-3342;
Kunio Iwatsuki, M. Kato, Gen Muarata & Y.P. Mogea B-1078; M. Kato, M.
Okamoto & E.B. Walujo B- s.n. Kalimantan Selatan: M. Kato, Gen Murata &
Y.P.Mogea B-3731; M. Kato, Gen Murata & Y.P. Mogea B-3739; B-3891; T.Ng.
Praptosuwiryo 1878, 1879, 1880, 1945, 1955, 1900, 1878, 1879, 1880.
NOTES. Kato (1994) stated that in the large leaves the pinnules are lobed
more than ½ to costule.
57. Diplazium squarrasum K. Iwats. & M. Kato
Diplazium squarrasum K. Iwats. & M. Kato, Acta Phytotax. Geobot. 31
(4-6): 179. 1980.
Rhizome stout, short, erect, 1.8 c, thick when dry. Stipe 33.5-36.5 cm
long, 5 mm thick, deeply grooved above, dark brown, densely scales throughout;
scales ovate-lanceolate, 2.5-12 mm long, 0.25 – 2 mm broad, concolour, light
brown, margin sharply toothed, without thickening black strands. Lamina simply
pinnate, oblong, 72 cm long, 24 cm broad, lateral pinnae 17 pairs below deltoid
deeply lobed apex of lamina; lower pinnae shortly stalked to 1.5 mm long, upper
ones adnate-sessile, 4.2-7.2 cm apart, patent (at right angle to rachise), oblong,
265
one pair basal pinnae reduced to 7.7 cm long, 2.8 cm broad, larger ones 11.3-15.5
cm long, 2.7-3.3 cm broad, suddenty norrowed at 2/3 part of base, base shallowly
auricle, lower truncate, upper broadly cuneate, base of upper pinnae broadly
cuneate, margin entire, or slightly waved, apex acuminate; texture thin, but firm
(subherbaceous), lower surface scatterly bearing stelate trichome; upper surface
glabrous; rachise densely scales, scales like those on upper part of stipes; costa
raised below, densely minutely scales; veins anastomousing, veins group forming
angle 50-60° to costa, each vein group of 3 veins, middle veins forked 4-5 times
again, outer veins uniting with the nearest of outer veins of other veins group ¼-
1/3 way of costa (veins anastomousing 2/3-3/4 way to costa), forming areola
more or less 1.5 mm wide, inner veins also uniting with nearest veins. Sori
elongate along veinlets, cover ¼-4/5 of their length, opening towards both
acroscopic and basiscopic, sometimes diplazioid on basal acroscopic or on uniting
veinlets; indusia narrow, light brown, concolour, margin entire, persistent, firm,
opening when mature.
DISTRIBUTION. Borneo.
ECOLOGY. On humus-rich slopes near stream in evergreen forest, at ca.
1200 a.s.l.
SPECIMENS EXAMINED. --- BORNEO: M. Kato & H. Wiriadinata B-
4822, 5030.
58. Diplazium subalternisegmentum Praptosuwiryo, sp. nov.
TYPE: BORNEO: Mt. Kinabalu, Nr. Camp in mossy low jungle, ca.1500
m, 19 May 1933, J. & M. S. Clemens 33169 (holotype, BO).
Rhizoma breve erectum crassum. Stipites (?) longae, 7 mm crassi fere
basi, nigris, muricatus, basin squamis lineary lanceolatus 5-15 mm longae 0.5-1.5
mm latae denticulatus cum filum niger spissescens densus vestitae. Lamina
tripinata; pinnae stipitae ad 3.7 longis, subtriangular, pinnulae liberis ad 29-
jugatae; pinnae stipitae ad 3.7 cm longae, subtriangulare, libere pinnulae ad 29
jugatae; inferiorae pinnulae stipitae ad 5 mm longae, pinnatae (7 jugatae),
subtriangulare, 11 cm longis, 2.4 cm latis, apice acuminatus, libere segmentae 4-6
jugatae; segmentae subalternate, ad 6 mm seorsum, adnatus, subhastatus, ad 12
266
mm longis 5 mm latis ad basim, apice acutus, margine lobatae ad ½ costam
versus. Venae pinnata in segmento, venulae ad 7-jugatae, simpliciter vel
bifurcates in uno lobo. Sori e basi ½-2/3 venulae occupants, venulae infimus
acroscopicus interdum diplazioideus. Indusia brunneus, latae, margine integrae.
Rhizome short, erect, stout. Stipe (?) long, 7 mm diam. near base, dark
brow, black, muricate, densely black scales at base; scales on stipe black, lineary
lanceolate, 5-15 mm long, 0.5-1.5 mm broad, margin sharply toothed, with
thickening black strand. Lamina tripinnate. Pinnae on stalk to 3.7 cm long,
subtriangular in outline, free pinnulae to 29 pair; lower pinnulae stalked to 5 mm
long, pinnate (7 pairs), subtriangular, 11 cm long, 2.4 cm broad, apex acuminate,
free segments 4-6 pairs below pinnatifi apex of pinnulae. Segments subalternate,
to 6 mm apart, adnate, subhastate, 12 mm long, 5 mm broad at base, apex acute,
margin lobed to ½ way to costulet. Veins pinnate in the segments, veinlets to 7
pairs; veinlet simple to 2 forked in lobes. Sori from basal elongate, covers ½-2/3
of their length, sometimes diplazioid on basal acroscopic veins of segments.
Indusia brown, broad, margin entire.
DISTRIBUTION. Borneo.
ECOLOGY. Light shady in primary forest.
NOTES. D. subalternisegmentum resembles D. megasegmentum. These
two species are sharing characters combination as follow: (1) lamina tripinnatifid-
tripinnate; (2) subalternate segments at middle part of pinnulae; (3) sori often
cover ½ of veinlets length. D. subalternisegmentum.differs from D.
megasegmentum in characters: scales lineary lanceolate, margin sharply toothed,
with thickening black strand; pinnae more deeply lobed to froms 4-6 pairs of free
segments; indusia entire. Whereas, D. megasegementum has rounded scales with
entire margin, pinnae lobed to within 2 mm of costa without forming free
segments; indusi laciniate.
ETYMOLOGY. The species epithet subalternisegmentum.is used to depict
the pinnulae that having subalternate segments at subbasal part.
267
59. Diplazium subintegrum Holttum
Diplazium subintegrum Holttum, Gard. Bull. S.S. 9: 125. 1940; Holtt.,
Gard. Bul. S.S. 11: 104. 1940. --- TYPE: R.E. Holttum SFN 31350 (holotype,
SING!), S. Burong, Cameron Highland, Pahang, Malay Peninsula.
Rhizome short, erect. Stipe 59 cm long, 4 cm diam. when dry, scales;
scales brown, entire, to about 1.5 cm by 2 mm. Lamina simply pinnate, 53,8 by
32 cm; pinnae 10 pairs, the apical lamina narrowly deltoid, lobed at the base;
pinnae ca. 22 by 3 cm, lowest stalked to 1.5 cm, upper 2 pinnae only sessile;
bases almost equal, of lower pinnae narrower, cuneate, edges toothed, apices
acuminate; texture softly chartaceous, glabrous; veins forked near the costa, the
upper branch always simple, the lower branch forked 1-4 times. Sori on upper
branch from base 1/3-1/2 of way towards margin, diplazioid, usually also on one
branch only of lower vein, simple; indusia pale brown, thin, persistent.
DISTRIBUTION. Malay Peninsula, Sumatra.
ECOLOGY. Terrestrial. In jungle, on humus rich soils, in deep shade in
valley at first over 4000- 5000 ft. (1600-1800 m sea level)
SPECIMENS EXAMINED. --- SUMATRA: K. Iwatsuki, G. Murata, J.
Dransfield, & D. Saerudin S-985. --- MALAY PENINSULA: R.E.Holttum
23338; R.E. Holttum SFN 31350 (20 May 1936) (HOLOTYPE); Ridley 13969;
R.E. Holttum s.n. (13 may 1936), A.G. Piggott 2425; T. Shimizu, K. Iwatsuki, F.
Fukuda & M. Hutoh 13242; C.F. Symington 36053; H.D Harvey s.n. (1889); B.
Scortechini 391; J.H. Burkill s.n. (5 March 1924); I.H. Burkill & R.E.Holttum
8818; R.E. Holttum s.n. (14 September 1923), 4000 FT., R.E. Holttum 21539;
Ridley 70144.
268
60. Diplazium subpolypodioides (Alderw.) Alderw.Diplazium subpolypodioides (Alderw.) Alderw., Bull. Btz. 20: 11. 1915;
Malayan Handb. Suppl. 1: 273. 1916. --- D. asperum var. subpolypodioides
Alderw. Bull. Btz. 17: 8. 1914. TYPE: Sumatra, Bukit Barisan, C.G. Matthew
653 (Holotype, BO!).
Rhizome stout, erect. Stipe (?) long, end glabrescent, nearly glossy.
Lamina bipinnate; pinnae oblong lanceolate, shortly stalked to 9 mm long, 40 by
14 cm; rachis glabrous, nearly glossy; pinnules 17 pairs below acuminate deeply
pinnatified apex of pinnae, one basal pinnule a little reduce, acrossopic ones
smaller than basiscopic, upper pinnule adnate to sessile with broadly cuneate base,
lower pinnules shorthly stalked to 2 mm long; larger pinnules to 8.2 by 1.9 cm,
oblong subtriangular with sharply acuminate apex, base truncate, margin deeply
lodeb to within 0.5-1 of the costule; costule distinctly winged above, tomentose
beneath, liniery small scales; lobes (segments) almost at right angle to costae,
basal basiscopic lobes the largest, oblong, to 12 mm long, 3.5 mm wide, apex
obliquely rounded to acute with crenate tip, margin crenate ¼-2/3 way to
costulets; costulets distinctly winged above, tomentose beneath; veins pinnate in
each segment 6-7 pairs, veinlets on larger lobes forked 1-2 times in each
crenation, one pair end simple, veinlets on smaller mostly simple. Sori elongate
along veinlets a half or less length from costulets, on one pair basal ones usually
diplazioid; indusia pale brown, thin fragile, margin sharply toothed, crisped when
opening.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar; polar outline (excluding perine) transversely elliptical, sides
convex-straight; equatorial longitudinal view (excluding perine) concave-convex;
equatorial tranverse view, proximal face convex, distal face biconvex; perinate.
E: 32.49(40.29)47.17±4.08, P: 17.15(21.54)28.28±2.90. Laesura: concealed by
perine ridge. Perine: alate, costate-alate, irregular envelope, reparated from exine;
costae or alae sometimes form loose reticulation; lacunae irregular polygons c.
3.3—12.5 µm, interior of lacunae smooth under SEM; muri, wing-like, smooth,
terminal margin entire. Exine: smooth under SEM.
269
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an uninterrupted U-shaped with an angle 100º, angles ridges outward, end
ridges outward and shallowly grooved.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 2292, BO).
DISTRIBUTION. Java.
ECOLOGY. Growing on on humus rich soil rather opened places of
secondary mountain forest.
SPECIMENS EXAMINED. --- JAVA: van Steenis 10979; T.Ng.
Praptosuwiryo 2292.
61. Diplazium subserratum (Blume) Moore
Diplazium subserratum (Blume) Moore. Index 338. 1862; Backer &
Posth., Varenfl. Java: 126. 1939; Holttum, Gard. Bull. S.S. 11: 104. 1940. ---
Asplenium subserratum Blume, En. Pl. Jav. 1828. TYPE: C.L. Blume s.n.
(holotype, L!), G. Parang, Java.
Rhizome slender, erect, suberect. Stipes slender, clustered, up to 22 cm
long, dark brown, blackish toward base, glabrescent, minutely scaly at base, more
dense when young, slightly grooved above; scales oblong subtriangular, about 1
mm long, less than 1 mm wide at base, apex acuminate, margin shortly toothed.
Lamina simple, lanceolate up to 39.5 cm long, 3.7 cm wide at middle, narrowed
gradually toward attenuate apex; margin entire or irregularly undulate in lower
part, toothed towards apex; herbaceous shoftly papyraceous; midrib grooved
above, prominent beneath, scatteredly dark brown oblong subtriangular scales 1
mm long or less; veins free, in small group at angle about 60 to midrib, up to 6
mm apart; veins group forked at midrib, upper branch simple, usually soriferous,
lower brach forked 2-4 times, sometimes soriferous also. Sori on acroscopic
veinlets diplazioid in various length, others simple, shorter; indusia very firm,
brown when dry, margin entire, rolled back when old, persistent.
SPORES. Monolete, bilaterally symmetrical, heteropolar; polar outline
elliptical, sides convex; equatorial longitudinal view concave-convex to plano-
270
convex; equatorial transverse view, proximal face planar to concave, distal face
hemispherical; perinate. Size: E:27.88(41.19)45.49±5.03, P: 16.54
(27.13)20±5.03. Laesure: concealed by perine ridge. Perine: costate-alate,
irregular envelope, separated from exine; costae form irregular loose reticulation;
lacunae irregular polygons c.to 23 µm across; interior of lacunae and costae
smooth. Exine: visible throught perine, smoothly granulate under LM.
ANATOMY. Transverse section of stipe near lamina: Vascular bundle
form an interrupted U-shape that formed by two ovale leaf traces, with an angle
about 110º.
CHROMOSOMES. 2n = 82 (T.Ng. Praptosuwiryo 1463, BO), 123 (T.Ng.
Praptosuwiryo 1072, BO), 164 (T.Ng. Praptosuwiryo 1380, BO).
DISTRIBUTION. Sumatra, Malaya, Borneo, Java.
ECOLOGY. On moist mountain slopes by stream in evergreen jungle at
about 700- m.
SPECIMENS EXAMINED. --- MALAY PENINSULA: H.N. Ridley s.n.
(1895); H.N. Ridley s.n. (May 1891); A.G. Piggott 1704. --- BORNEO: M. Kato,
G. Murata & Y.P. Mogea B-3804; M. Kato, G. Murata & Y.P. Mogea B-3506; J.
& M.S. Clemens 20406; M Kato, G. Murata & Y.P. Mogea B-3804; B-3506; J.
& M.S. Clemens 20406. --- SUMATRA: T.Ng. Praptosuwiryo 2538, 2538b,
2538c, 2538e. --- JAVA: T.Ng. Praptosuwiryo 1311, 1370, 1372, 1376, 1378,
1379, 1380, 1381, 1439, 1441, 1458, 1459, 1462, 1463, 1512, 1705, 1706, 1765,
2278, 2283, 2286, 2287, 2289, 2290, 2294, 2295, 2296, 2299, 2301.
NOTE: Alderwerlt (1920) recognized two forms united by intermediates.
Forma tytica with fronds bluntly crenulate to toothed at or towards the apex and
distributed in Java and Malacca. The second form, viz. Forma lobata, revealed
fronds coarsely toothed or even lobed throughout and found in Sumatra (Deli,
Sibolangit, W. Docters van Leeuwen No. 161). The intermediates forms is showed
J.A. Lörzing No. 5043 from Sumatra (Deli, Bandarbaru) is intermediate between
both forms.
271
62. Diplazium subvirescens Praptosuwiryo, sp. nov. Plate 13 & 14.
TYPE: Java. West Java, Mt. Gede, Cibodas Forest, behind Cibodas
Botanic Garden, ca. 1450 m, 28 March 2002, T.Ng. Praptosuwiryo 1178
(holotype, BO).
Rhizoma procumbent, c.1.5 cm. diametro, apice dense squamatus. Stipites
20-36 longi, 3-6 cassi fere basi, in vivo viridis, basi squamis lineari-lanceolatis, 5-
17 mm longis, 0.5-1.25 mm latis sparse vestitae. Lamina deltoideus, 49-54 cm
longae et latae, bipinnata vel tripinnata, pinnae ad 9-jugatae. Pinnae basin
maximae, oblongis-subtringularis, c.28-30 cm longae, in stipitibus ad 2.7 cm
longis stantes, pinnulae ad 10-jugatae. Pinnulae subalternatis; pinnulae maximae
7-9.2 cm longae, 2.6-3.2 cm latae, oblong subtriangular, in stipitatae ad 3.5 mm
longis, basi subcordatae vel truncatae, apice acuminatae dentatae, cetera margine
lobatae 2/3 costam versus; lobi oblongis, ad 6 mm lati, margine serratis apice
versus, apice subtruncati-rotundatis. Rachis et costae supra sulcata, infra sparse
squamulosus; venulae perspicuus in pagina inferiore et superiore, in uno lobo ad
6-jugatae, simpliciter vel furcatae, in venulis infimus basiscopicus potius dilatatio.
Textura tenua subcoriacea, colorae supra aeroginosus. Sori submedianus vel
medianus venularum occupantes, plerumque submedianus, sori infimus
acroscopicus obliquus, diplazioideus; indusia angusta, tenuis, laciniatus, ante
sporangiis maturis aperiens.
Rhizome long creeping, blackish, bearing weary black roots, apex densely
scales. Stipe 20-36 cm long, green when living, black at base, 3-6 mm diam near
scaly base, densely clothed with dark brown scales at base, upward scattered
scales, grooved on upper surface, scales liniery lanceolate, 5-17 mm long, 0.5-
1.25 mm broad at base, dark brown or blackish, margin toothed almost regular
with thickening black strand, teeth simple or sometimes forked near tips. Lamina
bipinnate to tripinnate, deltoid in outline with acuminate deeply lobed apex frond,
to 49-51 cm long, 49-54 cm wide, pinnae 8-9 pairs, basal pinnae the largest;
pinnae stalked to 2.7 cm long, oblong subtriangular with acuminate deeply lobed
apex, basal pinnae the largest, 28-30 cm long, pinnules subalternate, numerous, to
10 pairs; pinnules on stalk to 3.5 mm long, upper ones sessile, oblong
subtriangular, larger pinnules 7-9.2 cm long, 2.6-3.2 cm wide, base truncate-
272
subcordate, margin deeply lobed to 2/3 way to costule, apex serrate acuminate,
lobus numerous, to 12 pairs; lobes oblong with rounded apex, basal or subbasal
lobe the largest, to 6 mm wide, margin sharply serrate toward apex. Rachis
scattered minutely pale brown scales beneath, grooved on upper surface; costa
minutely scales towards base, shallowly grooved on upper surface; costules raised
beneath, glabrous, shallowly grooved on upper surface; veins prominent on both
surface, raised above, pinnate to 6 pairs, simple or once forked; basal basiscopic
vein rather dilated. Texture firm-herbaceous, deep green above when living. Sori
linier on medial or sub-medial veinlets, usually on basal-subbasal veins, obliquely
diplazioid on basal acroscopic veins. Indusia thin, margin laciniate, irregularly
opening at maturity.
PARATYPE. JAVA. West Java: G. Gede, Cibodas Forest, behind
Cibodas Botanic Garden, ca. 1450 m, 7 August 2001, T.Ng. Praptosuwiryo 1012,
1013; 28 March 2002, T.Ng. Praptosuwiryo 1177 ( BO).
ANATOMY. Vascular bundle form an uninterrupted U-shape with an
angle about 90º, end ridges outward to form an angle about 130º.
CHROMOSOMES. 2n=123 (Cytotype: T.Ng. Praptosuwiryo 1177, BO)
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropalar; polar outline (excluding perine) transversely elliptical, sides
convex; equatorial longitudinal view (excluding perine) plano convex, equatorial
transverse view, proximal face planar, distal view convex; perinate. Laesura:
concealed by wing-like alae spinous coralline perine. Perine: alate to costate
alate, alae or costae forming irregular reticulation, reticulation often incomplete;
lacunae with echinae and coralline within, wing-like muri project 3-11 µm,
terminating margins pappilate or echinate; pappilae project c.0.7-1.5 µm; surface
of perine echinae – coralline with densely irregular hole, windows-like. Exine:
not visible through perine under SEM.
DISTRIBUTION. Java.
ECOLOGY. In shady places in the forest on humus rich soil at about
1500-1600 m a.s.l.
NOTES. D. subvirescens has affinity to D. virescens in the characters
combination as follows: rhizome long creeping, black with densely scales on
273
younger part; scales on stipes lineary lanceolate, polish dark brown with toothed
margin; lamina deltoid, deep green, firm herbaceous with veinlets prominent on
both surface; sori oblong to linear, medial to supramedial; indusia thin-
membranaceous, laciniate at margin, irregurarly broken at maturity.
D. subvirescens differs from D. virescens in the following characters.
Lamina to tripinnate so that giving its pinnulae more divided to form adnate to
sessile free segments, while the lamina of D. virescens are to bipinnate with
pinnulae lobed to; pinnulae of bipinnate fronds subtriangular, much wider (2.6-3.2
cm ), margin lobed 2/3-3/4 way to costa; veins sometimes once forked.
This fern is very distinct among the bipinnate-tripinnate fronds of
Diplazium in West Malesia. The long creeping rhizome and deep green of firm-
herbaceous lamina with veinlets very prominent above as well as the position of
the sori at the middle or sub-middle are useful features which distinguish this
species from the other bipinnate species of Diplazium.
ETYMOLOGY. The species epithet virescens is chosen in illustrating the
closely related of D. subvirescens to D. virescens.
63. Diplazium tomentosum Blume
Diplazium tomentosum Blume, En. Pl. Jav.: 192. 1828; Bedd., Hand Ferns
Br. Ind.: 179. 1883; Backer & Posth., Varenfl. Java: 127. 1939. Holttum, Gard.
Bull S.S. 11: 105. 1940; Tagawa & K. Iwats., Fl. Thailand 3: 457. 1988. –
Athyrium tomentosum (Blume) Milde, Bot. Zeit; 354. 1870; Holttum, Rev. Fl.
Malaya 2: 551. f. 324. 1966. --- TYPE: C.L. Blume s.n. (L!), Java.
Rhizome short, erect. Stipe 9-43 cm long, 3-4 mm diam, grooved on upper
surface, brown when dry, black toward base, scales throughout; scales 2.5-5 mm,
0.5 mm wide, dark brown, margin entire, sometimes with glandulars cell near
base or at tip. Lamina pinnate, oblong lanceolate to nearly oblong subtriangular in
outline, to 26 cm long, 14 cm wide, pinnae to 20 pairs below indistinct acuminate
apex of frond; basal pinnae the largest, deflexed, slightly elliptical, widest about
1/3 from base, base narrowed, apex acuminate; middle and upper pinnae narrower
with almost parallel side, nearly at right angle to rachis, base truncate, auricled
upper side; pinnae of larger frond deeply lobed a half a way or nearly to costa,
274
lobes to 3 mm wide above base, at angle about 60º to costa; rachis and costa
densely covered with multicellulars hairs with pale and dark brown cros-walls;
veins pinnate in each lobe, veinlets to 5 pairs. Sori on acroscopic basal veinlets
the largest, nearly from the basal veins almost reach margin, diplazioid; indusia
firm, dark brown, persistent, margin entire, nearly crescentic.
SPORES. Monolete, bilaterally summetrical, heteropolar; polar outline
elliptical, sides convex, equatorial longitudinal view plano-concex-concave
convex, equatorial transverse view, proximal face planar to concave, distal face
convex; perinate. E: 34.64(41.06)47.19±3.88, P: 23.14(26.77)28.67±2.13.
Laesura: concealed by perine. Perine: alate to costate-alate, loosely reticulate;
irregular envelope separated from exine surrounds the spore in continuous
anastomosing wings, forming a loose reticulation, occasionally wing-like alae
only form 3 reticulation surraound the spore; lacunae large irregular polygons,
papillae within; thin wing–like muri projected 3-12 µm, terminating margin are
often ciliate, ciliae projected to ca. 0.5 µm. Exine: visible through perine,
granulate under LM.
ANATOMY. Tranverse section of stipe near lamina: Vascular bundle
form uninterrupted V-shaped, with an angle 90º, end simple.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 2336b,
BO), 164 (T.Ng. Praptosuwiryo 2066, BO), 205 (Cytotype: T.Ng. Praptosuwiryo
1722, BO).
DISTRIBUTION. Burma (Tenessarian), Malaya, Vietnam, Sumatra, Java
(type), Malaya, Borneo, Seram, and Mindanao.
ECOLOGY. On mountain slopes in deep shade of dense forest at altitudes
800 (1000-5000 ft).
SPECIMENS EXAMINED. --- MALAY PENINSULA: H.N. Ridley
9866; A.G. Piggott 1904; R.E. Holttum 9537; Smith E.S. 818, Md. Nur 11038,
G.H. Addison (10-8-1939), M. Shah & noor M.S. 734; I.H. Burkill & R.E.
Holttum 1886; R.E. Holttum s.n. (26-12-1939); R.E. Holttum s.n.; G.A. Best
SFN 14134; H.N. Ridley s.n. (1891); H.N. Ridley 16213; A.G. Piggott 2430;
H.N Ridley s.n (Dec. 1895); B. Allan 1571; H.N. Ridley s.n. (1904); H.D.
275
Harvey s.n. (1889); 800-1000 ft, King 10767; H.N. Ridley 13438; H.N. Ridley
s.n. (21 June 1889); R.E. Holttum s.n.; E.J.H. Corner 30104. --- BORNEO: M.
Kato, M. Okamoto, K. ueda, D. Darnaedi & E.B. Walujo B-8316. --- JAVA:
Backer 10704; Donk 24; Hodorleg 248; Raciborski s.n. (G. Salak); T.Ng.
Praptosuwiryo 1236, 1721, 1722, 1723, 1724, 1747, 1762, 1769, 1748, 1804, 1807,
2336. --- SUMATRA: T.Ng. Praptosuwiryo 2030a, 2030b, 2047, 2540a, 2048a,
2048b, 2048c, 2048d, 2050a, 2050b, 2050c, 2050d, 2066, 2087.
NOTES. Holttum (1940) stated that this species is closely related to D.
velutinum Holtt., a species that has hitherto only been found in the moist shady
valleys at Cameron Highlands, Malay Peninsula. Diplazium tomentosum seems to
be related also D.crenatoserratum.
64. Diplazium tricholepis C. Chr.
Diplazium tricholepis C. Chr., Gard. Bull. S.S. 7: 270 t.57. 1934. ---
TYPE: Borneo, Mt. Kinabalu, Komborangah, R.E. Holttum 25522 (Holotype,
SING!).
Rhizome short, erect, densely scales on youger part Stipe 25 cm long,
densely scales at base; scales lineary lanceolate, 3-10 mm long, 0.5-1 mm broad,
light brown, concolour, without thickening black strands, margin sharply toothed,
teeth simple. Lamina broadly lanceolate, 70 cm long, 25 cm wide, bipinnatifid,
pinnae 17-18 pairs, one pair basal bending ward; pinnae subfalcate, sessile,
oblong subtriangular, base truncate, acuminate, incised to a wing 2 mm broad;
segments patent, triangular-oblong, 6-7 mm wide at base, lower side usually
somewhat longer than the upper ones (1.4 against 1.2 cm), obtuse, toothed in the
outer half or occasionally acute on the lower pinnae; rachise grooved on upper
surface, minutely scattered scales, non gemmiferous; veins dark, free, pinnate in
the lobes or segment, veins group forming angle 45º to midrib, veinlets 6-10 pairs,
forked once to twice or simple. Sori narrow, straight, extending from costa nearly
to margin, cover to 6-7 of veinlets length (to 4 mm long), acroscopic basal ones
diplazioid; indusia brown, concolour, persistent, margin entire.
DISTRIBUTION. Borneo
276
ECOLOGY. In the forest, near streams or in the gorge at 1000-2100 m
above sea level.
SPECIMENS EXAMINED. BORNEO: J & M.S.Clemens 34476; RE
Holttum 25522.
NOTES. This species is similar to D. christii, but scales are very different.
D.christii has entire scales while D. tricholepis with sharply toothed scales.
65. Diplazium umbrosum (Smith) Bedd.
Diplazium umbrosum (Smith) Bedd., Hand. Ferns Br. Ind.: 188. 1883;
Backer & Posth., Varenfl. Java: 130. 1939. – Asplenium umbrosum Smith, Jo. B.
4: 174. 1841 (n.v.); Hook., Syn. Fil.: 229. 1874.
Rhizome stout, short, erect, scales densely on younger part. Stipe 31-61
cm long, 5 mm thick, pale brown, glabrescent, black densely scales at base; scales
narrowly triangular, dark brown, margin toothed, teeth mostly simple, without
thickening black strand. Lamina bipinate-tripinnatified, to 60 cm long, about 50
cm wide; pinnae numerous, to 18 pairs, basal ones a little reduce; larger pinnae
oblong lanceolate with deeply pinnatified acuminate apex, to 21 by 6.8 cm,
shortly stalked to 6 mm long; pinnules numerous, 17 pairs, lower one shotly
stalked to 1 mm long, upper ones sessile; costa winged, distinctly grooved above;
larger pinnules 7.5 by 1.8 cm, oblong subtriangular, base subtruncate to
subcordate, apex toothed acuminate, margin deeply lobed to one mm or less from
costules, on lower portion almost reach the costules; costules winged; segments
mostly 3.5-5 mm wide, one basal basiscopic lobes the largest, to 10 by 6 mm,
apex blunt or truncate, margin crenate or lobed 1/3 way to costulet of segments;
texture herbaceous; vein pinnate in each segments 4-6 pairs, mostly forked once in
each crenations, on larger crenation pinnate 2-3 pairs of second veinlets. Sori
elongated from near costulet of segments or on middle veinlets; indusia pale
brown, attachment side darker, thin but firm, broad, persistent margin entire.
ANATOMY. Transverse section of stipe near lamina: vascular strand
form an uninterrupted U-shaped with an angle 90º, base almost flat, angle thicker
277
than base, end ridges outward to form an angle about 130º. Stomata: polocytic
and copolocytic.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 1348, BO).
DISTRIBUTION. Madeira, Canaries, Azores, Guinea Coast, Himalayas
to Ceylon, Java, Australia, Tasmania, and New Zealand.
ECOLOGY. On mountain slopes forest at elevation 1400-2000 m sea
level.
SPECIMENS EXAMINED. --- JAVA: Backer 14650; Bruggeman 625;
Donk 343, 675, 775; Meijer 1632; Paidan 8; Sapiin 2727, 2743; v. Slooten 199,
T.Ng. Praptosuwiryo 1332, 1352, 1453, 1496. --- SUMATRA: H. Surbeck 146.
66. Diplazium velutinum Holttum
Diplazium velutinum Holttum, Gard. Bull. S.S. 9: 126. 1937; Gard. Bull.
S.S. 11: 106. 1940. --- TYPE: R.E. Holttum 31221 (holotype, SING!), Telom
Road, Cameron Highland, Pahang, Malay Peninsula.
Rhizome short, erect. Stipe to 40 cm long (commonly about 20 cm),
clothed throughout with scattered multicellilar brown hairs, and towards the base
with narrow almost black entire scales; scales to about 1 cm long and barely 1 mm
wide at the base. Lamina bipinnate, to about 25 cm long, 15 cm wide, the rachis
and costa beneath densely clothed with brown hairs like those on the stipe, with a
few small narrow scales; free pinnae 12-15 pairs, the sub-basal largest, the upper
ones gradually reduced and grading into the l.obed apical lamina; largest pinnae
ca. 8 cm long, 3 cm wide (commnonly about 6 by 2 cm), subsessile, narrowed
from truncate base to acute or acuminate apex, pinnate towards the base only, for
the rest lobed almost to the costa, the pinnules or segments at right angles to the
pinna-rachis at the base, gradually more oblique towards the apex; free pinnules
few, the rest more or less broadly adnate to the pinna-rachis; largest pinnules
about 12 mm. long and 4 mm wide, base unequally cuneate, margin serrate or
lobed as much as half-way to the costule, apex rounded, texture very firm; veins
in pinnules or pinna-lobes 7-10-jugate, mostly forked. Sori on basal ½-2/3 of
278
veinlets; in the largest pinnules several diplazioid sori, in smaller pinnules only
one such sorus; indusia thin, broad, persistent.
DISTRIBUTION. Malaya
ECOLOGY. Occurring on moist shady valley at 1460-1520 m sea level.
SPECIMENS EXAMINED. --- MALAY PENINSULA: R.E. Holttum
SFN 31221 (HOLOTYPE), SFN 23428, s.n..
NOTES. This species is very closely related to D. tomentosum but much
more deeply dissected. Hitherto it has only been found in the moist shady valleys
at Cameron Highlands.
67. Diplazium vestitum C. Presl
Diplazium vestitum C. Presl, Epimel. Bot. 87. 1849; Copel., Polypod.
Philipp.: 75. 1905. --- Athyrium vestitum (C. Presl) Milde; Copel., Philipp. Journ.,
IIIc: 295. 1908. --- TYPE: Philippines, Samar, Cuming 336 (Isosyntype, US!).
Key to the Variety
Lower pinnulae on stalk to 1.5 mm long, to 11.5 cm long; veinlets 4-5 pairs, all simple.Sori elongated from base of costulets to near margin,diplazioid on basal acroscopic veinlets ……………………………. a. var. vestitum
Lower pinnulae on stalk to 3 mm long, to 13 cm long; veinlets 6-8 pairs, simple or once forked.Sori elongated from base of costulets covers 1/3-3/4 of their length,diplazioid on one basal veinlets …………………………………… b. var. borneense
a. Var. vestitum C. Presl
Rhizome stout, short, erect. Stipe 50 cm long, 7 mm thick near base, light
brown when dry, black towards base, muricate. Lamina bipinnate, ovate in
outline, to about 75 cm long, 60 cm broad, pinnate pinnae to 4 pairs below 7 pairs
pinnatifid ones, apex deltoid with deeply lobed; pinnae stalked to 7 cm long, sub
basal pinnae the largest, elliptical in outline, to 33 cm long, 13 cm broad, pinnule
6-10 pairs; lower pinnule subssile, oblong subtriangular with suddently narrowed
towards acuminate apex, to 11.5 cm long, base broadly cuneate, margin lobed to
1/5-1/3 way to costule; lobes to 5 mm wide, ends truncate, entire; veins pinnate in
the lobe, main vein forming an angle about 50-75º to costule; veinlets 4-5 pairs,
279
simple, forming an angle about 15-30º to main vein. Sori elangated along veinlets
from near main vein, not reaching the margin, cover to about ½ veinlets length,
basal acroscopic usually diplazioid; indusia narrow, dark brown, margin entire,
opening when mature, persistent.
SPORES. Monolete, bilaterally symmetrical (made asymmetric by
perine), heteropolar, polar outline (excluding perine) transversely elliptical, sides
vonvex; equatorial longitudinal view (excluding perine) concave-convex;
equatorial tranverse view, proximal face convex, distal face convec to
hemispherical; perinate. E: 28.21(32.58)35.33±2.26, P: 16.55(19.94)22.73±1.64.
Laesura: concelated by perine. Perine: costate-alate, almost no reticulation
irregular envelop, separated from exine, surrounds the spore with alae-costae
ridge (occasionally) projected 2-5 µm; surface of perine smooth. Exine: visible
through perine, smooth under SEM.
DISTRIBUTION. Borneo, Philippines.
ECOLOGY: primary forest, lowland, shadowed place, river bank.
SPECIMENS EXAMINED. --- BORNEO: M. Kato, H. Okada, R.
Imaichi, H. Tsukaya, Y. Mori, K. Aso & Dian Komara No 63; ,J. & M.S.
Clemens 27478; M. Kato, M. Okamoto, K. ueda, D. Darnaedi & E.B. Walujo B-
8246; M. Kato, M. Okamoto, K . Ueda, D. Darnaedi & E.. Walujo B-8290; M.
Kato & H. Wiriadinata B-6097; T.Ng.Praptosuwiryo 2162, 2159.
b. Var. borneense C.Chr. in C.Chr. & Holttum, Gard. Bull. S.S. 7:
273. 1934. --- TYPE: Borneo, Koung-Dallas, Mt. Kinabalu, 240 m, R.E.
Holttum 25134 (holotype, SING!); Dallas/Tenompok, 1400 m, Clemens 27734
(syntype, BM n.v.; isosyntype US n.v.); Kabayau/Kaung, 300 m, Clemens 27478
(syntype, BM n.v.; isosyntipe, K n.v.).
Rhizome (?). Stipe cm long, 8 mm thick pale, spinouse or muricate, black
at base, (?). Scales on stipes brown shining, lanceolate, margin toothed with
thickening black strands, teeth forked. Lamina bipinnate, (?) cm long, (?) cm
broad; pinnae stalked to 1.5 cm long, oblong subtriangular, to 52 cm long, 25 cm
broad, pinnulae 10-12 pairs below deltoid deeply lobed apex of pinnae; lower
pinnulae on stalk to 3 mm long, oblong subtriangular with acuminate apex, to 13
cm long, lower base subcordate, upper base subtruncate; upper pinnae adnate-
280
subsessile with subtruncate-broadly cuneate base; larger pinnulae to 13 cm long, 3
cm wide, lobed to 1/3 way to costule, basal acroscopic lobes the largest, to 8 mm
wide, ends truncate-blunt, slightly toothed to entire; texture thinly herbaceous;
rachis caffy like the stipe; veins pinnate in the lobus, veinlets 6-8 pairs, simple or
once forked on lower lobes. Sori elongate from basal veinlets covers 1/3-3/4 of
their length, one pair basal acroscopic diplazioid; indusia thin, margin entire,
persistent.
DISTRIBUTION. Borneo.
ECOLOGY. On wet slope in deep shade.
SPECIMENS EXAMINED. --- BORNEO: M. Kato & H. Wiriadinata B-
5170; 350-850 m, M. Kato & H. Wiriadinata B-5065; M. Kato & H. Wiriadinata
B-5065; R.E. Holttum SFN 25134; R.H.Endert 2892; R.E. Holttum SFN 25134;
Hallier 3227; M. Kato, M. Okamoto & E.B. Walujo B-9258; M . Kato, M.
Okamoto & E.B. Walujo B-10415; M. Kato, M.Okamoto & E.B. Walujo B-
11219; B-11365; Endert 3814; M. Kato, Gen Murata & Y.P. Mogea B-3839; M.
Kato, G. Murata & Y. P. Mogea B-3768; K. Iwatsuki, M. Kato, G. Murata & Y.P.
Mogea B-1082.
68. Diplazium wahauense Kato, Darnaedi et K. IwatsukiDiplazium wahauense Kato, Darnaedi et K. Iwatsuki, J. Fac. Sci. Univ.
Tokyo III, 15 : 101. 1991.
Rhizome short, erect. Stipe black, 4-10 cm long, 1 mm diam. near base.
Scales on stipe lanceolate, 4 mm long, 1 mm broad, margin entire. Lamina simple
imparipinnate, oblong, 10-13 cm long, 5.5-6.5 cm broad, pinnae 2-4 pairs; pinnae
adnate – sessile, lineary oblong, 3.7 – 8 cm long, 0.8 – 1.0 cm broad, glabrous,
base cuneate, margin crenulate, apex acuminate; terminal pinnae conform to the
lateral ones; rachis glabrous; veins free, 1-3 times forked. Sori elongate along
veinlets, basal acroscopic diplazioid; indusia broad, dark brown, margin subentire.
NOTES. As stated by Kato et al (1991), this species is closely related to,
and may have been derived from D riparium which occurs in riparian and dryland
forest in Borneo. These two species share black, somewhat crisped, entire scales,
blackish stipes, dark brown, naked rachises, and imparipinnate leaves with 3-4
281
pairs of entire lateral pinnae. Diplazium wahauense differs from D riparium
mainly in its narrow pinnae, which are characteristic of rheophytes.
CHROMOSOMES. 2n = 164.
DISTRIBUTION. Borneo.
ECOLOGY. A long river side, lowland rain forest. 50-100 m.
SPECIMENS EXAMINED. BORNEO: F.H. Endert 5900; M. Kato & H.
Wiriadinata B-6871; T.Ng.Praptosuwiryo 1972, 2126.
69. Diplazium xiphophyllum (Baker) C.Chr.
Diplazium xiphophyllum (Baker) C.Chr., Ind. Fil. 241. 1905; Holttum,
Gard. Bull. S.S. 11; 106. 1940. ---Diplazium fraxinifolium var. grossum C. Chr.,
Gard. Bull. S.S. 7: 274. 1934. –-- Athyrium xiphophyllum Baker, J. Bot. 1879:
40. Holttum, Rev. Fl. Malaya 2: 553. 1966. --- Anisogonium grossum Presl, Epim.
Bot. 93. 1851, nom. nud.
Rhizome short, erect. Stipe to 73 cm long, 8 mm thick,stramineous when
dry, scales toward base; scales subulate, to 6-17 mm long, 2-4 mm wide at base,
pale brown, margin entire with glandular cells. Lamina oblong, 55 cm long, 45
cm broad, pinnate, pinnae 6-9 pairs, terminal one like the rest; basal pinnae reduce
to 12,5 by 2.5 cm; lower pinnae shortly stalked, upper pinnae slightly adnate to
decurent, smaller, without buds on their axils; larger pinnae to 26-36 cm long, 4.1-
6.4 cm wide, elliptical, narrowed gradually to slightly unequal cuneate base,
ubrubtly to acuminate-caudate apex, margin entire to irregularly toothed
throughout; texture thin, drying light brownish; rachis pale, glabrous; costa pale,
glabrous, prominent beneath; veins in small group, at about 45-55° to costa,
commonly of one basal pairs and one central veins which is forked 1-3 times,
scarcely 4 times, sometimes acroscopic basal veins anastomousing with
basiscopic basal veins of the nearest vein group or with the nearest branch of
central vein near margin. Sori on basal pair of vein in each group elongated from
near costa to near margin, or other vein shorter, commonly 2-5 sori on each vein-
group, sori on acroscopic basal vein diplazioid, indusi persistent, pale brown, not
so thin, margin entire.
282
SPORES. Monolete, bilaterally symmetrical, heteropolar; polar outline
(excluding perine) elliptical, sides convex; equatorial longitudinal view plano-
convex to concave-convex; equatorial transverse view, proximal face planar-
convex, distal view convex; perinate. Laesura: concealed with wing-like perine.
E: 36.49(40.76)47.14±3.92, P: 19.16(24.88)30.65±3.32. Perine: alate to costate-
alate, free of reticulation or occasionally loosely reticulate; irregular envelope,
separated from exine, surround the spore in free wings and anastomosing costae-
ridge, reticulation irregular and often incomplete; lacunae shallow, irregular
polygons 2.5-8 µm across; wing-like muri thin, project 2.5-10 m, terminating
margins often papillate. Exine: visible through perine, smooth under LM.
CHROMOSOMES. 2n = 82 (Cytotype: T.Ng. Praptosuwiryo 1841 , BO),
164 (Cytotype: T.Ng. Praptosuwiryo 2040b, BO), 246 (Cytotype: T.Ng.
Praptosuwiryo 1190, BO).
DISTRIBUTION. Java, Sumatra, Borneo, Philippines, Moluccas.
ECOLOGY. In shadowed places of secondary and primary forest, at
ca.50-1100 m sea level.
SPECIMENS EXAMINED. ---BORNEO: M. Kato & H. Wiriadinata B-
6161,B-7019, B-4672; T.Ng. Praptosuwiryo 1841, 1842, 2146. ---JAVA: Donk
s.n. (1941), 22. --- MALAY PENINSULA: B. Holesworth-Allen 1436; H.L.
Hume 8777; Md Nur 34310; C.F. Symington 24200; H.N. Ridley 7833; King
2698; B. Burtis 1361; M.R. Henderson 10833; H.N. Ridley 14227. ---JAVA: W.S.
Hoover, J.M. Hunter, H. Wiriadinata, D.Girmansyah & A. Ruskandi ARs 94,
T.Ng. Praptosuwiryo 1717, 1718, 1719, 1735, 1779, 1782, 1791, 1792, 1794. ---
SUMATRA: H.A.B. Bünnenmeijer 4168; C.G. Mathew 636; T.Ng.
Praptosuwiryo 2040a, 2040b, 2040c, 2040d, 2086, 2058.
NOTES. Kato (1994) state that this species seems to be related to D.
fraxinifolium Presl. With anastomosing veins its linear, brown, entire scales,
imparipinnate leaves, and gradually narrowed subtruncate pinna-base. Kato
(1977) also suggested an affinity of D. xiphophyllum to D. subserratum, on the
basis of similarities in having costae raised on the adaxial side of leaves and
notched pinna-margin. He inferred that the simple leaves of D. subserratum have
been derived from pinnte ones of an ancestral species like D. xiphophyllum.
283
CHAPTER 10
GENERAL DISCUSSION
10.1. Synthesis
Systematists have become increasingly aware that reliance on single data
set may result in insufficiency resolution or an erroneous picture of phylogenetic
relationship. Therefore, it is now common practice to use multiple data sets. Most
researchers use datasets combining both molecular and non molecular for
phylogenetic inference (Soltis & Soltis 2000). Data from morphological
characters are compared or combined with one or more molecular data and then
summarized for their phylogenetic concensus.
Combining different data sets into a single analysis defines the “total
evidence” approach (Kluge 1989). However, the “total evidence” approach is
still debated (Myamoto & Fitch 1995). One major objection to “total evindence”
is that if data heterogeneity is high, the risk of obtaining a wrong inference from a
combined analysis increases (Bull et al 1993). One argument in separated
analysis to detect heterogeneity simply and quickly based on agreement and
disagreement highlighting conflicts caused by natural selection, differential rates
of evolution, hybridization, horizontal transfer, or lineage sorting (de Queiroz et al
1995). Comparison of separately analysed trees has been seen as especially useful
in identifying hybrids, wherein, for example, one may see conflicts between
uniparentally inherited genomes (most mitochondria and chloroplast) and nuclear
genes and/ or morphological characters (e.g. Rieseberg & Soltis 1991). It has
been argued that combined analysis may obscure significant patterns of
congruence or conflict among characters (Bull et al 1993, de Queiroz 1993). An
alternative way of expressing the idea of heterogeneity is if characters within a
data set are less independent than characters in different data sets, then there is
heterogeneity among data sets (de Queiroz et al 1995). As reviewed by de
Queiroz et al (1995), a number of authors have pointed to independent between
data sets (and explicitly or implicitly, nonindependence within data sets) as the
basis for arguments in favour of separate analysis (de Queiroz 1993, Lanyon
284
1993, Myamoto & Fitch 1995, Shaffer et al 1991). In this context, as pointed out
by De Queiroz et al (1995), nonindependence within data sets does not necessarily
imply functional or physical linkage, but only that characters within a data set are
more likely to share some property relevant for phylogeny estimation than
characters in different data sets. Using the support for conflict among trees from
different data sets as a means of assessing such independence (de Queiroz 1993,
Shaffer et al 1991) can be seen as a test of heterogeneity.
This study showed that morphological datasets indicate contain
phylogenetic signal and gene rcbL sequences data showed more phylogenetic
informative characters than morphological data. However gene rbcL sequences
data are only provided on 29 species of 69 species of Diplazium that analyzed.
Combining morphological datasets of 69 species with gene rbcL sequences data of
29 species would result more than 50% of missing data on combined matrix data,
and consequently would depict unnatural relationship. Therefore this study did not
combine the two datasets and summarized for their phylogenetic concencus. The
short general discussion is given below.
10.2. General Discussion
Phylogenetic analysis of West Malesian Diplazium revealed that the
homoplasies characters in Diplazium are high, Homoplasy Index (HI = 0.75)
(Chapter 6). Homoplasies may be resulted from parallelisms, convergence and
reversals. Within a cladistic framework homoplasies are also synapomorphies,
they represent derived character transformations defining two or more unrelated
groups of taxa. Whereas the “unique” synapomorphies define only a single group.
In the topological tree depicted in Figure 6.8. showed some terminal clades are
formed from the unrelated of species or not closely related species, such as clade
that contain that comprises D. hewittii and D. umbrosum, D. esculentum and D.
insigne, D. aquibasale and D. angustipinna.
Phylogenetic tree inferred from morphological tree showed that many
terminal clades are comprising of the species that presumed to be closely related
by previously author and seems depicting natural relationship although without or
with weak statistical support. For example D. riparium and D. wahauense that
285
presumed to be closely related are in the same tereminal clade. The same
depiction was also showed in the clade that containing D. malaccense,D.
sorzogonense and D. tricholepis, and ‘D. porphyrachis group’. It is indicate that
morphological data of Diplazium contain phylogenetic signal although many
characters are polymorphic and homoplastic. Backer et al (1998) stated that
morphological datasets contain more phylogenetic signal per characters and can
swamp much larger molecular datasets. Therefore, study that include more
morphological characters from living materials as well as anatomical characters,
such as stipe or rachis, would give an inferred phylogenetic tree with strong
statically support. As showed at Chapter 4, anatomical characters of stipe give an
indication that the closely related species are showing the similar anatomical
characters.
Knowing the status of species, whether originally hybrid or not, that will
be included in the phylogenetic analysis of Diplazium is important as many
species of Diplazium are apogamous and they are presumed to be originally
hybrids. Because if hybridization has occurred among the species of a taxon
under cladistic analysis the results are varied. Hybridization results in incongruent
intersecting data that obscure the underlying hiearachy (Funk 1985). All
hyphotheses regarding hybrids identification of course must be corroborated by
chromosomal, distributional, and ecological data. Therefore some species of
Diplazium that presumed to be originally hybrids based on result of this, such as
D. procumbens, D. subvirescens, and D. asymmetricum, should be confirmed in
the future study.
10.3. Systematic Implications for the Genus Diplazium
It is too premature to propose a systematic frame within the genus
Diplazium because: (1) The most parsimonious tree generated from morphological
data is not supported by the objective statistical arguments; (2) Phylogenetic tree
generated from gene rbcL sequences data has not given robust phylogentic tree as
most of the key species inferred from morphological tree are not included in
analysing; (3) Phylogenetic analysis on data sets combining both molecular and
non molecular for inferring the phylogeny of Diplazium has not been conducted
286
yet due to the incomplete data on many taxa (the gene rbcL sequences data are
only provided on 29 species of the 69 West Malesian species). However, this
results of this study is interesting because it shows that some clades generated
from morphological data are congruence with those generated from gene rbcL
sequence data, such as the separation of D. porphyrorachis from the other species
of Diplazium and the consistency of some species that included in the terminal
clades of both morphological tree and gene rbcL tree.
287
CHAPTER 11
CONCLUSIONS
Systematic study on West Malesian Diplazium revealed that based on
morphological characters, 69 species can be recognized. Thirteen species of them
are proposed to be new species, namely Diplazium asymmetricum, D.
batuayauense, D. crameri, D. densisquamatum, D. halimunense, D. loerzingii, D.
megasegmentum, D. megasimplicifolium, D. meijeri, D. parallelivenium, D.
profluens, D. subalternisegmentum, and D. subvirescens. D. donianum is new
record for Java and Sumatra.
Six species are recognized having morphological variations that led to
divide the species into two or three varieties. Diplazium accedens are consiting
three varieties, namely var. accedens, var. spinosum and var. ridleyi. D.
cordifolium are recognized to have three varieties, namely var. cordifolium, var.
integrifolium and var. pariens. D. pallidum consist two varieties, namely var.
pallidum and var. montanum. D. sorzogonense have two varieties, namely var.
sorzogonense and var. major. D. silvaticum are recognized to have two varieties,
namely var. silvaticum and var. pinnae-ellipticum. D. vestitum are divided into
two variesties, namely var. vestitum and var. borneense. D. accedens var.
spinosum, D. silvaticum var. pinnae-ellipticum are new varieties proposed.
Whereas D. pallidum var. montanum and D. accedens var. ridleyi are new status
proposed.
Ecological study showed that most of species are terrestrial dryland ferns
and found at 20 – 3400 m above sea level in the primary and secondary forest on
moist humus-rich soil in light and deep shady places. Species diversity were
culminated at 1000-1500 m. The different genetic load in the same species
sometimes grows in the different habitat gradient.
Distributional study resulted that the total number of species for each main
island are 40, 30, 29, 28 for Borneo, Java, Sumatra, and Malay Peninsula,
respectively. Nineteen species of West Malesia distribute very wide. They are D.
accedens, D. bantamense, D. cordifolium, D. crenatoserratum, D. dilatatum, D.
donianum, D. esculentum, D. fuliginosum, D. malaccense, D. pallidum, D.
288
polypodioides, D. prescottianum, D. riparium, D. silvaticum, D. simplicivenium,
D. sorzogonense, D. subserratum, D. tomentosum and D. xiphophyllum. Twenty
three species are presumed as locally endemic: (1) Nine endemic species are
occurring in Borneo (D. atrosquamosum, D. batuayauense, D. beamanii, D.
crinitum, D. squarrosum, D. subalternisegmentum, D. tricholepis and D.
wahauense); (2) Two endemic species in Malaya Peninsula (D. christii and D.
velutinum); (3) Six species are endemic in Sumatra (D. albidosquamatum, D.
betimusense, D. crameri, D. densisquamatum, D. meijerii, and D. melanolepis);
and (4) Six endemic species are recorded for Java (D. asymmetricum, D.
halimunense, D. megasegmentum, D. parallelivenium, D. profluens, and D.
subvirescens).
Anatomical study on the transversal section of stipe showed that the leaf-
trace shape of Diplazium stipe is varying among species and constant among the
adult individuals in a species. The shapes are consisted of five types: (1)
uninterrupted V-shaped, (2) interrupted, (3) uninterrupted U-shaped, (4)
interrupted U-shaped, and (5) W-shaped. Each type seems to vary among the
species. Each type may diversify into some different derivative forms that enables
to determine a species among closely related species. Therefore the leaf-trace
shapes are important diagnostic features which support species delimitation in
Diplazium.
Cytological observations on somatic chromosomes number showed that
West Malesian species showed basic chromosome number x = 41. New
cytological information for science on 19 species are recorded. They are D.
aequibasale (tetraploid), D. angustipinna (triploid), D. asymmetricum (triploid),
D. batuayauense (tetraploid and pentaploid), D. crenatoserratum (triploid and
tetraploid), D. halimunense (triploid), D. hewittii (triploid), D. profluens
(tetraploid), D. loerzingii (diploid and triploid), D. pallidum (diploid), D. petiolare
(diploid), D. porphyrorachis (tetraploid), D. riparium (diploid and triploid), D.
spiniferum (diploid), D. subserratum (diploid, triploid, tetraploi), triploid D.
subvirescens, D. tomentosum (diploid and pentaploid), D. xiphophyllum (diploid,
hexaploid), and Diplazium wahauense (2n=164).
289
This study showed that intraspecific diversity on West Malesian Diplazium
are high enough. Twelve species of the 31 species successfully examined are
having series ploidy: D. angustipinna (triplioid and tetraploid), D. bantamense
(traploid and oktoploid). D. cordifolium (tetraploid, pentaploid, dan hexaploid),
D. pallidum (diploid and tetraploid), D. silvaticum (triploid and tetraploid), D.
tomentosum (diploid, tetraploid, pentaploid), D. subserratum (diploid, triploid,
and tetraploid), D. xiphophyllum (diploid, tetraploid and hexaploid). Thirteen
species showed only polyploidy race: D. aequibasale (2n=164, tetraploid), D.
profluens (tetraploid), D. porphyrorachis (ca. 164/tetraploid), D. procumbens
(triploid), D. simplicivenium (triploid). Whereas nine species revealed only
diploid race (D. accedens, D. esculentum, D. malaccense, D. petiolare, D.
polypodioides, D. speciosum, D. spiniferum, D. sorzogonense, D.
subpolypodioides, and D. umbrosum).
Study on the relationships between ploidy level and morphological
variation in some species, such as D. batuayauense, D. crenatoserratum, D.
loerzingii, D. riparium, D. tomentosum, and D. xiphophyllum, suggested that they
are autoploid. Triploid D. silvaticum was presumed alloploid. However further
cytological study are needed to verify the polyploidy mechanisme involved in the
species mentioned above and the others.
Study on the relationship between ploidy level and habitat gradient
showed that generally West Malesian Diplazium species are showing no any
strick correlation between ploidy level and altitude, but D. pallidum, D. riparium,
D. silvaticum and D. xiphophyllum. Reproductive studies revealed that many
species are apogamous triploid and both the sexual and apomictic species can
occur in the relatively moist habitats.
Phylogenetic study by using morphological datasets revealed that the
phylogenetic relationship among species in the genus Diplazium was very difficult
to explain due to the lack of or weak support Bootstrap value. Therefore the
monophyletic species groups in this genus could not be identified confidently.
The high homoplastic on morphological characters, parallel evolution and some
apogamus species that suggested as originally hybrids distort the inference of
phylogenetic relationship among species.
290
However the lack of or weak support for a phylogenetic tree does not
strictly indicate that the pattern observed is incorrect but it does limit the amount
of confidence that can be placed in the relationships between taxa.and the
conclusions can be drawn from them. This study showed that some terminal
clades formed are consisting of species that presumed to be closely related species
by formerly authors and congruence with topological tree generated from gene
rbcL sequences, such as the affinity of D. porphyrorachis group and
imparipinnate species group (D. hottae, D. crameri, D. xiphophyllym, D.
fraxinifolium, D. subintegrum, D. bantamense, D. lobbianum). This study also
revealed that the classification of van Alderwereld van Rosenburgh (1908) in
dividing Diplazium into two sections (Eudiplazium and Anisogonium) is not
natural. Moreover, this study gave indication that Kato’s classification (1977)
could not be applied on West Malesian Diplazium. The lack of phylogenetic
signal in morphological datasets shows the need for other more informative data,
such as molecular data, for inferring phylogentic relationships.
Spore morphology study revealed that Diplazium have monolete,
bilateraly symmetrical, heteropolar, polar outline elliptical, perinates with with
laesura concealed by its perine. Perine ornamentation varied from alate to costate
alate and rugulate with variation of additional decoration such as ciliae, coralline,
echinae, and fenestrae. Diplazium species. The value of spore characters is as
descriptive device and an aid to the identification of dispersed spores grains.
Therefore perine ornamentations support in delimitating species concept in
Diplazium. The phylogenetic analysis using parsimony revealed that
morphological variation of spore is inadequate to depict natural relationship
among Diplazium species. Superficial similarity of mature perispores would not
indicate on systematic relationship among species as similar surface pattern it
derived through different development pathways.
Gene rbcL sequences data showed that this gene is very well in supporting
species delimitation among species and revealing the intraspecific diversity within
species of Diplazium. This study showed that most of West Malesian species are
genetically varying, mainly those having polyploidy types or seri ploidy.
Therefore the division of D. pallidum into two varieties is supported with this
291
gene rbcL sequences. This study give an indication that species in which have
both morphological difference and seri ploidy, such D. accedens, D. cordifolium,
D. crenatoserratum, D. bantamens, D. subserratum and D. tomentosum, should be
more studied by encompassing much more sample from their range of
distribution.
This study revealed that gene rbcL is more informative in inferring
phylogeny of the genus Diplazium in West Malesia. Therefore the phylogenetic
analysis of gene rbcL sequences found that Diplazium in West Malesia is
monophyletic.
The position of D. porphyrorachis group at the basal clade of the
phylogenetic tree generated from morphological data and separated from other
species is supported by the phylogenetic tree generated from molecular data (gene
rbcL sequence). This result is in accordance with the statement of Price (1983)
that D. porphyrorachis and close related species are belonging to the different
lines within the genus Diplazium. This study also showed the congruence
between the clade of ‘riparium Group’ drawn by rbcL tree and clade of
‘imparipinnate frond group’ drawn by morphological tree.
The result of this study also showed that the classification of van
Alderwerelt van Rosenburgh (1908) is not natural and polyphyletic. The
classification of Kato (1977) in dividing Japanese Diplazium into six group can
not be referred to West Malesian Diplazium. Moreover the Diplazium groups of
Kato (1977) are not monophyletic.
The result of the molecular systematic study also indicates that some
uncertain relationships require further analysis in the future. More data, including
both more taxa that include all West Malesian species and from wider areas and
more molecular data, are needed before a well resolved phylogenetic hypothesis
for West Malesian Diplazium can be offered.
292
LITERATURES
Abraham A, Ninan CA, Mathew PM. 1962. Studies on the cytology and
phylogeny of the pteridophytes: VII. Observation on one hundred species
of south Indian ferns. J Indian Bot Soc 41: 339-421.
Adams KL, Cronn R, Percifield R, Wendel JF. 2003. Genes duplicated by
polyploidy show unequal contributions to the transcription and organ-specific
reciprocal silencing. Proc Natl Acad Sci 100: 4649-4654.
Alston AHG. 1956. The Subdivision of the Polypodiaceae. Taxon 5: 23-25.
Andrews SB. 1990. Ferns of Queensland. Brisbane: Queensland Department of
Primary Industries.
Backer CA en O Posthumus. 1939. Varenflora voor Java. Uitgave van’s Lands
Plantetuin, Buitenzorg.
Backer RH, Yu X, DeSalle R. 1998. Assessing the relationship contribution of
molecular and morphological characters in simultaneous analysis trees. Mol
Phylog Evol 9: 427 – 436.
Barrington DS, Paris CA, Ranker TA. 1986. Systematic inferences from spores
and stomata size in the ferns. Amer Fern J 76 (3): 149-159.
Baack EJ. 2004. Cytotype segregation on regional and microbiographic scales in
snow buttercups (Rannunculus adoneus: Ranunculaceae). Amer J Bot 91:
1783-1788.
Bhavanandan KV, Ammal LS. 1991. Cytological investigations on family
Aspidiaceae (sensu Copeland) from south India. Indian Fern J 8: 78-86.
Bir SS. 1962. Taxonomy of the Indian members of family “Aspleniaceae”. Bull
Bot Surv India 4: 1-16.
Bir SS. 1969. The stelar anatomy of Diplazium latifolium Moore. Amer Fern J
59: 23 – 26.
Blackmore S. 2000. The palynological compass: the contribution of palynology
to systematics. In: Nordenstan B, El-Ghozaly G, Kassas M, editors. Plant
Systematics for the 21st Century. London: Portland Press. Pp 161-177.
Bower FO. 1912. Studies in the phylogeny of the Filicales. II. Lophosoria and its
relation to the Cyatheoideae and other ferns. Annals Bot 26: 269-325.
293
Bower FO. 1913. Studies in the phylogeny of the Filicales. III. On Metaxya
and certain other relative primitive ferns. Annals Bot 27: 443-477.
Bower FO. 1923-1928. The Ferns (Filicales). 3 Vols. England: Cambridge.
Bremer K, Wanntorp HE. 1978. Phylogenetic systematic in botany. Taxon 27:
317-329.
Brummit RK 1992. Vascular Plant Families and Genera. Kew: Royal Botanic
Garden.
Bull JJ, Huelsenbeck JP, Cunningham CW, Swofford DL, Waddel PJ. 1993.
Partaining and combining data in phylogenetic analysis. Syst Biol 42: 384-
397.
Chase MW, Soltis DE, Olmstead RG, Morgan D, Les DH, Mishler BD, Duvall
MR, Price RA, Hill HG, Qiu Y-L, Kron KA, Rettig JH, Conti E, Palmer JD,
Manhart JR, Sytsma KJ, Michaels HJ, Kress WJ, Karol KG, Clark WD,
Hedrén M, Gaut BS, Jansen RK, Kim K-J, Wimpee CF, Smith JF, Furnier GR,
Strauss SH, Xiang Q-Y, Plunkett GM, Soltis PS, Swensen SM, Williams SE,
Gadek PA, Quinn CJ, Eguiarte LE, Golenberg E, Learn GH Jr, Graham SW,
Barrett SCH, Dayanandan S, Albert VA. 1993. Phylogenetics of seed plants:
an analysis of nucleotide sequences from the plastid gene rbcL. Ann Missouri
Bot Gard 80: 528-580.
Chin WY. 1997. Ferns of the Tropics. Singapore: Times Editions.
Ching R-C. 1940. On natural classification of the family ‘Polypodiaceae”.
Sunyatsenia 5: 201-286.
Ching R-C. 1954. A systematic arrangement of the Chinese fern families and
genera with corresponding names in Chinese. Acta Phytotax Sin 3: 93-98.
Ching R-C. 1964a. On some confused genera of the family Athyriaceae. Acta
Phytotax Sin 9 (1): 41-84.
Ching R-C. 1964b. On the genus Diplaziopsis C. Chr. Acta Phytotax Sin 9(1): 31-
36.
Christensen C. 1911. On the Genus Dryopteris. Amer Fern J 1: 33-37.
Christensen C, Holttum RE. 1934. The ferns of Mount Kinabalu. Gard Bull Sing
7: 191-279.
294
Colwell RK, Hurtt GC. 1994. Non biological gradients in species richness and a
spurious Rapoport effect. American Naturalist 144: 570-595.
Copeland EB. 1908. A revision of the Philippine species of Athyrium. Philip J
Sci IIIc: 285-230.
Copeland EB. 1929. The oriental genera of Polypodiaceae. Univ Cal Publ Bot
16: 45-128.
Copeland EB. 1947. Genera Filicum. MASS, USA: Waltham.
Crane EH. 1997. A revised circumscription of the genera of the fern family
Vittariaceae. Syst Bot 22 (3): 509-517.
Crane EH, Farrar DR, Wendel JH. 1995. Convergent simplification leads to a
polyphyletic Vittaria. Amer Fern J 85: 283-305.
Darnaedi D. 1991. Kromosom dalam Taksonomi. Makalah dalam Kursus
Singkat Metodologi Penelitian Taksonomi Tumbuhan, PAU Hayat IPB, 12
Desember 1990-12 Februari 1991. Bogor.
Darnaedi D. 1992. A preliminary cytologicalstudy of fern flora of Gede-
Pangrango National Park (West Java). Proc Sem Asian Pterid II: 73-78.
Darnaedi D, Kato M, Iwastsuki K. 1989. A cytotaxonomic study of Dryopteris
sparsa and closely related species (Dryoptridaceae). J Jap Bot 64: 330-
340.
Davis PH, Heywood VH. 1963. Principles of Angiosperm Taxonomy. Edinburg
and London: Oliver and Boyd.
de Vogel EF. 1987. Guidelines for the preparations of revisions. In de Vogel EF,editor. Manual of Herbarium Taxonomy Theory and Practice. Jakarta:UNESCO.
Dickason FG. 1946. A phylogenetic study of the ferns of Burma. Ohio J Sci 42(2): 73 – 108.
Doyle JJ, Doyle JL. 1987. A rapid DNA isolation procedure for small quantities
of fresh leaf tissue. Phytochem Bull 19: 11-15.
Dubuisson J-Y, Hebant-Mauri R, Galtier J. 1998. Molecules and morphology:
Conflicts and congruence within the fern genus Trichomanes
(Hymenophyllaceae). Mol Phylog & Evol 9 (3): 390-397.
295
Duvall MR, Clegg MT, Chase MW, Clark WD, Kress WJ, Hills HG, Eguiarte LE,
Smith JF, Gaut BS, Zimmer EA, Learn GH Jr. 2003. Phylogenetics
hyphothesis for the Monocotyledons constructed from rbcL sequence data.
Ann Missouri Bot Gard 80: 607-619.
Eastwood W, Cronk QCB, Vogel JC, Hemp A, Gibby M. 2004. Comparison of
molecular and morphological data on St. Helena: Elaphoglossum. Plant Syst
Evol 245: 93-106.
Edie, H. 1978. Ferns of Hongkong. Hong Kong: Hong Kong University Press.
Colwell RK, Hurtt GC. 1994. Nonbiologcal gradients in species richness and a
spurious Rapoport effect. Amer Naturalist 144: 570-595.
Endress PK, Bass P, Gregory M. 2000. Systematic plant morphology and
anatomy –50 years of progress. Taxon 49 (3): 401-434.
Estabrook GF. 1978. Some concept for estimation of evolutionary relationships
in Systematic Botany. Syst Bot 3: 146 – 158.
Felsenstein J. 1985. Confidence limits on phylogenies: an approach using the
bootstrap. Evolution 39: 783 -791.
Ferrarini E, Ciampolini F, Pichi Sermolli REG, Marchetti D. 1986. Iconographia
palynologica Pteridophytorum Italiae. Webbia 40(1): 1-202.
Funk VA. 1985. Phylogenetic pattern and hybridization. 1985. Ann Missouri
Bot Gard. 72: 681-715.
Gastony GJ. Spore morphology in the Dicksoniaceae. II. The genus Cibotium.
Cannad J Bot 60: 955-972.
Gastony GJ, Ungerer MC. 1997. Molecular systematics and a revised taxonomy
of the onocleoid ferns (Dryopteridaceae: Onocleeae). Amer J Bot 84: 840-
849.
Gastony GJ, Johnson WP. 2001. Phylogenetic placements of Loxoschape
thecifera (Aspleniaceae) and Actiniopteris radiate (Pteridaceae) based on
analysis of rbcL nucleotide sequences. Amer Fern Jour 91: 197-213.
Geiger JMO, Ranker TA. 2005. Molecular phylogenetic and historical
biogeography of Hawaiian Dryopteris (Dryopteridaceae). Mol Phylog & Evol
34: 392-407.
296
Gentry AH, Dodson CH. 1987. Diversity and biogeography of neotropical
vascular epiphytes. Annals Missouri Bot Gard 74: 205-233.
Givnish TJ, Sytsma KJ. 1997. Consistency, characters, and the likelihood of
correct phylogenetic inference. Mol Phylog & Evol 7: 320-330.
Golenberg EM, Giannasi DE, Clegg MT, Smiley CJ, Durbin M, Henderson D,
Zurawski G. 1990. Chloroplast DNA sequence from a Miocene Magnolia
species. Nature 344: 656-658.
Godfray HC, Lawton JH. 2001. Scale and species numbers. TRENDS in Ecol &
Evol. 16 (7): 400-404.
Grant V. 1981. Plant Speciation. 2nd Edition. New York: Columbia University
Press.
Harris WF. 1955. A Manual of the Spores of New Zealand Pteridophyta.
Department of Scientific and Industrial Research Bulletin 116. Government
Printer, Wellington.
Hasebe M et al. 1994. rbcL gene sequences provide evidence for the
evolutionary lineages of leptosporangiatae ferns. Proc Natl Acad Sci USA 91:
5730-5734.
Hasebe M et al. 1995. Fern phylogeny based on rbcL nucleotida sequences.
Amer Fern J 85: 134-181.
Hassler M, Brian S. 2002. Family Athyriaceae, genus Diplazium; world species
list. Published on the internet.
http://homepages.caverock.net.nz/~bj/fern/diplazium.htm (accessed 6
November 2005).
Haufler CH, Ranker TA. 1995. rbcL sequences provide phylogenetic insights
among sister species of the ferns genus Polypodium. Amer Fern J 85: 361-
374.
Haufler CH, Soltis DE. 1986. Genetic evidence suggests that homosporous fern
with high chromosome numbers are diploid. Proc Natl Acad Sci 83: 4389-
4393.
297
Haufler CH, Hooper EA, Therrien JP. 2000. Modes and mechanism of speciation
in pteridophytes: Implications of contrasting patterns in ferns representing
temperate and tropical habitats. Plant Spec Biol 15: 223-236.
Hauk WD, Parks CR, Chase MW. 2003. Phylogenetic studies of
Ophioglossaceae: evidence from rbcL and trnL-F plastid DNA sequences and
morphology. Mol Phylog & Evol 28: 131-151.
Hennequin S, Ebihara A, Ito M, Iwatsuki K, Dubuisson J-Y. 2006. New insights
into the phylogeny of the genus Hymenophyllum s.l. (Hymenophyllaceae):
Revealing the polyphyly of Mecodium. Syst Bot 31(2): 271-284.
Herter WG. 1949. Flora de Uruguay. I. Pteridophyta. Rev Sudamer Bot 9: 1-
28.
Hillis DM. 1987. Molecular versus morphological approaches to systematics.
Ann Rev Ecol Syst 18: 23-42.
Holbrook-Walker S, Lloyd RM. 1973. Reproductive biology and gametophyte
morphology of the Hawaiian fern genus Sadleria (Blechnaceae) relative to
habitat diversity and propensity for colonization. Bot J Linn Soc 67: 157-174.
Holttum, R.E. 1938. The Ecology of Tropical Pteridophytes. In: Verdoorn F,
editor. Manual of Pteridology. The Netherlands: Martinus Nijhoff. Pp. 420-
450.
Holttum RE. 1940. The fern genus Diplazium in the malay Peninsula. Gard Bull
Sing 9 (2): 74 – 107.
Holttum RE. 1947. A revised classification of Leptosporangiate ferns. J Linn
Soc Bot 53: 123-158.
Holttum RE. 1954. A Revised Flora of Malaya. II. Ferns of Malaya. Singapore:
Gov Print Office.
Holttum RE. 1966. A Revised Flora of Malaya. II. Ferns of Malaya. Singapore:
Gov Print Office. 658p. + Appendix.
Holttum RE, Sen U. 1961. Morphology and classification of the tree ferns.
Phytomorphology 11: 406-420.
Holttum RE, Roy SK. 1965. Cytological observation on some ferns from New
Guinea with descriptions of new species. Blumea 13:129-139.
298
Huang Y-M, Chou H-M, Wang J-C, Chiou W-L. 2007. The distribution and
habitats of the Pteris fauriei complex in Taiwan. Tawania 52(1): 49-58.
Hyrabayashi H. 1974. Cytogeographic Studies on Dryopteris of Japan. Tokyo:
Harashobo.
Imaichi R, Kato M. 1995. Speciation and morphological evolution in rheophytes.
In: Iwatsuki K, Raven PH, editors. Evolution and Diversification of Land
Plants. Tokyo: Springer-Verlag. Pp. 309- 318.
IPNI (The International Plant names Index). 2006. Published on the internet
http://www.ipni.org (accessed 17 July 2006).
Iwatsuki K, Kato M. 1984. Additions to the enumeration of East Kalimantan
Pteridophytes 3. Acta Phytotax Geobot 35 (1-3): 59 -67.
Jian-xiu LZ, Feng-Qin, Yu-long Z. 1988. Studies on the spores morphology of
Hypodematium in China. Pp. 269-272. In KH. Shing & K.U. Kramer (Eds.).
Proceeding of the International Symposium on Systematic Pteridology.
September 5-10. Beijing.
John RJ. 1995. Malesia-An Introduction. Curtis’s Bot Mag 12 (2): 52-62.
Kato M. 1972. The vascular structure and its taxonomic significant in the the
Athyriaceae. Acta Phyototax Geobot 25 (2-3): 79 – 91.
Kato M. 1973. Taxonomical evaluation of the articulate hairs found in the
Athyriaceae. Acta Phytotax Geobot 25 (4-6):119-125.
Kato M. 1977. Classification of Athyrium and allied genera of Japan. Bot Mag
(Tokyo) 90: 23-40.
Kato M. 1979. Taxonomic study of the genus Cornopteris (Athyriaceae). Acta
Phytotax Geobot 33(4-6): 101-118.
Kato M. 1984. A taxonomic study of the Athyrioid fern genus Deparia with
main reference to the Pacific species. J Fac Sci Univ Tokyo IV 13: 375-
430.
Kato M. 1987. A phylogenetic classification Ophioglossaceae. Gard Bull Sing
40 (1): 1 – 14.
Kato M. 1992. Taxonomic study of Pteridophytes of ambon and Seram
(Molluccas) collected by Indonesian – Japanese botanical expedition IX.
299
Woodsiaceae, Lindsaeaceae, and Adiantaceae. J Fac Sci Univ Tokyo Sect III
(15): 135-152.
Kato M. 1994. Taxonomic Studies of Pteridophytes of Ambon and Seram
(Mollucas) Collected by Indonesian-Japanese Botanical Expedition IX.
Woodsiaceae, Lindsaeaceae and Adiantaceae. J Fac Sci Univ Tokyo 15(4):
315-347.
Kato M. 1995. Woodsiaceae. In: Iwatsuki K, Yamasaki Y, Boufford DE, Ohba
F, editors. Flora of Japan. Volume I. Pteridophyta & Gymnospermae. Tokyo:
Kodhansha. Pp. 195-231.
Kato M, Setoguchi H. 1999. An rbcL-based phylogeny and heteroblastic leaf
morphology of Matoniaceae. Syst Bot 23(4): 391-400.
Kato M, Darnaedi D, Iwatsuki K. 1991. Fern rheophytes of Borneo. J Fac Sci
Univ Tokyo III 15: 91-110.
Kato M, Yatabe Y, Sahashi N, Murakami N. 2001. Taxonomic studies of
Cheiropleuria (Dipteridaceae). Blumea :
Kikkawa J, Williams EE. 1971. Vegetational structure and avian diversity in
several New World areas. American Naturalist 105: 425-435.
Kitching IJ, Forey PL, Humphries CJ, Williams DM. 1998. Cladistics: The
Theory and Practice of Parsimony Analysis. Ed.2. Oxford: Oxford University
Press.
Kilpper K. 1964. Über eine Rät/Lias-Flora aus dem Nördlichen Abfall des
Albursgebirges in Nordiran. 1. Bryophyta und Pteridophyta.
Palaeontographica, Abt B, Paläophytol 114: 1-78.
Kluge AG. 1989. A concern for evidence and a phylogenetic hyphothesis of
relationships among Epicrates (Boidae, Serpentes). Syst Zool 38: 7 – 25.
Kramer KU, Holttum RE, Moran RC, Smith AR. 1990. Dryopteridaceae. In:
Kramer KU and Green PS (eds.) Pteridophytes and Gymnosperms. Berlin:
Springer-Verlag. Pp 101-144.
Large MF, Braggins JE. 1991. Spore Atlas of New Zealand Ferns and Ferns
Allies. Wellington: SIR Publishing.
300
Lee MSY. 2004. Molecular and morphological datasets have similar numbers of
relevant phylogenetic characters. Taxon 53 (4): 1019-2022.
Levin DA. 2002. The Role of Chromosomal Change in Plant Evolution. Oxford:
Oxford University Press. 240pp.
Lewis WH. 1980. Polyploidy in species population. In: Lewis WH, editor.
Polyploidy: Biological Relevance. NY – USA: Plenum Press. Pp. 103-144.
Lincoln RJ, Boxshall GA, Clark PF. 1982. A Dictionary of Ecology, Evolution
and Systematics. New York: Cambridge University Press.
Linder HP. 2001. On areas of endemism, with an example from the
Restionaceae. Syst Biol 50 (6): 892-912.
Lloyd RM. 1974. Mating system and genetic load in pioneer and non-pioneer
Hawaiian Pteridophyta. Bot J Linn Soc 69: 23-35.
Lomolino MV. 2001. Elevational gradients of species-density: historical and
prospective views. Global Ecology & Biogeography 10: 3-13.
Löve A, Löve D, Pichi Sermoli REG. 1977. Cytotaxonomical Atlas of the
Pteridophyta. Vaduz: J Cramer. 398p.
Lovis JD. 1977. Evolutionary patterns and processes in ferns. Adv Bot Res 4:
229-415.
Lu J-M, Barrington DS, Li D-Z. 2007. Molecular phylogeny of the Polystichoid
ferns in Asia based on rbcL sequences. Syst Bot 32 (1): 26-33.
Maddison WP. 1993. Missing data versus missing characters in phylogenetic
analysis. Syst Biol 42: 576-581.
Manton I. 1950. Problems of Cytology and Evolution in the Pteridophyta.
Cambridge: Cambridge Univ. Press.
Manton I. 1953. The cytological evolution of the fern flora of Ceylon. Symp Soc
Expl Biol 7: 174-185.
Manton I. 1954. Cytological notes on one hundred species of Malayan ferns. In:
Holttum RE. A Revised Flora of Malaya. II. Appendix: 623 – 627.
Manton I, Sledge WA. 1954. Observations on the cytology and taxonomy of the
Pteridophyte flora of Ceylon. Phil Trans B. 238.
Maxted N. 1992. Towards defining a taxonomic revision methodology. Taxon
41(4): 653-660.
301
McCoy ED. 1990. The distribution of insects along elevational gradients. Oikos
58: 313-322.
Mickel JT. 1974. Phyletic lines in the modern fern. Annals Missouri Bot Gard 61
(2): 474 – 482.
Manen, J.-F., V. Savolainen, S. De Marchi, and B. Rion. 1995. Chloroplast DNA
sekuense from Mioece diatomite deposit in Ardeche (France) Comptes Rendu.
Academy of Science Paris 318: 971-975.
Matsumoto S. 2003. Species ecological study on reproductive systems and
speciation of Cyrtomium falcatum complex (Dryopteridaceae) in Japanese
Archipelago. Ann Tsukuba Bot Gard 22: 1-141.
Mitsui K. 1986. The development of the perispore in the fern family
Thelypteridaceae. In Blackmore S, Ferguson IK (eds). Pollen and spores:
form and function. Linnean Society Symposium Series No 12 : 401-403.
London: Academic Press.
Mitsuta S. 1985. List of West Sumatran ferns and fern allies collected by dr.
Hotta and others during 1980-83 (3). Acta Phytotax Geobot 36 (1-3): 73-86.
Moore S-J, Hsieh T-H, Huang YM, Chiou W-L. 2002. Diplazium maoenense
Ching, a poorly known species of the Athyriaceae (Pteridophyta) in Taiwan.
Taiwan J For Sci 17(1): 113-118.
Murakami N, Yokoyama J, Cheng X, Iwasaki H, Imaichi R, Iwatsuki K. 1998.
Molecular -taxonomy of Hymenasplenium obliquissimum complex
(Aspleniaceae) based on rbcL sequences. Plant Spec Biol 13: 51-56.
Murakami N, Nagomi S, Watanabe M, Iwatsuki K. 1999a. Phylogeny
Aspleniaceae inferred from rbcL nucleotide sequences. Amer Fern J 89: 232-
243.
Murakami N, Watanabe M, Yokoyama J, Yatabe Y, Iwasaki H, Serizawa S.
1999b. Molecular taxonomic study and revision of the three Japanese species
of Asplenium sect. Thamnopteris. J Plant Res 112: 15-25.
Myamoto MM, Fitch WM. 1995. Testing species phylogenies and phylogenetic
methods with congruence. Syst Biol 44: 64 – 76.
302
Nagalingum NS, Schneider H, Pryer KM. 2007. Molecular phylogenetic
relationship and morphological evolution in the heterosporous fern genus
Marsilea. Sys Bot 32 (1): 16 – 25.
Nakato I. 1981. Notes chromosomes of Japanese pteridophytes (1). J Jap Bot
56: 200-205.
Nayar B. 1970. A phylogenetic classification of the homosporous ferns. Taxon.
19: 229-236.
Nelson G, Platnick N. 1981. Systematics and Biogeography. New York:
Columbia Univ Press.
Nishida M, Nishida H. 1982. Histology of the rhizome of Loxsomopsis and
affinity of Solenostelopsis loxsomoides Ogura. Acta Phytotax Geobot 33: 302
– 307.
Ohta N, Takamiya M. 1999. Taxonomic studies of the Diplazium mettenianum
complex (Woodsiaceae; Pteridophyta) in Japan: Morphology, cylotogy and
taxonomy of plants with normal-shaped spores. J Plant Res 112: 67-86.
Pacheco L. 2004. A new species of Diplazium (Woodsiaceae) from Ecuador.
Brittonia 56(2): 121-123.
Palmer JD, Jansen RK, Michael HJ, Chase DW, Manhart JH. 1988. Chloroplast
DNA variation and plant phylogeny. Ann Missouri Bot Gard 75: 1180-1206.
Park CH, Kato M. 2003. Apomixis in the interspecific triploid hybrid fern
Cornopteris christenseniana (Woodsiaceae). J Plant Res 116: 93-103.
Parris BS, Beaman RS, Beaman, JH. 1992. The Plants of Mount Kinabalu. I.
Ferns and Fern Allies. Kew: Royal Botanic Gardens.
Parris BS. 2003. The distribution of Grammitidaceae (Filicales) inside and
outside Malesia. Telopea 10(1): 451-466.
Pettit JM. 1966. Exine stucture in some fossil and recent spores and pollen as
revealed by light and electron microscopy. Bull Brit Mus (Nat Hist) Geol 13:
221-257.
Pichi-Sermolli REG. 1977. Tentamen Pteridophytorum genera in taxonomicum
ordinem redigendi. Webbia 31 (2): 313-512.
Praptosuwiryo TNg. 1999. Biosystematic Study of Diplazium in Java. Program
Pasca Sarjana, Institut Pertanian Bogor. [Thesis]
303
Praptosuwiryo TNg. 2003. Studi Kromosom Somatik Beberapa Jenis Tumbuhan
Paku di Pulau Lombok dan Sumbawa. Widyariset 5 : 87-96.
Praptosuwiryo TNg, Darnaedi D. 1994. Cytological Study of the Ferns Genus
Diplazium in Gede-Pangrango National Park, Java. Floribunda 1(15): 57-
60.
Praptosuwiryo TNg, Darnaedi D. 2004. Cytological Study of Some Species of
Ferns Genus Diplazium in Java I. Floribunda 2(5): 128-137.
Price MG. 1983. Several unusual Malesian diplazia. Gard Bull Sing 36 : 25-29.
Qiu Y-J, White RA, Turner MD. 1995. The developmental anatomy of Metaxya
rostrata (Filicales: Metaxyaceae). Amer J Bot 82 (8): 969-981.
Queiroz A de, Donoghue MJ, Kim J. 1995. Separate versus combined analysis of
phylogenetic evidence. Annu Rev Ecol Syst 26: 657-681.
Rahbek C. 1995. The elevational gradient of species richness: a uniform pattern?
Ecography 18: 200-205.
Rangker TA. 1989. Spore morphology and generic delimitation of new world
Hemionitis, Gymnopteris, and Bommeria (Adiantaceae). Amer J Bot 76(2):
297-306.
Ranker TA, Floyd SK, Trapp PG. 1994. Multiple colonization of Asplenium
adiantum-nigrum onto the Hawaiian Archipelago. Evolution 48: 1364-1370.
Ramsey J, Schemske WD. 1998. Pathways, mechanism, and rates of polyploidy
formation in flowering plants. Ann Rev Ecol Syst 29: 467-501.
Rifai MA. 1976. Sendi-sendi Botani Sistematika. Bogor: Lembaga Biologi
Nasional – LIPI.
Roos MC. 1993. State of affairs regarding Flora Malesiana: progress in revision
work and publication schedule. Flora Malesiana Bulletin 11: 133-142.
Roos M. 1995. Flora Malesiana: progress, needs and prospects. In: J.
Dransfield et. al. Pp. 231-246. Plant Diversity in Malesia III. Proceedings of
the Third International Flora Malesiana Symposium.
Roubik, D.W. 2003. Pollen and spores of Barro Colorado Islands. 2003.
http://striweb.si.edu/roubik/ [24 Feb. 2007].
Rouhan G, Dubuisson J-Y, Rakotondrainibe F, Motley TJ, Mickel JT, Labat J-N,
Moran RC. 2004. Molecular phylogeny of the fern genus Elaphoglossum
304
(Elaphoglossaceae) based on chloroplast non-coding DNA sequences:
contributions of species from the Indian Ocean area. Mol Phylog & Evol 33:
745 – 763.
Sano R, Takamiya M, Kurita S, Ito M, Hasebe M. 2000a. Diplazium
subsinuatum and Di. Tomitaroanum should be moved to Deparia according to
molecular, morphological, and cytological characters. J Plant Res 113: 157-
163.
Sano R, Takamiya M, Ito M, Kurita S, Hasebe M. 2000b. Phylogeny of the lady
fern group, tribe Physematieae (Dryopteridaceae), based on chloroplast
rbcL gene sequences. Mol Phylog & Evol 15: 403-413.
Schölch A. 2000. Relations between sumbarginal and marginal sori in ferns. I.
The sori of selected Hypolepidaceae and Dennstaedtiaceae. Plant Syst Evol
220: 161 – 183.
Shmida A, Wilson MV. 1985. Biological determinants of species diversity.
Journal of Biogeography 12: 1-20.
Shivas MG. 1961. Contributions to the cytology and taxonomy of species of
Polypodium in Europe and America: I. Cytology. J Linn Soc (Bot) 58: 13-
25.
Skog JE, Mickel JT, Moran RC, Volovsek ME, Zimmer EA. 2004. Molecular
studies of representative species in the fern genus Elaphoglossum
(Dryopteridaceae) based on chloroplast DNA sequences rbcL, trnL-F, and
rps4-trnS. Int J Plant Sci 165 (6): ….
Sledge WA. 1962. The athyrioid ferns of Ceylon. Bull Brit Mus (Nat Hist) Bot
2: 277-323.
Smith AR. 1972. Comparison of fern and flowering plant distribution with some
evolutionary interpretation for ferns. Biotropica 4: 4-9.
Smith AR, Kreier H-P, Haufler CH, Ranker TA, Schneider H. 2006.
Serpocaulon (Polypodiaceae), a new genus segregated from Polypodium.
Taxon 55(4): 919-930.
Soltis DE, Soltis PS. 1998. Choosing an approach and an appropriate gene for
phylogenetic analysis. In DE Soltis, PS Soltis, JJ Doyle. Pp 1-42. Mollecular
305
Systematics of Plant II. DNA Sequencing. Boston: Kluwer Academic
Publishers.
Soltis PS, Soltis DE, Smiley CJ. 1992. An rbcL sequences from a Miocene
Taxodium (Bald Cypress). Proceeding of the National Academy of Sciences
U.S.A. 89:449-451.
Soltis PS and Soltis DE. 2000. The role of genetic and genomic attributes in
the success of polyploids. Proc Natl Acad Sci 97: 7051-7057.
Stebbins GL. 1971. Chromosomal Evolution in Higher Plants. MA – USA:
Addison – Wesley.
Steenis, C.G.G.J. van. 1957. Specific and Infraspecific Delimitation. In: Flora
Malesiana. Ser. I, Vol. 5.
Steenis CGGJ van. 1981. Rheophytes of the World. Sijthoff & Noordhoff,
Alpena an den Rjn: Sijthoff & Noordhoff.
Strong EE, Lipscomb D. 1999. Character coding and inapplicable data.
Cladistics 15: 363-371.
Swartz, O. 1801. Achrad. J. Bot. 1800(2): 61.
Swofford DL. 1998. PAUP*. Phylogenetic Analysis Using Parsimony (*and
Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts.
Tagawa M. 1972. Ferns of Borneo, collected by M Hirano and M. Hotta 6. Acta
Phytotax Geobot. 25 (2-3): 61-68.
Tagawa M, Iwatsuki K. 1988. Flora of Thailand. Vol. 3. Part 3: 436-466.
Takamiya M (ed). 1996. Index to Chromosomes Number of Japanese
Pteridophyta (1910-1996). Tokyo: Japan Pteridological Society.
Takamiya M. 1997. Current status and problems of cytotaxonomical studies of
Japanese pteridophytes. Bull Jap Pterid Soc 70: 1-23.
Takamiya M, Takaoka C, Ohta N. 1999. Cytological and reproductive studies on
Japanese Diplazium (Woodsiaceae; Pteridophyta): Apomictic
reproduction in Diplazium with evergreen bi- to tripinnate leaves. J Plant
Res 112: 419-436.
Takamiya M, Ohta N, Fujimaru-Takaoka C, Uki K. 2000. J Plant Res 113: 203-
215.
306
Takamiya M, Ohta N, Yatabe Y, Murakami N. 2001. Cytological,
morphological, genetic, and molecular phylogenetic studies on intraspecific
differentiation within Diplazium doederleinii (Woodsiaceae: Pteridophyta).
Int J Plant Sci 162 (3): 625-636.
Tardieu-Blot. 1932. Les Apleniees Du Tonkin. Theses. Prentees A la Faculte
des Science de Paris. Toulouse Imprimerie henri Basuyau & C’. 190p.
Thiele K. 1993. The holy grail of the perfect characters: the cladistic treatment
of morphometric data. Cladistics 9: 275-304.
Tiegham P van, Douliot H. 1886. Sur la polystelie. Ann Sci Nat Bot Ser 7 (3):
275 – 322.
Tindale MD & SK Roy. 2002. A Cytotaxonomic Survey of the Pteridophyta of
Australia. Australian Syst Bot15: 839-937.
Triono T. 2006. A Phylogenetic Study of Pouteria (Sapotaceae) in Malesia and
Australia using Morphology and Molecular Data. The Australian National
University. Canberra, Australia. [Doctoral Thesis]
Tryon A. 1986. Stasis, diversity andfunction in spores based on an electron
microscope survey of the Pterudophyta. In: Blackmore S, Ferguson IK,
editors. Pollen and spores: form and function. Linnean Society Symposium
Series No 12 : 233-249. London: Academic Press.
Tryon AF, Lugardon B. 1991. Spore of the Ptridophyta. New York: Springer
Verlag.
Tseng-Chieng, H. 1981. Spore Flora of Taiwan. 1981. Taiwan: National Taiwan
University.
Tryon RM, Stolze RG. 1991. Pteridophyta of Peru. Part IV 17.
Dryopteridaceae. Fieldiana No 17: 1-176.
Tsutsumi,C. and Kato,M. 2006. Evolution of Epiphytes in Davalliaceae and
Related Ferns. Bot J Linn Soc 151, 495-510.
Van Alderwerelt van Rosenburgh CRWK. 1908. Malayan Ferns. Batavia:
Landsdrukkerij.
Van Konijnenburg-van Cittert JHA. 1975. Some notes on Marattia anglica from
the Jurassic of Yorkshire. Rev Palaeobot Palynol 20: 205-214.
307
Van Konijnenburg-van Cittert JHA. 1999. On the evolution of fern spore
anchitecture. In: Kurmann MH, Hemsley AR (Eds). The Evolution of Plant
Architecture. Pp 179-287. Kew: Royal Botanic Garden.
Vida G. 1976. The role of polyploidy in evolution. In: Nivac VJA & Pacltova,
editors. Evolutionary Biology. Rechoslovak: Prague-CCzechoslovak
Acad Sci Prague. Pp.267-294.
Vogel JC, Rumsey FJ, Schneller JJ, Barret J, Gibby M. 1999. Where are the
glacial refugia in Europe? Evidence from pteridophytes. Bot J Linn Soc
66: 23-37.
Wang M-L, Chen Z-D, Zhang X-C, Lu S-G, Zhao G-F. 2003. Phylogeny of
Athyriaceae: evidence from chloroplast trnL-F region sequences. Acta
Phytotax Sin 41 (5): 416-426.
Wagner WH. 1961. Problems in the classification of ferns. Recent Advances in
Botany. Pp. 841-844. University of Toronto Press.
Wagner WH. 1972. Disjunctions in homosporous vascular plants. Ann Missouri
Bot Gard 59: 203-217.
Wagner WH. 1974. Structure of spores in relation to fern phylogeny. Ann
Missouri Bot Gard 61: 332-353.
White RA. 1974. Comparative anatomical studies of the ferns. Ann Missouri Bot
Gard 62(2): 379 – 387.
Wiley EO. 1980. Phylogenetic systematic and vicariance biogeography. Syst Bot
5: 194-220.
Wolf PG, Soltis PS, Soltis DE. 1994. Phylogenetic relationships of
Dennstaedtioid ferns: Evidence from rbcL sequences. Mol Phylog & Evol
3(4): 383 – 392.
Wolf PG, Conant DS. 1994. Use of Molecular Data in Evolutionary Studies of
Pteridophytes. Amer J Bot 81 (6): 119. (Abstract).
Yatabe Y, Masuyama S, Darnaedi D, Murakami N. 2001. Molecular systematics
of the Asplenium nidus complex from Mt. Halimun National park, Indonesia:
Evidence for reproductive isolation among three sympatric rbcL sequence
types. Amer J Bot 88 (8): 1517-1522.
308
Yatabe Y, Takamiya M, Murakami N. 1998. Variation in the rbcL sequence of
Stegnogramma pozoi subsp. mollisima (Thelypteridaceae) in Japan. J Plant
Res 111: 557-564.
Yatabe Y, Nishida H, Murakami N. 1999. Phylogeny of Osmundaceae inferred
from rbcL nucleotide sequence and comparison to the fossil evidences. J
Plant Res 112: 397-404.
309
Appendix 1. Matrix of 88 Morphological Characters for Maximum Parsimony
Characters and Character StatesNo. Species
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 251. Athyrium anisopterum 0 0 0 1 1 0 2 2 2 0 0 - 0 0 0 1 - 0 0 1 1 4 4 7 12. D. accedens 0 0 0 0 1 {05} 4 2 2 1 1 1 1 1 2 4 6 1 0 0 1 4 3 0 63. D. acuminatum 0 1 2 0 1 {03} 3 1 4 0 1 ? 1 0 0 2 1 0 0 0 1 2 3 1 14. D. aequibasale 0 0 0 0 1 {26} 1 2 4 0 0 0 0 1 0 1 0 0 0 0 1 4 2 0 ?5. D. albidosquamatum 0 0 2 4 1 {24} 4 1 4 0 1 0 0 1 2 3 4 0 0 1 2 5 5 8 46. D. angustipinna 0 0 1 0 1 {45} 2 1 3 0 0 0 0 1 0 2 4 0 0 0 1 2 2 0 47. D. asymmetricum 0 0 2 0 1 {45} 1 2 4 0 - - - 1 2 4 4 0 0 0 1 4 5 {10} 48. D. atrosquamosum 0 0 2 2 1 6 2 5 3 1 0 - 0 1 1 5 3 1 0 0 1 4 5 4 39 D. bantamense 0 {01} 1 2 1 2 5 2 4 1 2 1 0 1 1 4 3 0 0 0 2 4 2 0 310. D. barbatum 0 0 1 2 1 {02} 3 1 3 0 2 1 0 0 2 2 3 0 0 0 1 3 5 1 311. D. batuayauense 0 0 1 0 1 {02} 2 0 3 0 0 0 - 1 0 1 2 0 0 0 1 4 3 8 212. D. beamanii 0 0 ? 2 1 6 4 4 2 ? 0 ? 0 0 2 5 5 0 0 0 2 0 5 8 513. D. betimusense 0 0 2 0 0 {67} 1 4 4 1 0 0 - ? 2 3 3 2 0 0 1 5 5 ? 314. D. christii 0 0 1 0 1 ? 2 1 3 0 0 0 - 0 0 2 3 0 0 0 1 5 7 1 315. D. cordifolium 0 0 1 3 1 5 2 2 1 1 0 0 0 1 1 3 2 0 0 0 2 4 {02} 0 216. D. crameri 0 0 1 0 1 2 2 3 4 1 0 ? 0 0 1 3 3 0 0 0 1 4 2 0 317. D. crenatoserratum 0 0 1 0 1 1 1 0 4 1 1 0 0 0 0 2 1 0 0 0 1 4 3 2 218. D. crinitum 0 0 1 4 1 2 2 1 0 0 0 2 1 0 1 2 1 1 0 0 2 4 5 3 219. D. cumingii 0 0 1 0 1 4 2 1 4 1 0 0 - 0 1 2 2 0 0 0 1 5 2 0 220. D. densisquamatum 0 0 2 4 1 {26} 5 2 4 1 0 0 1 0 2 3 4 1 0 1 1 5 5 {10} 521. D. dilatatum 0 0 2 2 1 0 6 1 4 1 2 1 0 1 2 6 7 0 0 0 2 4 {56} 4 522. D. dolichosorum 0 ? 2 2 1 0 2 ? 4 0 ? ? ? ? 2 2 6 2 0 0 2 ? 5 8 723. D. donianum 1 3 1 0 1 {26} 2 3 3 0 1 1 0 1 0 3 2 0 0 0 1 0 2 0 224. D. esculentum 1 0 2 0 1 0 3 1 4 0 2 1 0 1 2 2 6 0 0 0 0 4 {45} 5 225. D. fraxinifolium 0 0 1 0 1 2 5 2 3 0 0 0 ? 0 1 4 3 0 0 0 1 0 2 0 226. D. fuliginosum 0 0 1 4 1 0 2 2 5 0 0 0 0 0 0 0 2 0 1 1 2 6 1 7 3
310
Appendix 1. Continued
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 2527. D.halimunense 0 0 1 2 1 2 2 1 4 1 2 1 0 1 1 2 - 0 0 0 1 0 2 0 ?28. D. hewittii 0 0 1 0 1 6 1 3 4 1 0 0 1 0 1 3 2 0 0 0 5 3 {45} 2 429. D. hottae 0 0 1 0 1 2 5 2 1 1 0 0 - 1 1 4 3 0 0 0 1 0 2 0 430. D. insigne 0 0 2 1 1 {05} 6 2 2 0 2 1 0 1 2 7 9 2 0 0 2 4 5 0 931. D. Kunstlerii 0 0 2 0 0 7 ? ? 4 1 0 0 1 0 2 4 5 1 0 0 1 5 5 4 632. D. laevipes 0 0 2 0 0 7 1 5 4 1 0 0 0 0 2 6 1 0 0 0 1 5 {45} 4 333. D. latisquamatum 0 0 2 1 1 7 5 5 4 1 0 0 2 0 2 7 1 0 0 1 1 5 {67} 2 734. D. lobbianum 0 0 1 2 1 4 5 9 3 1 2 1 0 1 1 5 1 0 0 0 4 5 2 3 335. D. lomariaceum 0 0 1 4 1 0 3 1 5 0 - - - - 0 1 0 0 0 1 2 6 1 7 336. D. Loerzingii 0 0 1 2 1 2 3 1 4 1 0 0 0 1 1 3 1 0 0 0 2 4 3 1 337. D. malaccense 0 0 1 2 1 3 4 1 3 0 0 0 0 - 1 3 0 0 0 0 2 4 3 2 338. D.megasegmentum 0 0 2 0 0 7 2 7 4 1 0 0 1 0 2 9 1 1 0 0 1 5 6 4 639. D.megasimplicifolium 1 0 0 0 1 ? ? ? ? ? ? ? - - 0 1 0 0 0 0 0 4 0 7 240. D. meijerii ? 3 2 0 1 {02} 3 1 5 0 1 1 0 1 2 4 0 0 0 0 1 4 5 9 341. D. melanolepis 0 0 2 0 1 2 3 2 5 1 1 1 0 0 2 4 1 0 0 1 2 5 {45} ? -42. D. moultonii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? - - ? ? ? ? ? 6 ? -43. D. pallidum 0 0 1 2 1 3 3 1 4 1 0 0 1 0 1 3 0 0 0 0 2 4 {23} 0 244. D. parallelivenium 0 0 2 0 1 2 3 1 4 1 1 1 0 1 2 4 1 0 0 0 1 2 5 4 545. D. petiolare 0 0 1 0 1 3 ? ? 4 0 2 2 0 0 0 3 0 0 0 0 1 5 3 2 246. D. poiense 0 0 1 2 1 {02} 2 1 3 0 0 0 0 0 1 2 0 0 0 0 2 4 {45} 1 247. D. polypodioides 0 0 2 2 1 0 6 1 4 1 2 2 0 1 2 5 1 0 0 0 2 4 {56} 9 648. D. porphyrorachis 0 0 1 2 1 0 2 0 4 1 ? ? ? ? 0 1 0 0 2 1 2 6 1 8 049. D. Prescottianum 0 0 1 0 1 {02} 4 2 4 1 0 0 0 0 1 2 1 0 0 0 0 0 3 0 250. D. procumbens 1 3 2 0 1 {02} ? ? 4 0 1 2 0 0 1 2 3 0 1 0 0 5 5 {10} 351. D. profluens 0 0 2 0 0 7 ? 1 4 1 0 0 0 0 2 5 5 1 0 0 1 5 5 4 -52. D. riparium 0 0 1 0 1 2 2 1 5 1 0 0 1 0 1 3 3 0 0 0 1 6 2 0 2
311
Appendix 1. Continued
Characters and Character StatesNo. Species
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 2553. D. silvaticum 0 0 1 0 1 {34} 4 1 4 0 2 2 0 0 0 2 2 0 0 0 1 5 3 2 154. D. simplicivenium 0 0 2 2 1 {02} 6 2 4 1 1 1 0 1 2 5 9 0 0 0 2 5 5 4 655. D. sorzogonense 0 0 2 2 1 0 5 5 4 1 0 0 0 1 1 3 7 0 1 1 2 5 3 1 456. D. speciosum 0 0 1 2 1 0 6 5 3 0 1 2 0 0 1 4 4 0 0 0 2 4 3 1 557. D. spiniferum 0 0 2 0 0 7 1 6 4 1 0 0 1 0 2 4 6 0 0 1 5 0 5 2 658. D. squarrasum 0 0 2 4 1 {26} 3 1 2 0 2 ? 0 0 2 2 3 1 1 1 2 5 3 0 459. D. subintegrum 0 0 1 0 1 {02} 6 3 3 1 0 0 - - 1 4 3 0 0 0 0 3 2 2 360. D.subalternisegmentum ? ? ? ? 1 2 4 1 5 1 2 2 0 0 2 - 5 2 0 0 2 5 7 ? ?61. D. subpolypodioides 0 0 2 0 1 0 ? 3 4 1 - - - - 2 - - 0 0 0 5 5 5 1 ?62. D. subserratum 0 0 1 0 1 1 0 0 4 0 1 0 0 0 0 2 1 0 1 0 1 0 0 8 263. D. subvirescens 0 3 2 0 1 2 4 1 4 1 1 2 0 1 2 2 1 0 0 1 1 5 5 {10} 364. D. tomentosum 0 0 1 0 1 1 1 0 4 0 0 0 1 0 0 2 1 0 2 1 2 5 3 2 165 D. tricholepis 0 0 1 2 1 {02} 2 1 2 0 2 1 0 0 1 1 2 0 0 0 2 4 4 7 466. D. umbrosum 0 0 2 2 1 3 - - 4 1 1 1 0 0 1 2 3 0 0 0 2 4 {56} 3 267. D. velutinum 0 0 1 4 1 0 4 1 5 1 0 0 0 0 0 2 2 0 2 1 2 5 5 2 168. D. vestitum 0 0 2 0 1 2 - - 3 1 2 2 0 1 2 6 7 2 0 0 1 2 5 3 669. D. wahauense 0 0 1 0 0 2 - - 5 1 0 0 0 0 0 0 1 0 0 0 1 6 2 0 070. D. xiphophyllum 0 0 1 0 1 4 9 4 2 1 0 0 1 0 2 4 4 0 0 0 1 0 3 0 3
312
Appendix 1. Continued
Species Characters & Character States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
1. Athyriumanisopterum
0 3 1 0 2 1 2 2 2 1 4 0 0 8 3 8 {13} 2 1 0 0 0 0 - -
2. D. accedens 2 3 1 0 3 2 1 2 3 1 0 3 1 {24} {24} {13} {14} 1 2 0 0 0 0 - -3. D. acuminatum 1 3 1 0 {23} 2 1 1 3 2 3 1 0 4 1 8 2 {23} 2 0 0 0 0 - -4. D. aequibasale ? 3 1 0 0 2 1 1 3 2 5 1 ? 4 3 1 4 1 2 0 0 0 0 - -5. D.
albidosquamatum? 3 1 0 1 2 1 1 3 2 5 2 2 5 4 9 5 3 2 {12} 0 0 0 2 2
6. D. angustipinna 5 3 1 0 {01} 2 1 1 3 3 2 2 0 1 1 1 1 1 2 0 0 0 0 0 -7. D. asymmetricum 5 3 1 0 {12} 2 1 1 3 2 4 7 7 5 4 9 5 6 3 2 1 2 3 8 98. D.
atrosquamosum4 3 1 0 1 2 1 1 3 2 5 3 3 5 4 9 5 6 2 2 1 2 4 4 3
9. D. bantamense 2 3 1 0 {01} 2 1 1 3 2 0 2 0 7 6 2 2 1 4 0 0 0 0 - -10. D. barbatum 2 3 1 0 1 2 1 1 3 2 5 1 1 5 7 9 5 7 2 1 1 1 2 - ?11. D. batuayauense 2 3 1 0 2 2 1 1 3 2 5 1 0 1 1 8 2 2 2 0 1 0 0 - -12. D. beamanii ? 3 1 0 ? 2 ? ? 3 {12} 1 5 4 5 4 9 5 6 3 3 3 2 {34} 4 313. D.betimusense ? 3 1 0 ? 2 1 1 3 2 4 5 4 5 4 9 5 6 2 2 3 2 4 4 414. D. christii 2 3 1 0 3 2 1 1 3 2 5 2 0 4 3 8 3 2 3 0 0 0 0 - -15. D. cordifolium 1 3 1 1 {01} 2 1 1 3 1 0 2 1 6 5 1 1 1 4 0 0 0 0 - -16. D. crameri 2 3 1 0 1 2 1 1 3 2 5 1 1 4 3 1 3 1 3 0 0 0 0 - -17. D.
crenatoserratum1 3 1 1 {23} 1 1 1 3 2 3 1 0 8 4 {58} {12} {12} 2 0 0 0 0 - -
18. D. crinitum 1 3 1 0 ? 2 1 1 3 2 0 2 2 5 4 9 6 6 2 3 3 1 2 - 219. D. cumingii 2 3 1 0 0 2 1 1 3 2 0 2 1 4 3 1 1 5 4 0 0 0 0 - -20. D.
densisquamatum6 3 1 0 2 1 1 1 3 2 5 5 3 5 4 9 5 6 2 4 3 2 3 3 3
21. D. dilatatum 7 3 1 0 {12} 2 1 1 3 3 0 0 9 5 4 9 5 6 3 3 3 2 3 9 622. D. dolichosorum ? 3 1 0 ? 2 1 2 3 2 2 6 5 1 1 9 5 6 3 3 3 2 2 5 323. D. donianum 1 3 1 0 0 2 1 1 3 2 5 2 1 4 3 1 1 1 2 0 0 0 0 - 2
313
Appendix 1. Continued
Species Character & Character States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
24. D. esculentum 2 3 1 0 {12} 2 1 1 3 3 0 4 3 5 4 9 5 6 2 3 1 2 2 4 325. D. fraxinifolium 4 3 1 0 {01} 1 1 1 3 2 2 3 1 4 3 {58} {24} 1 {35} 0 0 0 0 - -26 D. fuliginosum 0 3 1 0 5 1 1 3 1 1 5 6 0 1 1 2 3 1 1 0 0 0 0 - -27. D.halimunense ? 3 1 0 0 2 1 1 3 1 5 2 1 4 3 1 2 1 2 0 0 0 0 - -28. D. hewittii 2 3 1 0 {12} 2 1 1 3 2 6 2 1 5 4 9 5 6 2 1 2 1 7 1 129. D. hottae 3 3 1 0 {23} 1 1 1 3 2 6 3 0 4 3 1 4 1 2 0 0 0 0 - -30. D. insigne ? 3 1 0 ? 2 1 2 3 2 1 6 3 5 4 9 5 6 2 3 2 1 2 4 331. D. Kunstlerii 6 3 1 0 2 2 1 1 3 2 5 7 4 5 4 9 5 6 2 {23} 3 1 4 4 432. D. laevipes 3 3 1 0 {12} 2 1 2 3 2 7 3 3 5 4 9 5 6 2 2 3 1 3 4 333. D. latisquamatum 6 3 1 0 1 2 1 1 3 2 2 5 4 5 4 9 5 6 2 {23} 3 2 4 4 334. D. lobbianum 2 3 1 0 3 2 1 1 3 1 2 1 0 2 6 1 3 1 2 0 0 0 0 - -35. D. lomariaceum 1 3 - 0 - 0 0 0 0 0 0 - - 1 0 0 0 0 0 0 0 0 0 - -36. D. Lorzingii 2 3 1 0 {34} 2 1 1 3 2 6 2 0 1 1 7 3 0 2 0 0 0 0 - -37. D. malaccense 2 3 1 0 {23} 2 2 1 3 2 4 2 0 1 8 {04} {13} 2 2 0 0 0 0 - -38. D. megasegmentum 1 3 1 0 2 2 1 1 3 2 6 6 6 5 4 9 5 6 2 4 3 2 4 6 539. D.
megasimplicifolium1 0 - 2 - 0 0 0 0 0 0 - - 0 0 0 0 0 0 0 0 0 0 - -
40. D. meijerii 5 3 1 0 2 2 1 2 3 {12} 6 3 3 5 4 8 5 6 3 2 3 {12} 4 3 341. D. melanolepis ? 3 1 0 ? 2 1 1 3 2 6 4 2 1 1 9 5 6 2 3 2 1 4 2 042. D. moultonii 2 3 ? - ? - 1 1 3 2 5 3 2 2 2 9 5 6 2 4 3 1 3 3 243. D. pallidum 2 3 1 0 {34} 2 1 1 3 1 4 2 0 1 6 {67} 3 1 2 0 0 0 0 - -44. D. parallelivenium 9 3 1 0 1 2 1 1 3 2 3 6 7 5 7 9 5 6 2 {23} 3 2 4 7 545. D. petiolare 2 3 1 0 {23} 2 1 1 3 1 6 1 0 4 3 8 3 3 2 0 0 0 0 - -46. D. poiense 1 3 1 0 4 2 1 1 3 1 4 1 0 1 3 7 5 4 2 0 0 0 0 - -47. D. polypodioides 7 3 1 0 1 2 1 1 3 2 5 6 4 1 2 9 5 5 2 {34} 3 1 3 6 448. D. porphyrorachis 0 3 0 0 - 0 0 0 0 0 0 - - 0 0 0 0 0 0 0 0 0 0 - -49. D. Prescottianum 2 3 1 0 2 2 1 1 3 2 4 2 0 {78} 1 4 4 1 3 0 0 0 0 - -
314
Appendix 1. Continued
Species Character & Characters States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
50. D. procumbens ? 3 1 0 1 2 1 1 3 2 5 5 4 5 2 9 5 6 2 {23} 3 1 3 3 551. D. profluens ? 3 1 0 1 2 1 1 3 2 6 6 5 5 2 9 5 6 2 3 3 2 4 7 652. D. riparium 2 3 1 0 0 2 1 1 3 2 2 3 1 5 3 1 1 1 4 0 0 0 0 - 353. D. silvaticum 2 3 2 0 {12} 2 1 1 3 1 6 1 0 8 3 8 3 2 2 0 0 0 0 - 154. D. simplicivenium 7 3 1 0 1 2 1 1 3 2 5 4 3 5 4 9 5 6 2 {34} 3 1 3 5 355. D. sorzogonense 3 3 1 0 {23} 1 2 1 3 1 6 2 1 {25} {25} 8 3 {46} 2 0 0 0 0 - -56. D. speciosum 2 3 1 0 {23} 2 1 1 3 1 2 3 0 2 1 8 3 4 2 0 0 0 0 - -58. D. squarrasum ? 3 2 0 3 2 1 1 3 2 5 1 1 {48} {17} 1 1 1 2 0 0 0 1 - -59. D. subintegrum 2 3 1 0 1 2 1 1 3 1 4 3 1 4 3 7 3 1 2 0 0 0 1 0 -60. D.subalternisegmentum ? ? 1 0 ? 2 1 1 3 2 5 - ? 5 4 9 5 7 2 4 3 2 3 5 461. D. subpolypodioides ? 3 0 {12} 1 1 1 1 3 2 2 - ? 2 2 9 5 7 2 3 3 1 3 ? ?62. D. subserratum 0 1 0 - - 0 0 0 0 0 0 - ? 0 0 0 0 0 0 0 0 0 0 ? -63. D. subvirescens 4 3 0 1 1 1 1 1 3 2 5 3 ? 3 2 {89} 5 6 2 2 3 2 5 2 464. D. tomentosum 0 3 1 0 3 1 2 1 2 1 3 0 0 {18} 1 8 3 {12} 2 0 0 0 0 - -65. D. tricholepis 2 3 1 0 3 2 2 1 1 1 5 ? ? 1 1 8 2 6 3 0 0 0 0 - ?66. D. umbrosum 2 3 1 0 3 2 1 2 3 2 6 2 1 2 2 9 5 7 2 3 3 1 3 1 167. D. velutinum 1 3 2 1 2 2 1 1 2 2 2 1 1 1 1 {68} 2 2 2 1 2 1 3 0 068. D. vestitum 8 3 2 1 2 1 1 1 2 3 3 7 4 2 2 9 5 6 1 2 3 2 3 5 469. D. wahauense 0 3 1 1 0 2 1 1 2 1 2 0 0 4 3 1 4 1 {34} 0 0 0 0 - -70. D. xiphophyllum 3 3 1 0 1 2 1 1 3 2 3 4 1 4 3 1 {23} 1 2 0 0 0 0 - -
315
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
1. Athyrium anisopterum 0 0 0 0 2 ? 3 15 3 ? 0 12 0 1 0 0 1 0 1 1 0 ? 1 0 02. D. accedens 0 0 0 0 0 0 0 0 2 6 0 0 1 2 1 0 2 0 0 0 6 - 3 3 13. D. acuminatum 0 0 0 0 1 ? 1 5 3 5 0 0 1 0 0 0 1 0 1 0 0 1 0 2 14. D. aequibasale 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - - 2 0 2 3 3 05. D. albidosquamatum {23} 3 2 2 1 4 0 1 3 1 0 2 0 0 0 0 0 0 1 0 0 - 0 2 16. D. angustipinna 0 0 0 0 0 0 1 0 1 2 0 2 0 0 0 1 - 1 1 1 0 - 3 2 37. D. asymmetricum 1 4 4 3 2 7 3 5 3 5 0 2 0 2 0 0 1 1 1 1 0 - 2 1 08. D. atrosquamosum 0 4 4 3 3 1 2 5 3 - 0 0 1 2 0 0 1 0 1 0 0 - 0 0 09. D. bantamense 0 0 0 0 0 0 0 0 0 5 0 0 1 0 0 1 - 1 1 2 0 - 3 3 010. D. barbatum 1 1 1 2 0 - 0 0 3 1 0 4 0 2 0 0 2 1 1 1 0 - 0 2 011. D. batuayauense 0 0 0 0 0 3 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 - 2 1 112. D. beamanii 1 4 4 3 2 3 1 {56} 3 ? 0 ? 0 0 0 0 1 0 1 0 0 ? 0 1 113. D. betimusense {13} 4 2 3 1 4 1 3 3 6 0 0 0 0 0 0 - - 1 0 0 1 2 0 014. D. christii 0 0 0 0 0 4 0 0 3 1 0 0 0 0 0 0 1 0 1 2 0 - 0 2 015. D. cordifolium 0 0 0 0 0 0 0 0 1 3 0 0 1 0 1 1 - - 0 2 3 2 3 3 116. D. crameri 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 1 1 0 2 3 3 017. D. crenatoserratum 0 0 0 0 0 2 0 0 3 5 0 0 0 0 0 1 - - 1 1 2 - 3 3 118. D. crinitum 3 4 2 3 2 - 1 7 3 0 0 2 0 2 0 0 1 0 1 0 1 - 0 2 019. D. cumingii 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - 1 1 2 2 - 3 3 020. D. densisquamatum 1 4 3 3 1 1 1 5 3 4 0 2 0 2 0 0 1 0 1 1 1 - 0 1 121. D. dilatatum {14} 4 2 4 1 8 3 4 3 6 0 0 0 2 0 0 2 1 0 1 1 0 0 2 122. D. donianum 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 1 2 1 1 3 3 023. D. dolichosorum 1 2 2 3 1 ? 1 5 3 6 0 0 0 0 0 0 1 0 1 0 1 ? 0 2 024. D. esculentum {15} 4 2 3 1 1 3 4 3 6 0 0 0 0 1 0 2 - 0 0 1 - 3 3 125. D. fraxinifolium 0 0 0 0 1 0 0 0 1 2 0 0 1 0 1 1 - - 0 2 1 {12} 3 3 126. D. fuliginosum 0 0 0 0 1 ? 0 0 3 5 0 4 0 2 0 1 - - 1 1 0 - 3 3 027. D.halimunense 0 0 0 0 1 0 0 0 1 2 0 0 0 0 0 1 - - 1 2 0 2 3 3 0
316
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
28. D. hewittii {35} 3 1 3 2 ? 0 0 3 0 0 0 0 0 0 1 - - 1 1 0 - 0 3 029. D. hottae 0 0 0 0 1 0 0 0 1 2 0 0 1 0 0 1 - - 1 2 0 0 3 3 030. D. insigne 3 2 0 0 0 0 0 0 3 1 0 0 0 0 1 0 2 0 0 0 6 - 3 3 031. D. Kunstlerii {13} 4 3 1 2 4 3 5 3 6 0 0 1 0 0 0 0 1 1 0 1 - 2 1 032. D. laevipes 2 4 {23} 3 2 2 1 5 3 6 0 1 1 1 0 0 2 1 1 0 1 2 0 2 033. D. latisqua-matum {24} 3 4 3 1 2 1 2 3 6 0 0 0 0 0 1 1 1 0 1 1 1 0 1 034. D. lobbianum 0 0 0 0 0 0 0 0 2 3 0 0 0 0 0 1 - - 1 2 1 2 3 2 035. D. lomariaceum 0 0 0 0 0 9 0 0 0 6 0 3 0 - 0 1 - - 1 ? 1 - 0 3 036. D. Lorzingii 0 0 0 0 0 5 1 6 3 2 0 0 1 0 0 0 2 0 1 0 1 2 2 2 137. D. malaccense 0 0 0 0 0 4 3 5 3 1 0 0 0 0 0 0 2 0 1 0 1 - 3 3 038. D.megaseg-mentum 1 4 7 3 1 7 3 7 3 6 1 0 0 0 0 0 3 1 1 1 0 - 0 1 139. D.megasimpli-cifolium 0 0 0 0 1 0 0 0 0 2 - - 0 0 1 - - 2 1 2 5 1 3 3 040. D. meijerii 1 4 3 3 2 4 {13} 1 3 6 0 0 0 0 0 0 {01} {01} 1 1 0 - 2 {23} 141. D. melanolepis 1 4 2 2 2 0 1 5 3 5 0 2 0 2 0 1 - - 1 2 0 - 0 2 042. D. moultonii 1 4 7 3 1 2 3 6 3 1 0 0 0 0 0 0 1 0 - 1 0 0 0 0 043. D. pallidum 0 0 0 0 0 0 0 0 2 5 0 2 0 2 0 - - - 1 1 0 - 3 3 044. D.parallelive-nium 1 4 {12} 4 1 ? 1 5 3 6 0 0 0 0 0 0 1 1 1 0 0 2 0 1 045. D. petiolare 0 0 0 0 0 3 0 0 3 0 0 0 0 0 0 0 1 0 1 0 0 2 3 3 046. D. poiense 0 0 0 0 0 2 0 0 3 4 1 0 0 0 0 0 ? 0 1 0 0 - 2 2 147. D. polypodioides 2 4 0 3 2 2 3 4 3 6 0 0 0 0 0 0 2 1 1 0 0 - 3 3 048. D. porphyrorachis 0 0 0 0 0 6 2 0 0 2 1 3 0 0 0 1 - - 1 0 0 - 3 3 049. D. Prescottianum 0 0 0 0 0 0 0 0 2 5 0 0 0 0 0 1 - - 1 2 0 - 3 3 050. D. procumbens 1 4 3 3 1 5 3 5 3 1 0 2 0 2 0 0 1 1 1 0 0 2 0 1 051. D. profluens 1 4 6 3 2 - 4 5 3 4 1 0 0 0 0 0 2 1 1 0 0 2 0 1 052. D. riparium 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 0 2 1 2 3 3 053. D. silvaticum 0 0 0 0 0 5 0 0 3 1 0 2 1 1 0 0 0 0 0 0 0 - 3 3 154. D. simplicivenium 1 4 {12} 3 1 4 1 5 3 4 0 0 0 2 0 0 1 0 1 0 0 - 3 2 1
317
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
55. D. sorzogonense 0 0 0 3 1 4 3 7 3 6 1 3 0 2 0 0 2 1 - 0 0 - 2 2 056. D. speciosum 0 0 0 0 0 3 0 0 3 0 0 0 0 0 0 0 2 0 1 0 0 - 0 2 057. D. spiniferum 3 4 1 3 1 1 2 5 3 4 0 0 0 0 0 0 0 0 1 0 0 - 2 0 058. D. squarrasum 0 0 0 0 0 0 0 0 2 1 0 4 0 2 1 1 3 - 1 2 5 2 3 3 059. D. subintegrum 0 0 0 0 0 0 0 0 2 5 0 0 0 0 0 1 - - 1 2 0 - 0 1 060. D.subalternisegmentum 1 4 7 3 1 ? 4 7 3 - 0 0 0 0 0 0 2 1 1 2 0 - 0 2 061. D. subpolypodioides 1 4 6 3 2 - 4 6 3 2 0 0 0 0 0 0 2 1 1 1 0 - 0 1 162. D. subserratum 0 0 0 0 0 - 0 0 1 6 1 - - 2 0 1 - 1 1 2 0 2 3 3 063. D. subvirescens {14} 4 {67} 3 1 1 4 7 3 1 0 0 0 0 0 0 1 1 - 1 0 - 2 1 064. D. tomentosum 0 0 0 0 1 1 1 1 3 1 0 3 0 0 0 0 1 0 0 0 0 2 3 3 065. D. tricholepis 0 0 0 0 1 - {24} 5 3 1 0 2 0 2 0 {01} 2 1 1 1 0 0 3 3 066. D. umbrosum {24} 4 6 1 {12} 2 6 3 1 1 0 0 0 0 0 1 1 1 1 0 0 - 2 2 067. D. velutinum 3 4 0 1 2 - 2 3 0 0 0 0 0 0 1 2 1 1 1 1 0 - 3 2 168. D. vestitum 3 4 1 1 3 - 5 3 2 1 0 0 0 0 0 1 1 1 1 1 0 - 3 2 169. D. wahauense 0 0 0 0 1 - 0 0 1 1 0 0 0 0 0 1 0 - 1 2 0 - 3 3 070. D. xiphophyllum 0 0 0 0 1 - 0 0 1 2 0 2 1 1 0 1 - - 1 2 1 1 3 3 1
318
Appendix 1. Continued
76 77 78 79
80 81 82 83 84 85 86 87 88
1. Athyrium anisopterum {12} 0 0 1 1 1 0 0 0 0 1 1 ?2. D. accedens 2 2 2 0 1 1 0 1 0 0 1 0 03. D. acuminatum 2 2 2 0 1 1 1 1 0 0 1 1 04. D. aequibasale 2 1 2 0 0 1 1 0 1 0 1 0 05. D. albidosquamatum 2 2 2 0 1 1 0 2 0 - 1 1 06. D. angustipinna 1 1 2 0 0 1 1 0 0 0 0 1 07. D. asymmetricum 1 2 2 1 1 1 0 0 0 0 1 1 08. D. atrosquamosum 2 1 2 1 1 1 0 0 0 2 1 0 09 D. bantamense 1 1 2 1 1 1 0 1 0 0 1 0 010. D. barbatum 0 1 2 0 1 1 0 0 0 0 1 1 011. D. batuayauense 1 - 2 0 1 1 0 1 0 0 1 1 012. D. beamanii 2 2 2 1 1 1 0 ? ? 2 1 ? 013. D. betimusense 1 0 2 0 0 1 0 2 ? 2 0 0 014. D. christii 2 1 2 0 1 0 0 1 0 0 1 0 015. D. cordifolium 1 2 2 1 1 1 0 1 0 0 1 0 016. D. crameri 1 1 2 1 1 1 0 1 0 0 1 1 017. D. crenatoserratum 1 1 2 1 1 1 0 1 0 0 1 1 018. D. crinitum 2 1 2 1 1 1 0 2 1 0 1 1 019. D. cumingii 1 2 2 1 1 1 0 2 0 0 1 1 020. D. densisquamatum 2 0 2 0 1 1 0 2 0 2 1 0 021. D. dilatatum 2 2 2 0 1 1 0 1 0 0 1 0 022. D. dolichosorum 2 2 2 1 1 1 0 1 0 ? 1 1 023. D. donianum 1 1 2 1 1 1 1 1 0 1 1 1 024. D. esculentum 2 2 2 0 0 1 1 2 0 2 1 0 025. D. fraxinifolium 2 2 2 1 1 1 1 0 0 1 1 1 026. D. fuliginosum 0 0 2 1 1 1 0 2 0 0 1 1 027. D.halimunense 1 2 2 1 1 1 0 1 0 0 1 1 028. D. hewittii 1 1 2 0 1 1 0 1 0 0 1 0 0
319
Appendix 1. Continued
Species Character & Character States76 77 78 79 80 81 82 83 84 85 86 87 88
29. D. hottae 1 1 2 0 1 1 1 1 0 0 1 1 030. D. insigne 2 1 2 0 0 1 0 2 0 0 1 0 031. D. Kunstlerii 2 1 2 0 0 0 0 2 0 2 1 0 032. D. laevipes 1 2 2 0 1 1 0 2 0 2 1 0 033. D. latisquamatum 2 1 2 1 1 1 1 1 1 0 2 0 034. D. lobbianum 1 1 2 1 1 1 0 1 0 0 2 1 035. D. lomariaceum 0 0 2 0 1 1 0 2 0 1 1 0 036. D. Lorzingii 1 2 2 1 1 1 0 1 0 0 1 1 037. D. malaccense 2 1 2 0 1 1 0 0 0 0 1 1 038. D.megasegmentum 2 2 2 1 1 0 0 1 0 2 1 0 039. D.megasimplicifo-
lium1 1 2 1 1 1 1 1 0 0 1 0 0
40. D. meijerii 2 3 2 0 1 1 0 0 0 2 1 1 041. D. melanolepis 1 1 2 1 1 1 1 2 0 0 1 1 042. D. moultonii 1 1 2 1 1 1 0 1 0 2 1 0 043. D. pallidum 0 0 2 0 1 1 0 0 0 0 1 1 044. D. parallelivenium 1 1 2 0 1 1 1 1 1 2 1 0 045. D. petiolare 2 2 2 - - - - - - - - - 046. D. poiense 1 0 2 1 1 1 0 0 0 0 1 1 047. D. polypodioides 2 2 2 0 1 1 0 0 0 0 1 0 048. D. porphyrorachis 0 0 2 1 1 1 0 0 0 0 1 0 049. D. Prescottianum 0 0 2 0 1 1 0 0 0 0 1 1 050. D. procumbens 2 2 2 1 1 1 0 1 0 2 1 0 051. D. profluens 1 2 2 0 1 1 0 2 0 0 1 1 052. D. riparium 1 1 2 0 1 1 0 0 0 0 1 0 053. D. silvaticum 2 2 2 1 1 1 0 0 2 1 1 0 054. D. simplicivenium 2 2 2 1 1 1 0 1 0 2 1 0 055. D. sorzogonense 2 2 1 1 1 1 0 2 1 0 1 1 0
320
Appendix 1. Continued
Species Characters & character States76 77 78 79 8
081 82 83 84 85 86 87 88
56. D. speciosum 2 1 2 1 1 1 0 1 0 0 1 0 057. D. spiniferum 2 1 2 0 1 1 1 2 0 2 1 0 058. D. squarrasum 1 0 2 0 1 1 0 1 0 0 1 0 059. D. subintegrum 1 2 2 1 1 1 1 0 0 0 1 1 060. D. subalternisegmen-
tum2 0 2 1 1 1 0 1 0 0 1 0 0
61. D. subpolypodioides 2 1 2 0 1 1 0 2 0 0 1 1 062. D. subserratum 1 1 2 1 1 1 1 0 0 0 1 1 063. D. subvirescens 2 2 2 0 1 0 0 1 0 2 1 0 064. D. tomentosum 1 1 2 1 1 1 0 2 0 0 1 1 065. D. tricholepis 2 3 2 0 1 1 0 1 0 0 1 1 066. D. umbrosum 1 1 {12} 1 1 1 0 0 1 0 1 0 067. D. velutinum 2 2 2 1 1 1 0 1 0 0 1 0 068. D. vestitum 2 2 2 0 1 1 0 2 0 0 1 1 069. D. wahauense 1 1 2 0 1 1 1 2 0 0 1 0 070. D. xiphophyllum 1 2 2 1 1 1 0 0 0 0 1 1 0
309
Appendix 1. Matrix of 88 Morphological Characters for Maximum Parsimony
Characters and Character StatesNo. Species
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 251. Athyrium anisopterum 0 0 0 1 1 0 2 2 2 0 0 - 0 0 0 1 - 0 0 1 1 4 4 7 12. D. accedens 0 0 0 0 1 {05} 4 2 2 1 1 1 1 1 2 4 6 1 0 0 1 4 3 0 63. D. acuminatum 0 1 2 0 1 {03} 3 1 4 0 1 ? 1 0 0 2 1 0 0 0 1 2 3 1 14. D. aequibasale 0 0 0 0 1 {26} 1 2 4 0 0 0 0 1 0 1 0 0 0 0 1 4 2 0 ?5. D. albidosquamatum 0 0 2 4 1 {24} 4 1 4 0 1 0 0 1 2 3 4 0 0 1 2 5 5 8 46. D. angustipinna 0 0 1 0 1 {45} 2 1 3 0 0 0 0 1 0 2 4 0 0 0 1 2 2 0 47. D. asymmetricum 0 0 2 0 1 {45} 1 2 4 0 - - - 1 2 4 4 0 0 0 1 4 5 {10} 48. D. atrosquamosum 0 0 2 2 1 6 2 5 3 1 0 - 0 1 1 5 3 1 0 0 1 4 5 4 39 D. bantamense 0 {01} 1 2 1 2 5 2 4 1 2 1 0 1 1 4 3 0 0 0 2 4 2 0 310. D. barbatum 0 0 1 2 1 {02} 3 1 3 0 2 1 0 0 2 2 3 0 0 0 1 3 5 1 311. D. batuayauense 0 0 1 0 1 {02} 2 0 3 0 0 0 - 1 0 1 2 0 0 0 1 4 3 8 212. D. beamanii 0 0 ? 2 1 6 4 4 2 ? 0 ? 0 0 2 5 5 0 0 0 2 0 5 8 513. D. betimusense 0 0 2 0 0 {67} 1 4 4 1 0 0 - ? 2 3 3 2 0 0 1 5 5 ? 314. D. christii 0 0 1 0 1 ? 2 1 3 0 0 0 - 0 0 2 3 0 0 0 1 5 7 1 315. D. cordifolium 0 0 1 3 1 5 2 2 1 1 0 0 0 1 1 3 2 0 0 0 2 4 {02} 0 216. D. crameri 0 0 1 0 1 2 2 3 4 1 0 ? 0 0 1 3 3 0 0 0 1 4 2 0 317. D. crenatoserratum 0 0 1 0 1 1 1 0 4 1 1 0 0 0 0 2 1 0 0 0 1 4 3 2 218. D. crinitum 0 0 1 4 1 2 2 1 0 0 0 2 1 0 1 2 1 1 0 0 2 4 5 3 219. D. cumingii 0 0 1 0 1 4 2 1 4 1 0 0 - 0 1 2 2 0 0 0 1 5 2 0 220. D. densisquamatum 0 0 2 4 1 {26} 5 2 4 1 0 0 1 0 2 3 4 1 0 1 1 5 5 {10} 521. D. dilatatum 0 0 2 2 1 0 6 1 4 1 2 1 0 1 2 6 7 0 0 0 2 4 {56} 4 522. D. dolichosorum 0 ? 2 2 1 0 2 ? 4 0 ? ? ? ? 2 2 6 2 0 0 2 ? 5 8 723. D. donianum 1 3 1 0 1 {26} 2 3 3 0 1 1 0 1 0 3 2 0 0 0 1 0 2 0 224. D. esculentum 1 0 2 0 1 0 3 1 4 0 2 1 0 1 2 2 6 0 0 0 0 4 {45} 5 225. D. fraxinifolium 0 0 1 0 1 2 5 2 3 0 0 0 ? 0 1 4 3 0 0 0 1 0 2 0 226. D. fuliginosum 0 0 1 4 1 0 2 2 5 0 0 0 0 0 0 0 2 0 1 1 2 6 1 7 3
310
Appendix 1. Continued
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 2527. D.halimunense 0 0 1 2 1 2 2 1 4 1 2 1 0 1 1 2 - 0 0 0 1 0 2 0 ?28. D. hewittii 0 0 1 0 1 6 1 3 4 1 0 0 1 0 1 3 2 0 0 0 5 3 {45} 2 429. D. hottae 0 0 1 0 1 2 5 2 1 1 0 0 - 1 1 4 3 0 0 0 1 0 2 0 430. D. insigne 0 0 2 1 1 {05} 6 2 2 0 2 1 0 1 2 7 9 2 0 0 2 4 5 0 931. D. Kunstlerii 0 0 2 0 0 7 ? ? 4 1 0 0 1 0 2 4 5 1 0 0 1 5 5 4 632. D. laevipes 0 0 2 0 0 7 1 5 4 1 0 0 0 0 2 6 1 0 0 0 1 5 {45} 4 333. D. latisquamatum 0 0 2 1 1 7 5 5 4 1 0 0 2 0 2 7 1 0 0 1 1 5 {67} 2 734. D. lobbianum 0 0 1 2 1 4 5 9 3 1 2 1 0 1 1 5 1 0 0 0 4 5 2 3 335. D. lomariaceum 0 0 1 4 1 0 3 1 5 0 - - - - 0 1 0 0 0 1 2 6 1 7 336. D. Loerzingii 0 0 1 2 1 2 3 1 4 1 0 0 0 1 1 3 1 0 0 0 2 4 3 1 337. D. malaccense 0 0 1 2 1 3 4 1 3 0 0 0 0 - 1 3 0 0 0 0 2 4 3 2 338. D.megasegmentum 0 0 2 0 0 7 2 7 4 1 0 0 1 0 2 9 1 1 0 0 1 5 6 4 639. D.megasimplicifolium 1 0 0 0 1 ? ? ? ? ? ? ? - - 0 1 0 0 0 0 0 4 0 7 240. D. meijerii ? 3 2 0 1 {02} 3 1 5 0 1 1 0 1 2 4 0 0 0 0 1 4 5 9 341. D. melanolepis 0 0 2 0 1 2 3 2 5 1 1 1 0 0 2 4 1 0 0 1 2 5 {45} ? -42. D. moultonii ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? - - ? ? ? ? ? 6 ? -43. D. pallidum 0 0 1 2 1 3 3 1 4 1 0 0 1 0 1 3 0 0 0 0 2 4 {23} 0 244. D. parallelivenium 0 0 2 0 1 2 3 1 4 1 1 1 0 1 2 4 1 0 0 0 1 2 5 4 545. D. petiolare 0 0 1 0 1 3 ? ? 4 0 2 2 0 0 0 3 0 0 0 0 1 5 3 2 246. D. poiense 0 0 1 2 1 {02} 2 1 3 0 0 0 0 0 1 2 0 0 0 0 2 4 {45} 1 247. D. polypodioides 0 0 2 2 1 0 6 1 4 1 2 2 0 1 2 5 1 0 0 0 2 4 {56} 9 648. D. porphyrorachis 0 0 1 2 1 0 2 0 4 1 ? ? ? ? 0 1 0 0 2 1 2 6 1 8 049. D. Prescottianum 0 0 1 0 1 {02} 4 2 4 1 0 0 0 0 1 2 1 0 0 0 0 0 3 0 250. D. procumbens 1 3 2 0 1 {02} ? ? 4 0 1 2 0 0 1 2 3 0 1 0 0 5 5 {10} 351. D. profluens 0 0 2 0 0 7 ? 1 4 1 0 0 0 0 2 5 5 1 0 0 1 5 5 4 -52. D. riparium 0 0 1 0 1 2 2 1 5 1 0 0 1 0 1 3 3 0 0 0 1 6 2 0 2
311
Appendix 1. Continued
Characters and Character StatesNo. Species
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 2553. D. silvaticum 0 0 1 0 1 {34} 4 1 4 0 2 2 0 0 0 2 2 0 0 0 1 5 3 2 154. D. simplicivenium 0 0 2 2 1 {02} 6 2 4 1 1 1 0 1 2 5 9 0 0 0 2 5 5 4 655. D. sorzogonense 0 0 2 2 1 0 5 5 4 1 0 0 0 1 1 3 7 0 1 1 2 5 3 1 456. D. speciosum 0 0 1 2 1 0 6 5 3 0 1 2 0 0 1 4 4 0 0 0 2 4 3 1 557. D. spiniferum 0 0 2 0 0 7 1 6 4 1 0 0 1 0 2 4 6 0 0 1 5 0 5 2 658. D. squarrasum 0 0 2 4 1 {26} 3 1 2 0 2 ? 0 0 2 2 3 1 1 1 2 5 3 0 459. D. subintegrum 0 0 1 0 1 {02} 6 3 3 1 0 0 - - 1 4 3 0 0 0 0 3 2 2 360. D.subalternisegmentum ? ? ? ? 1 2 4 1 5 1 2 2 0 0 2 - 5 2 0 0 2 5 7 ? ?61. D. subpolypodioides 0 0 2 0 1 0 ? 3 4 1 - - - - 2 - - 0 0 0 5 5 5 1 ?62. D. subserratum 0 0 1 0 1 1 0 0 4 0 1 0 0 0 0 2 1 0 1 0 1 0 0 8 263. D. subvirescens 0 3 2 0 1 2 4 1 4 1 1 2 0 1 2 2 1 0 0 1 1 5 5 {10} 364. D. tomentosum 0 0 1 0 1 1 1 0 4 0 0 0 1 0 0 2 1 0 2 1 2 5 3 2 165 D. tricholepis 0 0 1 2 1 {02} 2 1 2 0 2 1 0 0 1 1 2 0 0 0 2 4 4 7 466. D. umbrosum 0 0 2 2 1 3 - - 4 1 1 1 0 0 1 2 3 0 0 0 2 4 {56} 3 267. D. velutinum 0 0 1 4 1 0 4 1 5 1 0 0 0 0 0 2 2 0 2 1 2 5 5 2 168. D. vestitum 0 0 2 0 1 2 - - 3 1 2 2 0 1 2 6 7 2 0 0 1 2 5 3 669. D. wahauense 0 0 1 0 0 2 - - 5 1 0 0 0 0 0 0 1 0 0 0 1 6 2 0 070. D. xiphophyllum 0 0 1 0 1 4 9 4 2 1 0 0 1 0 2 4 4 0 0 0 1 0 3 0 3
312
Appendix 1. Continued
Species Characters & Character States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 5
01. Athyrium
anisopterum0 3 1 0 2 1 2 2 2 1 4 0 0 8 3 8 {13} 2 1 0 0 0 0 - -
2. D. accedens 2 3 1 0 3 2 1 2 3 1 0 3 1 {24} {24} {13} {14} 1 2 0 0 0 0 - -3. D. acuminatum 1 3 1 0 {23} 2 1 1 3 2 3 1 0 4 1 8 2 {23} 2 0 0 0 0 - -4. D. aequibasale ? 3 1 0 0 2 1 1 3 2 5 1 ? 4 3 1 4 1 2 0 0 0 0 - -5. D. albidos-
quamatum? 3 1 0 1 2 1 1 3 2 5 2 2 5 4 9 5 3 2 {12} 0 0 0 2 2
6. D. angustipinna 5 3 1 0 {01} 2 1 1 3 3 2 2 0 1 1 1 1 1 2 0 0 0 0 0 -7. D. asymmetricum 5 3 1 0 {12} 2 1 1 3 2 4 7 7 5 4 9 5 6 3 2 1 2 3 8 98. D.
atrosquamosum4 3 1 0 1 2 1 1 3 2 5 3 3 5 4 9 5 6 2 2 1 2 4 4 3
9. D. bantamense 2 3 1 0 {01} 2 1 1 3 2 0 2 0 7 6 2 2 1 4 0 0 0 0 - -10. D. barbatum 2 3 1 0 1 2 1 1 3 2 5 1 1 5 7 9 5 7 2 1 1 1 2 - ?11. D. batuayauense 2 3 1 0 2 2 1 1 3 2 5 1 0 1 1 8 2 2 2 0 1 0 0 - -12. D. beamanii ? 3 1 0 ? 2 ? ? 3 {12} 1 5 4 5 4 9 5 6 3 3 3 2 {34} 4 313. D.betimusense ? 3 1 0 ? 2 1 1 3 2 4 5 4 5 4 9 5 6 2 2 3 2 4 4 414. D. christii 2 3 1 0 3 2 1 1 3 2 5 2 0 4 3 8 3 2 3 0 0 0 0 - -15. D. cordifolium 1 3 1 1 {01} 2 1 1 3 1 0 2 1 6 5 1 1 1 4 0 0 0 0 - -16. D. crameri 2 3 1 0 1 2 1 1 3 2 5 1 1 4 3 1 3 1 3 0 0 0 0 - -17. D.
crenatoserratum1 3 1 1 {23} 1 1 1 3 2 3 1 0 8 4 {58} {12} {12} 2 0 0 0 0 - -
18. D. crinitum 1 3 1 0 ? 2 1 1 3 2 0 2 2 5 4 9 6 6 2 3 3 1 2 - 219. D. cumingii 2 3 1 0 0 2 1 1 3 2 0 2 1 4 3 1 1 5 4 0 0 0 0 - -20. D.
densisquamatum6 3 1 0 2 1 1 1 3 2 5 5 3 5 4 9 5 6 2 4 3 2 3 3 3
21. D. dilatatum 7 3 1 0 {12} 2 1 1 3 3 0 0 9 5 4 9 5 6 3 3 3 2 3 9 622. D. dolichosorum ? 3 1 0 ? 2 1 2 3 2 2 6 5 1 1 9 5 6 3 3 3 2 2 5 323. D. donianum 1 3 1 0 0 2 1 1 3 2 5 2 1 4 3 1 1 1 2 0 0 0 0 - 2
313
Appendix 1. Continued
Species Character & Character States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
24. D. esculentum 2 3 1 0 {12} 2 1 1 3 3 0 4 3 5 4 9 5 6 2 3 1 2 2 4 325. D. fraxinifolium 4 3 1 0 {01} 1 1 1 3 2 2 3 1 4 3 {58} {24} 1 {35} 0 0 0 0 - -26 D. fuliginosum 0 3 1 0 5 1 1 3 1 1 5 6 0 1 1 2 3 1 1 0 0 0 0 - -27. D.halimunense ? 3 1 0 0 2 1 1 3 1 5 2 1 4 3 1 2 1 2 0 0 0 0 - -28. D. hewittii 2 3 1 0 {12} 2 1 1 3 2 6 2 1 5 4 9 5 6 2 1 2 1 7 1 129. D. hottae 3 3 1 0 {23} 1 1 1 3 2 6 3 0 4 3 1 4 1 2 0 0 0 0 - -30. D. insigne ? 3 1 0 ? 2 1 2 3 2 1 6 3 5 4 9 5 6 2 3 2 1 2 4 331. D. Kunstlerii 6 3 1 0 2 2 1 1 3 2 5 7 4 5 4 9 5 6 2 {23} 3 1 4 4 432. D. laevipes 3 3 1 0 {12} 2 1 2 3 2 7 3 3 5 4 9 5 6 2 2 3 1 3 4 333. D. latisquamatum 6 3 1 0 1 2 1 1 3 2 2 5 4 5 4 9 5 6 2 {23} 3 2 4 4 334. D. lobbianum 2 3 1 0 3 2 1 1 3 1 2 1 0 2 6 1 3 1 2 0 0 0 0 - -35. D. lomariaceum 1 3 - 0 - 0 0 0 0 0 0 - - 1 0 0 0 0 0 0 0 0 0 - -36. D. Lorzingii 2 3 1 0 {34} 2 1 1 3 2 6 2 0 1 1 7 3 0 2 0 0 0 0 - -37. D. malaccense 2 3 1 0 {23} 2 2 1 3 2 4 2 0 1 8 {04} {13} 2 2 0 0 0 0 - -38. D. megasegmentum 1 3 1 0 2 2 1 1 3 2 6 6 6 5 4 9 5 6 2 4 3 2 4 6 539. D.
megasimplicifolium1 0 - 2 - 0 0 0 0 0 0 - - 0 0 0 0 0 0 0 0 0 0 - -
40. D. meijerii 5 3 1 0 2 2 1 2 3 {12} 6 3 3 5 4 8 5 6 3 2 3 {12} 4 3 341. D. melanolepis ? 3 1 0 ? 2 1 1 3 2 6 4 2 1 1 9 5 6 2 3 2 1 4 2 042. D. moultonii 2 3 ? - ? - 1 1 3 2 5 3 2 2 2 9 5 6 2 4 3 1 3 3 243. D. pallidum 2 3 1 0 {34} 2 1 1 3 1 4 2 0 1 6 {67} 3 1 2 0 0 0 0 - -44. D. parallelivenium 9 3 1 0 1 2 1 1 3 2 3 6 7 5 7 9 5 6 2 {23} 3 2 4 7 545. D. petiolare 2 3 1 0 {23} 2 1 1 3 1 6 1 0 4 3 8 3 3 2 0 0 0 0 - -46. D. poiense 1 3 1 0 4 2 1 1 3 1 4 1 0 1 3 7 5 4 2 0 0 0 0 - -47. D. polypodioides 7 3 1 0 1 2 1 1 3 2 5 6 4 1 2 9 5 5 2 {34} 3 1 3 6 448. D. porphyrorachis 0 3 0 0 - 0 0 0 0 0 0 - - 0 0 0 0 0 0 0 0 0 0 - -49. D. Prescottianum 2 3 1 0 2 2 1 1 3 2 4 2 0 {78} 1 4 4 1 3 0 0 0 0 - -
314
Appendix 1. Continued
Species Character & Characters States26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50
50. D. procumbens ? 3 1 0 1 2 1 1 3 2 5 5 4 5 2 9 5 6 2 {23} 3 1 3 3 551. D. profluens ? 3 1 0 1 2 1 1 3 2 6 6 5 5 2 9 5 6 2 3 3 2 4 7 652. D. riparium 2 3 1 0 0 2 1 1 3 2 2 3 1 5 3 1 1 1 4 0 0 0 0 - 353. D. silvaticum 2 3 2 0 {12} 2 1 1 3 1 6 1 0 8 3 8 3 2 2 0 0 0 0 - 154. D. simplicivenium 7 3 1 0 1 2 1 1 3 2 5 4 3 5 4 9 5 6 2 {34} 3 1 3 5 355. D. sorzogonense 3 3 1 0 {23} 1 2 1 3 1 6 2 1 {25} {25} 8 3 {46} 2 0 0 0 0 - -56. D. speciosum 2 3 1 0 {23} 2 1 1 3 1 2 3 0 2 1 8 3 4 2 0 0 0 0 - -58. D. squarrasum ? 3 2 0 3 2 1 1 3 2 5 1 1 {48} {17} 1 1 1 2 0 0 0 1 - -59. D. subintegrum 2 3 1 0 1 2 1 1 3 1 4 3 1 4 3 7 3 1 2 0 0 0 1 0 -60. D.subalternisegmentum ? ? 1 0 ? 2 1 1 3 2 5 - ? 5 4 9 5 7 2 4 3 2 3 5 461. D. subpolypodioides ? 3 0 {12} 1 1 1 1 3 2 2 - ? 2 2 9 5 7 2 3 3 1 3 ? ?62. D. subserratum 0 1 0 - - 0 0 0 0 0 0 - ? 0 0 0 0 0 0 0 0 0 0 ? -63. D. subvirescens 4 3 0 1 1 1 1 1 3 2 5 3 ? 3 2 {89} 5 6 2 2 3 2 5 2 464. D. tomentosum 0 3 1 0 3 1 2 1 2 1 3 0 0 {18} 1 8 3 {12} 2 0 0 0 0 - -65. D. tricholepis 2 3 1 0 3 2 2 1 1 1 5 ? ? 1 1 8 2 6 3 0 0 0 0 - ?66. D. umbrosum 2 3 1 0 3 2 1 2 3 2 6 2 1 2 2 9 5 7 2 3 3 1 3 1 167. D. velutinum 1 3 2 1 2 2 1 1 2 2 2 1 1 1 1 {68} 2 2 2 1 2 1 3 0 068. D. vestitum 8 3 2 1 2 1 1 1 2 3 3 7 4 2 2 9 5 6 1 2 3 2 3 5 469. D. wahauense 0 3 1 1 0 2 1 1 2 1 2 0 0 4 3 1 4 1 {34} 0 0 0 0 - -70. D. xiphophyllum 3 3 1 0 1 2 1 1 3 2 3 4 1 4 3 1 {23} 1 2 0 0 0 0 - -
315
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
1. Athyrium anisopterum 0 0 0 0 2 ? 3 15 3 ? 0 12 0 1 0 0 1 0 1 1 0 ? 1 0 02. D. accedens 0 0 0 0 0 0 0 0 2 6 0 0 1 2 1 0 2 0 0 0 6 - 3 3 13. D. acuminatum 0 0 0 0 1 ? 1 5 3 5 0 0 1 0 0 0 1 0 1 0 0 1 0 2 14. D. aequibasale 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - - 2 0 2 3 3 05. D. albidosquamatum {23} 3 2 2 1 4 0 1 3 1 0 2 0 0 0 0 0 0 1 0 0 - 0 2 16. D. angustipinna 0 0 0 0 0 0 1 0 1 2 0 2 0 0 0 1 - 1 1 1 0 - 3 2 37. D. asymmetricum 1 4 4 3 2 7 3 5 3 5 0 2 0 2 0 0 1 1 1 1 0 - 2 1 08. D. atrosquamosum 0 4 4 3 3 1 2 5 3 - 0 0 1 2 0 0 1 0 1 0 0 - 0 0 09. D. bantamense 0 0 0 0 0 0 0 0 0 5 0 0 1 0 0 1 - 1 1 2 0 - 3 3 010. D. barbatum 1 1 1 2 0 - 0 0 3 1 0 4 0 2 0 0 2 1 1 1 0 - 0 2 011. D. batuayauense 0 0 0 0 0 3 0 0 0 1 0 0 0 0 0 0 1 0 1 0 0 - 2 1 112. D. beamanii 1 4 4 3 2 3 1 {56} 3 ? 0 ? 0 0 0 0 1 0 1 0 0 ? 0 1 113. D. betimusense {13} 4 2 3 1 4 1 3 3 6 0 0 0 0 0 0 - - 1 0 0 1 2 0 014. D. christii 0 0 0 0 0 4 0 0 3 1 0 0 0 0 0 0 1 0 1 2 0 - 0 2 015. D. cordifolium 0 0 0 0 0 0 0 0 1 3 0 0 1 0 1 1 - - 0 2 3 2 3 3 116. D. crameri 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 1 1 0 2 3 3 017. D. crenatoserratum 0 0 0 0 0 2 0 0 3 5 0 0 0 0 0 1 - - 1 1 2 - 3 3 118. D. crinitum 3 4 2 3 2 - 1 7 3 0 0 2 0 2 0 0 1 0 1 0 1 - 0 2 019. D. cumingii 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - 1 1 2 2 - 3 3 020. D. densisquamatum 1 4 3 3 1 1 1 5 3 4 0 2 0 2 0 0 1 0 1 1 1 - 0 1 121. D. dilatatum {14} 4 2 4 1 8 3 4 3 6 0 0 0 2 0 0 2 1 0 1 1 0 0 2 122. D. donianum 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 1 2 1 1 3 3 023. D. dolichosorum 1 2 2 3 1 ? 1 5 3 6 0 0 0 0 0 0 1 0 1 0 1 ? 0 2 024. D. esculentum {15} 4 2 3 1 1 3 4 3 6 0 0 0 0 1 0 2 - 0 0 1 - 3 3 125. D. fraxinifolium 0 0 0 0 1 0 0 0 1 2 0 0 1 0 1 1 - - 0 2 1 {12} 3 3 126. D. fuliginosum 0 0 0 0 1 ? 0 0 3 5 0 4 0 2 0 1 - - 1 1 0 - 3 3 027. D.halimunense 0 0 0 0 1 0 0 0 1 2 0 0 0 0 0 1 - - 1 2 0 2 3 3 0
316
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
28. D. hewittii {35} 3 1 3 2 ? 0 0 3 0 0 0 0 0 0 1 - - 1 1 0 - 0 3 029. D. hottae 0 0 0 0 1 0 0 0 1 2 0 0 1 0 0 1 - - 1 2 0 0 3 3 030. D. insigne 3 2 0 0 0 0 0 0 3 1 0 0 0 0 1 0 2 0 0 0 6 - 3 3 031. D. Kunstlerii {13} 4 3 1 2 4 3 5 3 6 0 0 1 0 0 0 0 1 1 0 1 - 2 1 032. D. laevipes 2 4 {23} 3 2 2 1 5 3 6 0 1 1 1 0 0 2 1 1 0 1 2 0 2 033. D. latisqua-matum {24} 3 4 3 1 2 1 2 3 6 0 0 0 0 0 1 1 1 0 1 1 1 0 1 034. D. lobbianum 0 0 0 0 0 0 0 0 2 3 0 0 0 0 0 1 - - 1 2 1 2 3 2 035. D. lomariaceum 0 0 0 0 0 9 0 0 0 6 0 3 0 - 0 1 - - 1 ? 1 - 0 3 036. D. Lorzingii 0 0 0 0 0 5 1 6 3 2 0 0 1 0 0 0 2 0 1 0 1 2 2 2 137. D. malaccense 0 0 0 0 0 4 3 5 3 1 0 0 0 0 0 0 2 0 1 0 1 - 3 3 038. D.megaseg-mentum 1 4 7 3 1 7 3 7 3 6 1 0 0 0 0 0 3 1 1 1 0 - 0 1 139. D.megasimpli-cifolium 0 0 0 0 1 0 0 0 0 2 - - 0 0 1 - - 2 1 2 5 1 3 3 040. D. meijerii 1 4 3 3 2 4 {13} 1 3 6 0 0 0 0 0 0 {01} {01} 1 1 0 - 2 {23} 141. D. melanolepis 1 4 2 2 2 0 1 5 3 5 0 2 0 2 0 1 - - 1 2 0 - 0 2 042. D. moultonii 1 4 7 3 1 2 3 6 3 1 0 0 0 0 0 0 1 0 - 1 0 0 0 0 043. D. pallidum 0 0 0 0 0 0 0 0 2 5 0 2 0 2 0 - - - 1 1 0 - 3 3 044. D.parallelive-nium 1 4 {12} 4 1 ? 1 5 3 6 0 0 0 0 0 0 1 1 1 0 0 2 0 1 045. D. petiolare 0 0 0 0 0 3 0 0 3 0 0 0 0 0 0 0 1 0 1 0 0 2 3 3 046. D. poiense 0 0 0 0 0 2 0 0 3 4 1 0 0 0 0 0 ? 0 1 0 0 - 2 2 147. D. polypodioides 2 4 0 3 2 2 3 4 3 6 0 0 0 0 0 0 2 1 1 0 0 - 3 3 048. D. porphyrorachis 0 0 0 0 0 6 2 0 0 2 1 3 0 0 0 1 - - 1 0 0 - 3 3 049. D. Prescottianum 0 0 0 0 0 0 0 0 2 5 0 0 0 0 0 1 - - 1 2 0 - 3 3 050. D. procumbens 1 4 3 3 1 5 3 5 3 1 0 2 0 2 0 0 1 1 1 0 0 2 0 1 051. D. profluens 1 4 6 3 2 - 4 5 3 4 1 0 0 0 0 0 2 1 1 0 0 2 0 1 052. D. riparium 0 0 0 0 0 0 0 0 1 2 0 0 0 0 0 1 - - 0 2 1 2 3 3 053. D. silvaticum 0 0 0 0 0 5 0 0 3 1 0 2 1 1 0 0 0 0 0 0 0 - 3 3 154. D. simplicivenium 1 4 {12} 3 1 4 1 5 3 4 0 0 0 2 0 0 1 0 1 0 0 - 3 2 1
317
Appendix 1. Continued
Species Character & Characters States51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 67 68 69 70 71 72 73 74 75
55. D. sorzogonense 0 0 0 3 1 4 3 7 3 6 1 3 0 2 0 0 2 1 - 0 0 - 2 2 056. D. speciosum 0 0 0 0 0 3 0 0 3 0 0 0 0 0 0 0 2 0 1 0 0 - 0 2 057. D. spiniferum 3 4 1 3 1 1 2 5 3 4 0 0 0 0 0 0 0 0 1 0 0 - 2 0 058. D. squarrasum 0 0 0 0 0 0 0 0 2 1 0 4 0 2 1 1 3 - 1 2 5 2 3 3 059. D. subintegrum 0 0 0 0 0 0 0 0 2 5 0 0 0 0 0 1 - - 1 2 0 - 0 1 060. D.subalternisegmentum 1 4 7 3 1 ? 4 7 3 - 0 0 0 0 0 0 2 1 1 2 0 - 0 2 061. D. subpolypodioides 1 4 6 3 2 - 4 6 3 2 0 0 0 0 0 0 2 1 1 1 0 - 0 1 162. D. subserratum 0 0 0 0 0 - 0 0 1 6 1 - - 2 0 1 - 1 1 2 0 2 3 3 063. D. subvirescens {14} 4 {67} 3 1 1 4 7 3 1 0 0 0 0 0 0 1 1 - 1 0 - 2 1 064. D. tomentosum 0 0 0 0 1 1 1 1 3 1 0 3 0 0 0 0 1 0 0 0 0 2 3 3 065. D. tricholepis 0 0 0 0 1 - {24} 5 3 1 0 2 0 2 0 {01} 2 1 1 1 0 0 3 3 066. D. umbrosum {24} 4 6 1 {12} 2 6 3 1 1 0 0 0 0 0 1 1 1 1 0 0 - 2 2 067. D. velutinum 3 4 0 1 2 - 2 3 0 0 0 0 0 0 1 2 1 1 1 1 0 - 3 2 168. D. vestitum 3 4 1 1 3 - 5 3 2 1 0 0 0 0 0 1 1 1 1 1 0 - 3 2 169. D. wahauense 0 0 0 0 1 - 0 0 1 1 0 0 0 0 0 1 0 - 1 2 0 - 3 3 070. D. xiphophyllum 0 0 0 0 1 - 0 0 1 2 0 2 1 1 0 1 - - 1 2 1 1 3 3 1
318
Appendix 1. Continued
76 77 78 79
80 81 82 83 84 85 86 87 88
1. Athyrium anisopterum {12} 0 0 1 1 1 0 0 0 0 1 1 ?2. D. accedens 2 2 2 0 1 1 0 1 0 0 1 0 03. D. acuminatum 2 2 2 0 1 1 1 1 0 0 1 1 04. D. aequibasale 2 1 2 0 0 1 1 0 1 0 1 0 05. D. albidosquamatum 2 2 2 0 1 1 0 2 0 - 1 1 06. D. angustipinna 1 1 2 0 0 1 1 0 0 0 0 1 07. D. asymmetricum 1 2 2 1 1 1 0 0 0 0 1 1 08. D. atrosquamosum 2 1 2 1 1 1 0 0 0 2 1 0 09 D. bantamense 1 1 2 1 1 1 0 1 0 0 1 0 010. D. barbatum 0 1 2 0 1 1 0 0 0 0 1 1 011. D. batuayauense 1 - 2 0 1 1 0 1 0 0 1 1 012. D. beamanii 2 2 2 1 1 1 0 ? ? 2 1 ? 013. D. betimusense 1 0 2 0 0 1 0 2 ? 2 0 0 014. D. christii 2 1 2 0 1 0 0 1 0 0 1 0 015. D. cordifolium 1 2 2 1 1 1 0 1 0 0 1 0 016. D. crameri 1 1 2 1 1 1 0 1 0 0 1 1 017. D. crenatoserratum 1 1 2 1 1 1 0 1 0 0 1 1 018. D. crinitum 2 1 2 1 1 1 0 2 1 0 1 1 019. D. cumingii 1 2 2 1 1 1 0 2 0 0 1 1 020. D. densisquamatum 2 0 2 0 1 1 0 2 0 2 1 0 021. D. dilatatum 2 2 2 0 1 1 0 1 0 0 1 0 022. D. dolichosorum 2 2 2 1 1 1 0 1 0 ? 1 1 023. D. donianum 1 1 2 1 1 1 1 1 0 1 1 1 024. D. esculentum 2 2 2 0 0 1 1 2 0 2 1 0 025. D. fraxinifolium 2 2 2 1 1 1 1 0 0 1 1 1 026. D. fuliginosum 0 0 2 1 1 1 0 2 0 0 1 1 027. D.halimunense 1 2 2 1 1 1 0 1 0 0 1 1 028. D. hewittii 1 1 2 0 1 1 0 1 0 0 1 0 0
319
Appendix 1. Continued
Species Character & Character States76 77 78 79 80 81 82 83 84 85 86 87 88
29. D. hottae 1 1 2 0 1 1 1 1 0 0 1 1 030. D. insigne 2 1 2 0 0 1 0 2 0 0 1 0 031. D. Kunstlerii 2 1 2 0 0 0 0 2 0 2 1 0 032. D. laevipes 1 2 2 0 1 1 0 2 0 2 1 0 033. D. latisquamatum 2 1 2 1 1 1 1 1 1 0 2 0 034. D. lobbianum 1 1 2 1 1 1 0 1 0 0 2 1 035. D. lomariaceum 0 0 2 0 1 1 0 2 0 1 1 0 036. D. Lorzingii 1 2 2 1 1 1 0 1 0 0 1 1 037. D. malaccense 2 1 2 0 1 1 0 0 0 0 1 1 038. D.megasegmentum 2 2 2 1 1 0 0 1 0 2 1 0 039. D.megasimplicifo-
lium1 1 2 1 1 1 1 1 0 0 1 0 0
40. D. meijerii 2 3 2 0 1 1 0 0 0 2 1 1 041. D. melanolepis 1 1 2 1 1 1 1 2 0 0 1 1 042. D. moultonii 1 1 2 1 1 1 0 1 0 2 1 0 043. D. pallidum 0 0 2 0 1 1 0 0 0 0 1 1 044. D. parallelivenium 1 1 2 0 1 1 1 1 1 2 1 0 045. D. petiolare 2 2 2 - - - - - - - - - 046. D. poiense 1 0 2 1 1 1 0 0 0 0 1 1 047. D. polypodioides 2 2 2 0 1 1 0 0 0 0 1 0 048. D. porphyrorachis 0 0 2 1 1 1 0 0 0 0 1 0 049. D. Prescottianum 0 0 2 0 1 1 0 0 0 0 1 1 050. D. procumbens 2 2 2 1 1 1 0 1 0 2 1 0 051. D. profluens 1 2 2 0 1 1 0 2 0 0 1 1 052. D. riparium 1 1 2 0 1 1 0 0 0 0 1 0 053. D. silvaticum 2 2 2 1 1 1 0 0 2 1 1 0 054. D. simplicivenium 2 2 2 1 1 1 0 1 0 2 1 0 055. D. sorzogonense 2 2 1 1 1 1 0 2 1 0 1 1 0
320
Appendix 1. Continued
Species Characters & character States76 77 78 79 8
081 82 83 84 85 86 87 88
56. D. speciosum 2 1 2 1 1 1 0 1 0 0 1 0 057. D. spiniferum 2 1 2 0 1 1 1 2 0 2 1 0 058. D. squarrasum 1 0 2 0 1 1 0 1 0 0 1 0 059. D. subintegrum 1 2 2 1 1 1 1 0 0 0 1 1 060. D. subalternisegmen-
tum2 0 2 1 1 1 0 1 0 0 1 0 0
61. D. subpolypodioides 2 1 2 0 1 1 0 2 0 0 1 1 062. D. subserratum 1 1 2 1 1 1 1 0 0 0 1 1 063. D. subvirescens 2 2 2 0 1 0 0 1 0 2 1 0 064. D. tomentosum 1 1 2 1 1 1 0 2 0 0 1 1 065. D. tricholepis 2 3 2 0 1 1 0 1 0 0 1 1 066. D. umbrosum 1 1 {12} 1 1 1 0 0 1 0 1 0 067. D. velutinum 2 2 2 1 1 1 0 1 0 0 1 0 068. D. vestitum 2 2 2 0 1 1 0 2 0 0 1 1 069. D. wahauense 1 1 2 0 1 1 1 2 0 0 1 0 070. D. xiphophyllum 1 2 2 1 1 1 0 0 0 0 1 1 0
351
Plate 1. Diplazium asymmetricum Praptosuwiryo. Holotype(T.Ng. Praptosuwiryo 1728, BO). a. Bipinnate lamina; b. A part of pinna showingfertile pinnulae.
352
Plate 2. Diplazium batuayauense Praptosuwiryo. Paratype(T.Ng. Praptosuwiryo 1972e, BO). a. Frond. b. Part of pinnae with detailvenation.
353
Plate 3. Diplazium crameri Praptosuwiryo. Holotype (DR. Cramer 41, BO)a. Frond. b. Part of pinna bearing sori.
354
Plate 4. Diplazium densisquamatum Praptosuwiryo. Holotype(T.Ng. Praptosuwiryo 2491, BO). a. Lamina; b. Rhizome and base of stipes. c.Lower part of pinnae with pinnulae bearing mature sori; d. Middle part of pinnaewith pinnulae bearing immature sori.
355
Plate 5. Diplazium halimunense Praptosuwiryo. Holotype(T.Ng. Praptosuwiryo 2341, BO). a. Rhizome with stipes bearing scales at base; b.Fertile lamina; c. Part of lamina with pinnae; d. Part of pinna showing fertile freeveins.
356
Plate 6. Diplazium loerzingii Praptosuwiryo. Paratype (T.Ng. Praptosuwiryo2519, BO). a. Frond; b. Part of fertile pinnae on rachise bearing scales; c. Pinnaefrom the middle part of lamina; d. Young plant bearing at the adjacent betweenrachis and costa
357
Plate 7. Diplazium megasegmentum Praptosuwiryo. Holotype (T.Ng.Praptosuwiryo 1382, BO). a. Rhizome with base of stipes, scales fallen; b.Pinnae on middle part of lamina; c. Pinnulae on middle part of pinnae; d. Lowerpart of pinnule with segments bearing mature sori.
358
Plate 8. Diplazium megasimplicifolium Praptosuwiryo. Holotype (Veldkamp7998, BO). a. Whole part of plant. B. Part of lamina with reticulate fertile veins.
359
Plate 9. Diplazium meijerii Praptosuwiryo. Holotype (W. Meijer 3772, BO).a. whole part of plant; b. Rhizome; c. Pinnulae; d. A part of fertile pinna.
360
Plate 10. Diplazium parallelivenium Praptosuwiryo, Holotype(T.Ng. Praptosuwiryo 2338, BO). a. Lamina; b. Rhizome with base of stipesbearing brown scales; c. Part of young pinna; d. Fertile pinnulae
361
Plate 11. Diplazum profluens Praptosuwiryo. Holotype (T.Ng. Praptosuwiryo1820, BO). a. Pinna; b. Pinnulae. c. Middle part of pinnule with detail venationand sori
362
Plate 12. Diplazium subalternisegmentum Praptosuwiryo, Holotype (J. & M. S.Clemens 33169, BO). a. Stipe and part of lamina; b. Base of stipe bearing scales;c. Part of pinnae. d. Part of pinnulae with fertile segments.
363
Plate 13. Diplazium subvirescens Praptosuwiryo. Paratype (T.Ng.Praptosuwiryo 1013, BO). a. Bipinnate frond; b. A part of punnule bearing sori.c. Long creeping rhizome with base of stipes bearing brown scales.
364
Plate 14. Diplazium subvirescens Praptosuwiryo. Holotype (T.Ng.Praptosuwiryo 1178, BO). a. A part of tripinnate lamina. b. A part of pinnae withsegmented fertile pinnulae; c. Stipe with a part of lower pinnae; d. A part ofpinnule.