Revision of Rhynchoglossum (Gesneriaceae) in Malesia
REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN
MALESIA
ABDULROKHMAN KARTONEGORO
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
LETTER OF STATEMENT
I express that thesis entitled:
REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN MALESIA
is true represent result of my own research and have never been published. All
information and data that used have been expressed clearly and can be checked its
truth.
Bogor,
June 2011
Abdulrokhman Kartonegoro
NRP. G353080061
ABSTRACT
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in
Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Revision on Rhynchoglossum in Malesia has been done. This research
conducted based on morphological characters of 233 sheets of Herbarium
specimen and Herbarium specimen photographs from Herbarium Bogoriense
(BO), Singapore Botanic Gardens (SING), National Herbarium Netherlands (L),
Royal Botanic Garden Kew (K), Natural History Museum London (BM), Royal
Botanic Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian
Institute (US), and Harvard University (A). The result shows that six species are
recognized in Malesia and five are endemic. Five species are known previously
namely R. borneense, R. klugioides, R. medusothrix, R. obliquum, and
R. spumosum; one species is proposed as a new and newly described namely
R. celebicum. Rhynchoglossum borneense and R. medusothrix endemic to Borneo,
R. celebicum endemic to Sulawesi, while R. spumosum endemic to Philippine
Islands. Rhynchoglossum klugioides and R. obliquum distributed wider in
Malesia. Rhynchoglossum papuae and R. merrilliae are placed under synonym of
R. obliquum and R. spumosum. A phylogenetic analysis using PAUP version 4.0
program based on 60 morphological characters with Stauranthera coerulea and
Loxonia hirsuta as an outgroup was undertaken. The analysis resulted a
parsimonious tree which shown that Rhynchoglossum in Malesia can be divided in
two groups. First clade consist R. obliquum, R. spumosum and R. celebicum while
the second clade consist R. klugioides, R. borneense and R. medusothrix.
Taxonomic treatment was giving with a identification key to the species,
description, nomenclature, distribution, and notes each species.
Keywords: Gesneriaceae,
Rhynchoglossum
Malesia,
phylogenetic
analysis,
revision,
ABSTRAK
Abdulrokhman Kartonegoro. Revisi Rhynchoglossum (Gesneriaceae) di
Malesia. Dibimbing oleh Dr. Sri S. Tjitrosoedirdjo, M.Sc. dan Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Revisi marga Rhynchoglossum di Malesia telah dilakukan. Studi ini
dilakukan berdasarkan pendekatan karakter morfologi pada 233 lembar spesimen
herbarium dan foto spesimen herbarium dari Herbarium Bogoriense (BO),
Singapore Botanic Gardens (SING), National Herbarium Netherlands (L), Royal
Botanic Garden Kew (K), Natural History Museum London (BM), Royal Botanic
Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian Institute
(US), dan Harvard University (A). Hasil revisi menunjukkan ada enam jenis yang
diketahui di Malesia dan lima jenis endemik. Lima jenis merupakan jenis-jenis
yang sudah dikenal sebelumnya yaitu R. borneense, R. klugioides, R. medusothrix,
R. obliquum, dan R. spumosum; satu jenis diusulkan sebagai jenis baru dan baru
dipertelakan yaitu R. celebicum. Rhynchoglossum borneense dan R. medusothrix
endemik di Borneo, R. celebicum endemik di Sulawesi, sedangkan R. spumosum
endemik di Filipina. Rhynchoglossum klugioides dan R. obliquum tersebar lebih
luas di Malesia. Rhynchoglossum papuae dan R. merrilliae dimasukkan ke dalam
sinonim dari R. obliquum dan R. spumosum. Analisis filogenetik menggunakan
program PAUP versi 4.0 berdasarkan pada 60 karakter morfologi dengan
Stauranthera coerulea dan Loxonia hirsuta sebagai outgroup. Dari hasil analisis
dihasilkan pohon parsimoni yang menunjukkan bahwa Rhynchoglossum di
Malesia dapat dipisahkan menjadi dua kelompok. Kelompok pertama terdiri atas
R. obliquum, R. spumosum dan R. celebicum sedangkan kelompok kedua terdiri
atas R. klugioides, R. borneense dan R. medusothrix. Bagian taksonomi
disampaikan dengan kunci identifikasi jenis, pertelaan, tata nama, distribusi dan
catatan tiap-tiap jenis.
Kata kunci: Gesneriaceae, Malesia, filogenetik analisa, revisi, Rhynchoglossum
SUMMARY
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in
Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Rhynchoglossum Blume is a genus of fleshy herb belongs to family
Gesneriaceae. It is comprised about 10 species all over the world. The genus
Rhynchoglossum was established by Blume with one spesies of Rynchoglossum
obliquum. Rhynchoglossum can be recognized morphologically as fleshy herb
plant with anisophyllous decussate leaf or alternate leaf arrangement,
asymmetrically leaf blade and the unilaterally inflorescence. The genus has a little
economic value but until now still planted almost 130 years in several Botanic
Gardens and private gardens around the world.
Several taxonomic study of Rhynchoglossum has done since the genus
proposed by Blume. From many studies and revisions, there was no specific
taxonomic revision for Malesian region that known as the center of biodiversity of
the genus. Revision on this genus in Malesia has not been done before. Based on
that information, the revision of the genus Rhynchoglossum is needed.
The research objective of this study was to revise taxonomic of
Rhynchoglossum in Malesia. Taxonomic revision was done to find the
information about generic and species delimitation, by providing data on its
diversity, identification key for species, analytic description for genus and species,
distribution pattern, and phylogenetic of Rhynchoglossum in Malesia.
The study was carried out at the Systematic Laboratory of the Herbarium
Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI)
at Cibinong, West Java and the Herbarium of Singapore Botanic Gardens,
Singapore National Parks from January 2010 to February 2011. It is based on 233
sheets of Herbarium specimens (including spirit collections) and herbarium
specimen photographs from the Herbarium Bogoriense (BO), Herbarium of
Singapore Botanic Gardens (SING), National Herbarium of NederlandUniversiteit Leiden Branch (L), Natural History Museum of London (BM), Royal
Botanic Garden of Kew (K), Royal Botanic Garden of Edinburgh (E), Harvard
University (A), Smithsonian Institute (US), and New York Botanical Gardens
(NY) were also studied. Phylogenetic analysis of Malesian Rhynchoglossum was
studied based on sixty morphological characters which collected during
morphological observation and analyzed based on Maximum Parsimony using
PAUP version 4.0b.10 with the Heuristic search settings vegetative and floral
characters were selected for the analysis.
Based on the revision of Rhynchoglossum in Malesia, there were six
species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B.L.Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no
infraspecific status decided in this revision. Rhynchoglossum celebicum is new
species that newly described here beside the other five species.
In Malesia this genus consist of six species and five of them are endemic
to the region. Rhynchoglossum klugioides was previously only known from
Philippines now distributed also to Seram Island, Moluccas. Four species are
known endemic in each region. Rhynchoglossum spumosum is endemic to
Philippines known from Negros and Mindanao, while the R. celebicum is endemic
to Sulawesi. Rhynchoglossum borneense and R. medusothrix known endemic to
Borneo which is known only found in the eastern part of the island.
Rhynchoglossum can be found from lowland areas to mountain areas of 1500 m
above sea level at the edge of the forest, close to water shaded or in the rock of
limestone area.
The phylogenetic analysis using 60 morphological characters based on
Maximum Parsimony method. The 128 equally most parsimonious tree of 138
steps produce consistency index (CI) = 0.6562, homoplasy index (HI) = 0.3438
and retention index (RI) = 0.5769. The higher CI value was shown that the
number of homoplasy rather low. The separation of the ingroup and outgroup was
supported with a high bootstrap value. The ingroup was separated in two sister
clades, one consist R. obliquum, R. spumosum and R. celebicum, and the other
consist of R. klugioides, R. borneense and R. medusothrix.
Keywords: Gesneriaceae,
Rhynchoglossum
Malesia,
phylogenetic
analysis,
revision,
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REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN
MALESIA
ABDULROKHMAN KARTONEGORO
Thesis submitted
As partial fulfillment requirement for the Master Degree
In Plant Taxonomy
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
External examiner : Dr. Nunik Sri Ariyanti, M.Si.
Title
: Revision of Rhynchoglossum (Gesneriaceae) in Malesia
Name
: Abdulrokhman Kartonegoro
NRP
: G353080061
Study Program: Plant Biology
Approved by
Supervisor Committee
Dr. Sri S. Tjitrosoedirdjo, M.Sc.
(Chairman)
Head of Plant Biology Study Program
Dr. Ir. Miftahudin, M.Si.
Date of Examination: 10 June 2011
Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
(Member)
Dean of Graduate School
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Graduation:
ACKNOWLEDGEMENT
I would like to thank to my supervisors, Dr. Sri S. Tjitrosoedirdjo (IPB)
and Prof.(R) Dr. Elizabeth A. Widjaja (LIPI) for their valuable advices and
guidance in completing this thesis. I am gratefully thanked to Dr. Teguh Triono
(BO), Prof. Dr. Eko Baroto Walujo (BO), Dr. Harry Wiriadinata (BO), W.S.
Hoover (NETC) for their helpful giving advises for completing this thesis. I also
would like thank to Dr. David Middleton (E), Dr. Laurence Skog (US), Dr. Ruth
Kiew (KEP), and Dr. John R. Clark (SEL) for their helpful advices and critics to
my thesis draft especially in Gesneriaceae. Further thanked to Dr. J.F. Veldkamp
for helping in nomenclature taxonomy and latin diagnose for the new species.
Special thank to The Indonesian Botanical Exploration and Taxonomy
Project (IBETP) of New England Tropical Conservatory (NETC), Nellie Sleeth
Scholarships Grant of Gesneriad Society USA and Singapore Botanic Gardens
Fellowship Grant for financing my research and study at the Graduate School of
Bogor Agricultural University (IPB).
I would like to express my gratitude to Herbarium Bogoriense (BO) of
Research Center for Biology (LIPI) and Singapore Botanic Gardens for granting
permission to conduct my research work and providing some facilities. I would
like to thanks especially to the Director of Herbaria A, BO, BM, E, L, K, SING,
US, and NY for examined their valuable specimens. Special thanks giving to my
friends Lim Chung Lu and Yao Tze Liong from Forest Research Institue Malaysia
and Mr. Gerard Thijsse from National Herbarium Netherland for sending some
image of Rhynchoglossum Type from Kew and Leiden.
I thanks to my colleagues and friends in Herbarium Bogoriense and Plant
Biology Major IPB especially Alex Sumadijaya, Ina Erlinawati, Yessi Santika,
Arief Hidayat, Destri, Wina Dyah Puspitasari, Sukiman, Rina, Adeel, Sanou Faye
for their help, support and opportunity to cement a good friendship.
My very special thanks go to my wife Nur Rohmatin Isnaningsih and my
son Hilmi Ahmad Asqalani for their spirit, patience and support in my study.
CURRICULUM VITAE
The author was born in Jakarta on 4 January 1980, the fifth son of five of Mr.
Bunyamin (Alm) and Mrs. Muhanah.
In 1999, he became a student at the Department of Biology, Faculty of
Mathematic and Natural Science, University of Indonesia, Depok and graduated at
2005.
Since April 2006 he was employed by the Research Center for Biology,
Indonesian Institute of Sciences (LIPI) in Cibinong, West Java and still active till
now.
In 2008 he received a scholarship from the Indonesian Botanical Exploration
and Taxonomy Project (IBETP), New England Tropical Conservatory (NETC) to
continue his study at the Department of Biology, Faculty of Mathematic and
Natural Science, Bogor Agricultural University with a specification on Plant
Taxonomy.
CONTENTS
Page
LIST OF TABLES
…………..………………………………...…….......
xix
LIST OF FIGURES
……….………………………………….................
xxi
LIST OF APPENDICES
…………..………………………………........ xxiii
………………………..………………………..........
1
………………..……………………………..
3
…………..…………………………….
9
RESULT AND DISCUSSION …………..………………………...……
General morphology of Rhynchoglossum in Malesia …………...
Habit …...................................................................................
Leaves …...............................................................................
Inflorescences …...................................................................
Flowers ….............................................................................
Fruits ….................................................................................
Distribution ……………...………………………………………
Phylogenetic analysis ….......………………………………..…...
Taxonomic Treatment ……….....………………………..………
Rhynchoglossum Blume …....................................................
Key to Species of Rhynchoglossum in Malesia ….................
Rhynnchoglossum borneense Merr. …...................................
Rhynchoglossum celebicum Karton. sp.nov. …......................
Rhynchoglossum klugioides C.B. Clarke …...........................
Rhynchoglossum medusothrix B.L. Burtt …..........................
Rhynchoglossum obliquum Blume ….....................................
Rhynchoglossum spumosum Elmer …....................................
13
14
14
14
16
17
18
19
21
25
25
26
27
30
33
36
39
45
CONCLUSION
…………..………………………………………...……
49
REFERENCES
…………..………………………………………...……
51
APPENDICES
…………...………………………………………...……
55
INTRODUCTION
LITERATURE REVIEW
MATERIAL AND METHODS
LIST OF TABLES
Page
1 The characters and characters state used in the phylogenetic analysis ....
10
2 Distribution of Rhynchoglossum in Malesia …………………………....
20
LIST OF FIGURES
Page
1 Distribution of Rhynchoglossum in yellow circle
….......……………….. 4
2 Illustration of Loxotis obliqua = Rhynchoglossum obliquum
(from Brown 1838)
……………………………...……………………..
5
3
Leave shape. A. Ovate of R. obliquum B. Ovate of R. klugioides
C. Oblong of R. celebicum …………………………………………...…. 15
4
Rhynchoglossum obliquum A. Inflorescence; B. Flowers ………..……... 17
5
Fruit shape A.Ovoid of R. obliquum B. Capsule-like of R. celebicum
C. Elongate of R. klugioides ……………………………………….…...... 19
6
Cladogram of phylogenetic tree of Rhynchoglossum in Malesia.
Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769.
The number shown above line showed supported the Bootstrap values …. 22
7
Rhynchoglossum borneense Merr., Elmer 21467 from Tawao, Malaysian
Borneo (isotype SING)
...…………………………...………………... 29
8 Distribution map of Rhynchoglossum borneense in Malesia in the circle …. 30
9 Distribution map of Rhynchoglossum celebicum in Malesia in the circle ... 31
10 Rhynchoglossum celebicum Karton., Eyma 1572 from Maraowa, Sulawesi
(holotype BO) …………………………………………...……………… 32
11 Distribution map of Rhynchoglossum klugioides in Malesia in the circle .... 34
12 Rhynchoglossum klugioides C.B.Clarke, Cuming 824 from Tayabas, Luzon
(holotype K) …………………………………………………………….. 35
13 Distribution map of Rhynchoglossum medusothrix in Malesia in the circle .. 37
14 Rhynchoglossum medusothrix B.L.Burtt, Kostermans 13994 from Berau,
East Kalimantan, Borneo (isotype BO) .………………………………… 38
15 Distribution map of Rhynchoglossum obliquum in Malesia in the circle ... 43
16 Rhynchoglossum obliquum Blume, Blume 52 from Java (lectotype L) …. 44
17 Distribution map of Rhynchoglossum spumosum in Malesia in the circle .. 46
18 Rhynchoglossum spumosum Elmer, Elmer 9929 from Cuernos Mts.,
Negros (Isotype US) …………..…………………………..……….…..... 47
LIST OF APPENDICES
Page
1
List of species Rhynchoglossum proposed in Malesia (Burtt 1962) …...
57
2
Matrix of 60 Morphological Characters for Maximum Parsimony
Analysis ………..………………………………………………………...
58
INTRODUCTION
Rhynchoglossum Blume was a genus of fleshy herb belongs to family
Gesneriaceae. It comprised of 10 species all over the world. It is classified in the
tribe Epithemateae or Epithematoid Gesneriaceae together with Epithema Blume,
Gyrogyne W.T. Wang, Loxonia Jack, Monophyllaea R. Br., Stauranthera Benth.,
dan Whytockia W.W. Sm. Rhynchoglossum was mainly found from India, Ceylon,
Nepal, South China, Formosa, Indochina, Thailand, and Malesia, with only one
species grow in Southern America (Mayer et.al. 2003; Weber 2004b).
The genus Rhynchoglossum was established by Blume (1826) with one
spesies of Rynchoglossum obliquum. The type specimen was collected from Java.
The species has a wide distribution from India, Ceylon, South China, Indochina,
and Malay Archipelago through New Guinea. Some genus allied like Antonia
R. Br., Glossanthus Benth., Klugia Schltdl., and Loxotis R. Br. are proposed later
as the synonym of Rhynchoglossum (Burtt 1962).
Rhynchoglossum can be recognized morphologically as fleshy herb plant
with anisophyllous decussate leaf or alternate leaf arrangement, asymmetrically
leaf blade and the unilaterally inflorescence. The genus has a little economic value
but several Botanic Gardens and private gardens are growing the plant as an
ornamental (Skog 1985). One of the most ornamental planted is Rhynchoglossum
gardneri Theobald & Grupe from India and Ceylon. The epithet name of
Rhynchoglossum is from Greek whereas Rhynchos means beak; and glossa means
tongue. The second part of the name clearly alludes to the broad, tongue-like
lower lip of the corolla, the first part perhaps to the narrow corolla tube or to the
pointed petal tips (Weber 2004b).
Weber (2004b) put Rhynchoglossum under Epithematoid Gesneriaceae.
This group has unique and special characters of unequal cotyledons,
asymmetrically leaf blade and capsule fruit. The distribution of the group was
mostly in tropical Asia, few from Africa, and one species from southern America.
The disjunction distribution from this group was very unique especially in
Rhynchoglossum.
Several taxonomic study of Rhynchoglossum has done since the genus
proposed by Blume. Most of the studies generally observed the family
Gesneriaceae,
whereas
the
Rhynchoglossum
was
included.
Some
of
comprehensive studied were by Bentham (1876) and Clarke (1883). The latest
study was a little revision done by Burtt (1962) which is stated that
Rhynchoglossum has 13 species and with very limited material specimens. From
many studies and revisions, there was no specific taxonomic revision for Malesian
region that known as the center of biodiversity for the genus. Revision about
species biodiversity and the number of species in Malesia has not been done
before. Based on that information, the revision of the genus Rhynchoglossum is
needed.
The research objective of this study was to revise taxonomic of
Rhynchoglossum in Malesia. Taxonomic revision was done to find the
information about generic and species delimitation, by providing data on its
biodiversity, identification key for species, analytic description for genus and
species, distribution pattern, and phylogenetic of Rhynchoglossum in Malesia.
LITERATURE REVIEW
Rhynchoglossum taxonomically included in Epithematoid Gesneriaceae
group or tribe Epithemateae along with others genera like Whytockia W.W. Sm.,
Gyrogyne W.T. Wang, Epithema Blume, Monophyllaea R. Br., Loxonia Jack, and
Stauranthera Benth. (Weber 2004b). Generally these groups have a similarity in
unequal cotyledons, unequal leaf arrangement in one node, asymmetrically leaf
blade and capsule fruits. Rhynchoglossum is allied with Loxonia and Stauranthera
in the shape of leaf arrangement. Rhynchoglossum has similarity with
Monophyllaea in the shape of inflorescences with unilateral cymose shape (Weber
2004b).
In the earlier status, Rhynchoglossum distributed from India, Ceylon,
China, Taiwan, Indochina to New Guinea in the tropical Asia (Weber 2004b).
Burtt (1962) included Klugia Schltdl. as synonymy of Rhynchoglossum, which is
Klugia has a distribution in Southern America from Mexico to Peru. The united of
Klugia as a synonym makes distribution of Rhynchoglossum wider and disjunct to
America (Figure 1). These disjunction distributions on the genus within
Australasia to America make Rhynchoglossum very unique and have an
interesting distribution history. Burtt (1998) give preliminary hypothesis that
Rhynchoglossum reach America from Asia via Africa from where it now has
completely disappeared or no species are found.
Weber (2004a; 2004b) assumed that the genus spreads from Asia to
America recently, probably via transpacific trips or migrations of early
Polynesians. This is also suggested by the current localities which are usually near
former populations centers of ancient dwellers along the pacific coast. Mayer
et.al. (2003) indicate that American species (R. azureum (Schltdl.) B.L. Burtt) is a
recent introduction to the neotropics and that it does not represent an ancient relic.
It seem R. azureum more allied to R. notonianum (Wall.) B.L.Burtt that known
from India than other species from Malesian region. The phylogenetic showed
that R. azureum allied in one clade with R. notonianum and reflects clearly
morphological not geographical pattern in the genus (Mayer et.al. 2003).
Figure 1 Distribution of Rhynchoglossum in yellow circle
Rhynchoglossum obliquum Blume is the common and widespread species,
found from India, China to Malesia (Bakhuizen van den Brink Jr. 1965; Burtt
1962). In the Southern America only have one species R. azureum (Schltdl.) B.L.
Burtt which has disjunctions distribution with its allies in Asia especially in
Southern India (Mayer et.al. 2003). Some of the other species (e.g. R. borneense
Merr., R. medusothrix B.L.Burtt, and R. spumosum Elmer) only have restricted
distributions and even with endemic pattern. Its preferably habitat is on the wet
and shady (especially limestone) rocks, in the forest or open vegetation or shady
places; usually grow in the lowlands (Weber 2004b).
Rhynchoglossum in general have specific characters like perennial or
annual, monocarpic herbs, stems with sparse hairs indumentum from often
branched, many-celled, to glandular hairs. Stems terete, fleshy-succulent, usually
branched. Leaves alternate, nearly distichous, short petiolate; lamina obliquely
ovate-cordate; base asymmetrical, one half cordate, the other sinuate-attenuate;
texture thin and delicate. Inflorescences terminal on main and side branches,
unilateral racemes with two rows of flowers, subtending bracts small, linear,
displaced onto the pedicels in most species; a few sterile bracts often present on
the flowerless side of the inflorescence axis. Sepals connate in the lower half, with
or without wings at the lines of fusion, lobes narrow triangular-acute. Corolla
strongly zygomorphic; tube cylindrical, white; limb strongly bilabiate, upper lip of
two short, upright or reclined lobes, lower lip a large, roundish or elongate, not or
only slightly three-partite tongue of azure colour. Fertile stamens four (the former
genus Klugia) or two (Rhynchoglossum s.str.). Ovary globose-ovoid. Fruit a
globose or ovoid capsule included in the calyx, dehiscing loculicidally by 2 valves
(Weber 2004b). The illustration of the genus show on Figure 2.
Figure 2 Illustration of Loxotis obliqua = Rhynchoglossum obliquum
(from Brown 1838)
Brown (1832) proposed a new genus of Antonia with the species of
A. obliqua (Wall.) R. Br. which is combination of Wulfenia obliqua Wall. (Weber
2004a). Three years later Brown moved the species into the genus Loxotis R. Br.
ex Benth. and made a new combination of L. obliqua (Wall.) R. Br.. The species
is surviving for a long time as well as Miquel (1855) use it. Later on Clarke
(1883) realizes that Antonia and Loxotis are superfluous because it is similar and
identical to Rhynchoglossum and then he put both genera under synonym of
Rhynchoglossum. Recently Burtt (1962) also included Klugia to a synonym
because it is only a variation of Rhynchoglossum.
The latest study by Burtt (1962) sign 13 species of Rhynchoglossum all
over the world comprising 10 species from Asiatic and 3 from America. After that
the number species of Rhynchoglossum are decrease because reducing some
species into synonym. Rhynchoglossum grandiflorum (Fritsch) B.L. Burtt and
R. violaceum (Fritsch) B.L. Burtt are turn into synonym of R. azureum because of
the similarity in morphology and geography (Wiehler 1983). Wang et.al. (1998)
reduce R. hologlosum Hayata into synonym of R. obliquum. Since the reduction of
some species into synonim, total number species of Rhynchoglossum known only
10 species all over the world (Mayer et.al. 2003; Weber 2004b).
History of the number species proposed in Malesia start when Blume
(1826) established the R. obliquum as the type of the genus from Java. After
Blume published the type then followed by de Candolle (1845) which established
another species R. blumei DC. from Java. Clarke (1883) included R. blumei to be
a synonym of R. obliquum and established a new species of R. klugioides C.B.
Clarke from Luzon, Philippines. The species epithet is given by Clarke as a
species which have similarity with Klugia. In the early of twentieth century,
Elmer (1908) proposed a new R. spumosum Elmer from Negros Philippines and
followed by Kraenzlin (1913) proposed R. merrilliae Kraenzl. from Mindanao,
Philippines. Ten years later, a new species of Rhynchoglossum proposed by
Schlechter (1923) which is from New Guinea and named as R. papuae Schltr.
Another species from Borneo proposed by Merrill (1929) and named as
R. borneense Merr. The last species that ever found is R. medusothrix which is
proposed by Burtt (1962) also from Borneo and completely sign seven species
number in Malesia (Appendix 1).
Phylogenetic analysis of Rhynchoglossum using molecular DNA was
studied together with another member of tribe Epithemateae. Phylogenetic tree
showed that Rhynchoglossum separate in different clade with another
Epithemateae and act as sister position. This separation is in accordance with
many and strong morphological differences (e.g., alternate leaf arrangement,
strongly asymmetrical leaves, terminal inflorescences in the form of unilateral
racemes, enlarged lower lip of corolla). The genus has many special characters in
common with the other Epithemateae, except Monophyllaea and Whytockia. In
the Rhynchoglossum group reflected in the phylogram by two clades;
R. notonianum (India) and the neotropical R. azureum vs. R. obliquum (Malesia).
The separation clearly indicated the affinity with morphological characters and
not the geographical pattern, which can be seen as an indication that R. azureum is
a recent introduction to the neotropics and that it does not represent an ancient
relict. The groups can be roughly classified in two: (1) perennials with large
flowers and four stamens (the former genus Klugia) and (2) annuals with small
flowers and two stamens (Mayer et.al. 2003).
MATERIALS AND METHODS
This study was carried out at the Systematic Laboratory of the Herbarium
Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI)
at Cibinong, West Java and the Herbarium of Singapore Botanic Gardens,
Singapore National Parks from January 2010 to February 2011. It is based on 233
sheets of Herbarium specimens (including spirit collections) of Rhynchoglossum
from the Herbarium Bogoriense (BO) and the Herbarium of Singapore Botanic
Gardens (SING). A number of herbarium specimen photographs from National
Herbarium of Nederland-Universiteit Leiden Branch (L), Natural History
Museum of London (BM), Royal Botanic Garden of Kew (K), Royal Botanic
Garden of Edinburgh (E), Harvard University (A), Smithsonian Institute (US),
and New York Botanical Gardens (NY) were also studied.
The materials were studied based on morphological characters using 10 x
40 binocular microscope and stereo microscope. The information was taken under
the information data of location, general habitat, habit, stem, leaves,
inflorescences and infructescences. Morphological description was following
Veldkamp (1987). The revision methods were based on Rifai (1976, 2008) and
Vogel (1987).
Phylogenetic analysis of Malesian Rhynchoglossum was studied based on
60 morphological characters which collected during morphological observation
and analyzed based on Maximum Parsimony using PAUP version 4.0b.10
(Swofford 1998) with the Heuristic search settings vegetative and floral characters
were selected for the analysis, of which 47 were scored as binary and 13 as multistate (Table 1). In a few cases, some characters were scored as missing data when
the data were unavailable.
Table 1 The characters and characters state used in the phylogenetic analysis
NO
CHARACTERS
CHARACTER STATE
1
Growing habit
shrub (0); erect herb (1); creeping herb (2)
2
Habit
perennial (0); annual (1)
3
Height
≤ 30 cm (0); > 30 cm (1)
4
Stem diameter
≤ 3 mm (0); > 3 mm (1)
5
Leaf arrangement
opposite (0); alternate (1)
6
Leaf shape
ovate-elliptic (0); oblong (1)
7
Lamina
membranous (0); subcoriaceous (1)
8
Leaf color
pale green (0); dark green (1)
9
Leaf apex
acuminate (0); acute(1)
10
Apex tip
< 1 cm (0); 1 cm (1)
11
Leaf margin
entire (0); dentate or serrate (1)
12
Leaf base
rounded-cuneate (0); rounded-angustate (1);
cordate-cuneate (2); cordate-angustate (3)
13
Midrib indument
glabrous (0); puberulous (1); pubescent (2)
14
Abaxial leaf indument
glabrous (0); glabrescent (1); pubescent (2)
15
Petiole indument
glabrous (0); glabrescent (1); pubescent (2)
16
Petiole length
< 10 cm (0); 10-15 cm (1); > 15 cm (2)
17
Inflorescence position
axillar only (0); axillar and terminal (1)
18
Inflorescence growth
erect (0); spreading (1)
19
Terminal infl. length
≤ 15 cm (0); > 15 cm (1)
20
Axillar infl. length
≤ 5 cm (0); > 5 cm (1)
21
Inflorescence side
multilateral (0); unilateral (1)
22
Flower number
≤ 10 (0); > 10 (1)
23
Main axis indument
glabrous (0); puberulous (1); pubescent (2)
24
Peduncle length
≤ 3 cm (0); > 3 cm (1)
25
Bract shape
linier (0); oblong (1)
26
Bract length
≤ 4 mm (0); > 4 mm (1)
27
Pedicel indument
glabrous (0); glabrescent (1); puberulous (2);
pubescent (3)
__________________________________________________________________
Table 1 continued
NO
CHARACTERS
CHARACTER STATE
28
Pedicel length
≤ 3 mm (0); > 3 mm (1)
29
Bracteole length
≤ 5 mm (0); > 5 mm (1)
30
Bracteole position
rise from base pedicel (0); rise from middle
pedicel (1)
31
Flower length
≤ 15 mm (0); > 15 mm (1)
32
Calyx tube shape
campanulate (0); urceolate (1)
33
Calyx tube color
white (0); pale green (1); dark green (2)
34
Calyx tube adnate length
≤ 3 mm (0); > 3 mm (1)
35
Calyx lobe apex
acute (0); acuminate (1)
36
Calyx lobe length
≤ 3 mm (0); > 3 mm (1)
37
Corolla tube color
white (0); blue (1); green (2)
38
Corolla tube length
≤ 15 mm (0); > 15 mm (1)
39
Tube length
≤ 10 mm (0); > 10 mm (1)
40
Limb length
≤ 5 mm (0); > 5 mm (1)
41
Adaxial lip shape
2-lobed (0); emarginated (1)
42
Adaxial lip length
≤ 3 mm (0); > 3 mm (1)
43
Abaxial lip shape
tongue-like (0); 3-lobed (1)
44
Abaxial lip apex
rounded (0); obtuse (1)
45
Abaxial lip length
≤ 5 mm (0); > 5 mm (1)
46
Stamens number
2 (0); 4 (1)
47
Smaller stamen fil. shape
terete (0); flat (1)
48
Smaller stamen fil. length
≤ 2 mm (0); > 2 mm (1)
49
Smaller anther shape
ovoid (0); divaricate (1); reniform (2)
50
Smaller anther diameter
≤ 1 mm (0); > 1 mm (1)
51
Larger stamen fil. length
≤ 3 mm (0); > 3 mm (1)
52
Larger anther shape
oblong (0); reniform (1)
53
Ovary shape
globose (0); ovoid (1); ellipsoid (2);
oblong (3)
54
Ovary length
≤ 2 mm (0); > 2 mm (1)
__________________________________________________________________
Table 1 continued
NO
CHARACTERS
CHARACTER STATE
55
Style length
≤ 10 mm (0); > 10 mm (1)
56
Fruit shape
globose (0); ovoid (1); ellipsoid (2);
capsule-like (3); elongate (4)
57
Stalk length
≤ 10 mm (0); > 10 mm (1)
58
Stalk indument
glabrous (0); puberulous (1); pubescent (2)
59
Seed surface
tessellate (0); smooth (1)
60
Seed length
≤ 0.4 mm (0); > 0.4 mm (1)
All characters are threat as unorder data and have equal weight. The
“missing” data are symbolized with “?”. Starting tree(s) was obtained via stepwise
addition. Number of trees held at each step during stepwise addition = 1. Branch
swapping algorithm was run by using tree-bisection-reconnection (TBR),
evaluated using 100 bootstrap replicates and a complete heuristic search. The
complete data matrix is shown in appendix 2.
RESULT AND DISCUSSION
Based on this revision work of Rhynchoglossum in Malesia, there were six
species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton. spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B. L. Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no
infraspecific status decided in this revision. Rhynchoglossum celebicum is one of
new species that newly described here beside the other five species.
New status has been given for some species in this revision.
Rhynchoglossum papuae Schltr. from New Guinea now turn into synonym of
R. obliquum. Another species is R. merrilliae Kraenzl. from Philippine now under
synonym of R. spumosum. This new status decreasing the species number of
Rhynchoglossum in Malesia that giving by Burtt (1962) from seven to six here
with new species that newly described (R. celebicum).
Based on preliminary revision, Burtt (1962) proposed a seven species of
Rhynchoglossum comprising R. borneense, R. klugioides, R. medusothrix,
R. merrilliae, R, obliquum, R. papuae, and R. spumosum (Appendix 1). In this
recent revision, the number species are decreased because the status of two
species changed. Rhynchoglossum merrilliae is synonym of R. spumosum due to
its similarity on their morphological characters such as dwarf habit, small leaf
blade, few flower in the inflorescence and two fertile stamens. Based on the
observation to type specimens, it is realized that both are actually one species and
R. spumosum is used because this species name used earlier.
Rhynchoglossum papuae was proposed by Schlechter (1923) based on the
small corolla lip and globose or ovoid capsules comparing with R obliquum.
When more specimens were examined from New Guinea in this revision, it is
seen that most characters from each specimen have no different with R. obliquum
in other islands of Malesia. This consideration makes R. papuae is closely related
with R. obliquum and placed as a synonym. By placing two species as synonym in
this revision, the number of species became decreasing than the number species
that given by Burtt (1962) from seven species into five species. By adding the
new species (R. celebicum) in this revision, it showed that the total number of
Rhynchoglossum in Malesia become six species.
General Morphology of Rhynchoglossum in Malesia
Habit
Rhynchoglossum are erect and creeeping fleshy-succulent herbs plant from
5-150 cm height. The roots are adventitious with rhizomatous or non-rhizomatous.
The only rhizomatous species is R. spumosum is being creeping in habit and
dwarfing about 5-8 cm height. The stems usually are terete and wrinkled when
dried. They have a variable indument, the hairs being absent or simple puberulous
to pubescent. Most of the species have glabrous and smooth stem surface and
have swollen nodes.
Rhynchoglossum in broad sense have a growing habit as an herb with
perennial or annual growth. All Malesian species known have annual growth with
fleshy succulent herb. Rhynchoglossum spumosum is the only species have
creeping habit with rhizomatous root system. This habit makes the species has
only maximum height 8 cm known as a dwarf Rhynchoglossum. Another species
that reporting have rhizomatous habit is R. omeiense W.T.Wang from China, but
different in having perennial growth (Wang et.al. 1998). Because of that it is
necessary to study further by their molecular systematic to understand the
uncontinous charcters occurs in several species.
Leaves
Leaf arrangements are alternate and exstipulate. The leaf blade of all
Rhynchoglossum is oblique with asymmetrical shape between two sides.
Sometimes the bases of both size are in equal position or different in length till 2.5
cm in R. borneense. The leaf shape varies from ovate to oblong. Rhynchoglossum
celebicum is the only species which have oblong leaf (Figure 3B) while the other
species varies from ovate to elliptic (Figure 3A). The leaf bases are unequal which
one side has cuneate to acute shape and the other side has rounded to cordate
shape. The leaf margins are mostly entire only several species are serrate. The leaf
blades texture are mostly membranous, only R. klugioides have subcoriaceous leaf
blade. Most of the species have glabrous and smooth leaf surface. Some have
pubescent or puberulous indumentum when juvenile.
A
B
C
Figure 3 Leave shape. A. Elliptic of R. obliquum B. Ovate of R. klugioides
C. Oblong of R. celebicum
Rhynchoglossum have alternate leaf arrangement with no stipules. Different
from other Gesneriaceae which have opposite leaf arrangement Rhynchoglossum
have unique leaves development where the opposite leaf is reduced and become
alternate. Weber (2004b) gives hypothesis that the alternate leaf arrangement is
evolution trend in the genus because of the reduction of the opposite side. The
other allied genera like Stauranthera and Loxonia also have similar development
in leaf arrangement with anisophylly leaf arrangement, not reducing like
Rhynchoglossum. The opposite sides in the leaf arrangement are usually free or
with the inflorescences.
Most of the species of Rhynchoglossum known have ovate to elliptic leaf
shape. In this revision a new character of leaf which is oblong belongs to
R. celebicum. This different on its leaf characters made R. celebicum distinct and
realized as a new species. The leaf texture of Rhynchoglossum has two types, one
usually membranous (most of Malesian species) and the other one is subcoriceous
(R. klugioides). The subcoriaceous texture is related with the species that formerly
belongs to Klugia (known as R. azureum and R. notonianum recently).
Inflorescences
The inflorescence in Rhynchoglossum is raceme with terminally and axillary
grow. The axillary inflorescence are usually opposite with leaf. The inflorescences
are spreading or secundiflorous which is not erect (Figure 4A). Number of flower
in inflorescence varies from less than 3 in R. spumosum to 50 in R. obliquum.
Flowers in inflorescence are unilateral because the reduction of other side flowers
and arrange in two rows. The bracts usually absent in R. spumosum or occur in
linier shape in R. obliquum. Sometimes in inflorescences have bracts that not
supported flower or known as sterile bracts. Bracteoles in Rhynchoglossum are
mostly linier which rise from base of pedicel in R. klugioides and R. spumosum;
and rise from the middle of pedicel in other species.
Inflorescences of Rhynchoglossum are known as raceme with secundiflorous
form. These inflorescences are usually with only one side flowering or unilateral
with two rows. This unilaterally growth is obtained because of the reduction on
flower growth in other pairs of inflorescences (Weber 2004b). The reduction side
of the flower is sometimes has bracts which is not subtending and known as sterile
bracts. The prominent difference in inflorescences is observed on the rise of
bracteoles. Rhynchoglossum klugioides and R. spumosum known have bracteoles
from the base of pedicels while the other species have bracteoles rise from the
middle of pedicel.
A
B
Figure 4 Rhynchoglossum obliquum A. Inflorescence; B. Flowers
Flowers
The flowers are zygomorphic, tube-like and vary from less than 1 cm long
till more than 3 cm long. Calyx tubes are infundibuliformis with campanulate
shape in most species or urceolate shape in R. klugioides. The calyx adnate at base
sometimes winged with line fushion; white to green colour. Calyx lobes are
triangular and entire, glabrous, 5-merous. Corolla tubes are glabrous to
puberulent, with tube and limb. The limb has two lipped which is from the petal
fused (Figure 4B). The adaxial lip usually smaller with two lobes and the abaxial
lip larger 3-lobed or undivided bearing a rounded elongate or tongue–like. The
petals usually have a bluish to dark purple color and a few is white. Some species
like R. borneense, R. medusothrix and R. klugioides have a yellow dot pilose in
the throat of abaxial limb. The pilose dot of limb in R. medusothrix usually being
medussoid and dense.
Two enlarged lips corolla with enlarge lower one in the flower of
Rhynchoglossum are common in the genus. The variation among the species is
usually shown in the lower lip characters. The species that having small flower or
corolla (R. obliquum, R. spumosum and R. celebicum) have tongue-like lower lips
and the species with large flower or corolla (R. klugioides, R. borneense and
R. mrdusothrix) have 3-lobes, broader and elongate. In the species that have large
corollas has pilose dot in throat of lower lip but vary from less to dense pilose,
while in R. medusothrix with medussoid pilose hair.
Rhynchoglossum in Malesia have 2 or 4 stamens. Rhynchoglossum
obliquum, R. klugioides, R. celebicum and R. spumosum have 2 stamens.
Otherwise, R borneense and R. medusothrix have 4 stamens. All stamens in
flowers are arranged coherent in pairs. The stamens are adnate to corolla tube in
adaxial limb if only 2 stamens or in abaxial limb if there 4 stamens. The anthers
are basifixed, thecae parallel and dehiscing longitudinally. Filaments are usually
flat. The ovaries in most Rhynchoglossum are ovoid with one loculed. Stigma are
capitates or undivided and the style are usually glabrous and still remnant in
fruiting time.
Four numbers of stamens in Malesian Rhynchoglossum allied with another
species of Rhynchoglossum (R. notonianum (Wall.) B.L.Burtt and R. azureum
(Schltd.) B.L.Burtt) was previously from the genus Klugia. The different with the
species outside Malesia is in the arrangement of the stamens. R. notonianum and
R. azureum have four equal stamens that coherent together while R. borneense
and R. medusothrix have four unequal stamens (two larger and other two smaller
stamens). The arrangement of the stamens is also coherent but in pair not together
(dydinamous).
Fruits
The fruits in all Rhynchoglossum are capsule type with one loculed. The
shape of the fruits may vary from capsule-like, ovoid to elongate with prominent
calyx remnants (Figure 5). The calyx remnants are half closing the fruit in
R. celebicum (Figure 5C) and fully closing the fruit in other species (Figure 5A &
B). The fruits are dehiscing loculicidally to base with two valvate, not twisted.
The seeds are cuneate and unappendaged.
The shape of the fruits of Rhynchoglossum is varying. The ovoid fruit
shape is known oly from R. obliquum, R. borneense, R. medusothrix and R.
spumosum with calyx remnant closing fully the fruit and persistent.
Rhynchoglossum klugioides has elongate fruit shape and also closed by the
persistent calyx. The elongate shape is common in other species outside Malesia
that have large corollas (e.g. R. notonianum and R. gardneri). The only species
that not fully closed by the calyx is R. celebicum and also has capsule-like shape.
A
B
C
Figure 5
Fruit shape A. Ovoid of R. obliquum B. Elongate of R. klugioides
C. Capsule-like of R. celebicum
Distribution
Rhynchoglossum approximately have 10 species of fleshy herbs that found
throughout South Asia, China, South East Asia and Malesia to Southern America
(Weber, 2004). The Malesian region has about six species and five of them are
endemic to the region (Table 2). Rhynchoglossum obliquum is a widespread and
very common species distributed from Sumatra to New Guinea. Rhynchoglossum
klugioides was previously only known from Philippines currently distributed also
to Seram Island, Moluccas. The other four species are endemic in each region.
Rhynchoglossum spumosum was endemic to Philippines from Negros Island and
Mindanao Island, while the R. celebicum endemic to Sulawesi. Rhynchoglossum
borneense and R. medusothrix are endemic to Borneo which is recorded only
found in the eastern part of the island.
Table 2 Distribution of Rhynchoglossum in Malesia
Species
Sumatra
Peninsular
Borneo
Java
Sulawesi
Malaya
Lesser
Moluccas
Philippines
Sunda
New
Guinea
√
R. borneense
√
R. celebicum
R. klugioides
√
R. obliquum
√
√
√
√
√
R. medusothrix
√
√
√
√
√
√
√
R. spumosum
Based on the distribution pattern, the center of the diversity
Rhynchoglossum in Malesia is found in Borneo, Philippine and Sulawesi. Borneo
has three species (R. borneense, R. medusothrix, and R. obliquum), Philippine has
three species (R. klugioides, R. spumosum, and R. obliquum) while Sulawesi has
two species (R celebicum and R. obliquum). The regions shows the high number
of Rhynchoglossum species found comparing to the other islands like Sumatra,
Peninsular Malaya, Java, Lesser Sunda, Moluccas and New Guinea. The
endemicity of Rhynchoglossum in the center of diversity is also high. Beside
R. obliquum, the rest five species known endemic in that area while a few
R. klugioides also observed in Moluccas.
The phytogeography pattern for Rhynchoglossum is still related with the
pattern of all Asiatic Rhynchoglossum. Burtt (1998) said that Gesneriaceae is a
family
of
Gondwanic
origin.
Epithematoid
Gesneriaceae
which
is
Rhynchoglossum inside is a relict group that was once much larger and had much
wider distribution. The ancient of Rhynchoglossum is known from Asiatic
mainlad which now has the most species occur. Based on that hypothesis is
predicted that the Malesian species are spreading from west Wallace line to east
line. The species that found in the east Malesia are R. obliquum and R. klugioides
which known from Moluccas and New Guinea. In fact, both species also known
distributed in the west of Malesia like Sumatra, Java, Borneo and Philippine.
Rhynchoglossum celebicum is the only species that found in Sulawesi which is the
intermediate region between west and east Malesia. Distribution pattern of
Rhynchoglossum in Malesia made the genus is typically Sunda shelf or West
Malesia taxon.
Rhynchoglossum can be found from lowland areas to mountain areas of
1500 m above sea level at the edge of the forest, close to water shaded or in the
rock of limestone area. Most of the species like R. obliquum, R. celebicum,
R. klugioides and R. spumosum found in the forest with canopy or edge forest with
wet condition near the river. Rhynchoglossum borneense and R. medusothrix are
known from the limestone habitat in Borneo. It is looks that the preferable habitat
impacts the distribution of species.
Phylogenetic analysis
The phylogenetic relationships of the genus Rhynchoglossum in Malesia
was analyzed in the present study based on morphological characters. Sixty
characters were used in this analysis (Table 1). The sixty characters were selected
for their apparent taxonomic values that are good to separate one taxon with the
others.
Phylogenetic analysis indicated that all characters are parsimony
informative. The 128 equally most parsimonious tree of 138 steps produce
consistency index (CI) = 0.6562, homoplasy index (HI) = 0.3438 and retention
index (RI) = 0.5769. The higher CI value was resulted that the value of
homoplasy rather low. The value of RI above 0.5 indicated that these characters
could contribute some synapomorphy to the cladogram. Figure 8 showed tree
number 1 of 128 the most parsimonious trees.
Figure 6 Cladogram of phylogenetic tree of Rhynchoglossum in Malesia.
Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769.
The number shown above line showed supported the Bootstrap values.
The separation of the ingroup and outgroup in the phylogenetic analysis
was supported with a high bo
MALESIA
ABDULROKHMAN KARTONEGORO
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
LETTER OF STATEMENT
I express that thesis entitled:
REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN MALESIA
is true represent result of my own research and have never been published. All
information and data that used have been expressed clearly and can be checked its
truth.
Bogor,
June 2011
Abdulrokhman Kartonegoro
NRP. G353080061
ABSTRACT
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in
Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Revision on Rhynchoglossum in Malesia has been done. This research
conducted based on morphological characters of 233 sheets of Herbarium
specimen and Herbarium specimen photographs from Herbarium Bogoriense
(BO), Singapore Botanic Gardens (SING), National Herbarium Netherlands (L),
Royal Botanic Garden Kew (K), Natural History Museum London (BM), Royal
Botanic Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian
Institute (US), and Harvard University (A). The result shows that six species are
recognized in Malesia and five are endemic. Five species are known previously
namely R. borneense, R. klugioides, R. medusothrix, R. obliquum, and
R. spumosum; one species is proposed as a new and newly described namely
R. celebicum. Rhynchoglossum borneense and R. medusothrix endemic to Borneo,
R. celebicum endemic to Sulawesi, while R. spumosum endemic to Philippine
Islands. Rhynchoglossum klugioides and R. obliquum distributed wider in
Malesia. Rhynchoglossum papuae and R. merrilliae are placed under synonym of
R. obliquum and R. spumosum. A phylogenetic analysis using PAUP version 4.0
program based on 60 morphological characters with Stauranthera coerulea and
Loxonia hirsuta as an outgroup was undertaken. The analysis resulted a
parsimonious tree which shown that Rhynchoglossum in Malesia can be divided in
two groups. First clade consist R. obliquum, R. spumosum and R. celebicum while
the second clade consist R. klugioides, R. borneense and R. medusothrix.
Taxonomic treatment was giving with a identification key to the species,
description, nomenclature, distribution, and notes each species.
Keywords: Gesneriaceae,
Rhynchoglossum
Malesia,
phylogenetic
analysis,
revision,
ABSTRAK
Abdulrokhman Kartonegoro. Revisi Rhynchoglossum (Gesneriaceae) di
Malesia. Dibimbing oleh Dr. Sri S. Tjitrosoedirdjo, M.Sc. dan Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Revisi marga Rhynchoglossum di Malesia telah dilakukan. Studi ini
dilakukan berdasarkan pendekatan karakter morfologi pada 233 lembar spesimen
herbarium dan foto spesimen herbarium dari Herbarium Bogoriense (BO),
Singapore Botanic Gardens (SING), National Herbarium Netherlands (L), Royal
Botanic Garden Kew (K), Natural History Museum London (BM), Royal Botanic
Garden Edinburgh (E), New York Botanical Gardens (NY), Smithsonian Institute
(US), dan Harvard University (A). Hasil revisi menunjukkan ada enam jenis yang
diketahui di Malesia dan lima jenis endemik. Lima jenis merupakan jenis-jenis
yang sudah dikenal sebelumnya yaitu R. borneense, R. klugioides, R. medusothrix,
R. obliquum, dan R. spumosum; satu jenis diusulkan sebagai jenis baru dan baru
dipertelakan yaitu R. celebicum. Rhynchoglossum borneense dan R. medusothrix
endemik di Borneo, R. celebicum endemik di Sulawesi, sedangkan R. spumosum
endemik di Filipina. Rhynchoglossum klugioides dan R. obliquum tersebar lebih
luas di Malesia. Rhynchoglossum papuae dan R. merrilliae dimasukkan ke dalam
sinonim dari R. obliquum dan R. spumosum. Analisis filogenetik menggunakan
program PAUP versi 4.0 berdasarkan pada 60 karakter morfologi dengan
Stauranthera coerulea dan Loxonia hirsuta sebagai outgroup. Dari hasil analisis
dihasilkan pohon parsimoni yang menunjukkan bahwa Rhynchoglossum di
Malesia dapat dipisahkan menjadi dua kelompok. Kelompok pertama terdiri atas
R. obliquum, R. spumosum dan R. celebicum sedangkan kelompok kedua terdiri
atas R. klugioides, R. borneense dan R. medusothrix. Bagian taksonomi
disampaikan dengan kunci identifikasi jenis, pertelaan, tata nama, distribusi dan
catatan tiap-tiap jenis.
Kata kunci: Gesneriaceae, Malesia, filogenetik analisa, revisi, Rhynchoglossum
SUMMARY
Abdulrokhman Kartonegoro. Revision of Rhynchoglossum (Gesneriaceae) in
Malesia. Supervised by Dr. Sri S. Tjitrosoedirdjo, M.Sc. and Prof.(R) Dr.
Elizabeth A. Widjaja, M.Sc.
Rhynchoglossum Blume is a genus of fleshy herb belongs to family
Gesneriaceae. It is comprised about 10 species all over the world. The genus
Rhynchoglossum was established by Blume with one spesies of Rynchoglossum
obliquum. Rhynchoglossum can be recognized morphologically as fleshy herb
plant with anisophyllous decussate leaf or alternate leaf arrangement,
asymmetrically leaf blade and the unilaterally inflorescence. The genus has a little
economic value but until now still planted almost 130 years in several Botanic
Gardens and private gardens around the world.
Several taxonomic study of Rhynchoglossum has done since the genus
proposed by Blume. From many studies and revisions, there was no specific
taxonomic revision for Malesian region that known as the center of biodiversity of
the genus. Revision on this genus in Malesia has not been done before. Based on
that information, the revision of the genus Rhynchoglossum is needed.
The research objective of this study was to revise taxonomic of
Rhynchoglossum in Malesia. Taxonomic revision was done to find the
information about generic and species delimitation, by providing data on its
diversity, identification key for species, analytic description for genus and species,
distribution pattern, and phylogenetic of Rhynchoglossum in Malesia.
The study was carried out at the Systematic Laboratory of the Herbarium
Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI)
at Cibinong, West Java and the Herbarium of Singapore Botanic Gardens,
Singapore National Parks from January 2010 to February 2011. It is based on 233
sheets of Herbarium specimens (including spirit collections) and herbarium
specimen photographs from the Herbarium Bogoriense (BO), Herbarium of
Singapore Botanic Gardens (SING), National Herbarium of NederlandUniversiteit Leiden Branch (L), Natural History Museum of London (BM), Royal
Botanic Garden of Kew (K), Royal Botanic Garden of Edinburgh (E), Harvard
University (A), Smithsonian Institute (US), and New York Botanical Gardens
(NY) were also studied. Phylogenetic analysis of Malesian Rhynchoglossum was
studied based on sixty morphological characters which collected during
morphological observation and analyzed based on Maximum Parsimony using
PAUP version 4.0b.10 with the Heuristic search settings vegetative and floral
characters were selected for the analysis.
Based on the revision of Rhynchoglossum in Malesia, there were six
species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B.L.Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no
infraspecific status decided in this revision. Rhynchoglossum celebicum is new
species that newly described here beside the other five species.
In Malesia this genus consist of six species and five of them are endemic
to the region. Rhynchoglossum klugioides was previously only known from
Philippines now distributed also to Seram Island, Moluccas. Four species are
known endemic in each region. Rhynchoglossum spumosum is endemic to
Philippines known from Negros and Mindanao, while the R. celebicum is endemic
to Sulawesi. Rhynchoglossum borneense and R. medusothrix known endemic to
Borneo which is known only found in the eastern part of the island.
Rhynchoglossum can be found from lowland areas to mountain areas of 1500 m
above sea level at the edge of the forest, close to water shaded or in the rock of
limestone area.
The phylogenetic analysis using 60 morphological characters based on
Maximum Parsimony method. The 128 equally most parsimonious tree of 138
steps produce consistency index (CI) = 0.6562, homoplasy index (HI) = 0.3438
and retention index (RI) = 0.5769. The higher CI value was shown that the
number of homoplasy rather low. The separation of the ingroup and outgroup was
supported with a high bootstrap value. The ingroup was separated in two sister
clades, one consist R. obliquum, R. spumosum and R. celebicum, and the other
consist of R. klugioides, R. borneense and R. medusothrix.
Keywords: Gesneriaceae,
Rhynchoglossum
Malesia,
phylogenetic
analysis,
revision,
Copyright©2011, Bogor Agricultural University
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2. It is prohibited to republish and reproduce all parts of this thesis without
the written permission from Bogor Agricultural University.
REVISION OF RHYNCHOGLOSSUM (GESNERIACEAE) IN
MALESIA
ABDULROKHMAN KARTONEGORO
Thesis submitted
As partial fulfillment requirement for the Master Degree
In Plant Taxonomy
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2011
External examiner : Dr. Nunik Sri Ariyanti, M.Si.
Title
: Revision of Rhynchoglossum (Gesneriaceae) in Malesia
Name
: Abdulrokhman Kartonegoro
NRP
: G353080061
Study Program: Plant Biology
Approved by
Supervisor Committee
Dr. Sri S. Tjitrosoedirdjo, M.Sc.
(Chairman)
Head of Plant Biology Study Program
Dr. Ir. Miftahudin, M.Si.
Date of Examination: 10 June 2011
Prof.(R) Dr. Elizabeth A. Widjaja, M.Sc.
(Member)
Dean of Graduate School
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Graduation:
ACKNOWLEDGEMENT
I would like to thank to my supervisors, Dr. Sri S. Tjitrosoedirdjo (IPB)
and Prof.(R) Dr. Elizabeth A. Widjaja (LIPI) for their valuable advices and
guidance in completing this thesis. I am gratefully thanked to Dr. Teguh Triono
(BO), Prof. Dr. Eko Baroto Walujo (BO), Dr. Harry Wiriadinata (BO), W.S.
Hoover (NETC) for their helpful giving advises for completing this thesis. I also
would like thank to Dr. David Middleton (E), Dr. Laurence Skog (US), Dr. Ruth
Kiew (KEP), and Dr. John R. Clark (SEL) for their helpful advices and critics to
my thesis draft especially in Gesneriaceae. Further thanked to Dr. J.F. Veldkamp
for helping in nomenclature taxonomy and latin diagnose for the new species.
Special thank to The Indonesian Botanical Exploration and Taxonomy
Project (IBETP) of New England Tropical Conservatory (NETC), Nellie Sleeth
Scholarships Grant of Gesneriad Society USA and Singapore Botanic Gardens
Fellowship Grant for financing my research and study at the Graduate School of
Bogor Agricultural University (IPB).
I would like to express my gratitude to Herbarium Bogoriense (BO) of
Research Center for Biology (LIPI) and Singapore Botanic Gardens for granting
permission to conduct my research work and providing some facilities. I would
like to thanks especially to the Director of Herbaria A, BO, BM, E, L, K, SING,
US, and NY for examined their valuable specimens. Special thanks giving to my
friends Lim Chung Lu and Yao Tze Liong from Forest Research Institue Malaysia
and Mr. Gerard Thijsse from National Herbarium Netherland for sending some
image of Rhynchoglossum Type from Kew and Leiden.
I thanks to my colleagues and friends in Herbarium Bogoriense and Plant
Biology Major IPB especially Alex Sumadijaya, Ina Erlinawati, Yessi Santika,
Arief Hidayat, Destri, Wina Dyah Puspitasari, Sukiman, Rina, Adeel, Sanou Faye
for their help, support and opportunity to cement a good friendship.
My very special thanks go to my wife Nur Rohmatin Isnaningsih and my
son Hilmi Ahmad Asqalani for their spirit, patience and support in my study.
CURRICULUM VITAE
The author was born in Jakarta on 4 January 1980, the fifth son of five of Mr.
Bunyamin (Alm) and Mrs. Muhanah.
In 1999, he became a student at the Department of Biology, Faculty of
Mathematic and Natural Science, University of Indonesia, Depok and graduated at
2005.
Since April 2006 he was employed by the Research Center for Biology,
Indonesian Institute of Sciences (LIPI) in Cibinong, West Java and still active till
now.
In 2008 he received a scholarship from the Indonesian Botanical Exploration
and Taxonomy Project (IBETP), New England Tropical Conservatory (NETC) to
continue his study at the Department of Biology, Faculty of Mathematic and
Natural Science, Bogor Agricultural University with a specification on Plant
Taxonomy.
CONTENTS
Page
LIST OF TABLES
…………..………………………………...…….......
xix
LIST OF FIGURES
……….………………………………….................
xxi
LIST OF APPENDICES
…………..………………………………........ xxiii
………………………..………………………..........
1
………………..……………………………..
3
…………..…………………………….
9
RESULT AND DISCUSSION …………..………………………...……
General morphology of Rhynchoglossum in Malesia …………...
Habit …...................................................................................
Leaves …...............................................................................
Inflorescences …...................................................................
Flowers ….............................................................................
Fruits ….................................................................................
Distribution ……………...………………………………………
Phylogenetic analysis ….......………………………………..…...
Taxonomic Treatment ……….....………………………..………
Rhynchoglossum Blume …....................................................
Key to Species of Rhynchoglossum in Malesia ….................
Rhynnchoglossum borneense Merr. …...................................
Rhynchoglossum celebicum Karton. sp.nov. …......................
Rhynchoglossum klugioides C.B. Clarke …...........................
Rhynchoglossum medusothrix B.L. Burtt …..........................
Rhynchoglossum obliquum Blume ….....................................
Rhynchoglossum spumosum Elmer …....................................
13
14
14
14
16
17
18
19
21
25
25
26
27
30
33
36
39
45
CONCLUSION
…………..………………………………………...……
49
REFERENCES
…………..………………………………………...……
51
APPENDICES
…………...………………………………………...……
55
INTRODUCTION
LITERATURE REVIEW
MATERIAL AND METHODS
LIST OF TABLES
Page
1 The characters and characters state used in the phylogenetic analysis ....
10
2 Distribution of Rhynchoglossum in Malesia …………………………....
20
LIST OF FIGURES
Page
1 Distribution of Rhynchoglossum in yellow circle
….......……………….. 4
2 Illustration of Loxotis obliqua = Rhynchoglossum obliquum
(from Brown 1838)
……………………………...……………………..
5
3
Leave shape. A. Ovate of R. obliquum B. Ovate of R. klugioides
C. Oblong of R. celebicum …………………………………………...…. 15
4
Rhynchoglossum obliquum A. Inflorescence; B. Flowers ………..……... 17
5
Fruit shape A.Ovoid of R. obliquum B. Capsule-like of R. celebicum
C. Elongate of R. klugioides ……………………………………….…...... 19
6
Cladogram of phylogenetic tree of Rhynchoglossum in Malesia.
Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769.
The number shown above line showed supported the Bootstrap values …. 22
7
Rhynchoglossum borneense Merr., Elmer 21467 from Tawao, Malaysian
Borneo (isotype SING)
...…………………………...………………... 29
8 Distribution map of Rhynchoglossum borneense in Malesia in the circle …. 30
9 Distribution map of Rhynchoglossum celebicum in Malesia in the circle ... 31
10 Rhynchoglossum celebicum Karton., Eyma 1572 from Maraowa, Sulawesi
(holotype BO) …………………………………………...……………… 32
11 Distribution map of Rhynchoglossum klugioides in Malesia in the circle .... 34
12 Rhynchoglossum klugioides C.B.Clarke, Cuming 824 from Tayabas, Luzon
(holotype K) …………………………………………………………….. 35
13 Distribution map of Rhynchoglossum medusothrix in Malesia in the circle .. 37
14 Rhynchoglossum medusothrix B.L.Burtt, Kostermans 13994 from Berau,
East Kalimantan, Borneo (isotype BO) .………………………………… 38
15 Distribution map of Rhynchoglossum obliquum in Malesia in the circle ... 43
16 Rhynchoglossum obliquum Blume, Blume 52 from Java (lectotype L) …. 44
17 Distribution map of Rhynchoglossum spumosum in Malesia in the circle .. 46
18 Rhynchoglossum spumosum Elmer, Elmer 9929 from Cuernos Mts.,
Negros (Isotype US) …………..…………………………..……….…..... 47
LIST OF APPENDICES
Page
1
List of species Rhynchoglossum proposed in Malesia (Burtt 1962) …...
57
2
Matrix of 60 Morphological Characters for Maximum Parsimony
Analysis ………..………………………………………………………...
58
INTRODUCTION
Rhynchoglossum Blume was a genus of fleshy herb belongs to family
Gesneriaceae. It comprised of 10 species all over the world. It is classified in the
tribe Epithemateae or Epithematoid Gesneriaceae together with Epithema Blume,
Gyrogyne W.T. Wang, Loxonia Jack, Monophyllaea R. Br., Stauranthera Benth.,
dan Whytockia W.W. Sm. Rhynchoglossum was mainly found from India, Ceylon,
Nepal, South China, Formosa, Indochina, Thailand, and Malesia, with only one
species grow in Southern America (Mayer et.al. 2003; Weber 2004b).
The genus Rhynchoglossum was established by Blume (1826) with one
spesies of Rynchoglossum obliquum. The type specimen was collected from Java.
The species has a wide distribution from India, Ceylon, South China, Indochina,
and Malay Archipelago through New Guinea. Some genus allied like Antonia
R. Br., Glossanthus Benth., Klugia Schltdl., and Loxotis R. Br. are proposed later
as the synonym of Rhynchoglossum (Burtt 1962).
Rhynchoglossum can be recognized morphologically as fleshy herb plant
with anisophyllous decussate leaf or alternate leaf arrangement, asymmetrically
leaf blade and the unilaterally inflorescence. The genus has a little economic value
but several Botanic Gardens and private gardens are growing the plant as an
ornamental (Skog 1985). One of the most ornamental planted is Rhynchoglossum
gardneri Theobald & Grupe from India and Ceylon. The epithet name of
Rhynchoglossum is from Greek whereas Rhynchos means beak; and glossa means
tongue. The second part of the name clearly alludes to the broad, tongue-like
lower lip of the corolla, the first part perhaps to the narrow corolla tube or to the
pointed petal tips (Weber 2004b).
Weber (2004b) put Rhynchoglossum under Epithematoid Gesneriaceae.
This group has unique and special characters of unequal cotyledons,
asymmetrically leaf blade and capsule fruit. The distribution of the group was
mostly in tropical Asia, few from Africa, and one species from southern America.
The disjunction distribution from this group was very unique especially in
Rhynchoglossum.
Several taxonomic study of Rhynchoglossum has done since the genus
proposed by Blume. Most of the studies generally observed the family
Gesneriaceae,
whereas
the
Rhynchoglossum
was
included.
Some
of
comprehensive studied were by Bentham (1876) and Clarke (1883). The latest
study was a little revision done by Burtt (1962) which is stated that
Rhynchoglossum has 13 species and with very limited material specimens. From
many studies and revisions, there was no specific taxonomic revision for Malesian
region that known as the center of biodiversity for the genus. Revision about
species biodiversity and the number of species in Malesia has not been done
before. Based on that information, the revision of the genus Rhynchoglossum is
needed.
The research objective of this study was to revise taxonomic of
Rhynchoglossum in Malesia. Taxonomic revision was done to find the
information about generic and species delimitation, by providing data on its
biodiversity, identification key for species, analytic description for genus and
species, distribution pattern, and phylogenetic of Rhynchoglossum in Malesia.
LITERATURE REVIEW
Rhynchoglossum taxonomically included in Epithematoid Gesneriaceae
group or tribe Epithemateae along with others genera like Whytockia W.W. Sm.,
Gyrogyne W.T. Wang, Epithema Blume, Monophyllaea R. Br., Loxonia Jack, and
Stauranthera Benth. (Weber 2004b). Generally these groups have a similarity in
unequal cotyledons, unequal leaf arrangement in one node, asymmetrically leaf
blade and capsule fruits. Rhynchoglossum is allied with Loxonia and Stauranthera
in the shape of leaf arrangement. Rhynchoglossum has similarity with
Monophyllaea in the shape of inflorescences with unilateral cymose shape (Weber
2004b).
In the earlier status, Rhynchoglossum distributed from India, Ceylon,
China, Taiwan, Indochina to New Guinea in the tropical Asia (Weber 2004b).
Burtt (1962) included Klugia Schltdl. as synonymy of Rhynchoglossum, which is
Klugia has a distribution in Southern America from Mexico to Peru. The united of
Klugia as a synonym makes distribution of Rhynchoglossum wider and disjunct to
America (Figure 1). These disjunction distributions on the genus within
Australasia to America make Rhynchoglossum very unique and have an
interesting distribution history. Burtt (1998) give preliminary hypothesis that
Rhynchoglossum reach America from Asia via Africa from where it now has
completely disappeared or no species are found.
Weber (2004a; 2004b) assumed that the genus spreads from Asia to
America recently, probably via transpacific trips or migrations of early
Polynesians. This is also suggested by the current localities which are usually near
former populations centers of ancient dwellers along the pacific coast. Mayer
et.al. (2003) indicate that American species (R. azureum (Schltdl.) B.L. Burtt) is a
recent introduction to the neotropics and that it does not represent an ancient relic.
It seem R. azureum more allied to R. notonianum (Wall.) B.L.Burtt that known
from India than other species from Malesian region. The phylogenetic showed
that R. azureum allied in one clade with R. notonianum and reflects clearly
morphological not geographical pattern in the genus (Mayer et.al. 2003).
Figure 1 Distribution of Rhynchoglossum in yellow circle
Rhynchoglossum obliquum Blume is the common and widespread species,
found from India, China to Malesia (Bakhuizen van den Brink Jr. 1965; Burtt
1962). In the Southern America only have one species R. azureum (Schltdl.) B.L.
Burtt which has disjunctions distribution with its allies in Asia especially in
Southern India (Mayer et.al. 2003). Some of the other species (e.g. R. borneense
Merr., R. medusothrix B.L.Burtt, and R. spumosum Elmer) only have restricted
distributions and even with endemic pattern. Its preferably habitat is on the wet
and shady (especially limestone) rocks, in the forest or open vegetation or shady
places; usually grow in the lowlands (Weber 2004b).
Rhynchoglossum in general have specific characters like perennial or
annual, monocarpic herbs, stems with sparse hairs indumentum from often
branched, many-celled, to glandular hairs. Stems terete, fleshy-succulent, usually
branched. Leaves alternate, nearly distichous, short petiolate; lamina obliquely
ovate-cordate; base asymmetrical, one half cordate, the other sinuate-attenuate;
texture thin and delicate. Inflorescences terminal on main and side branches,
unilateral racemes with two rows of flowers, subtending bracts small, linear,
displaced onto the pedicels in most species; a few sterile bracts often present on
the flowerless side of the inflorescence axis. Sepals connate in the lower half, with
or without wings at the lines of fusion, lobes narrow triangular-acute. Corolla
strongly zygomorphic; tube cylindrical, white; limb strongly bilabiate, upper lip of
two short, upright or reclined lobes, lower lip a large, roundish or elongate, not or
only slightly three-partite tongue of azure colour. Fertile stamens four (the former
genus Klugia) or two (Rhynchoglossum s.str.). Ovary globose-ovoid. Fruit a
globose or ovoid capsule included in the calyx, dehiscing loculicidally by 2 valves
(Weber 2004b). The illustration of the genus show on Figure 2.
Figure 2 Illustration of Loxotis obliqua = Rhynchoglossum obliquum
(from Brown 1838)
Brown (1832) proposed a new genus of Antonia with the species of
A. obliqua (Wall.) R. Br. which is combination of Wulfenia obliqua Wall. (Weber
2004a). Three years later Brown moved the species into the genus Loxotis R. Br.
ex Benth. and made a new combination of L. obliqua (Wall.) R. Br.. The species
is surviving for a long time as well as Miquel (1855) use it. Later on Clarke
(1883) realizes that Antonia and Loxotis are superfluous because it is similar and
identical to Rhynchoglossum and then he put both genera under synonym of
Rhynchoglossum. Recently Burtt (1962) also included Klugia to a synonym
because it is only a variation of Rhynchoglossum.
The latest study by Burtt (1962) sign 13 species of Rhynchoglossum all
over the world comprising 10 species from Asiatic and 3 from America. After that
the number species of Rhynchoglossum are decrease because reducing some
species into synonym. Rhynchoglossum grandiflorum (Fritsch) B.L. Burtt and
R. violaceum (Fritsch) B.L. Burtt are turn into synonym of R. azureum because of
the similarity in morphology and geography (Wiehler 1983). Wang et.al. (1998)
reduce R. hologlosum Hayata into synonym of R. obliquum. Since the reduction of
some species into synonim, total number species of Rhynchoglossum known only
10 species all over the world (Mayer et.al. 2003; Weber 2004b).
History of the number species proposed in Malesia start when Blume
(1826) established the R. obliquum as the type of the genus from Java. After
Blume published the type then followed by de Candolle (1845) which established
another species R. blumei DC. from Java. Clarke (1883) included R. blumei to be
a synonym of R. obliquum and established a new species of R. klugioides C.B.
Clarke from Luzon, Philippines. The species epithet is given by Clarke as a
species which have similarity with Klugia. In the early of twentieth century,
Elmer (1908) proposed a new R. spumosum Elmer from Negros Philippines and
followed by Kraenzlin (1913) proposed R. merrilliae Kraenzl. from Mindanao,
Philippines. Ten years later, a new species of Rhynchoglossum proposed by
Schlechter (1923) which is from New Guinea and named as R. papuae Schltr.
Another species from Borneo proposed by Merrill (1929) and named as
R. borneense Merr. The last species that ever found is R. medusothrix which is
proposed by Burtt (1962) also from Borneo and completely sign seven species
number in Malesia (Appendix 1).
Phylogenetic analysis of Rhynchoglossum using molecular DNA was
studied together with another member of tribe Epithemateae. Phylogenetic tree
showed that Rhynchoglossum separate in different clade with another
Epithemateae and act as sister position. This separation is in accordance with
many and strong morphological differences (e.g., alternate leaf arrangement,
strongly asymmetrical leaves, terminal inflorescences in the form of unilateral
racemes, enlarged lower lip of corolla). The genus has many special characters in
common with the other Epithemateae, except Monophyllaea and Whytockia. In
the Rhynchoglossum group reflected in the phylogram by two clades;
R. notonianum (India) and the neotropical R. azureum vs. R. obliquum (Malesia).
The separation clearly indicated the affinity with morphological characters and
not the geographical pattern, which can be seen as an indication that R. azureum is
a recent introduction to the neotropics and that it does not represent an ancient
relict. The groups can be roughly classified in two: (1) perennials with large
flowers and four stamens (the former genus Klugia) and (2) annuals with small
flowers and two stamens (Mayer et.al. 2003).
MATERIALS AND METHODS
This study was carried out at the Systematic Laboratory of the Herbarium
Bogoriense, Research Center for Biology, Indonesian Institute of Sciences (LIPI)
at Cibinong, West Java and the Herbarium of Singapore Botanic Gardens,
Singapore National Parks from January 2010 to February 2011. It is based on 233
sheets of Herbarium specimens (including spirit collections) of Rhynchoglossum
from the Herbarium Bogoriense (BO) and the Herbarium of Singapore Botanic
Gardens (SING). A number of herbarium specimen photographs from National
Herbarium of Nederland-Universiteit Leiden Branch (L), Natural History
Museum of London (BM), Royal Botanic Garden of Kew (K), Royal Botanic
Garden of Edinburgh (E), Harvard University (A), Smithsonian Institute (US),
and New York Botanical Gardens (NY) were also studied.
The materials were studied based on morphological characters using 10 x
40 binocular microscope and stereo microscope. The information was taken under
the information data of location, general habitat, habit, stem, leaves,
inflorescences and infructescences. Morphological description was following
Veldkamp (1987). The revision methods were based on Rifai (1976, 2008) and
Vogel (1987).
Phylogenetic analysis of Malesian Rhynchoglossum was studied based on
60 morphological characters which collected during morphological observation
and analyzed based on Maximum Parsimony using PAUP version 4.0b.10
(Swofford 1998) with the Heuristic search settings vegetative and floral characters
were selected for the analysis, of which 47 were scored as binary and 13 as multistate (Table 1). In a few cases, some characters were scored as missing data when
the data were unavailable.
Table 1 The characters and characters state used in the phylogenetic analysis
NO
CHARACTERS
CHARACTER STATE
1
Growing habit
shrub (0); erect herb (1); creeping herb (2)
2
Habit
perennial (0); annual (1)
3
Height
≤ 30 cm (0); > 30 cm (1)
4
Stem diameter
≤ 3 mm (0); > 3 mm (1)
5
Leaf arrangement
opposite (0); alternate (1)
6
Leaf shape
ovate-elliptic (0); oblong (1)
7
Lamina
membranous (0); subcoriaceous (1)
8
Leaf color
pale green (0); dark green (1)
9
Leaf apex
acuminate (0); acute(1)
10
Apex tip
< 1 cm (0); 1 cm (1)
11
Leaf margin
entire (0); dentate or serrate (1)
12
Leaf base
rounded-cuneate (0); rounded-angustate (1);
cordate-cuneate (2); cordate-angustate (3)
13
Midrib indument
glabrous (0); puberulous (1); pubescent (2)
14
Abaxial leaf indument
glabrous (0); glabrescent (1); pubescent (2)
15
Petiole indument
glabrous (0); glabrescent (1); pubescent (2)
16
Petiole length
< 10 cm (0); 10-15 cm (1); > 15 cm (2)
17
Inflorescence position
axillar only (0); axillar and terminal (1)
18
Inflorescence growth
erect (0); spreading (1)
19
Terminal infl. length
≤ 15 cm (0); > 15 cm (1)
20
Axillar infl. length
≤ 5 cm (0); > 5 cm (1)
21
Inflorescence side
multilateral (0); unilateral (1)
22
Flower number
≤ 10 (0); > 10 (1)
23
Main axis indument
glabrous (0); puberulous (1); pubescent (2)
24
Peduncle length
≤ 3 cm (0); > 3 cm (1)
25
Bract shape
linier (0); oblong (1)
26
Bract length
≤ 4 mm (0); > 4 mm (1)
27
Pedicel indument
glabrous (0); glabrescent (1); puberulous (2);
pubescent (3)
__________________________________________________________________
Table 1 continued
NO
CHARACTERS
CHARACTER STATE
28
Pedicel length
≤ 3 mm (0); > 3 mm (1)
29
Bracteole length
≤ 5 mm (0); > 5 mm (1)
30
Bracteole position
rise from base pedicel (0); rise from middle
pedicel (1)
31
Flower length
≤ 15 mm (0); > 15 mm (1)
32
Calyx tube shape
campanulate (0); urceolate (1)
33
Calyx tube color
white (0); pale green (1); dark green (2)
34
Calyx tube adnate length
≤ 3 mm (0); > 3 mm (1)
35
Calyx lobe apex
acute (0); acuminate (1)
36
Calyx lobe length
≤ 3 mm (0); > 3 mm (1)
37
Corolla tube color
white (0); blue (1); green (2)
38
Corolla tube length
≤ 15 mm (0); > 15 mm (1)
39
Tube length
≤ 10 mm (0); > 10 mm (1)
40
Limb length
≤ 5 mm (0); > 5 mm (1)
41
Adaxial lip shape
2-lobed (0); emarginated (1)
42
Adaxial lip length
≤ 3 mm (0); > 3 mm (1)
43
Abaxial lip shape
tongue-like (0); 3-lobed (1)
44
Abaxial lip apex
rounded (0); obtuse (1)
45
Abaxial lip length
≤ 5 mm (0); > 5 mm (1)
46
Stamens number
2 (0); 4 (1)
47
Smaller stamen fil. shape
terete (0); flat (1)
48
Smaller stamen fil. length
≤ 2 mm (0); > 2 mm (1)
49
Smaller anther shape
ovoid (0); divaricate (1); reniform (2)
50
Smaller anther diameter
≤ 1 mm (0); > 1 mm (1)
51
Larger stamen fil. length
≤ 3 mm (0); > 3 mm (1)
52
Larger anther shape
oblong (0); reniform (1)
53
Ovary shape
globose (0); ovoid (1); ellipsoid (2);
oblong (3)
54
Ovary length
≤ 2 mm (0); > 2 mm (1)
__________________________________________________________________
Table 1 continued
NO
CHARACTERS
CHARACTER STATE
55
Style length
≤ 10 mm (0); > 10 mm (1)
56
Fruit shape
globose (0); ovoid (1); ellipsoid (2);
capsule-like (3); elongate (4)
57
Stalk length
≤ 10 mm (0); > 10 mm (1)
58
Stalk indument
glabrous (0); puberulous (1); pubescent (2)
59
Seed surface
tessellate (0); smooth (1)
60
Seed length
≤ 0.4 mm (0); > 0.4 mm (1)
All characters are threat as unorder data and have equal weight. The
“missing” data are symbolized with “?”. Starting tree(s) was obtained via stepwise
addition. Number of trees held at each step during stepwise addition = 1. Branch
swapping algorithm was run by using tree-bisection-reconnection (TBR),
evaluated using 100 bootstrap replicates and a complete heuristic search. The
complete data matrix is shown in appendix 2.
RESULT AND DISCUSSION
Based on this revision work of Rhynchoglossum in Malesia, there were six
species of Rhynchoglossum. The six species were Rhynchoglossum borneense
Merr., R. celebicum Karton. spec. nov., R. klugioides C.B.Clarke, R. medusothrix
B. L. Burtt, R. obliquum Blume, and R. spumosum Elmer. There was no
infraspecific status decided in this revision. Rhynchoglossum celebicum is one of
new species that newly described here beside the other five species.
New status has been given for some species in this revision.
Rhynchoglossum papuae Schltr. from New Guinea now turn into synonym of
R. obliquum. Another species is R. merrilliae Kraenzl. from Philippine now under
synonym of R. spumosum. This new status decreasing the species number of
Rhynchoglossum in Malesia that giving by Burtt (1962) from seven to six here
with new species that newly described (R. celebicum).
Based on preliminary revision, Burtt (1962) proposed a seven species of
Rhynchoglossum comprising R. borneense, R. klugioides, R. medusothrix,
R. merrilliae, R, obliquum, R. papuae, and R. spumosum (Appendix 1). In this
recent revision, the number species are decreased because the status of two
species changed. Rhynchoglossum merrilliae is synonym of R. spumosum due to
its similarity on their morphological characters such as dwarf habit, small leaf
blade, few flower in the inflorescence and two fertile stamens. Based on the
observation to type specimens, it is realized that both are actually one species and
R. spumosum is used because this species name used earlier.
Rhynchoglossum papuae was proposed by Schlechter (1923) based on the
small corolla lip and globose or ovoid capsules comparing with R obliquum.
When more specimens were examined from New Guinea in this revision, it is
seen that most characters from each specimen have no different with R. obliquum
in other islands of Malesia. This consideration makes R. papuae is closely related
with R. obliquum and placed as a synonym. By placing two species as synonym in
this revision, the number of species became decreasing than the number species
that given by Burtt (1962) from seven species into five species. By adding the
new species (R. celebicum) in this revision, it showed that the total number of
Rhynchoglossum in Malesia become six species.
General Morphology of Rhynchoglossum in Malesia
Habit
Rhynchoglossum are erect and creeeping fleshy-succulent herbs plant from
5-150 cm height. The roots are adventitious with rhizomatous or non-rhizomatous.
The only rhizomatous species is R. spumosum is being creeping in habit and
dwarfing about 5-8 cm height. The stems usually are terete and wrinkled when
dried. They have a variable indument, the hairs being absent or simple puberulous
to pubescent. Most of the species have glabrous and smooth stem surface and
have swollen nodes.
Rhynchoglossum in broad sense have a growing habit as an herb with
perennial or annual growth. All Malesian species known have annual growth with
fleshy succulent herb. Rhynchoglossum spumosum is the only species have
creeping habit with rhizomatous root system. This habit makes the species has
only maximum height 8 cm known as a dwarf Rhynchoglossum. Another species
that reporting have rhizomatous habit is R. omeiense W.T.Wang from China, but
different in having perennial growth (Wang et.al. 1998). Because of that it is
necessary to study further by their molecular systematic to understand the
uncontinous charcters occurs in several species.
Leaves
Leaf arrangements are alternate and exstipulate. The leaf blade of all
Rhynchoglossum is oblique with asymmetrical shape between two sides.
Sometimes the bases of both size are in equal position or different in length till 2.5
cm in R. borneense. The leaf shape varies from ovate to oblong. Rhynchoglossum
celebicum is the only species which have oblong leaf (Figure 3B) while the other
species varies from ovate to elliptic (Figure 3A). The leaf bases are unequal which
one side has cuneate to acute shape and the other side has rounded to cordate
shape. The leaf margins are mostly entire only several species are serrate. The leaf
blades texture are mostly membranous, only R. klugioides have subcoriaceous leaf
blade. Most of the species have glabrous and smooth leaf surface. Some have
pubescent or puberulous indumentum when juvenile.
A
B
C
Figure 3 Leave shape. A. Elliptic of R. obliquum B. Ovate of R. klugioides
C. Oblong of R. celebicum
Rhynchoglossum have alternate leaf arrangement with no stipules. Different
from other Gesneriaceae which have opposite leaf arrangement Rhynchoglossum
have unique leaves development where the opposite leaf is reduced and become
alternate. Weber (2004b) gives hypothesis that the alternate leaf arrangement is
evolution trend in the genus because of the reduction of the opposite side. The
other allied genera like Stauranthera and Loxonia also have similar development
in leaf arrangement with anisophylly leaf arrangement, not reducing like
Rhynchoglossum. The opposite sides in the leaf arrangement are usually free or
with the inflorescences.
Most of the species of Rhynchoglossum known have ovate to elliptic leaf
shape. In this revision a new character of leaf which is oblong belongs to
R. celebicum. This different on its leaf characters made R. celebicum distinct and
realized as a new species. The leaf texture of Rhynchoglossum has two types, one
usually membranous (most of Malesian species) and the other one is subcoriceous
(R. klugioides). The subcoriaceous texture is related with the species that formerly
belongs to Klugia (known as R. azureum and R. notonianum recently).
Inflorescences
The inflorescence in Rhynchoglossum is raceme with terminally and axillary
grow. The axillary inflorescence are usually opposite with leaf. The inflorescences
are spreading or secundiflorous which is not erect (Figure 4A). Number of flower
in inflorescence varies from less than 3 in R. spumosum to 50 in R. obliquum.
Flowers in inflorescence are unilateral because the reduction of other side flowers
and arrange in two rows. The bracts usually absent in R. spumosum or occur in
linier shape in R. obliquum. Sometimes in inflorescences have bracts that not
supported flower or known as sterile bracts. Bracteoles in Rhynchoglossum are
mostly linier which rise from base of pedicel in R. klugioides and R. spumosum;
and rise from the middle of pedicel in other species.
Inflorescences of Rhynchoglossum are known as raceme with secundiflorous
form. These inflorescences are usually with only one side flowering or unilateral
with two rows. This unilaterally growth is obtained because of the reduction on
flower growth in other pairs of inflorescences (Weber 2004b). The reduction side
of the flower is sometimes has bracts which is not subtending and known as sterile
bracts. The prominent difference in inflorescences is observed on the rise of
bracteoles. Rhynchoglossum klugioides and R. spumosum known have bracteoles
from the base of pedicels while the other species have bracteoles rise from the
middle of pedicel.
A
B
Figure 4 Rhynchoglossum obliquum A. Inflorescence; B. Flowers
Flowers
The flowers are zygomorphic, tube-like and vary from less than 1 cm long
till more than 3 cm long. Calyx tubes are infundibuliformis with campanulate
shape in most species or urceolate shape in R. klugioides. The calyx adnate at base
sometimes winged with line fushion; white to green colour. Calyx lobes are
triangular and entire, glabrous, 5-merous. Corolla tubes are glabrous to
puberulent, with tube and limb. The limb has two lipped which is from the petal
fused (Figure 4B). The adaxial lip usually smaller with two lobes and the abaxial
lip larger 3-lobed or undivided bearing a rounded elongate or tongue–like. The
petals usually have a bluish to dark purple color and a few is white. Some species
like R. borneense, R. medusothrix and R. klugioides have a yellow dot pilose in
the throat of abaxial limb. The pilose dot of limb in R. medusothrix usually being
medussoid and dense.
Two enlarged lips corolla with enlarge lower one in the flower of
Rhynchoglossum are common in the genus. The variation among the species is
usually shown in the lower lip characters. The species that having small flower or
corolla (R. obliquum, R. spumosum and R. celebicum) have tongue-like lower lips
and the species with large flower or corolla (R. klugioides, R. borneense and
R. mrdusothrix) have 3-lobes, broader and elongate. In the species that have large
corollas has pilose dot in throat of lower lip but vary from less to dense pilose,
while in R. medusothrix with medussoid pilose hair.
Rhynchoglossum in Malesia have 2 or 4 stamens. Rhynchoglossum
obliquum, R. klugioides, R. celebicum and R. spumosum have 2 stamens.
Otherwise, R borneense and R. medusothrix have 4 stamens. All stamens in
flowers are arranged coherent in pairs. The stamens are adnate to corolla tube in
adaxial limb if only 2 stamens or in abaxial limb if there 4 stamens. The anthers
are basifixed, thecae parallel and dehiscing longitudinally. Filaments are usually
flat. The ovaries in most Rhynchoglossum are ovoid with one loculed. Stigma are
capitates or undivided and the style are usually glabrous and still remnant in
fruiting time.
Four numbers of stamens in Malesian Rhynchoglossum allied with another
species of Rhynchoglossum (R. notonianum (Wall.) B.L.Burtt and R. azureum
(Schltd.) B.L.Burtt) was previously from the genus Klugia. The different with the
species outside Malesia is in the arrangement of the stamens. R. notonianum and
R. azureum have four equal stamens that coherent together while R. borneense
and R. medusothrix have four unequal stamens (two larger and other two smaller
stamens). The arrangement of the stamens is also coherent but in pair not together
(dydinamous).
Fruits
The fruits in all Rhynchoglossum are capsule type with one loculed. The
shape of the fruits may vary from capsule-like, ovoid to elongate with prominent
calyx remnants (Figure 5). The calyx remnants are half closing the fruit in
R. celebicum (Figure 5C) and fully closing the fruit in other species (Figure 5A &
B). The fruits are dehiscing loculicidally to base with two valvate, not twisted.
The seeds are cuneate and unappendaged.
The shape of the fruits of Rhynchoglossum is varying. The ovoid fruit
shape is known oly from R. obliquum, R. borneense, R. medusothrix and R.
spumosum with calyx remnant closing fully the fruit and persistent.
Rhynchoglossum klugioides has elongate fruit shape and also closed by the
persistent calyx. The elongate shape is common in other species outside Malesia
that have large corollas (e.g. R. notonianum and R. gardneri). The only species
that not fully closed by the calyx is R. celebicum and also has capsule-like shape.
A
B
C
Figure 5
Fruit shape A. Ovoid of R. obliquum B. Elongate of R. klugioides
C. Capsule-like of R. celebicum
Distribution
Rhynchoglossum approximately have 10 species of fleshy herbs that found
throughout South Asia, China, South East Asia and Malesia to Southern America
(Weber, 2004). The Malesian region has about six species and five of them are
endemic to the region (Table 2). Rhynchoglossum obliquum is a widespread and
very common species distributed from Sumatra to New Guinea. Rhynchoglossum
klugioides was previously only known from Philippines currently distributed also
to Seram Island, Moluccas. The other four species are endemic in each region.
Rhynchoglossum spumosum was endemic to Philippines from Negros Island and
Mindanao Island, while the R. celebicum endemic to Sulawesi. Rhynchoglossum
borneense and R. medusothrix are endemic to Borneo which is recorded only
found in the eastern part of the island.
Table 2 Distribution of Rhynchoglossum in Malesia
Species
Sumatra
Peninsular
Borneo
Java
Sulawesi
Malaya
Lesser
Moluccas
Philippines
Sunda
New
Guinea
√
R. borneense
√
R. celebicum
R. klugioides
√
R. obliquum
√
√
√
√
√
R. medusothrix
√
√
√
√
√
√
√
R. spumosum
Based on the distribution pattern, the center of the diversity
Rhynchoglossum in Malesia is found in Borneo, Philippine and Sulawesi. Borneo
has three species (R. borneense, R. medusothrix, and R. obliquum), Philippine has
three species (R. klugioides, R. spumosum, and R. obliquum) while Sulawesi has
two species (R celebicum and R. obliquum). The regions shows the high number
of Rhynchoglossum species found comparing to the other islands like Sumatra,
Peninsular Malaya, Java, Lesser Sunda, Moluccas and New Guinea. The
endemicity of Rhynchoglossum in the center of diversity is also high. Beside
R. obliquum, the rest five species known endemic in that area while a few
R. klugioides also observed in Moluccas.
The phytogeography pattern for Rhynchoglossum is still related with the
pattern of all Asiatic Rhynchoglossum. Burtt (1998) said that Gesneriaceae is a
family
of
Gondwanic
origin.
Epithematoid
Gesneriaceae
which
is
Rhynchoglossum inside is a relict group that was once much larger and had much
wider distribution. The ancient of Rhynchoglossum is known from Asiatic
mainlad which now has the most species occur. Based on that hypothesis is
predicted that the Malesian species are spreading from west Wallace line to east
line. The species that found in the east Malesia are R. obliquum and R. klugioides
which known from Moluccas and New Guinea. In fact, both species also known
distributed in the west of Malesia like Sumatra, Java, Borneo and Philippine.
Rhynchoglossum celebicum is the only species that found in Sulawesi which is the
intermediate region between west and east Malesia. Distribution pattern of
Rhynchoglossum in Malesia made the genus is typically Sunda shelf or West
Malesia taxon.
Rhynchoglossum can be found from lowland areas to mountain areas of
1500 m above sea level at the edge of the forest, close to water shaded or in the
rock of limestone area. Most of the species like R. obliquum, R. celebicum,
R. klugioides and R. spumosum found in the forest with canopy or edge forest with
wet condition near the river. Rhynchoglossum borneense and R. medusothrix are
known from the limestone habitat in Borneo. It is looks that the preferable habitat
impacts the distribution of species.
Phylogenetic analysis
The phylogenetic relationships of the genus Rhynchoglossum in Malesia
was analyzed in the present study based on morphological characters. Sixty
characters were used in this analysis (Table 1). The sixty characters were selected
for their apparent taxonomic values that are good to separate one taxon with the
others.
Phylogenetic analysis indicated that all characters are parsimony
informative. The 128 equally most parsimonious tree of 138 steps produce
consistency index (CI) = 0.6562, homoplasy index (HI) = 0.3438 and retention
index (RI) = 0.5769. The higher CI value was resulted that the value of
homoplasy rather low. The value of RI above 0.5 indicated that these characters
could contribute some synapomorphy to the cladogram. Figure 8 showed tree
number 1 of 128 the most parsimonious trees.
Figure 6 Cladogram of phylogenetic tree of Rhynchoglossum in Malesia.
Tree number 1 of 128 the most parsimonious trees of the genus
Rhynchoglossum in Malesia. CI= 0.6562, HI= 0.3438, and RI= 0.5769.
The number shown above line showed supported the Bootstrap values.
The separation of the ingroup and outgroup in the phylogenetic analysis
was supported with a high bo