Merremia dennstedt ex endlicher (Convolvulaceae) in Sumatra.
MERREMIA DENNSTEDT EX ENDLICHER
(CONVOLVULACEAE)
IN SUMATRA
HAFNI RAHMADANI
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2013
LETTER OF STATEMENT
I express that this thesis entitled:
MERREMIA DENNSTEDT EX ENDLICHER (CONVOLVULACEAE) IN
SUMATRA
Has been compiled and written by myself and it is true represent result of my own
research and has never been previously submitted in whole or part for another degree. All
information and data that used have been expressed clearly and can be checked its truth.
Bogor, October 2013
Hafni Rahmadani
NRP. G353100131
SUMMARY
HAFNI RAHMADANI. Merremia Dennstedt ex Endlicher (Convolvulaceae) in
Sumatra. Supervised by Dr. SRI SUDARMIYATI TJITROSODIRDJO and Dr. HARRY
WIRIADINATA
Merremia is a small genus in the family Convolvulaceae. Merremia is a
perennial or annual herb or woody liana with stems creeping or twinning on trees
or rocks, put out milky white sap when injured and some species with tubers.
Merremia is often confused with genus Ipomoea, due to they have resemblance in
funnel shaped or campanulate corolla and straight anther. Phenetically, some
members of Merremia have yellow flower whereas Ipomoea has white, purple or
purplish white. Merremia is also confused with Operculina due both have nonspinulose pollen and dehiscent capsule. Nevertheless, Operculina characterized by
circumciselli capsule and Merremia characterized by valve capsule. The important
character in Merremia is the spread of the hairs on corolla that can be easily
observed at the mature flower bud. The genus Merremia is found in the tropical
regions of both hemispheres. Merremia wildly distributed throughout the tropical
regions of Asia, Africa, Australia, North and South America and approximately
there were 80 species of Merremia in the world.
A Total of 175 sheets (164 collection numbers) which are deposited in
Herbarium Bogoriense, 97 sheets (95 collection numbers) which are deposited
Andalas University Herbarium (ANDA) and 9 sheets (4 collection numbers)
which are deposited in BIOTROP Herbarium (BIOT) were studied. It composed
of all specimens of Merremia in Sumatra. Additional specimens were collected by
the author 28 sheets (15 collection numbers) from North Sumatra, West Sumatra
and Lampung, which are stored in BO and BIOT. The digital data that consist of
type specimens and distributions of species from JSTOR Plant Science and
Convolvulaceae Unlimited were also used here.
All the hebarium specimens were studied based on their morphological
characters following de Vogel (1987) and Rifai (1976). The terminology of
morphological characters followed Lawrence (1955), Veldkamp (1987), Harris
and Harris (1994), Hickey and King (2005) and Stearn (1983). Information on the
habitat, ecology, distribution, vernacular names and uses were noted from the
specimens label and literatures.
Taxonomic studies in Sumatra were conducted. There are eight species,
one subspecies, one varietas and two forma from nine previously species (M.
cissoides, M. dissecta, M. emarginata, M. hirta, M. peltata, M. quinquefolia, M.
tridentata, M. tuberosa and M. vitifolia), two subspecies (M. umbellata ssp.
orientalis and M. tridentata ssp. hastata), one varietas (M. boisiana var.
sumatrana) and two forma (M. hederacea f. pubescens and M. hederacea f.
barbata) can be recognized. One species (M. tridentata) and subspecies (M.
tridentata ssp. hastata) are excluded because they have been proposed to different
new taxa Xenostegia by Austin and Staples. The existence of M. emarginata was
noted as new record in Sumatra and M. cissoides was noted as new record in
Indonesia.
A phylogenetic analysis using PAUP*4.Ob10 was undertaken based on
morphological characters, using Evolvulus nummularius and Porana volubis as
the out group. The phylogenetic analysis used 35 morphological characters was
resulted consistency index (CI) = 0.60, homoplasy index (HI) = 0.40, retention
index (RI) = 0.56 and Rescaled consistency index (RC) = 0.34.
Phylogenetic tree used morphological characters divided taxa into two
groups. The separation support M. boisiana var. sumatrana, M. cissoides, M.
dissecta, M. emarginata, M. hederacea f. pubescens, M. hederacea f. barbata, M.
hirta, M. peltata, M. quinquefolia, M. tuberosa, M. vitifolia and M. umbellata ssp.
orientalis as ingroup with E. nummularius and P. volubis as outgroup. This study
support the monophyly, paraphyly and metaphyly of genus Merremia in Sumatra.
Key words: Convolvulaceae, Merremia, morphology, phylogenetic, Sumatra
RINGKASAN
HAFNI RAHMADANI. Merremia Dennstedt ex Endlicher (Convolvulaceae) di
Sumatera. Dibimbing oleh Dr. SRI SUDARMIYATI TJITROSODIRDJO dan Dr.
HARRY WIRIADINATA
Merremia merupakan marga dari Convolvulaceae yang mempunyai ciri-ciri
merupakan herba atau semak dengan batang memanjang, menjalar atau merambat
sampai 20 m, membelit pada pucuknya, beberapa jenis dengan perakaran yang
mempunyai umbi, mengeluarkan cairan putih seperti susu ketika dilukai (Stone
1970). Marga Merremia sering rancu dengan marga Ipomoea yang mempunyai
kemiripan pada mahkota bunga yang berbentuk funnel atau campanulate dan
benang sari yang berbentuk straight. Bunga Merremia biasanya berwarna kuning
sedangkan Ipomoea berwarna putih, ungu ataupun putih keunguan. Marga ini juga
rancu dengan Marga Operculina, yang sama-sama memiliki polen non-spinulose
dan buah dengan kapsul yang secara teratur akan pecah. Akan tetapi pada
Operculina kapsul berbentuk circumcisseli dan pada Merremia berbentuk valve.
Penyebaran bulu-bulu yang terdapat di mahkota bunga adalah ciri yang penting
dalam Merremia dan dapat dilihat dengan mudah pada kuncup bunga yang telah
dewasa. Di dunia, Marga Merremia diperkirakan terdapat 80 Jenis dan menyebar
luas di kawasan tropis mulai dari Afrika, Asia, Australia, Amerika Utara dan
Amerika Selatan.
Penelitian ini berdasarkan spesimen herbarium Merremia yang dikoleksi
dari Sumatera. Sebanyak 175 spesimen yang tersimpan di Herbarium Bogoriense
(BO), 9 spesimen di Herbarium Biotrop (Biot), 97 spesimen di Herbarium
Universitas Andalas (ANDA) dan 28 Spesimen dari hasil eksplorasi di Sumatera
Utara, Sumatera Barat dan Lampung. Data digital yang terdiri dari spesimen tipe
dan penyebaran spesies dari JSTOR Plant Science dan Convolvulaceae Unlimited juga
digunakan.
Semua spesimen herbarium dilihat berdasarkan karakter-karakter
morfologinya yang mengacu pada metode Rifai (1976) dan Vogel (1987).
Terminologi karakter morfologi mengacu pada Lawrence (1955), Veldkamp
(1987), Harris dan Harris (1994), Hickey dan King (2005) dan Stearn (1983).
Informasi habitat, ekologi, distribusi, nama daerah dan lain sebagainya di catat
berdasarkan label dan literatur spesimen.
Kajian taksonomi mengenai marga Merremia Dennstedt ex Endlicher
(Convolvulaceae) di Sumatera yang berdasarkan pada karakter morfologi telah
dilakukan. Dari studi ini terdapat 8 jenis Merremia, 1 anak jenis, 1 varitas dan 2
forma dari 9 jenis sebelumnya (M. cissoides, M. dissecta, M. emarginata, M.
hirta, M. peltata, M. quinquefolia, M. tridentata, M. tuberosa dan M. vitifolia), 2
anak jenis (M. umbellata ssp. orientalis dan M. tridentata ssp. hastata), 1 varitas
(M. boisiana var. sumatrana) dan 2 forma (M. hederacea f. pubescens dan M.
hederacea f. barbata) telah diperoleh 1 jenis (M. tridentata) dan anak jenis (M.
tridentata ssp. hastata) dikeluarkan karena telah diajukan menjadi takson baru
yaitu Xenostegia oleh Austin dan Staples. M. emarginata merupakan catatan baru
di Sumatera dan M. cissoides merupakan catatan baru di Indonesia.
Analisa hubungan kekerabatan menggunakan Program PAUP*4.Ob10
dengan E. nummularifolius dan P. volubis sebagai outgroup. Analisis
phylogenetic menggunakan 35 karakter morfologi menghasilkan consistensi index
(CI)=0.60, homoplasi index (HI)=0.40, retention index (RI)=0.56 dan rescaled
consistency index (RC)=0.34.
Pohon filogenetik berdasarkan karakter morfologi membagi takson dalam
dua kelompok. Pemisahan itu mendukung M. boisiana var. sumatrana, M.
cissoides, M. dissecta, M. emarginata, M. hederacea f. pubescens, M. hederacea
f. barbata, M. hirta, M. peltata, M. quinquefolia, M. tuberosa, M. vitifolia and M.
umbellata ssp. orientalis sebagai ingroup dengan E. nummularius dan P. volubis
sebagai outgroup. Studi marga Merremia ini memperlihatkan terjadinya
monophyly, paraphyly dan metaphyly.
Kata kunci: Convolvulaceae, Merremia, morfologi, phylogenetic, Sumatera
Copyright©2013, Bogor Agricultural University
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the written permission from Bogor Agricultural University.
MERREMIA DENNSTEDT EX ENDLICHER
(CONVOLVULACEAE)
IN SUMATRA
HAFNI RAHMADANI
Thesis submitted
As partial fulfillment requirement for the Master Degree
In Plant Taxonomy
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2013
External examiner: Dr.Rugayah, M.Si.
Title
Name
NRP
: Merremia Dennstedt ex Endlicher (Convolvulaceae) in Sumatra
: Hafni Rahmadani
: G353100131
Certified by
Supervisor Committee
Dr. Sri Sudarmiyati Tjitrosoedirdjo. M.Sc
Chairman
Dr. Harry Wiriadinata
Member
Approved by
Head of Plant Biology
Study Program
Dr. Ir Mifiahudin, M.si
Date of Examination: 30 July 2013
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Graduation:
o3
OCT 2013
Title
Name
NRP
: Merremia Dennstedt ex Endlicher (Convolvulaceae) in Sumatra
: Hafni Rahmadani
: G353100131
Certified by
Supervisor Committee
Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc
Chairman
Dr. Harry Wiriadinata
Member
Approved by
Head of Plant Biology
Study Program
Dean of Graduate School
Dr. Ir Miftahudin, M.Si
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Examination: 30 July 2013
Date of Graduation:
ACKNOWLEDGMENT
Thanks to Allah that I have finished my thesis. I would like to express my
gratitude to my supervisors, Dr. Sri Sudarmiyati Tjitrosoedirdjo and Dr. Harry
Wiriadinata for their valuable advices, guidance, and encouragement throughtout
my study. I gratefully acknowledge the DIPA Project 2011 through Dr. Sri
Sudarmiyati Tjitrosoedirdjo from Southeast Asean Regional Centre of Tropical
Biologi (SEAMEO BIOTROP).
I would like to express my gratitude to Herbarium Bogoriense (LIPI),
Herbarium Andalas University (ANDA) and Herbarium BIOTROP (BIOT), Bukit
Barisan Selatan National Park for granting me permission to conduct this research
and providing me some facilities. In particularly I would like to thanks all
taxonomist and technicians who gave their support and assisted me when
conducted this study.
I am also very grateful to Dr. George Staples (Singapore Botanical
Garden), Dr. Rugayah and Lulut Dwi Sulistyaningsih M.Si (BO), Budi Setiabudi
S.Hut (SEAMEO BIOTROP), for their interest, discussion and valuable comment
on the manuscript. Many thanks to my friends Azwar Haris Siregar, Indah
Wahyuni, Rahmanida and Alfiah Rahmi.
Last but not the least, my special thanks to my family, my father H. Edi
Syahputra, my mother Hj. Faridawati Siregar, and my brother Riza Afriandi for
their spirit, patience, and supporting of my study.
Bogor,
October 2013
Hafni Rahmadani
TABLE OF CONTENTS
LIST OF TABLES
vi
LIST OF FIGURES
vi
LIST OF APPENDIX
vi
1
INTRODUCTION
1
2
LITERATURE REVIEW
Taksonomy of Merremia Dennst. ex Endlicher
Distribution species of Merremia
Economic value of Merremia
2
3
4
MATERIALS AND METHODS5
Materials
Methods
Phylogenetic analysis.
4
4
4
3
4
RESULT AND DISCUSSIONS
Delimitation of Genus and Species Diversity
General Morpholgy Merremia Dennstedt ex Endlicher of Sumatra
Habit
Stem
Leaves
Inflorescences
Flower buds
Flower
Calyx
Corolla
Stamen
Pistill
Fruit
Seed
Distribution of Merremia in Sumatra
Phylogenetic analysis
Generic description
Key to the Sumatran species of Merremia
Taxonomic treatment
Merremia boisiana var. sumatrana
Merremia cissoides
Merremia dissecta
Merremia emarginata
Merremia hederacea
Merremia hirta
Merremia peltata
5
5
5
5
6
7
7
8
8
8
9
10
10
10
10
12
17
18
21
23
24
27
29
30
Merremia quiquefolia
Merremia tuberosa
Merremia umbellata
Merremia vitifolia
5
33
34
36
38
CONCLUSION
41
REFERENCES
41
APPENDIX
44
CURRICULUM VITAE
45
LIST OF TABLES
1. Distribution of Merremia in Sumatra
2. Morphological characters state used in phylogenetic analysis
12
13
LIST OF FIGURES
1. Distribution of Merremia
2. Palmately compound leaves
3. Variation blade of simple leaves
4. Variation ramified inflorecences
5. Variation corollas
6. Variation shapes anther
7. Variation capsules
8. Distribution of Merremia in Sumatra
9. Strict consensus tree resulted from morphological characters
10. Distribution of Merremia var. sumatrana in Sumatra
11. Merremia boisiana var. sumatrana
12. Merremia cissoides
13. Distribution of Merremia cissoides in Sumatra
14. Distribution of Merremia dissecta in Sumatra
15. Distribution of Merremia emarginata in Sumatra
16. Merremia emarginata
17. Distribution of Merremia hederacea in Sumatra
18. Distribution of Merremia hirta in Sumatra
19. Merremia peltata
20. Distribution of Merremia peltata in Sumatra
21. Distribution of Merremia quinquefolia in Sumatra
22. Distribution of Merremia tuberosa in Sumatra
23. Distribution of Merremia umbellata ssp. orientalis in Sumatra
24. Distribution of Merremia vitifolia in Sumatra
25. Merremia vitifolia
3
6
7
8
9
9
10
11
14
19
20
22
23
24
25
26
28
30
32
33
34
35
37
39
40
LIST OF APPENDIX
1. Matrix of morphological characters for phylogenetic analysis
44
1 INTRODUCTION
Invasive plant is a plant that can successfully adapt in its new habitat and
spread very widely, which would then occupy and replace the native plants and
finally would threaten the diversity, structure and function of ecosystem
(Radosevich et al. 2007). The invasion begins of migration, aggregation and then
competition. The characteristics of invasive plants are able to grow quickly,
reproduce vegetatively, high spread, tolerance to environmental condition, have
not natural enemies and usually associated with humans.
The twinning or creeping species with fast growth rates can become
troublesome weeds of agriculture and forestry. Several Merremia species have
become serious weeds of forestry plantations in Sumatra. Merremia peltata is a
big problem in Bukit Barisan Selatan National Park, due the National park is
habitat of protected animal such as Tiger, Elephant and Sumatran Rhino. This
National park has area with 356.800 ha (BBTNBBS 2009) and the invasion of
Merremia peltata has reached more than 7.008 ha even has caused animals life
were interrupted, so the animals migrate to more northern areas, which are the
village, estate and inflict the conflict with local people (Irianto and Tjitrosoedirdjo
2010).
Merremia is a small genus belonging to the family Convolvulaceae which
perennial or annual herb or woody with stems creeping or twinning on trees or
rocks, put out milky white sap when injured and some species with tubers (Stone
1970). Merremia is often confused with genus Ipomoea, due to a resemblance in
funnel shaped or campanulate corolla and straight anther. Phenetically, some
members of Merremia have yellow flowers whereas Ipomoea has white, purple or
purplish white. Merremia is also confused with Operculina because both have
non-spinulose pollen and dehiscent capsule. Nevertheless, Operculina is
characterized by circumciselli capsule, while Merremia is characterized by valve
capsule (Staples 2010). The important character in Merremia is the spread of the
hairs on corolla that can be easily observed at the mature flower bud (Fang and
Staples 1995).
The genus Merremia was recognized by Hallier (1893). He separated it
from Ipomoea almost entirely on the basis of its non-spinulose pollen grains and
five distict mid-petaline bands of corolla. He recognized several sections of
Merremia, namely Skinneria, Xanthips, Streptandra (Hallier 1894) and also
adding one more section that is Hailale ( Hallier 1913). Van Ooststroom and
Hoogland (1953) replaced the name Skinneria by Eu-Merremia and established
Wavula as the fifth section.
Taxonomic research of Merremia has been done by Ooststroom and
Hoogland (1953) who recorded 23 species Merremia in Malesian region and
Backer and Backhuizen Jr. (1965) recorded 12 species occur in Java.
Unfortunately taxonomic studies of Merremia in Sumatra is lacking since there
are inconsistencies in number of species proposed for inclusion in Merremia of
Sumatra e.g. Ooststroom and Hoogland (1953) mentioned there were 3 species
Merremia and on the other hand Staples (2010) mentioned 5 species.
The relationship between species of Merremia has been studied but there
were still difference of opinion. Phylogenetic studies based on molecular data of
Convolvulaceae (Stefanovic 2002) with 6 species of Merremia (M. aegyptia, M.
2
dissecta, M. hastata, M. peltata, M. umbellata and M. vitifolia) have indicated that
tribe is not monopohyletic, and that its largest genus Merremia is polyphyletic.
Relationship studies of Convolvulacae based on chemical markers reported that
Merremia containing flavonoids, quinonones, saponins, phenolic acids, and
alkaloids. Based of the distribution flavonoids, indicated that no close relationship
within the genus (Nair 2012).
The aim of this study is to understand the species diversity of Merremia
and their distribution pattern, to provide an identification key to the species, and to
determine relationship among the species of Merremia in Sumatera using
morphological character for cladistic analysis.
2 LITERATURE REVIEW
Taksonomy Merremia Dennstedt ex Endlicher
The Genus name of Merremia firstly published by Dennstedt (1818) which
the combination of Merremia convolvulacea. The name was appeared based on
plate of Rheede in Hortus Malabaricus without description and delimitation of
genus so the name is not validly published. After that some botanists mentioned
the genus in many publications with more detail to describe this plant, either by
the introduction of a new species or displacements that caused change of the
names among other species of genus.
Choisy (1833) renames Merremia to be Skinneria and Bojer (1837)
renames Merremia to be Spiranthera. However both of them were rejected
because the name Skinnera has been used on Onagraceae by Forster (1775) and
the name of Spiranthera has been used by St.Hil (1823) on Rutaceae. Endlicher
(1841) had accomplished the description Merremia in which the name became
validly published and recorded in International Code of Botanical Nomenclatur
and the status is set as Nomina Conservanda (Art. 14.8.), unfortunately the
description was still incorporated with Ipomoea.
Hallier (1893) had made genus Merremia delimitation and separated it
from Ipomoea based on pollen character in which Ipomoea has spinulose pollen
while Merremia has non-spinulose pollen. Furthermore, Hallier (1894) had
divided genus Merremia into 3 sections consist of Xanthips, Streptandra and
Skinneria. In 1913 Wingkler complements writings of section by adding type of
species at each section and furthermore he adding one more section that is
Hailale. Van Ooststroom and Hoogland (1953) adding one more section that is
Wavula and renames of Skinneria to be Eu-Merremia because the name of
Skinneria has been used as genus Skinnera Forst.
Morphological studies of pollen first done by Hallier (1893) where the
character is very important to recognize members of Convolvulaceae. After that
some botanists such as Erdtman (1952), Manitz (1969) and Huang (1972) worked
with pollen although limited in some regions. Sengupta (1972) conducted a study
of pollen extensively and discovered some type of pollen and evolutionary
relationship, although the work was not covering all species of Convolvulaceae.
The genus Merremia is confused with Operculina that has same nonspinulose pollen and capsule will break regularly. The pollen from 55 species
Merremia and Operculina have been studied and groups into five types:
3
tricolpate, 5-6 colpate, 9-12 colpate, 12 rugate and pantoporate. The study showed
no closed relationship between Merremia and Operculina (Ferguson et al 1977).
Austin and Staples (1980) have proposed M. tridentata (L.) Hallier to be a
new genus namely Xenostegia, due to M. tridentata has porate pollen, since all
Merremia have colpate pollen.
Distribution species of Merremia
The genus Merremia is found in the tropical regions of both hemispheres.
Merremia wildly distributed throughout the tropical regions of Asia, Africa,
Australia, North and South America (Fang and Staples 1995) and approximately
there were 80 species of Merremia in the world (Ooststroom and Hoogland 1953).
In Mexico, there are 12 species (Mc Donal 1991), Micronesia 7 species (Fosberg
& Sachet 1977), Australasia and Pasific 49 species (Staples 2010), Malesiana 23
species (Ooststroom and Hoogland 1953), China 19 species (Huang 1995), Malay
Paninsula 8 species, Malesia 23 species (Ooststroom 1939) and Java 12 species
(Backer and Bakhuizen van den Brink Jr. 1965)
Figure 1 Distribution of Merremia
(George Staples 2012)
Economic value of Merremia
Some species of the genus Merremia have been used by human kind for a
long time especially for food and medicinal purposes (roots, leaves and tubers)
and it has economic value. The tubers, roots or stems of several Merremia species
are used as a purgative. A decoction of the root M. peltata is used to treat stomach
muscular rigidity. The tubers of M. tuberosa in Java and India are known as a
drastic purgative, M. umbellata in India and M. peltata in Philippines are mildly
laxative and widely taken for dysentery. The sap from the stems of M. peltata in
Philippines mixed with lumps of sugar is a remedy for cough and in Indonesia,
Fiji and India, diluted sap from the young stems of M. peltata is used as eye or ear
drops.
The leaf of M. emarginata in Indonesia has applied as a laxative and
mixed with lumps of sugar is a remedy for cough. An infusion of the leaves of M.
4
dissecta in Africa and M. peltata in Philippines are taken as a sedative for chest
complaints, and a poultice of fresh, crushed leaves is applied as a resolutive), in
Argentina M. dissecta var. edentata used as food by local people (Austin 2007). In
Fiji, a decoction of the leaves of M. peltata is used to treat boils, infections and
appendicitis and in Florida, M. tuberosa used as laxatin (Austin 1998). The beauty
of shape and color of flowers are very attractive, may chance to make this plant as
ornamental plant.
3 MATERIALS AND METHODS
Materials
A Total of 175 sheets (164 collection numbers) which are deposited in
Herbarium Bogoriense, 97 sheets (95 collection numbers) which are deposited
Andalas University Herbarium (ANDA) and 9 sheets (4 collection numbers)
which are deposited in BIOTROP Herbarium (BIOT) were studied. It composed
of all specimens of Merremia in Sumatra. Additional specimens were collected by
the author 28 sheets (15 collection numbers) from North Sumatra, West Sumatra
and Lampung, which are stored in BO and BIOT. The digital data that consist of
type specimens and distributions of species from JSTOR Plant Science and
Convolvulaceae Unlimited were also used here.
Methods
All the hebarium specimens were studied based on their morphological
character following de Vogel (1987) and Rifai (1976). The terminology of
morphological characters followed Lawrence (1955), Veldkamp (1987), Harris
and Harris (1994), Hickey and King (2005) and Stearn (1983). Information on the
habitat, ecology, distribution, vernacular names and uses were noted from the
specimens label and literatures.
Phylogenetic Analysis.
Thirty five morphological characters selected during morphological
observation of field and herbarium specimens were used in this study (table 1).
Evolvulus nummularius and Porana volubis were selected as an outgroup.
Morphological characters were analyzed based on Maximum Parsimony using
PAUP*4.0b4 (Swofford, 2002).
4 RESULT AND DISCUSSIONS
By examined 175 sheets specimens at 3 Herbaria (BO, BIOT and ANDA),
8 species of Merremia (M. dissecta, M. emarginata, M. hirta, M. peltata, M.
quinquefolia, M. tridentata, M. tuberosa and M. vitifolia), 2 subspecies (M.
umbellata ssp. orientalis and M. tridentata ssp. hastata), 1 varietas (M. boisiana
var. sumatrana) and 2 forma (M. hederacea f. pubescens and M. hederacea f.
barbata) recognized. In addition, M. cissoides was found in North Sumatra as a
new record. M. tridentata and M. tridentata ssp. hastata are excluded because
they have been changed to be a new genus Xenostegia by Austin and Staples. So
5
that, as a result there are 8 species, 1 subspecies, 1 varietas and 2 forma of
Merremia found in Sumatra.
Delimitation of Genus and Species Diversity
Morphological Species Concept (MSC) was used in recognizing the taxa
Merremia in Sumatra. The taxa delimitation in the genus or species level of
Merremia in Sumatra based on their morphological characters. This
morphological delimitation has developed based on morphological variation of
generative and vegetative organs like leaf morphology, infloresences form,
capsule form, seed and also hairs in each part of plant. The genus Merremia can
be distinguished from other genera of Convolvulaceae by the following
characters: variation of leaves, corolla funnel-shape or campanulate which are
white or yellow, hairs on corolla, 5 nerved midpetaline bands distinctly, stigma
globular, anther straight, spiral or coil, capsule valve dehiscent regularly, seeds 24 glabrous, pubescent or pillose. Futher full description and delimitation of each
species can be seen in taxonomic treatment.
General Morphology Merremia Dennstedt ex Endlicher of Sumatra
Habit
Most species Merremia in Sumatra are annual or perennial herbaceous
vines (M. emarginata, M. hirta, M. hederacea, M. cissoides, M. umbellata) and
woody vines (M. boisiana var. Sumatrana, M. peltata, M. tuberosa and M.
vitifolia).
Stem
Stem shape Merremia is terete. They are creeping or twinning and often
low-growing on grassy or rocky places and upon trees. All of Merremia have
milky white sap and a few has fragrant (M. cissoides). The stems of Merremia
have unique character, due to they can be twin on trees up to 30 m without tendril.
They have active mechanisms to climb on and attach themselves to the host plant.
They present a circumnutational movement in which their stems, somewhat
arching in the distal portion, rotate on their own axis, rather like the hands of a
clock. This movement is essential so that the vine can locate a structure on which
it can climb on the canopy of the shrubs and young trees and thus use as a source
of support. They have long and flexible stems that depend on external support to
maintain themselve erect to reach illuminated areas in their habitat. Their stems
are characterized by the scarcity of supporting cells (fibers) and an increase in the
diameter of the xylem vessels, which may be visible to the naked eye (Herbaceous
stems). For the woody vines (M. peltata, M. vitifolia, M. boisiana var. sumatrana)
they have stems to 3-8 cm in diameter, the stems have a non-circular cross section
which oval and somewhat furrowed. Woody tissues can be isolated by more
flexible ones, the stems then become more flexible and even when it becomes
very woody and it is still elastic. This kind of wood anatomy is particularly well
developed in vine species that are subject to important growth constraints like
torsion and tears. Some species of Merremia have tuberous roots and these tubers
are formed from the main roots (M. peltata, M. tuberosa and M. cissoides). These
tubers occur in both wet and dry environments. In extremely dry conditions, stems
6
and roots can become a caudex that is one of kind of tuber indicating an
adaptation to extreme environment.
Leaves
The leaves of Merremia are alternate and have variable shapes and sizes.
Some species usually have small leaves (1-5 cm long) for example in M.
emarginata and M. hirta but the other has bigger and broader leave to 40 cm in M.
peltata. Almost the Sumatra M. have simple leaves except M. quinquefolia and M.
cissoides which have compound leaves. The blade shapes variable from ovate (M.
umbellata ssp. orientalis and M. boisiana var. sumatrana), linear (M. hirta),
reniform (M. emarginata), peltate (M. peltata), trilobe (M. hederacea), shallow
lobe (M. vitifolia), deeply lobes (M. cissoides, M. dissecta, M. tuberosa and M.
quinquefolia).
The leaf base, usually are cordate or attenuate but sometimes peltate (M.
peltata). The margin are variable from entire to crenate (M. hederacea, M. peltata,
M. tuberosa, M. umbellata ssp. orientalis and M. emarginata) and dentate (M.
vitifolia, M. dissecta, M. cissoides and M. quinquefolia). The apex also have
variable shapes from emarginate (M. emarginata), mucronulate (M. peltata, M.
umbellata ssp. orientalis and M. vitifolia), acute to acuminate (M. cissoides, M.
tuberosa) and caudate (M. boisiana var. sumatrana). The leaf surfaces are often
glabrous and pubescent on both sides but sometimes hirsute (M. vitifolia) or
sparsely pubescent to glabrous above and glabrous below (M. umbellata ssp.
orientalis). The nerves are variable from pinnate (M. boisiana var. sumatrana, M.
emarginata, M. hirta, M. peltata, and M. umbellata ssp. orientalis) and palmate
(M. cissoides, M. hedeacea, M. tuberosa, M. vitifolia and M. quinquefolia).
Figure 3 Palmately compound leaves (M. cissoides and M. quinquefolia)
7
a
c
b
e
f
d
g
Figure 2 The variation blade of simple leaves a. linear (M. hirta); b. ovate (M.
umbellata ssp. orientalis and M. boisiana var. sumatrana); c. reniforme
(M. emarginata); d. peltate (M. peltata); e. trilobe (M. hederacea), f.
shallow lobe (M. vitifolia); g. deeply lobes (M. dissecta and M.
tuberosa).
Infloresences
The inflorescences are axillary with few to many flowered and usually
cymose. The ramified inflorescences are variable, usually cyme but sometime
panicle, (M. boisiana var. sumatrana), cincinnus (M. vitifolia) and umbel (M.
umbellata ssp. orientalis). The peduncle thick or thin, sub terete. They have
variable peduncle sizes but sometime M. emarginata has 1 mm or nearly absent.
The peduncle surface is usually glabrous or pubescent, but sometime hirsute (M.
vitifolia and M. cissoides). The bracts of Merremia are short, occasionally
caducous and have variable shapes that is ovate or triangularis or ovoid.
Flower buds
The important character in Merremia is the spread of the hairs on corolla
that can be easily observed at the mature flower bud. The flower bud shapes
variable from ovoid (M. quinquefolia, M. dissecta, M. hirta, M. peltata, M.
umbellata ssp. orientalis and M. vitifolia), ovoid to conical (M. tuberosa),
subglobose (M. boisiana var. sumatrana) and subrhombhoid (M. cissoides).
8
a
b
Figure 4
c
d
The variation ramified inflorescence a. Cincinus (M. vitifolia); b.
cyme (M. hirta, M. emarginata, M. peltata, M. hederacea, M. dissecta
and M. tuberosa); d. umbel (M. umbellata ssp. orientalis); e. panicle
(M. boisiana var. sumatrana)
Flower
Generally, Flower is arranged in a stalk head and form a flowering head
with the calyx, corolla tubes which are joined together. The flowers are
pentamerous, actinomorphic and bisexual. The species have variable flower sizes.
Some species usually have small (1-2 cm long) at M. hederacea, M. emarginata,
and M. cissoides, slightly big (2-3 cm long) at M. hirta, M. quinquefolia, M.
boisiana, M. umbellata ssp. orientalis and M. vitifolia) and big (>3 cm long) at M.
dissecta, M. peltata and M. tuberosa).
Calyx – The calyx have five free sepals, overlapping in a quincuncial
arrangement with outer and inner sepal unequal or almost equal length, concave,
persistent, usually have glabrous or pubescent surfaces but sometime have hirsute
(M. cissoides and M. vitifolia) and enlarged in fruit. The sepal is an important
morphological character to distinguish between species. The outer and inner
sepals have variable sizes and shapes. Sometimes outer more long than inner sepal
or reverse. The sepal shapes are orbicular to transverse elliptic (M. boisiana var.
sumatrana), elliptic (M. hirta), rhomboid (M. cissoides), oval lanceolate (M.
dissecta), obovate (M. emarginata, M. hederacea, and M. umbellata ssp.
orientalis), ovate (M. peltata, M. tuberosa, M. vitifolia and M. quinquefolia).
Corolla - The corolla is gamopetalous or petals are fused into a tube. They
have 5 bands in the middle of each petal. These bands are clearly seen in the
opened flower and are generally known as the mid-petaline bands and it which
help the corolla to coil on self when fading. They usually have glabrous
midpetaline bands but sometimes pillose (M. boisiana var. sumatrana). The
corolla limbs are slightly 5-lobe or sometimes 10-lobe (M. boisiana var
sumatrana) with pubescent surface.
9
a
Figure 5
b
The variation corollas a. campanulate (M. boissiana var. sumatrana,
M. hederacea, M. emarginata, M. peltata, and M. hirta); b. Funnelshaped (M. cissoides, M. dissecta, M. umbellata ssp. orientalis, M.
tuberosa, M. vitifolia, and M. quinquefolia).
There are 2 corolla shapes, that is campanulate (Merremia boissiana var.
sumatrana, M. hederacea, M. emarginata, M. peltata and M. hirta) and funnelshaped (M. cissoides, M. umbellata ssp. orientalis, M. tuberosa, M. vitifolia, and
M. quinquefolia). The corolla colours variable such as yellow or bright yellow (M.
dissecta, M. peltata, M. tuberosa, M. hederacea, M. vitifolia and M. hirta),
whitish yellow or orange (M. umbellata ssp. orientalis) and white (M. cissoides
and M. boisiana var. sumatrana.
Stamen – The number of stamens are 5. Usually, they are inserted in the
corolla. The filament is sticky at tube, slender, thin but broader and hairy at the
basal and dorsifixed below. The anther shape various from straight (M.
emarginata, M. hederacea, and M. umbellata ssp. orientalis), spiral twisted (M.
cissoides, M. dissecta, M. hirta, M. peltata and M. quinquefolia), and coil (M.
boisiana var. sumatrana, M. tuberosa and M. vitifolia). The indument of anther
usually glabrous but sometimes hairy (M. peltata)
a
Figure 6
b
c
The variation shapes anther a. straight (M. emarginata, M.
hederacea, and M. umbellata ssp. orientalis); b. coil (M. boisiana
var. sumatrana, M. tuberosa and M. vitifolia); c. spiral twisted (M.
cissoides, M. dissecta, M. hirta, M. peltata and M. quinquefolia.
10
Pistill – The stigma of all species are always macrostylous, biglobular
with smooth and glabrous and the colour of stigma is white. The style shape is
terete and has glabrous surface. The ovaries of all species are always superior and
glabrous.
Fruit
The fruit is typically irregularly dehiscing capsule with 4 cells and 4
valves. The capsule shapes are variable from sub globose (M. dissecta, M.
tuberosa, M. emarginata and M. vitifolia), ovoid (M. hirta and M. hederacea,
ovoid to conical (M. peltata, M. dissecta and M. umbellata ssp. orientalis). The
stalks are subterete and then thicked towards to the top.
a
Figure 7
b
The variation capsules a. ovoid to conical (M. peltata, M. dissecta
and M. umbellata ssp. orientalis); b. sub globose (M. dissecta, M.
tuberosa, M. emarginata and M. vitifolia).
Seed
The seed shapes variable from late elliptic (M. emarginata and M. hirta),
ovoid to conical (M. hederacea f. pubescent, M. peltata, M. tuberosa and M.
dissecta), transverse ovate (M. hederacea f. barbata) and obovate (M. cissoides
and M. vitifolia). Merremia have various seed sizes. Some species have small
sizes (M. hederacea, M. emarginata, M. cissoides, M. quinquefolia), slightly big
sizes (M. umbellata ssp. orientalis, M. vitifolia, M. quinquefolia and M. peltata)
and big sizes (M. tuberosa and M. dissecta). The seed surfaces are variable from
glabrous (M. vitifolia, M. dissecta), pubescent (M. boisiana var. sumatrana, M.
cissoides), bearded (M. hederaceae f. barbata), pillose (M. umbellata ssp.
orientalis and M. peltata), pubescent at margin and hylum (M. hederacea f.
pubescens and M. tuberosa), pillose at margin and hylum (M. hirta). The colours
of seed are brown, dark brown and black.
Distribution Merremia in Sumatra
The distribution of Merremia in Sumatra (Figure 8) covered Nangro Aceh
Darussalam, North Sumatra, West Sumatra, Riau and Riau Archipelago, Jambi,
Bengkulu, South Sumatra, Bangka Belitung and Lampung. The highest
concentration of species is found in North Sumatra 10 species (M. hederacea f.
11
pubescent, M. peltata, M. umbellata ssp. orientalis, M. tuberosa, M. vitifolia and
M. boissiana var. sumatrana, M. dissecta, M. emarginata, M. hirta and M.
cissoides) and then followed by West Sumatra 7 species (M. boisiana var.
sumatrana, M. hederacea f. barbata, M. hirta, M. peltata, M. umbellata ssp.
orientalis, M. tuberosa and M. vitifolia), Lampung 5 species (M. peltata, M.
umbellata ssp. orientalis, M. vitifolia, M. hirta and M. quinquefolia), 3 species
South Sumatra (M. hederacea f. barbata, M. umbellata ssp. orientalis, and M.
vitifolia) and NAD (M. vitifolia, M. peltata and M. umbellata ssp. orientalis), 2
species in Bangka Belitung (M. umbellata ssp. orientalis and M hirta), Bengkulu
(M. umbellata ssp. orientalis and M. peltata), and Jambi (M. hederacea f.
pubescens, M. umbellata ssp. orientalis), 1 species in Riau Archipelago (M.
umbellata ssp. orientalis) and Riau (M. biosiana var. sumatrana).
Figure 8 Distribution of Merremia in Sumatra.
M. umbellata ssp. orientalis found widespread in all of Sumatra province
except Riau from 20 mdpl until 1000 mdpl. This plant is native in China and
naturalized in Sumatra. M. peltata was found in NAD, North Sumatra west
Sumatra, Bengkulu and Lampung at 20-600 mdpl. This plant is origin of pacific
12
region and has been invaded at Bukit Barisan Selatan National Park (Lampung).
M. boisiana var. Sumatra is endemic in Sumatra since they are found in this island.
This plant firstly found by Lorzing at 1916 in Sibolangit (North Sumatra) at 500600 mdpl which deposited in Herbarium Bogoriense. After a few time, this plant
widespread to Mountain Leuser (North Sumatra) at 50-100 mdpl, Kampar (Riau) at
400-550 mdpl and West Sumatra at 50-780 mdpl which deposited in Andalas
Herbarium. M. emarginata was new record in Sumatra while M. cissoides was new
record in Indonesia. The occurrence of M. emarginata in Sumatra was proven by a
specimen collected from North Sumatra (J.A Lorzing, 15891) at 15-16 mdpl. This
plant is native in Indonesia and widespread in tropical Africa and tropical Asia. M.
cissoides found Hafni Rahmadani (035) at 30 mdpl in Perbaungan (North
Sumatra). This species is native and weedy in tropical America. M. dissecta, M.
quinquefolia and M. tuberosa are cultivated in Sumatra (Table 1)
Table 1 Distribution of Merremia in Sumatra
NO
Species
NAD NS
1
M. boisiana var.
+
Sumatrana
2
M. cissoides
+
3
M. dissecta
+
4
M. emarginata
+
5
M. hederacea
+
f. pubescens
6
M. hederacea
f. barbata
7
M. hirta
+
8
M. peltata
+
+
9
M. quinquefolia
10
M. tuberosa
+
11
M. umbellata
+
+
ssp. orientalis
12
M. vitifolia
+
+
Total
3
10
NAD : Nangro Aceh Darussalam.
NS
: North Sumatra.
WS
: West Sumatra.
R
: Riau
RA
: Riau Archipelago.
JA
-
SS
-
BB
-
BE
LA
-
-
+
-
-
-
-
-
-
+
-
-
-
+
-
-
+
+
+
+
-
+
+
+
+
+
+
+
+
+
+
7
1
1
2
+
5
WS
+
R
+
RA
-
-
JA
SS
BB
BE
LA
-
+
+
+
2
3
2
: Jambi.
: South Sumatra
: Bangka Belitung
: Bengkulu
: Lampung
Phylogenetic Analysis
A phylogenetic analysis using PAUP vers.4.0 was undertaken based on
morphological characters, using E. nummularius and P. volubis as the out group.
The phylogenetic analysis used 35 morphological characters (Table 2) and data
matrix of those characters which were used for the analysis showed in appendix 1.
13
Table 2 Morphological characters states used in the phylogenetic analysis
No
1
2
3
4
5
6
7
8
9
10
11
12
Characters
Habit
Leaf attacment
Leaf
Leaf Notch
Venation major
leaves
Blade shape
Leaf surface
Leaf margin
Leaf base
Leaf apex
Petiole
Infloresece
0.
0.
0.
0.
0.
woody
distichous
simple
absent
actinodromous
0. ovate
0. hairy
0. entire to crenate
0. cordate
0. mucronulate
0. < 0.5 cm
0. terminal and
axillary
0. present
0. cyme
15
16
Peduncle
Inflorescences
ramified
Pedicel
Outer sepal
17
18
Flower
Corolla lobed
0. 0.5 cm
1. axillary
1. absent
1.panicle
1. > 0.5
1.shorter than
inner one
1. 1/ leaf axil
1. not depply
lobed
1. funnel
1. hairy
1. yellow
1. biglobular
1. < 2
1. > 1
1. < 2
0. absent
1. inserted
1. equal
1. not straight
1. present
1. hairy
1. globose
1. >1
1.long hairy
1. margin seed
2. cincinnus
2. rotate
2. capitates
2. glabrous
3.linear
3. umbel
14
Morphological characters were analyzed based on Maximum Parsimony
using PAUP*4.0b10 (Swofford 2002) with Heuristic Search settings vegetative
and floral characters were selected for the analysis of which 27 were scored as
binary and 8 as multistate. Some characters were scored as missing data when the
data were unavailable. All characters are treated as unorder data and have an 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) and was evaluated using 100 bootstrap replicates and a complete Hsearch.
Clade support values were obtained by using bootstrap.
Maximum Parsimony (MP) analysis of the morphology dataset produced
17 shortest trees of 68 steps with consistency index (CI) = 0.60, homoplasy index
(HI) = 0.40, retention index (RI) = 0.56 and Rescaled consistency index (RC) =
0.34. Of 35 characters used in this analysis, 21 characters (60%) were potentially
parsimony-informative, and 14 characters (40%) were parsimony-uninformative.
Evolvulus nummularius
Porana volubilis
M. boisiana var. sumatrana
M. cissoides
M. quinquefolia
M. tuberosa
M.dissecta
M. vitifolia
M. emarginata
M. hederacea barbata
M. hederacea pubescens
M. hirta
M. peltata
M. umbellata ssp. orientalis
Figure 9 Strict consensus tree resulted morphological characters. CI = 0.60, HI =
0.4, RI = 0.56, RC = 0.34. The number above the branches showed the
bootsrap value
15
The cladogram devided the taxa into two groups. The separation support
M. boisiana var. sumatrana, M. cissoides, M. dissecta, M. emarginata, M.
hederacea f. pubescens, M. hederacea f. barbata, M. hirta, M. peltata, M.
quinquefolia, M. tuberosa, M. vitifolia and M. umbellata ssp. orientalis as ingroup
with E. nummularius and P. volubis as outgroup as strengthened (BS=77%)
(figure 9). It was separated base on some character such as leaf attachment
(character 2), petiole (character 11), infloreseces (character 12), pedicel (character
13), flower (character 17), corolla lobed (character 18), corolla surface (character
20), corolla colour (character 21), ovary (character 24), style number (character
25), style branch (character 26), stamen (character 27), stamen length (character
28), filament (character 31) and seed number (character 33).
The topology of strict consensus tree resulting from morphological and
strongly support the monophyletic, paraphyletic species and metaspecies of
Merremia. M. boisiana var. sumatrana is metaphyletic (unresolved) and it is a
sister group of monophyletic M. cissoides, M. dissecta, M. hederacea f.
pubescens, M. hederacea f. barbata, M. quinquefolia, M. tuberosa and M.
vitifolia. M. boisiana var. sumatrana is paraphyletic species with M. umbellata
ssp. oriental, M. hirta, and M. peltata which shared by two populations of the
metaspecies and monophyletic. While M. emarginata is metaspecies (unresolved).
The morphological phylogenetic analysis for monophyletic showed that M.
cissoides closed relationship with M. quinquefolia. Both species were placed in
the same clade with strong support 79%. While, M. tuberosa become as the basal
sister of M. cissoides and M. quinquefolia with weakened 61%. Phenetically, M.
tuberosa separated from M. cissoides and M. quinquefolia based on some
character such as leaf (character 3) venation major leaves (character 5) and margin
leaf (character 8). M. vitifolia and M. dissecta are closed relationship with
weakened 58 % but separated based on Ramified (character 14) and outer sepal
(character 16). M. hederacea f. barbata closed relationship with M. hederacea f.
pubescens strong support 51% and separated base on seed surface (character 34)
and position of seed surface (character 35. Both of each species were placed in the
same clade.
16
Generic Description
Merremia Dennstedt ex Endlicher
Merremia Dennst., Schlüss., Hort. Malab. 1818, 34, (nomen nudum); Hall. f. in
Engl., Bot. Jahrb. XVI (1893) 581; Baker & Rendle in This.-Dyes, Fl. Trop. Afr.
IV, 2 (1905) 101; Ridl., Fl. Mal. Penin. II (1923) 456; Merrill, Enum. Philipp. Fl.
Pl. III (1923) 360; Ooststr., Blumea III (1939) 292; Ooststr., FM I.4.4 (1953) 450;
Backer and Bakhuizen f., FJ. II (1965) 488. Type species: Merremia
convolvulacea (= M. hederacea (Burm. f.) Hall f.) (illustration!)
.─ Skinneria Choisy, in Mem. Soc. Phys. Genève VI (1833) 487. Type species:
not seen.
.─ Spiranthera Boj., Hort. Maurit. (1837) 226; Griseb., Fl. Brit. West Ind. Isl.
(1864) 470. Type species: not seen.
Annual or perennial herbs or woody twinning and creeping at trees or
stones. Stems terete, prostate, sometimes stems come out from tuber, glabrous or
hairy, with milky juice. Leaves simple or palmately compound, alternate: petiole
terete, glabrous or pubescent or glabrescent or hirsute; blade variable shape and
size, ovate or orbicular or emarginate in outline, base cordate or peltate or
attenuate or truncate or rounded, margin entire or crenate or dentate, apex
mucronulate or acute or acuminate or emarginated or caudate, primer vine pinnate
or palmate. Inflorescence axillary, solitary or in few to many flower, cymose,
ramified variable, compound cyme or cincinnus or umbel or panicle; peduncle
terete, glabrous or pubescent or hirsute. Bract small and caducous, ovate or
triangularis or ovoid. Flower buds ovoid or ovoid to conical or subglobose or
rhombhoid. Flower bisexual: stalk terete, glabrous or pubescent or hirsute sepals 5
concave, outer and inner sepal unequal or almost equal length, orbicular to
transverse elliptic or elliptic or subrhomboid or oval lanceolate or ovate, glabrous
or pubescent or hirsute, in several species enlarged in fruit; corolla funnel or
campanulate, yellow or white or whitish yellow or bright yellow or orange or
white deep purple, corolla limb slightly 5-lobed or 10-lobed, pubescent,
midpetaline bands 5, glabrous or pillose; stamens 5, anther variable shape, straight
or spiral twisted or coil, glabrous or pillose, filament unequal, dorsifixed below,
sticky at tube, slender, thin but broader and hairy at the basal pistill 1, stigma
macrostylous, biglobular, smooth and glabrous, white; styles terete, glabrous
ovary superior, glabrous. Fruit a capsule, valve, dehiscent, globose or ovoid or
ovoid to conical, glabrous. Seeds 2-4, variable shape and size, ovate or elliptic or
ovoid to conical or transverse ovate or obovate, glabrous or pubescent or sericious
around or at margin and hylum, brown or dark brown or black.
17
Key to the Sumatran Species of Merremia
1.
2.
3.
4.
5.
6.
7.
8.
9.
Leaves simple…………………………………………………….…........2
Leaves palmately compound with 5 leaflets………………………….….3
Blade palmate with 3-7 segment shallow or deeply lobes…….….....…....4
Blade not palmate, peltate, ovate, linear, reniform, ……………...……….7
Leaflets elliptic, apex mucronulate, margin pubescent; bract linear; sepal
subrhomboid, hirsute; capsule globose; seed 2 mm long;
brown………………………………………………………..2. M. cissoides
b. Leaflets narrowly oblong, apex acute, margin glabrous; bract narrowly
triangular; sepal ovate to oblong, glabrous; capsule ovoid to conical; seed
4 mm long, black………………………………..…..….8. M. quinquefolia
a. Blade deeply lobed into 7 segments, inflorescense ramified cyme, corolla
funnel shaped…………………………………………………………...….5
b. Blade shallow lobed with 3-7 segments, inflorescense ramified cyme or
cincinus, corolla funnel shaped or campanulate……………………….....6
a. Petiole hirsute; blade lanceolate-elliptic, glabrous, one upper and two
lateral blade segment larger than two lateral and two bottom ones, margin
dentate, apex mucronulate; sepal ovoid; anther spiral twisted; capsule
globose; seeds 7-7.5 mm long, glabrous, black……………...3. M. dissecta
b. Petiole pubescent; blade oblong-lanceolate, pubescent, one upper blade
segment larger than lateral and bottom one, margin entire, apex acute to
acuminate; sepal ovate; anther coil; capsule globose; seed 13-14 mm long,
pu
(CONVOLVULACEAE)
IN SUMATRA
HAFNI RAHMADANI
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2013
LETTER OF STATEMENT
I express that this thesis entitled:
MERREMIA DENNSTEDT EX ENDLICHER (CONVOLVULACEAE) IN
SUMATRA
Has been compiled and written by myself and it is true represent result of my own
research and has never been previously submitted in whole or part for another degree. All
information and data that used have been expressed clearly and can be checked its truth.
Bogor, October 2013
Hafni Rahmadani
NRP. G353100131
SUMMARY
HAFNI RAHMADANI. Merremia Dennstedt ex Endlicher (Convolvulaceae) in
Sumatra. Supervised by Dr. SRI SUDARMIYATI TJITROSODIRDJO and Dr. HARRY
WIRIADINATA
Merremia is a small genus in the family Convolvulaceae. Merremia is a
perennial or annual herb or woody liana with stems creeping or twinning on trees
or rocks, put out milky white sap when injured and some species with tubers.
Merremia is often confused with genus Ipomoea, due to they have resemblance in
funnel shaped or campanulate corolla and straight anther. Phenetically, some
members of Merremia have yellow flower whereas Ipomoea has white, purple or
purplish white. Merremia is also confused with Operculina due both have nonspinulose pollen and dehiscent capsule. Nevertheless, Operculina characterized by
circumciselli capsule and Merremia characterized by valve capsule. The important
character in Merremia is the spread of the hairs on corolla that can be easily
observed at the mature flower bud. The genus Merremia is found in the tropical
regions of both hemispheres. Merremia wildly distributed throughout the tropical
regions of Asia, Africa, Australia, North and South America and approximately
there were 80 species of Merremia in the world.
A Total of 175 sheets (164 collection numbers) which are deposited in
Herbarium Bogoriense, 97 sheets (95 collection numbers) which are deposited
Andalas University Herbarium (ANDA) and 9 sheets (4 collection numbers)
which are deposited in BIOTROP Herbarium (BIOT) were studied. It composed
of all specimens of Merremia in Sumatra. Additional specimens were collected by
the author 28 sheets (15 collection numbers) from North Sumatra, West Sumatra
and Lampung, which are stored in BO and BIOT. The digital data that consist of
type specimens and distributions of species from JSTOR Plant Science and
Convolvulaceae Unlimited were also used here.
All the hebarium specimens were studied based on their morphological
characters following de Vogel (1987) and Rifai (1976). The terminology of
morphological characters followed Lawrence (1955), Veldkamp (1987), Harris
and Harris (1994), Hickey and King (2005) and Stearn (1983). Information on the
habitat, ecology, distribution, vernacular names and uses were noted from the
specimens label and literatures.
Taxonomic studies in Sumatra were conducted. There are eight species,
one subspecies, one varietas and two forma from nine previously species (M.
cissoides, M. dissecta, M. emarginata, M. hirta, M. peltata, M. quinquefolia, M.
tridentata, M. tuberosa and M. vitifolia), two subspecies (M. umbellata ssp.
orientalis and M. tridentata ssp. hastata), one varietas (M. boisiana var.
sumatrana) and two forma (M. hederacea f. pubescens and M. hederacea f.
barbata) can be recognized. One species (M. tridentata) and subspecies (M.
tridentata ssp. hastata) are excluded because they have been proposed to different
new taxa Xenostegia by Austin and Staples. The existence of M. emarginata was
noted as new record in Sumatra and M. cissoides was noted as new record in
Indonesia.
A phylogenetic analysis using PAUP*4.Ob10 was undertaken based on
morphological characters, using Evolvulus nummularius and Porana volubis as
the out group. The phylogenetic analysis used 35 morphological characters was
resulted consistency index (CI) = 0.60, homoplasy index (HI) = 0.40, retention
index (RI) = 0.56 and Rescaled consistency index (RC) = 0.34.
Phylogenetic tree used morphological characters divided taxa into two
groups. The separation support M. boisiana var. sumatrana, M. cissoides, M.
dissecta, M. emarginata, M. hederacea f. pubescens, M. hederacea f. barbata, M.
hirta, M. peltata, M. quinquefolia, M. tuberosa, M. vitifolia and M. umbellata ssp.
orientalis as ingroup with E. nummularius and P. volubis as outgroup. This study
support the monophyly, paraphyly and metaphyly of genus Merremia in Sumatra.
Key words: Convolvulaceae, Merremia, morphology, phylogenetic, Sumatra
RINGKASAN
HAFNI RAHMADANI. Merremia Dennstedt ex Endlicher (Convolvulaceae) di
Sumatera. Dibimbing oleh Dr. SRI SUDARMIYATI TJITROSODIRDJO dan Dr.
HARRY WIRIADINATA
Merremia merupakan marga dari Convolvulaceae yang mempunyai ciri-ciri
merupakan herba atau semak dengan batang memanjang, menjalar atau merambat
sampai 20 m, membelit pada pucuknya, beberapa jenis dengan perakaran yang
mempunyai umbi, mengeluarkan cairan putih seperti susu ketika dilukai (Stone
1970). Marga Merremia sering rancu dengan marga Ipomoea yang mempunyai
kemiripan pada mahkota bunga yang berbentuk funnel atau campanulate dan
benang sari yang berbentuk straight. Bunga Merremia biasanya berwarna kuning
sedangkan Ipomoea berwarna putih, ungu ataupun putih keunguan. Marga ini juga
rancu dengan Marga Operculina, yang sama-sama memiliki polen non-spinulose
dan buah dengan kapsul yang secara teratur akan pecah. Akan tetapi pada
Operculina kapsul berbentuk circumcisseli dan pada Merremia berbentuk valve.
Penyebaran bulu-bulu yang terdapat di mahkota bunga adalah ciri yang penting
dalam Merremia dan dapat dilihat dengan mudah pada kuncup bunga yang telah
dewasa. Di dunia, Marga Merremia diperkirakan terdapat 80 Jenis dan menyebar
luas di kawasan tropis mulai dari Afrika, Asia, Australia, Amerika Utara dan
Amerika Selatan.
Penelitian ini berdasarkan spesimen herbarium Merremia yang dikoleksi
dari Sumatera. Sebanyak 175 spesimen yang tersimpan di Herbarium Bogoriense
(BO), 9 spesimen di Herbarium Biotrop (Biot), 97 spesimen di Herbarium
Universitas Andalas (ANDA) dan 28 Spesimen dari hasil eksplorasi di Sumatera
Utara, Sumatera Barat dan Lampung. Data digital yang terdiri dari spesimen tipe
dan penyebaran spesies dari JSTOR Plant Science dan Convolvulaceae Unlimited juga
digunakan.
Semua spesimen herbarium dilihat berdasarkan karakter-karakter
morfologinya yang mengacu pada metode Rifai (1976) dan Vogel (1987).
Terminologi karakter morfologi mengacu pada Lawrence (1955), Veldkamp
(1987), Harris dan Harris (1994), Hickey dan King (2005) dan Stearn (1983).
Informasi habitat, ekologi, distribusi, nama daerah dan lain sebagainya di catat
berdasarkan label dan literatur spesimen.
Kajian taksonomi mengenai marga Merremia Dennstedt ex Endlicher
(Convolvulaceae) di Sumatera yang berdasarkan pada karakter morfologi telah
dilakukan. Dari studi ini terdapat 8 jenis Merremia, 1 anak jenis, 1 varitas dan 2
forma dari 9 jenis sebelumnya (M. cissoides, M. dissecta, M. emarginata, M.
hirta, M. peltata, M. quinquefolia, M. tridentata, M. tuberosa dan M. vitifolia), 2
anak jenis (M. umbellata ssp. orientalis dan M. tridentata ssp. hastata), 1 varitas
(M. boisiana var. sumatrana) dan 2 forma (M. hederacea f. pubescens dan M.
hederacea f. barbata) telah diperoleh 1 jenis (M. tridentata) dan anak jenis (M.
tridentata ssp. hastata) dikeluarkan karena telah diajukan menjadi takson baru
yaitu Xenostegia oleh Austin dan Staples. M. emarginata merupakan catatan baru
di Sumatera dan M. cissoides merupakan catatan baru di Indonesia.
Analisa hubungan kekerabatan menggunakan Program PAUP*4.Ob10
dengan E. nummularifolius dan P. volubis sebagai outgroup. Analisis
phylogenetic menggunakan 35 karakter morfologi menghasilkan consistensi index
(CI)=0.60, homoplasi index (HI)=0.40, retention index (RI)=0.56 dan rescaled
consistency index (RC)=0.34.
Pohon filogenetik berdasarkan karakter morfologi membagi takson dalam
dua kelompok. Pemisahan itu mendukung M. boisiana var. sumatrana, M.
cissoides, M. dissecta, M. emarginata, M. hederacea f. pubescens, M. hederacea
f. barbata, M. hirta, M. peltata, M. quinquefolia, M. tuberosa, M. vitifolia and M.
umbellata ssp. orientalis sebagai ingroup dengan E. nummularius dan P. volubis
sebagai outgroup. Studi marga Merremia ini memperlihatkan terjadinya
monophyly, paraphyly dan metaphyly.
Kata kunci: Convolvulaceae, Merremia, morfologi, phylogenetic, Sumatera
Copyright©2013, Bogor Agricultural University
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MERREMIA DENNSTEDT EX ENDLICHER
(CONVOLVULACEAE)
IN SUMATRA
HAFNI RAHMADANI
Thesis submitted
As partial fulfillment requirement for the Master Degree
In Plant Taxonomy
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
2013
External examiner: Dr.Rugayah, M.Si.
Title
Name
NRP
: Merremia Dennstedt ex Endlicher (Convolvulaceae) in Sumatra
: Hafni Rahmadani
: G353100131
Certified by
Supervisor Committee
Dr. Sri Sudarmiyati Tjitrosoedirdjo. M.Sc
Chairman
Dr. Harry Wiriadinata
Member
Approved by
Head of Plant Biology
Study Program
Dr. Ir Mifiahudin, M.si
Date of Examination: 30 July 2013
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Graduation:
o3
OCT 2013
Title
Name
NRP
: Merremia Dennstedt ex Endlicher (Convolvulaceae) in Sumatra
: Hafni Rahmadani
: G353100131
Certified by
Supervisor Committee
Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc
Chairman
Dr. Harry Wiriadinata
Member
Approved by
Head of Plant Biology
Study Program
Dean of Graduate School
Dr. Ir Miftahudin, M.Si
Dr. Ir. Dahrul Syah, M.Sc. Agr.
Date of Examination: 30 July 2013
Date of Graduation:
ACKNOWLEDGMENT
Thanks to Allah that I have finished my thesis. I would like to express my
gratitude to my supervisors, Dr. Sri Sudarmiyati Tjitrosoedirdjo and Dr. Harry
Wiriadinata for their valuable advices, guidance, and encouragement throughtout
my study. I gratefully acknowledge the DIPA Project 2011 through Dr. Sri
Sudarmiyati Tjitrosoedirdjo from Southeast Asean Regional Centre of Tropical
Biologi (SEAMEO BIOTROP).
I would like to express my gratitude to Herbarium Bogoriense (LIPI),
Herbarium Andalas University (ANDA) and Herbarium BIOTROP (BIOT), Bukit
Barisan Selatan National Park for granting me permission to conduct this research
and providing me some facilities. In particularly I would like to thanks all
taxonomist and technicians who gave their support and assisted me when
conducted this study.
I am also very grateful to Dr. George Staples (Singapore Botanical
Garden), Dr. Rugayah and Lulut Dwi Sulistyaningsih M.Si (BO), Budi Setiabudi
S.Hut (SEAMEO BIOTROP), for their interest, discussion and valuable comment
on the manuscript. Many thanks to my friends Azwar Haris Siregar, Indah
Wahyuni, Rahmanida and Alfiah Rahmi.
Last but not the least, my special thanks to my family, my father H. Edi
Syahputra, my mother Hj. Faridawati Siregar, and my brother Riza Afriandi for
their spirit, patience, and supporting of my study.
Bogor,
October 2013
Hafni Rahmadani
TABLE OF CONTENTS
LIST OF TABLES
vi
LIST OF FIGURES
vi
LIST OF APPENDIX
vi
1
INTRODUCTION
1
2
LITERATURE REVIEW
Taksonomy of Merremia Dennst. ex Endlicher
Distribution species of Merremia
Economic value of Merremia
2
3
4
MATERIALS AND METHODS5
Materials
Methods
Phylogenetic analysis.
4
4
4
3
4
RESULT AND DISCUSSIONS
Delimitation of Genus and Species Diversity
General Morpholgy Merremia Dennstedt ex Endlicher of Sumatra
Habit
Stem
Leaves
Inflorescences
Flower buds
Flower
Calyx
Corolla
Stamen
Pistill
Fruit
Seed
Distribution of Merremia in Sumatra
Phylogenetic analysis
Generic description
Key to the Sumatran species of Merremia
Taxonomic treatment
Merremia boisiana var. sumatrana
Merremia cissoides
Merremia dissecta
Merremia emarginata
Merremia hederacea
Merremia hirta
Merremia peltata
5
5
5
5
6
7
7
8
8
8
9
10
10
10
10
12
17
18
21
23
24
27
29
30
Merremia quiquefolia
Merremia tuberosa
Merremia umbellata
Merremia vitifolia
5
33
34
36
38
CONCLUSION
41
REFERENCES
41
APPENDIX
44
CURRICULUM VITAE
45
LIST OF TABLES
1. Distribution of Merremia in Sumatra
2. Morphological characters state used in phylogenetic analysis
12
13
LIST OF FIGURES
1. Distribution of Merremia
2. Palmately compound leaves
3. Variation blade of simple leaves
4. Variation ramified inflorecences
5. Variation corollas
6. Variation shapes anther
7. Variation capsules
8. Distribution of Merremia in Sumatra
9. Strict consensus tree resulted from morphological characters
10. Distribution of Merremia var. sumatrana in Sumatra
11. Merremia boisiana var. sumatrana
12. Merremia cissoides
13. Distribution of Merremia cissoides in Sumatra
14. Distribution of Merremia dissecta in Sumatra
15. Distribution of Merremia emarginata in Sumatra
16. Merremia emarginata
17. Distribution of Merremia hederacea in Sumatra
18. Distribution of Merremia hirta in Sumatra
19. Merremia peltata
20. Distribution of Merremia peltata in Sumatra
21. Distribution of Merremia quinquefolia in Sumatra
22. Distribution of Merremia tuberosa in Sumatra
23. Distribution of Merremia umbellata ssp. orientalis in Sumatra
24. Distribution of Merremia vitifolia in Sumatra
25. Merremia vitifolia
3
6
7
8
9
9
10
11
14
19
20
22
23
24
25
26
28
30
32
33
34
35
37
39
40
LIST OF APPENDIX
1. Matrix of morphological characters for phylogenetic analysis
44
1 INTRODUCTION
Invasive plant is a plant that can successfully adapt in its new habitat and
spread very widely, which would then occupy and replace the native plants and
finally would threaten the diversity, structure and function of ecosystem
(Radosevich et al. 2007). The invasion begins of migration, aggregation and then
competition. The characteristics of invasive plants are able to grow quickly,
reproduce vegetatively, high spread, tolerance to environmental condition, have
not natural enemies and usually associated with humans.
The twinning or creeping species with fast growth rates can become
troublesome weeds of agriculture and forestry. Several Merremia species have
become serious weeds of forestry plantations in Sumatra. Merremia peltata is a
big problem in Bukit Barisan Selatan National Park, due the National park is
habitat of protected animal such as Tiger, Elephant and Sumatran Rhino. This
National park has area with 356.800 ha (BBTNBBS 2009) and the invasion of
Merremia peltata has reached more than 7.008 ha even has caused animals life
were interrupted, so the animals migrate to more northern areas, which are the
village, estate and inflict the conflict with local people (Irianto and Tjitrosoedirdjo
2010).
Merremia is a small genus belonging to the family Convolvulaceae which
perennial or annual herb or woody with stems creeping or twinning on trees or
rocks, put out milky white sap when injured and some species with tubers (Stone
1970). Merremia is often confused with genus Ipomoea, due to a resemblance in
funnel shaped or campanulate corolla and straight anther. Phenetically, some
members of Merremia have yellow flowers whereas Ipomoea has white, purple or
purplish white. Merremia is also confused with Operculina because both have
non-spinulose pollen and dehiscent capsule. Nevertheless, Operculina is
characterized by circumciselli capsule, while Merremia is characterized by valve
capsule (Staples 2010). The important character in Merremia is the spread of the
hairs on corolla that can be easily observed at the mature flower bud (Fang and
Staples 1995).
The genus Merremia was recognized by Hallier (1893). He separated it
from Ipomoea almost entirely on the basis of its non-spinulose pollen grains and
five distict mid-petaline bands of corolla. He recognized several sections of
Merremia, namely Skinneria, Xanthips, Streptandra (Hallier 1894) and also
adding one more section that is Hailale ( Hallier 1913). Van Ooststroom and
Hoogland (1953) replaced the name Skinneria by Eu-Merremia and established
Wavula as the fifth section.
Taxonomic research of Merremia has been done by Ooststroom and
Hoogland (1953) who recorded 23 species Merremia in Malesian region and
Backer and Backhuizen Jr. (1965) recorded 12 species occur in Java.
Unfortunately taxonomic studies of Merremia in Sumatra is lacking since there
are inconsistencies in number of species proposed for inclusion in Merremia of
Sumatra e.g. Ooststroom and Hoogland (1953) mentioned there were 3 species
Merremia and on the other hand Staples (2010) mentioned 5 species.
The relationship between species of Merremia has been studied but there
were still difference of opinion. Phylogenetic studies based on molecular data of
Convolvulaceae (Stefanovic 2002) with 6 species of Merremia (M. aegyptia, M.
2
dissecta, M. hastata, M. peltata, M. umbellata and M. vitifolia) have indicated that
tribe is not monopohyletic, and that its largest genus Merremia is polyphyletic.
Relationship studies of Convolvulacae based on chemical markers reported that
Merremia containing flavonoids, quinonones, saponins, phenolic acids, and
alkaloids. Based of the distribution flavonoids, indicated that no close relationship
within the genus (Nair 2012).
The aim of this study is to understand the species diversity of Merremia
and their distribution pattern, to provide an identification key to the species, and to
determine relationship among the species of Merremia in Sumatera using
morphological character for cladistic analysis.
2 LITERATURE REVIEW
Taksonomy Merremia Dennstedt ex Endlicher
The Genus name of Merremia firstly published by Dennstedt (1818) which
the combination of Merremia convolvulacea. The name was appeared based on
plate of Rheede in Hortus Malabaricus without description and delimitation of
genus so the name is not validly published. After that some botanists mentioned
the genus in many publications with more detail to describe this plant, either by
the introduction of a new species or displacements that caused change of the
names among other species of genus.
Choisy (1833) renames Merremia to be Skinneria and Bojer (1837)
renames Merremia to be Spiranthera. However both of them were rejected
because the name Skinnera has been used on Onagraceae by Forster (1775) and
the name of Spiranthera has been used by St.Hil (1823) on Rutaceae. Endlicher
(1841) had accomplished the description Merremia in which the name became
validly published and recorded in International Code of Botanical Nomenclatur
and the status is set as Nomina Conservanda (Art. 14.8.), unfortunately the
description was still incorporated with Ipomoea.
Hallier (1893) had made genus Merremia delimitation and separated it
from Ipomoea based on pollen character in which Ipomoea has spinulose pollen
while Merremia has non-spinulose pollen. Furthermore, Hallier (1894) had
divided genus Merremia into 3 sections consist of Xanthips, Streptandra and
Skinneria. In 1913 Wingkler complements writings of section by adding type of
species at each section and furthermore he adding one more section that is
Hailale. Van Ooststroom and Hoogland (1953) adding one more section that is
Wavula and renames of Skinneria to be Eu-Merremia because the name of
Skinneria has been used as genus Skinnera Forst.
Morphological studies of pollen first done by Hallier (1893) where the
character is very important to recognize members of Convolvulaceae. After that
some botanists such as Erdtman (1952), Manitz (1969) and Huang (1972) worked
with pollen although limited in some regions. Sengupta (1972) conducted a study
of pollen extensively and discovered some type of pollen and evolutionary
relationship, although the work was not covering all species of Convolvulaceae.
The genus Merremia is confused with Operculina that has same nonspinulose pollen and capsule will break regularly. The pollen from 55 species
Merremia and Operculina have been studied and groups into five types:
3
tricolpate, 5-6 colpate, 9-12 colpate, 12 rugate and pantoporate. The study showed
no closed relationship between Merremia and Operculina (Ferguson et al 1977).
Austin and Staples (1980) have proposed M. tridentata (L.) Hallier to be a
new genus namely Xenostegia, due to M. tridentata has porate pollen, since all
Merremia have colpate pollen.
Distribution species of Merremia
The genus Merremia is found in the tropical regions of both hemispheres.
Merremia wildly distributed throughout the tropical regions of Asia, Africa,
Australia, North and South America (Fang and Staples 1995) and approximately
there were 80 species of Merremia in the world (Ooststroom and Hoogland 1953).
In Mexico, there are 12 species (Mc Donal 1991), Micronesia 7 species (Fosberg
& Sachet 1977), Australasia and Pasific 49 species (Staples 2010), Malesiana 23
species (Ooststroom and Hoogland 1953), China 19 species (Huang 1995), Malay
Paninsula 8 species, Malesia 23 species (Ooststroom 1939) and Java 12 species
(Backer and Bakhuizen van den Brink Jr. 1965)
Figure 1 Distribution of Merremia
(George Staples 2012)
Economic value of Merremia
Some species of the genus Merremia have been used by human kind for a
long time especially for food and medicinal purposes (roots, leaves and tubers)
and it has economic value. The tubers, roots or stems of several Merremia species
are used as a purgative. A decoction of the root M. peltata is used to treat stomach
muscular rigidity. The tubers of M. tuberosa in Java and India are known as a
drastic purgative, M. umbellata in India and M. peltata in Philippines are mildly
laxative and widely taken for dysentery. The sap from the stems of M. peltata in
Philippines mixed with lumps of sugar is a remedy for cough and in Indonesia,
Fiji and India, diluted sap from the young stems of M. peltata is used as eye or ear
drops.
The leaf of M. emarginata in Indonesia has applied as a laxative and
mixed with lumps of sugar is a remedy for cough. An infusion of the leaves of M.
4
dissecta in Africa and M. peltata in Philippines are taken as a sedative for chest
complaints, and a poultice of fresh, crushed leaves is applied as a resolutive), in
Argentina M. dissecta var. edentata used as food by local people (Austin 2007). In
Fiji, a decoction of the leaves of M. peltata is used to treat boils, infections and
appendicitis and in Florida, M. tuberosa used as laxatin (Austin 1998). The beauty
of shape and color of flowers are very attractive, may chance to make this plant as
ornamental plant.
3 MATERIALS AND METHODS
Materials
A Total of 175 sheets (164 collection numbers) which are deposited in
Herbarium Bogoriense, 97 sheets (95 collection numbers) which are deposited
Andalas University Herbarium (ANDA) and 9 sheets (4 collection numbers)
which are deposited in BIOTROP Herbarium (BIOT) were studied. It composed
of all specimens of Merremia in Sumatra. Additional specimens were collected by
the author 28 sheets (15 collection numbers) from North Sumatra, West Sumatra
and Lampung, which are stored in BO and BIOT. The digital data that consist of
type specimens and distributions of species from JSTOR Plant Science and
Convolvulaceae Unlimited were also used here.
Methods
All the hebarium specimens were studied based on their morphological
character following de Vogel (1987) and Rifai (1976). The terminology of
morphological characters followed Lawrence (1955), Veldkamp (1987), Harris
and Harris (1994), Hickey and King (2005) and Stearn (1983). Information on the
habitat, ecology, distribution, vernacular names and uses were noted from the
specimens label and literatures.
Phylogenetic Analysis.
Thirty five morphological characters selected during morphological
observation of field and herbarium specimens were used in this study (table 1).
Evolvulus nummularius and Porana volubis were selected as an outgroup.
Morphological characters were analyzed based on Maximum Parsimony using
PAUP*4.0b4 (Swofford, 2002).
4 RESULT AND DISCUSSIONS
By examined 175 sheets specimens at 3 Herbaria (BO, BIOT and ANDA),
8 species of Merremia (M. dissecta, M. emarginata, M. hirta, M. peltata, M.
quinquefolia, M. tridentata, M. tuberosa and M. vitifolia), 2 subspecies (M.
umbellata ssp. orientalis and M. tridentata ssp. hastata), 1 varietas (M. boisiana
var. sumatrana) and 2 forma (M. hederacea f. pubescens and M. hederacea f.
barbata) recognized. In addition, M. cissoides was found in North Sumatra as a
new record. M. tridentata and M. tridentata ssp. hastata are excluded because
they have been changed to be a new genus Xenostegia by Austin and Staples. So
5
that, as a result there are 8 species, 1 subspecies, 1 varietas and 2 forma of
Merremia found in Sumatra.
Delimitation of Genus and Species Diversity
Morphological Species Concept (MSC) was used in recognizing the taxa
Merremia in Sumatra. The taxa delimitation in the genus or species level of
Merremia in Sumatra based on their morphological characters. This
morphological delimitation has developed based on morphological variation of
generative and vegetative organs like leaf morphology, infloresences form,
capsule form, seed and also hairs in each part of plant. The genus Merremia can
be distinguished from other genera of Convolvulaceae by the following
characters: variation of leaves, corolla funnel-shape or campanulate which are
white or yellow, hairs on corolla, 5 nerved midpetaline bands distinctly, stigma
globular, anther straight, spiral or coil, capsule valve dehiscent regularly, seeds 24 glabrous, pubescent or pillose. Futher full description and delimitation of each
species can be seen in taxonomic treatment.
General Morphology Merremia Dennstedt ex Endlicher of Sumatra
Habit
Most species Merremia in Sumatra are annual or perennial herbaceous
vines (M. emarginata, M. hirta, M. hederacea, M. cissoides, M. umbellata) and
woody vines (M. boisiana var. Sumatrana, M. peltata, M. tuberosa and M.
vitifolia).
Stem
Stem shape Merremia is terete. They are creeping or twinning and often
low-growing on grassy or rocky places and upon trees. All of Merremia have
milky white sap and a few has fragrant (M. cissoides). The stems of Merremia
have unique character, due to they can be twin on trees up to 30 m without tendril.
They have active mechanisms to climb on and attach themselves to the host plant.
They present a circumnutational movement in which their stems, somewhat
arching in the distal portion, rotate on their own axis, rather like the hands of a
clock. This movement is essential so that the vine can locate a structure on which
it can climb on the canopy of the shrubs and young trees and thus use as a source
of support. They have long and flexible stems that depend on external support to
maintain themselve erect to reach illuminated areas in their habitat. Their stems
are characterized by the scarcity of supporting cells (fibers) and an increase in the
diameter of the xylem vessels, which may be visible to the naked eye (Herbaceous
stems). For the woody vines (M. peltata, M. vitifolia, M. boisiana var. sumatrana)
they have stems to 3-8 cm in diameter, the stems have a non-circular cross section
which oval and somewhat furrowed. Woody tissues can be isolated by more
flexible ones, the stems then become more flexible and even when it becomes
very woody and it is still elastic. This kind of wood anatomy is particularly well
developed in vine species that are subject to important growth constraints like
torsion and tears. Some species of Merremia have tuberous roots and these tubers
are formed from the main roots (M. peltata, M. tuberosa and M. cissoides). These
tubers occur in both wet and dry environments. In extremely dry conditions, stems
6
and roots can become a caudex that is one of kind of tuber indicating an
adaptation to extreme environment.
Leaves
The leaves of Merremia are alternate and have variable shapes and sizes.
Some species usually have small leaves (1-5 cm long) for example in M.
emarginata and M. hirta but the other has bigger and broader leave to 40 cm in M.
peltata. Almost the Sumatra M. have simple leaves except M. quinquefolia and M.
cissoides which have compound leaves. The blade shapes variable from ovate (M.
umbellata ssp. orientalis and M. boisiana var. sumatrana), linear (M. hirta),
reniform (M. emarginata), peltate (M. peltata), trilobe (M. hederacea), shallow
lobe (M. vitifolia), deeply lobes (M. cissoides, M. dissecta, M. tuberosa and M.
quinquefolia).
The leaf base, usually are cordate or attenuate but sometimes peltate (M.
peltata). The margin are variable from entire to crenate (M. hederacea, M. peltata,
M. tuberosa, M. umbellata ssp. orientalis and M. emarginata) and dentate (M.
vitifolia, M. dissecta, M. cissoides and M. quinquefolia). The apex also have
variable shapes from emarginate (M. emarginata), mucronulate (M. peltata, M.
umbellata ssp. orientalis and M. vitifolia), acute to acuminate (M. cissoides, M.
tuberosa) and caudate (M. boisiana var. sumatrana). The leaf surfaces are often
glabrous and pubescent on both sides but sometimes hirsute (M. vitifolia) or
sparsely pubescent to glabrous above and glabrous below (M. umbellata ssp.
orientalis). The nerves are variable from pinnate (M. boisiana var. sumatrana, M.
emarginata, M. hirta, M. peltata, and M. umbellata ssp. orientalis) and palmate
(M. cissoides, M. hedeacea, M. tuberosa, M. vitifolia and M. quinquefolia).
Figure 3 Palmately compound leaves (M. cissoides and M. quinquefolia)
7
a
c
b
e
f
d
g
Figure 2 The variation blade of simple leaves a. linear (M. hirta); b. ovate (M.
umbellata ssp. orientalis and M. boisiana var. sumatrana); c. reniforme
(M. emarginata); d. peltate (M. peltata); e. trilobe (M. hederacea), f.
shallow lobe (M. vitifolia); g. deeply lobes (M. dissecta and M.
tuberosa).
Infloresences
The inflorescences are axillary with few to many flowered and usually
cymose. The ramified inflorescences are variable, usually cyme but sometime
panicle, (M. boisiana var. sumatrana), cincinnus (M. vitifolia) and umbel (M.
umbellata ssp. orientalis). The peduncle thick or thin, sub terete. They have
variable peduncle sizes but sometime M. emarginata has 1 mm or nearly absent.
The peduncle surface is usually glabrous or pubescent, but sometime hirsute (M.
vitifolia and M. cissoides). The bracts of Merremia are short, occasionally
caducous and have variable shapes that is ovate or triangularis or ovoid.
Flower buds
The important character in Merremia is the spread of the hairs on corolla
that can be easily observed at the mature flower bud. The flower bud shapes
variable from ovoid (M. quinquefolia, M. dissecta, M. hirta, M. peltata, M.
umbellata ssp. orientalis and M. vitifolia), ovoid to conical (M. tuberosa),
subglobose (M. boisiana var. sumatrana) and subrhombhoid (M. cissoides).
8
a
b
Figure 4
c
d
The variation ramified inflorescence a. Cincinus (M. vitifolia); b.
cyme (M. hirta, M. emarginata, M. peltata, M. hederacea, M. dissecta
and M. tuberosa); d. umbel (M. umbellata ssp. orientalis); e. panicle
(M. boisiana var. sumatrana)
Flower
Generally, Flower is arranged in a stalk head and form a flowering head
with the calyx, corolla tubes which are joined together. The flowers are
pentamerous, actinomorphic and bisexual. The species have variable flower sizes.
Some species usually have small (1-2 cm long) at M. hederacea, M. emarginata,
and M. cissoides, slightly big (2-3 cm long) at M. hirta, M. quinquefolia, M.
boisiana, M. umbellata ssp. orientalis and M. vitifolia) and big (>3 cm long) at M.
dissecta, M. peltata and M. tuberosa).
Calyx – The calyx have five free sepals, overlapping in a quincuncial
arrangement with outer and inner sepal unequal or almost equal length, concave,
persistent, usually have glabrous or pubescent surfaces but sometime have hirsute
(M. cissoides and M. vitifolia) and enlarged in fruit. The sepal is an important
morphological character to distinguish between species. The outer and inner
sepals have variable sizes and shapes. Sometimes outer more long than inner sepal
or reverse. The sepal shapes are orbicular to transverse elliptic (M. boisiana var.
sumatrana), elliptic (M. hirta), rhomboid (M. cissoides), oval lanceolate (M.
dissecta), obovate (M. emarginata, M. hederacea, and M. umbellata ssp.
orientalis), ovate (M. peltata, M. tuberosa, M. vitifolia and M. quinquefolia).
Corolla - The corolla is gamopetalous or petals are fused into a tube. They
have 5 bands in the middle of each petal. These bands are clearly seen in the
opened flower and are generally known as the mid-petaline bands and it which
help the corolla to coil on self when fading. They usually have glabrous
midpetaline bands but sometimes pillose (M. boisiana var. sumatrana). The
corolla limbs are slightly 5-lobe or sometimes 10-lobe (M. boisiana var
sumatrana) with pubescent surface.
9
a
Figure 5
b
The variation corollas a. campanulate (M. boissiana var. sumatrana,
M. hederacea, M. emarginata, M. peltata, and M. hirta); b. Funnelshaped (M. cissoides, M. dissecta, M. umbellata ssp. orientalis, M.
tuberosa, M. vitifolia, and M. quinquefolia).
There are 2 corolla shapes, that is campanulate (Merremia boissiana var.
sumatrana, M. hederacea, M. emarginata, M. peltata and M. hirta) and funnelshaped (M. cissoides, M. umbellata ssp. orientalis, M. tuberosa, M. vitifolia, and
M. quinquefolia). The corolla colours variable such as yellow or bright yellow (M.
dissecta, M. peltata, M. tuberosa, M. hederacea, M. vitifolia and M. hirta),
whitish yellow or orange (M. umbellata ssp. orientalis) and white (M. cissoides
and M. boisiana var. sumatrana.
Stamen – The number of stamens are 5. Usually, they are inserted in the
corolla. The filament is sticky at tube, slender, thin but broader and hairy at the
basal and dorsifixed below. The anther shape various from straight (M.
emarginata, M. hederacea, and M. umbellata ssp. orientalis), spiral twisted (M.
cissoides, M. dissecta, M. hirta, M. peltata and M. quinquefolia), and coil (M.
boisiana var. sumatrana, M. tuberosa and M. vitifolia). The indument of anther
usually glabrous but sometimes hairy (M. peltata)
a
Figure 6
b
c
The variation shapes anther a. straight (M. emarginata, M.
hederacea, and M. umbellata ssp. orientalis); b. coil (M. boisiana
var. sumatrana, M. tuberosa and M. vitifolia); c. spiral twisted (M.
cissoides, M. dissecta, M. hirta, M. peltata and M. quinquefolia.
10
Pistill – The stigma of all species are always macrostylous, biglobular
with smooth and glabrous and the colour of stigma is white. The style shape is
terete and has glabrous surface. The ovaries of all species are always superior and
glabrous.
Fruit
The fruit is typically irregularly dehiscing capsule with 4 cells and 4
valves. The capsule shapes are variable from sub globose (M. dissecta, M.
tuberosa, M. emarginata and M. vitifolia), ovoid (M. hirta and M. hederacea,
ovoid to conical (M. peltata, M. dissecta and M. umbellata ssp. orientalis). The
stalks are subterete and then thicked towards to the top.
a
Figure 7
b
The variation capsules a. ovoid to conical (M. peltata, M. dissecta
and M. umbellata ssp. orientalis); b. sub globose (M. dissecta, M.
tuberosa, M. emarginata and M. vitifolia).
Seed
The seed shapes variable from late elliptic (M. emarginata and M. hirta),
ovoid to conical (M. hederacea f. pubescent, M. peltata, M. tuberosa and M.
dissecta), transverse ovate (M. hederacea f. barbata) and obovate (M. cissoides
and M. vitifolia). Merremia have various seed sizes. Some species have small
sizes (M. hederacea, M. emarginata, M. cissoides, M. quinquefolia), slightly big
sizes (M. umbellata ssp. orientalis, M. vitifolia, M. quinquefolia and M. peltata)
and big sizes (M. tuberosa and M. dissecta). The seed surfaces are variable from
glabrous (M. vitifolia, M. dissecta), pubescent (M. boisiana var. sumatrana, M.
cissoides), bearded (M. hederaceae f. barbata), pillose (M. umbellata ssp.
orientalis and M. peltata), pubescent at margin and hylum (M. hederacea f.
pubescens and M. tuberosa), pillose at margin and hylum (M. hirta). The colours
of seed are brown, dark brown and black.
Distribution Merremia in Sumatra
The distribution of Merremia in Sumatra (Figure 8) covered Nangro Aceh
Darussalam, North Sumatra, West Sumatra, Riau and Riau Archipelago, Jambi,
Bengkulu, South Sumatra, Bangka Belitung and Lampung. The highest
concentration of species is found in North Sumatra 10 species (M. hederacea f.
11
pubescent, M. peltata, M. umbellata ssp. orientalis, M. tuberosa, M. vitifolia and
M. boissiana var. sumatrana, M. dissecta, M. emarginata, M. hirta and M.
cissoides) and then followed by West Sumatra 7 species (M. boisiana var.
sumatrana, M. hederacea f. barbata, M. hirta, M. peltata, M. umbellata ssp.
orientalis, M. tuberosa and M. vitifolia), Lampung 5 species (M. peltata, M.
umbellata ssp. orientalis, M. vitifolia, M. hirta and M. quinquefolia), 3 species
South Sumatra (M. hederacea f. barbata, M. umbellata ssp. orientalis, and M.
vitifolia) and NAD (M. vitifolia, M. peltata and M. umbellata ssp. orientalis), 2
species in Bangka Belitung (M. umbellata ssp. orientalis and M hirta), Bengkulu
(M. umbellata ssp. orientalis and M. peltata), and Jambi (M. hederacea f.
pubescens, M. umbellata ssp. orientalis), 1 species in Riau Archipelago (M.
umbellata ssp. orientalis) and Riau (M. biosiana var. sumatrana).
Figure 8 Distribution of Merremia in Sumatra.
M. umbellata ssp. orientalis found widespread in all of Sumatra province
except Riau from 20 mdpl until 1000 mdpl. This plant is native in China and
naturalized in Sumatra. M. peltata was found in NAD, North Sumatra west
Sumatra, Bengkulu and Lampung at 20-600 mdpl. This plant is origin of pacific
12
region and has been invaded at Bukit Barisan Selatan National Park (Lampung).
M. boisiana var. Sumatra is endemic in Sumatra since they are found in this island.
This plant firstly found by Lorzing at 1916 in Sibolangit (North Sumatra) at 500600 mdpl which deposited in Herbarium Bogoriense. After a few time, this plant
widespread to Mountain Leuser (North Sumatra) at 50-100 mdpl, Kampar (Riau) at
400-550 mdpl and West Sumatra at 50-780 mdpl which deposited in Andalas
Herbarium. M. emarginata was new record in Sumatra while M. cissoides was new
record in Indonesia. The occurrence of M. emarginata in Sumatra was proven by a
specimen collected from North Sumatra (J.A Lorzing, 15891) at 15-16 mdpl. This
plant is native in Indonesia and widespread in tropical Africa and tropical Asia. M.
cissoides found Hafni Rahmadani (035) at 30 mdpl in Perbaungan (North
Sumatra). This species is native and weedy in tropical America. M. dissecta, M.
quinquefolia and M. tuberosa are cultivated in Sumatra (Table 1)
Table 1 Distribution of Merremia in Sumatra
NO
Species
NAD NS
1
M. boisiana var.
+
Sumatrana
2
M. cissoides
+
3
M. dissecta
+
4
M. emarginata
+
5
M. hederacea
+
f. pubescens
6
M. hederacea
f. barbata
7
M. hirta
+
8
M. peltata
+
+
9
M. quinquefolia
10
M. tuberosa
+
11
M. umbellata
+
+
ssp. orientalis
12
M. vitifolia
+
+
Total
3
10
NAD : Nangro Aceh Darussalam.
NS
: North Sumatra.
WS
: West Sumatra.
R
: Riau
RA
: Riau Archipelago.
JA
-
SS
-
BB
-
BE
LA
-
-
+
-
-
-
-
-
-
+
-
-
-
+
-
-
+
+
+
+
-
+
+
+
+
+
+
+
+
+
+
7
1
1
2
+
5
WS
+
R
+
RA
-
-
JA
SS
BB
BE
LA
-
+
+
+
2
3
2
: Jambi.
: South Sumatra
: Bangka Belitung
: Bengkulu
: Lampung
Phylogenetic Analysis
A phylogenetic analysis using PAUP vers.4.0 was undertaken based on
morphological characters, using E. nummularius and P. volubis as the out group.
The phylogenetic analysis used 35 morphological characters (Table 2) and data
matrix of those characters which were used for the analysis showed in appendix 1.
13
Table 2 Morphological characters states used in the phylogenetic analysis
No
1
2
3
4
5
6
7
8
9
10
11
12
Characters
Habit
Leaf attacment
Leaf
Leaf Notch
Venation major
leaves
Blade shape
Leaf surface
Leaf margin
Leaf base
Leaf apex
Petiole
Infloresece
0.
0.
0.
0.
0.
woody
distichous
simple
absent
actinodromous
0. ovate
0. hairy
0. entire to crenate
0. cordate
0. mucronulate
0. < 0.5 cm
0. terminal and
axillary
0. present
0. cyme
15
16
Peduncle
Inflorescences
ramified
Pedicel
Outer sepal
17
18
Flower
Corolla lobed
0. 0.5 cm
1. axillary
1. absent
1.panicle
1. > 0.5
1.shorter than
inner one
1. 1/ leaf axil
1. not depply
lobed
1. funnel
1. hairy
1. yellow
1. biglobular
1. < 2
1. > 1
1. < 2
0. absent
1. inserted
1. equal
1. not straight
1. present
1. hairy
1. globose
1. >1
1.long hairy
1. margin seed
2. cincinnus
2. rotate
2. capitates
2. glabrous
3.linear
3. umbel
14
Morphological characters were analyzed based on Maximum Parsimony
using PAUP*4.0b10 (Swofford 2002) with Heuristic Search settings vegetative
and floral characters were selected for the analysis of which 27 were scored as
binary and 8 as multistate. Some characters were scored as missing data when the
data were unavailable. All characters are treated as unorder data and have an 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) and was evaluated using 100 bootstrap replicates and a complete Hsearch.
Clade support values were obtained by using bootstrap.
Maximum Parsimony (MP) analysis of the morphology dataset produced
17 shortest trees of 68 steps with consistency index (CI) = 0.60, homoplasy index
(HI) = 0.40, retention index (RI) = 0.56 and Rescaled consistency index (RC) =
0.34. Of 35 characters used in this analysis, 21 characters (60%) were potentially
parsimony-informative, and 14 characters (40%) were parsimony-uninformative.
Evolvulus nummularius
Porana volubilis
M. boisiana var. sumatrana
M. cissoides
M. quinquefolia
M. tuberosa
M.dissecta
M. vitifolia
M. emarginata
M. hederacea barbata
M. hederacea pubescens
M. hirta
M. peltata
M. umbellata ssp. orientalis
Figure 9 Strict consensus tree resulted morphological characters. CI = 0.60, HI =
0.4, RI = 0.56, RC = 0.34. The number above the branches showed the
bootsrap value
15
The cladogram devided the taxa into two groups. The separation support
M. boisiana var. sumatrana, M. cissoides, M. dissecta, M. emarginata, M.
hederacea f. pubescens, M. hederacea f. barbata, M. hirta, M. peltata, M.
quinquefolia, M. tuberosa, M. vitifolia and M. umbellata ssp. orientalis as ingroup
with E. nummularius and P. volubis as outgroup as strengthened (BS=77%)
(figure 9). It was separated base on some character such as leaf attachment
(character 2), petiole (character 11), infloreseces (character 12), pedicel (character
13), flower (character 17), corolla lobed (character 18), corolla surface (character
20), corolla colour (character 21), ovary (character 24), style number (character
25), style branch (character 26), stamen (character 27), stamen length (character
28), filament (character 31) and seed number (character 33).
The topology of strict consensus tree resulting from morphological and
strongly support the monophyletic, paraphyletic species and metaspecies of
Merremia. M. boisiana var. sumatrana is metaphyletic (unresolved) and it is a
sister group of monophyletic M. cissoides, M. dissecta, M. hederacea f.
pubescens, M. hederacea f. barbata, M. quinquefolia, M. tuberosa and M.
vitifolia. M. boisiana var. sumatrana is paraphyletic species with M. umbellata
ssp. oriental, M. hirta, and M. peltata which shared by two populations of the
metaspecies and monophyletic. While M. emarginata is metaspecies (unresolved).
The morphological phylogenetic analysis for monophyletic showed that M.
cissoides closed relationship with M. quinquefolia. Both species were placed in
the same clade with strong support 79%. While, M. tuberosa become as the basal
sister of M. cissoides and M. quinquefolia with weakened 61%. Phenetically, M.
tuberosa separated from M. cissoides and M. quinquefolia based on some
character such as leaf (character 3) venation major leaves (character 5) and margin
leaf (character 8). M. vitifolia and M. dissecta are closed relationship with
weakened 58 % but separated based on Ramified (character 14) and outer sepal
(character 16). M. hederacea f. barbata closed relationship with M. hederacea f.
pubescens strong support 51% and separated base on seed surface (character 34)
and position of seed surface (character 35. Both of each species were placed in the
same clade.
16
Generic Description
Merremia Dennstedt ex Endlicher
Merremia Dennst., Schlüss., Hort. Malab. 1818, 34, (nomen nudum); Hall. f. in
Engl., Bot. Jahrb. XVI (1893) 581; Baker & Rendle in This.-Dyes, Fl. Trop. Afr.
IV, 2 (1905) 101; Ridl., Fl. Mal. Penin. II (1923) 456; Merrill, Enum. Philipp. Fl.
Pl. III (1923) 360; Ooststr., Blumea III (1939) 292; Ooststr., FM I.4.4 (1953) 450;
Backer and Bakhuizen f., FJ. II (1965) 488. Type species: Merremia
convolvulacea (= M. hederacea (Burm. f.) Hall f.) (illustration!)
.─ Skinneria Choisy, in Mem. Soc. Phys. Genève VI (1833) 487. Type species:
not seen.
.─ Spiranthera Boj., Hort. Maurit. (1837) 226; Griseb., Fl. Brit. West Ind. Isl.
(1864) 470. Type species: not seen.
Annual or perennial herbs or woody twinning and creeping at trees or
stones. Stems terete, prostate, sometimes stems come out from tuber, glabrous or
hairy, with milky juice. Leaves simple or palmately compound, alternate: petiole
terete, glabrous or pubescent or glabrescent or hirsute; blade variable shape and
size, ovate or orbicular or emarginate in outline, base cordate or peltate or
attenuate or truncate or rounded, margin entire or crenate or dentate, apex
mucronulate or acute or acuminate or emarginated or caudate, primer vine pinnate
or palmate. Inflorescence axillary, solitary or in few to many flower, cymose,
ramified variable, compound cyme or cincinnus or umbel or panicle; peduncle
terete, glabrous or pubescent or hirsute. Bract small and caducous, ovate or
triangularis or ovoid. Flower buds ovoid or ovoid to conical or subglobose or
rhombhoid. Flower bisexual: stalk terete, glabrous or pubescent or hirsute sepals 5
concave, outer and inner sepal unequal or almost equal length, orbicular to
transverse elliptic or elliptic or subrhomboid or oval lanceolate or ovate, glabrous
or pubescent or hirsute, in several species enlarged in fruit; corolla funnel or
campanulate, yellow or white or whitish yellow or bright yellow or orange or
white deep purple, corolla limb slightly 5-lobed or 10-lobed, pubescent,
midpetaline bands 5, glabrous or pillose; stamens 5, anther variable shape, straight
or spiral twisted or coil, glabrous or pillose, filament unequal, dorsifixed below,
sticky at tube, slender, thin but broader and hairy at the basal pistill 1, stigma
macrostylous, biglobular, smooth and glabrous, white; styles terete, glabrous
ovary superior, glabrous. Fruit a capsule, valve, dehiscent, globose or ovoid or
ovoid to conical, glabrous. Seeds 2-4, variable shape and size, ovate or elliptic or
ovoid to conical or transverse ovate or obovate, glabrous or pubescent or sericious
around or at margin and hylum, brown or dark brown or black.
17
Key to the Sumatran Species of Merremia
1.
2.
3.
4.
5.
6.
7.
8.
9.
Leaves simple…………………………………………………….…........2
Leaves palmately compound with 5 leaflets………………………….….3
Blade palmate with 3-7 segment shallow or deeply lobes…….….....…....4
Blade not palmate, peltate, ovate, linear, reniform, ……………...……….7
Leaflets elliptic, apex mucronulate, margin pubescent; bract linear; sepal
subrhomboid, hirsute; capsule globose; seed 2 mm long;
brown………………………………………………………..2. M. cissoides
b. Leaflets narrowly oblong, apex acute, margin glabrous; bract narrowly
triangular; sepal ovate to oblong, glabrous; capsule ovoid to conical; seed
4 mm long, black………………………………..…..….8. M. quinquefolia
a. Blade deeply lobed into 7 segments, inflorescense ramified cyme, corolla
funnel shaped…………………………………………………………...….5
b. Blade shallow lobed with 3-7 segments, inflorescense ramified cyme or
cincinus, corolla funnel shaped or campanulate……………………….....6
a. Petiole hirsute; blade lanceolate-elliptic, glabrous, one upper and two
lateral blade segment larger than two lateral and two bottom ones, margin
dentate, apex mucronulate; sepal ovoid; anther spiral twisted; capsule
globose; seeds 7-7.5 mm long, glabrous, black……………...3. M. dissecta
b. Petiole pubescent; blade oblong-lanceolate, pubescent, one upper blade
segment larger than lateral and bottom one, margin entire, apex acute to
acuminate; sepal ovate; anther coil; capsule globose; seed 13-14 mm long,
pu