MERREMIA DENNSTEDT EX ENDLICHER
(CONVOLVULACEAE)
IN SUMATRA
HAFNI RAHMADANI
THE GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
BOGOR
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
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 non-spinulose 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
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.
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.
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.
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.
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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
Title : Merremia Dennstedt ex Endlicher (Convolvulaceae) in Sumatra
Name : Hafni Rahmadani
NRP : G353100131
Certified by Supervisor Committee
Dr. Sri Sudarmiyati Tjitrosoedirdjo, M.Sc Dr. Harry Wiriadinata
Chairman Member
Approved by
Head of Plant Biology Dean of Graduate School
Study Program
Dr. Ir Miftahudin, M.Si Dr. Ir. Dahrul Syah, M.Sc. Agr.
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
TABLE OF CONTENTS
Taksonomy of Merremia Dennst. ex Endlicher 2
Distribution species of Merremia 3
Economic value of Merremia 4
Delimitation of Genus and Species Diversity 5
General Morpholgy Merremia Dennstedt ex Endlicher of Sumatra 5
Habit 5
Key to the Sumatran species of Merremia
Merremia quiquefolia 33
Merremia tuberosa 34
Merremia umbellata 36
Merremia vitifolia 38
5 CONCLUSION 41
REFERENCES 41
APPENDIX 44
LIST OF TABLES
1. Distribution of Merremia inSumatra 12
2. Morphological characters state used in phylogenetic analysis 13
LIST OF FIGURES
1. Distribution of Merremia 3
2. Palmately compound leaves 6
3. Variation blade of simple leaves 7
4. Variation ramified inflorecences 8
5. Variation corollas 9
6. Variation shapes anther 9
7. Variation capsules 10
8. Distribution of Merremia in Sumatra 11
9. Strict consensus tree resulted from morphological characters 14 10.Distribution of Merremia var. sumatrana in Sumatra 19
11.Merremia boisiana var. sumatrana 20
12.Merremiacissoides 22
13.Distribution of Merremia cissoides in Sumatra 23
14.Distribution of Merremia dissecta in Sumatra 24
15.Distribution of Merremia emarginata in Sumatra 25
16.Merremia emarginata 26
17.Distribution of Merremia hederacea in Sumatra 28
18.Distribution of Merremia hirta in Sumatra 30
19.Merremia peltata 32
20.Distribution of Merremia peltata in Sumatra 33
21.Distribution of Merremia quinquefolia in Sumatra 34
22.Distribution of Merremia tuberosa in Sumatra 35
23.Distribution of Merremia umbellata ssp. orientalis in Sumatra 37
24.Distribution of Merremia vitifolia in Sumatra 39
25.Merremia vitifolia 40
LIST OF APPENDIX
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 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 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
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 MerremiaDennstedt 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 non-spinulose pollen and capsule will break regularly. The pollen from 55 species
Merremia and Operculina have been studied and groups into five types:
4
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.
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.
5
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 2-4 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
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
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).
7
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).
a b c
e
d
Figure 4 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) andbig (>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.
a b c d
9
Figure 5 The variation corollas a. campanulate (M. boissiana var. sumatrana, M. hederacea, M. emarginata, M. peltata, and M. hirta); b. Funnel-shaped (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 funnel-shaped (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)
Figure 6 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.
a b c
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. stalks are subterete and then thicked towards to the top.
Figure 7 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.
a b
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.
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 500-600 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 plantis native in Indonesiaand 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
NAD : Nangro Aceh Darussalam. JA : Jambi. matrix of those characters which were used for the analysis showed in appendix 1.
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Table 2 Morphological characters states used in the phylogenetic analysis
No Characters Character & State
1 Habit 0.woody 1.herbs
2 Leaf attacment 0.distichous 1.alternate
3 Leaf 0.simple 1.compound
10 Leaf apex 0.mucronulate 1.acuminate 2.emarginate
11 Petiole 0. < 0.5 cm 1. > 0.5 cm
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.
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.
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. a. Leaves simple……….…...2 b. Leaves palmately compound with 5 leaflets……….….3 2. a. Blade palmate with 3-7 segment shallow or deeply lobes…….…...…....4
b. Blade not palmate, peltate, ovate, linear, reniform, ………...……….7 3. a. 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
4. 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 5. 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, pubescent at margin and hilum, dull black……….9. M. tuberosa
6. a. Blade shallow lobed into 3 segments, ovate in outline; stem pubescent; inflorescense ramified cyme; sepal obovate; corolla campanulate; anther straight; seed ovoid to conical or transverse ovate, 2.2-3 mm long, pubescent or barbata...5. M. hederacea
b. Blade shallow lobed into 7 segments, orbicular in outline; stem hirsute; inflorescense ramified cincinus; sepal ovate; corolla funnel shaped; anther coil; seed 6 mm long, glabrous …..………...11. M. vitifolia
7. a. Blade peltate, ovate………..………...………..…...8
b. Blade linear, reniform………...………..…10
8. a. Blade peltate, glabrous, apex mucronulate; stems with tuber; inflorescense ramified cyme; sepal ovate; corolla funnel; anther spiral twisted, pillose; seeds 4, sericious, black………...7. M. peltata
b. Blade ovate..………..………9
9. a. Blade apex caudate; inflorescense ramified panicle; corolla campanulate, white, midpetaline bands pillose; sepal elliptic; anther coil; capsule ovoid; seeds 2, pubescent...1. M. boisiana var. sumatrana
b. Blade apex mucronulate; inflorescense ramified umbel; corolla funnel, yellow or orange or whitish yellow, midpetaline bands glabrous; sepal obovate; anther straight, glabrous; seeds 4, sericious, black
10. a. Blade linear, apex mucronulate; peduncle 1.5-3.4 cm; sepal elliptic; corolla broad funnel; anther spiral twisted; capsule ovoid; seed pubescent at
margin and hylum………..……….………..……....6. M. hirta
b. Blade reniform, apex emarginated; peduncle < 1 cm or nearly absent; corolla campanulate; anther straight; capsule globose; seed glabrous...4. M. emarginata
Taxonomic Treatment
1. Merremia boisiana (Gagnep.) Ooststr. var. sumatrana Ooststr.
Merremia boisiana (Gagnep.) Ooststr., Blumea III (1939) 343; Ooststr., FM I.4.4 (1953) 450.─ Ipomoea boisiana Gagnep., in Notul. System III (1914) 141. Type: Indochina, Tonkin 138 (Holotype P) (by photo).
Merremia boisiana (Gagnep.) Ooststr. var. sumatrana Ooststr., Blumea III (1939) 344; Ooststr., FM I.4.4 (1953) 450. Type: Sumatra, Sibolangit, J.A. Lorzing 4723 (Isotype BO!).
Parennial herbs or woody liana, twinning until 30 m long . Stem terete, pubescent, purplish green, milky juice. Leaves simple: petiole terete, glabrous, 5.3-9.8 cm long, purple; blade cordate or ovate in outline, 7.5-17.6 cm long and 5.3-15.5 cm wide, base cordate or truncate, margin entire to crenate, apex cuspidate, vine pinnate, purple. Inflorescense axillary, racemose, solitary, ramified panicle; peduncle terete, pubescent 5.3-16.7 cm long, green. Bract small, triangular, ovoid, apex acute to acuminate, 1-3 mm long. Flower bud subglobose, 5-10 mm long. Flower: stalk terete, thicked toward at top, surface pubescent, 4-9 mm long; sepals 5 concave, unequal or outer sepals shorter than inner, outer sepals 2, orbicular to transverse elliptic, 4-6 mm long, inner sepals 3, transverse elliptic, 5-7 mm long; apex rounded or slightly retuse, glabrous, green; corolla campanulate, white, 1-2 cm long, corolla limb lobed; midpetaline bands 5, pillose; stamens 5, anther straight and then coil, pubscent,filament unequal, 4-5 mm long, dorsifixed below, sticky at tube, slender, thin but broader and hairy at the basal at tube, yellow; pistill 1, stigma biglobular, smooth and glabrous, style terete, 6 mm long, glabrous, ovary superior, glabrous. Fruit a capsule, valve, dehiscent, ovoid to conical. Seeds 2, pubescent, black.
Distribution. Known in North Sumatra (Sibolangit, leuser Mountain), West Sumatra and Riau.
Habitat and Ecology. Grows in secondary forest at ca 50-600 m.
19
Note.Merremia boisiana var. sumatrana is endemic in Sumatra and potentially to be invasive. This species was described by Ooststroom (1939) based on the specimens collected from Lorzing 4723 but the description not complete. This species has changed to be Decalobanthus boisianus var sumatrana by G. Staples and Ana R Simose (person communication) but it is not published yet.
Specimen examined. NORTH SUMATRA: Medan, Sibolangit 17 Feb 1916 J.A. Lorzing 4234 (BO); ibid, 18 Nov. 1927 J.A Lorzing 12281 (BO); ibid Aug 2013
Hafni Rahmadani 030 2013 (BIOT); Mountain Leuser 8 Aug. 1979 W.J.J.O &
BEE de Wilde-Duyfjes 19591 (BO). RIAU: Kawasan lindung Bukit Suligi Kec. Tambun Kab. Kampar alt 400-550 m 30 April 1999 Dayar Arbain & Rusdi Tamin
119 (ANDA). WEST SUMATRA: Dareh river alt 50-100 m, 19 Aug. 1981 M. Hotta, H. Okada & Tamin 03 (ANDA), Taram Tanjung atas about 14 km from Payakumbuh alt 420 m, 1 April 1988 R. Tamin, H. Hasnah & Haryanti 255 (ANDA), Taram Kapalo Banda about 14 km from Payakumbuh alt 440 m, 3 April 1988 Desni 27/88 (ANDA); ibid Yanosel vosri & Sri Suhartini 09 (ANDA), Bonjol about 16 km south east from Lubuk Sikaping alt 520-780 m, 3 June 1984
Aida Eka Putri & Nursyirwani 012 (ANDA), Desa Kulampi Kec, Sijunjung, Kab. Sawahlunto Sijunjung about 6 km East From Sijunjung Alt 200-300 m, 21-23 April 2000 Rusydi, Tuti, Rita, Yati, Rina & Dian 35 (ANDA); ibid Selvi, Nova, Feni, Novel, Novi & Putri 44 (ANDA); ibid group taxo 06 (ANDA), ibid Las, emi, Iin & Serly 28 (ANDA).
n
Figure 10 Merremia Boisiana var. sumatrana (Lam.) Hall. f. A. Habit; B. Blade; C; Infloresence & Flower; D. Calyx; E. Outer sepal; F. Inner sepal
C
B
A D
E F
21
2. Merremiacissoides (Lam.) Hallier f.
Merremia cissoides (Lam.) Hall. f. in Engl., Bot. Jahrb. Syst. XVI (1893) 552.─ Convolvulus cissoides Lam., Tabl. Encycl. Meth. Bot. I (1791) 462.─
Batatas cissoides (Lam.) Choisy., Mém. Soc. Phys. Genève VI (1834) 438.─
Ipomoea cissoides (Lam.) Griseb., Fl. Brit. W. Ind. Isl. (1861) 473 Type: French Guiana, Cayenne, Leblond s.n (holotype, P-Lamarck) (by photo)
Parennial herbs, twinning or creeping on grass or rock until 6 m long. Tuber 30 cm long. Stem terete, hirsute, milky juice, fragrant. Leaves palmately compound with 5 leaflets, orbicular in outline: petiole terete, 6-12 mm, hirsute; leaflets elliptic, one in the upper 1.6-3.1 cm long by 0.6-1.2 cm wide and two in the middle 1.4-1.1 cm long by 0.5-1.1 cm wide and two bottom 1-2.1 cm long by 0.4-1 cm wide, pubescent, basal cordate, margin dentate, apex mucronulate. Inflorescense axillary, 1 or few flowers, cymose, ramified compound cyme: peduncle terete, 1.1-1.5 cm long, surface hirsute. Bract linear, 2-13 mm long. Flower bud subrhomboid, 9-11 mm long. Flower: stalk terete, 3-4 mm long, surface pubescent; sepals 5 concave, surface hirsute, subrhomboid, unequal or outer sepal longer than inner, outer sepals 2, 1.2 cm long, inner sepals 3, 1.1 cm long; corolla funnel, white, 1.7 cm long, corolla limb 5-lobed, 1 cm wide, midpetaline bands 5, glabrous; stamens 5, anther straight and then spiral twisted, glabrous, 1 mm long, filament unequal, dorsifixed below, sticky at tube, slender, thin but broader and hairy at the basal, white; pistill 1, stigma macrostylous, biglobular, white, smooth and glabrous, style terete, glabrous, ovary superior, glabrous. Fruit a capsule, valve, dehiscent, globose, 11 mm long. Seeds 4, surface pubescent, 2 mm long, black.
Distribution
. Perbaungan, North Sumatra.
Habitat and Ecology. Grows in open grassy or in along riversides and association with Imperata at ca. 30 m above sea level.
Note. This species is native in tropical America. This is weedy in tropical American. Widespread in tropical America, middle America (Mexico) and South America and also naturalized in several place in the old world tropics such West Africa. In Asia known in India, Srilanka, Thailand, Taiwan. In Indonesia is a new record and potentially as an invasive alien species. It will likely increase its range as it has done elsewhere in the Old World tropics.
Figure 12 Merremia cissoides (Lam.) Hallier f. A. Tuber; B. liana habit; C. blade; D. Flower; E. Corolla limb; F. Stamen; G. Outer sepal; H. Inner sepal; I. Fruit capsule; J. Seeds
D
G H
I 1.2 1.7
1 cm
1 cm
1.1
2 mm
30 cm
J
A
B
C
E F
23
Figure 13 Distribution of Merremiacissoides in Sumatra
3. Merremia dissecta (Jacq.) Hallier f.
Merremia dissecta (Jacq.) Hall. f. in Engl., Bot. Jahrb. XVI (1893) 552; id. in Engl., Bot. Jahrb. XVIII (1894) 114; Baker & Rendle in This.-Dyes, Fl. Trop. Afr. IV, 2 (1905) 104; Ooststr., Blumea III (1939) 328;Ooststr., FM I.4.4 (1953) 448; Backer and Bakhuizen f., FJ. II (1965) 489.─ Convolvulus dissectus Jacq. Obs. II (1767) 4. Type: Amerika, Jacquin, plate #28 in protologue (lectotype, designated by Austin) (by photo).
Distribution. Medan, North Sumatra.
Habitat and Ecology. Found in cultivated gardens as ornamental plant. altitude 20 m above sea level.
Note. This species native in tropical and subtropical America. Widespread in North America, U.S.A (Texas, Lousiana, Florida, Georgia), middle America (Mexico), West Indies, South America and Northern South America and distjunct to Southern Brazil, Bolivia and Paraguay, also widely introduced and naturalized throughout the old world tropics such as Asia, Pakistan, India, China (Guangdong), Myanmar, Sri Lanka, Indonesia: Sumatra (Medan), Java, Borneo. Australia, Western Australia, Northern Territory and Queensland.
Specimen examined. NORTH SUMATRA: Medan alt 20 m, 18 April 1930 J.A. Lorzing 16126 (BO).
Figure 14 Distribution of Merremia dissecta in Sumatra
4. Merremia emarginata (Burm f .) Hallier. f
Merremia emarginata (Burm.f.) Hall. f. in Engl., Bot. Jahrb. XVI (1893) 552; id. in Engl. Bot. Jahrb. VIII (1894) 118; Baker & Rendle in This.-Dyes, Fl. Trop. Afr. IV, 2 (1905) 113; Merrill, Enum. Philipp. Fl. Pl. III (1923) 360; Ooststr., Blumea III (1939) 312; Ooststr., FM I.4.4 (1953) 444; Backer and Bakhuizen f., FJ. II (1965) 489.─Evolvulusmarginatus Burm. f. Fl. Ind. (1768) 77.Type: Indonesia, Java, Anno, 1757, Kleinhof 85 (Lectotype, G-Burman.) (by photo)
25
Annual herbs, twinning or creeping on grass. Stem terete, thin, surface pubescent or glabrescent. Leaves simple: petiole terete, 7-40 mm long, surface pubescent; blade reniforme, kidney to broad ovate in outline, surface glabrous to pubescent 8-17 mm long by 10-15 mm wide, basal cordate, margin entire to crenate and with dentate at lobe, apex emarginate or sometimes rounded, vein pinnate. Inflorescense axillary, 1 or few flower, cymose, ramified compound cyme; peduncle terete, 1 mm long or near absent, surface glabrous. Bract small and caducous. Flower buds ovoid, apex acute, 6-14 mm long. Flower: stalk terete, thick toward to the upper 1-3 mm long, surface glabrous; sepals 5 concave, obovate or sometimes orbicular or subquadrate, unequal or outer sepal shorter than inner, 2 outer sepals and 1- 2 mm longer, 3 inner sepals and 3 mm long, surface pubescent; corolla campanulate, yellow; anther straight, surface glabrous; pistill 1, biglobular, ovary superior, glabrous. Fruit a capsule, valve, dehiscent, globose, 5 mm long. Seeds 4, 2 mm long, surface glabrous, black.
Distribution. Medan, North Sumatra
Habitat and Ecology. Grows in fields and open grasslands, roadsides at altitude ca. 15-16 m above sea level.
Note. This plant widespread in tropical Africa and tropical Asia. In Malesia: Philippines (Luzon), Indonesia (Sumatra (Medan), Java, Lesser Sunda (Sumbawa, Flores, Timor) and Celebes.
Specimen examined. NORTH SUMATRA: Medan alt 15-16 m, 5 June 1929 J.A. Lorzing 15891 (BO).
Figure 15 Merremia emarginata (Burm f.) Hall. f. A. Tuber; B. liana habit; C. Blade; D. Flower; E. Corolla limb; F. Stamen
27
5. Merremia hederacea (Burm. f.) Hallier f.
Merremia hederacea (Burm. f.) Hall. f. in Engl., Bot. Jahrb. XVIII (1894) 118; Ooststr., Blumea III (1939) 302; Ooststr., FM. I.4.4 (1953) 442; Backer and Bakhuizen f., FJ. II (1965) 490.─ Evolvulus hederaceus Burm. f. Fl. Ind. (1768) 77. Type: Indonesia. Java, D. Pryon s.n. (G-Burman)
.─ Merremia convolvulacea Dennst., Schlüss. Hort. Mal. (1818) 12, 23,34, nomen nudum; Hall. f. in Engl., Bot. Jahrb. XVI (1893) 552 Type: plate of Rheede in Hortus Malabaricus (Illustration!)
Merremia hederacea (Burm. f.) Hall. f. forma pubescens Ooststr., Blumea III (1939) 307; Ooststr., FM I.4.4 (1953) 442. Type: Sumatra, Teysmann s.n (Holotype L) (not seen).
Annual herb vines. Stem terete, twinning or creeping, surface pubescent or glabrescent except hirsuta at node. Leaves simple: petiole terete, 8-45 mm long, thin, surface pubescent; blade palmately 3-lobed or ovate in outline, surface pubescent, 2.3 cm long, apex mucronulate, basal cordate, vein palmate. Inflorescense axillary, 1 or few flower, cymose, ramified compound cyme; peduncle terete, thicker than petiole, 3-8.9 cm long, surface glabrous. Bract small and caducous. Flower bud obtuse. Flower: stalk terete, thick toward to the upper 2-4 mm long; sepals 5 concave, broad ovate, unequal or outer sepal shorter than inner, 2 outer sepals and 3 mm long, 3 inner sepals and 4 mm long; corolla campanulate, yellow; anther straight, surface glabrous; pistill 1, biglobular, ovary superior. Fruit a capsule, valve, dehiscent, ovoid to conical. Seeds 4, ovoid to conical, 2 mm long, surface pubescent at margin and hylum, black.
Distribution. Asahan and Medan (North Sumatra), Jambi.
Habitat and Ecology. Found on beach sides and road sides. Altitude ca. 0-30 m above sea level.
Merremiahederacea f. barbata Merremia hederacea f. pubescens
Specimen examined. NORTH SUMATRA: Medan alt ±25 m, 25 June 1928 J.A. Lorzing 996/13271 (BO), Lubuk Pakam, 26 Okt.1914 J.A. Lorzing 3299 (BO!), Serdang alt 8 m, 20 Nov. 1914 J.A. Lorzing 3386. JAMBI: Pahoe alt 30 m, Okt. 1929 Dr.O.Posthumus 1072 (BO). WEST SUMATRA: East Coast alt 20 m, Dec 1922 Jochems 3166 (BO), Pariaman, Teysman s.n. (BO).
Merremia hederacea (Burm. f.) Hall. f. forma barbata Ooststr., Blumea III (1939) 307; Ooststr., FM I.4.4 (1953) 442. Type: Sumatra, Korthals 273 (Holotype L) (by photo).
Different with forma pubescens, shape transverse ovate, seed size 2.5-3 mm, bearded at all surface of seed.
Distribution. Padang (West Sumatra) and Palembang (South Sumatra).
Habitat and Ecology. Grows on beach sides and road sides at altitude ca. 0-30 m above sea level.
Specimen examined. WESTSUMATRA: Padang March 1870, Anon, 226 (BO), ibid Korthals 271 (BO), SOUTH SUMATRA: Palembang alt 0 m, Jan. 1939
T.W.Rutten-Kooistra 109 (BO).
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6. Merremia hirta (L.) Merr.
Merremia hirta (L.) Merrill. in Philipp. Journ. Sc. VII (1912) Bot. 244; id., Enum. Philipp. Fl. Pl. III (1923) 361; id., in Philipp. Journ. Sc. LIX (1936) 452, pl. 1; Ooststr., Blumea III (1936) 307, Ooststr.FM. I. 4.4 (1953) 441.─ M. caespitosa (Roxb.) Hall. f. in Engl., Bot. Jahrb. XVI (1893) p. 552; id., Hall. f. in Engl., Bot. Jahrb. XLIX (1913) 378; Ridl., Fl. Mal. Penins. II (1923) 458; Backer and Bakhuizen f., FJ. II (1965) 490 Type: Philppine, Luzon, Merrill 3615 (Holotype L).
.─ Skinneria caespitosa (Roxb.) Choisy, in Mem. Soc. Phys. Genève VI (1833) 487 Type: not seen.
Annual herbs, twinning or creeping on grass until 4 meter long. Stem terete, surface puberulent or sometime hirsute, purple, milky juice. Leave simple: petiole terete, 0.2-1.9 cm long, surface glabrous or pubescent, purple; blade linear, linear to oblong, ovate in outline, 2.2-4.6 cm long by 0.15-0.16 cm wide, base rounded or auricularis, margin entire or entire to crenate, apex mucronulate, vein pinnate, surface glabrous. Inflorescense axillary, 2-few flower, cymose, ramified compound cyme; peduncle terete, 1.5- 3.4 cm long, glabrous. Bract small, ovate, 1 mm long. Flower bud, ovate, 0.2-0.6 mm long. Flower: stalk terete, thick toward to the upper 5-10 mm long, surface glabrous; sepals 5 concave, green, elliptic, unequal or outer sepal shorter than inner, outer sepals 2, 4-5 mm long by 2-3 mm wide and inner sepals 3, 5-6 mm long by 2-3 mm wide, surface glabrous, enlarge in fruit;corolla funnel, bright yellow, 1.5-2 cm long, corolla limb slightly 5-lobed, pubescent, midpetaline bands 5, glabrous, purplish; stamens 5, anther straight and then spiral twisted, glabrous, filament unequal, dorsifixed below, sticky at tube, slender, thin but broader and hairy at the basal pada tube, white; stigma macrostylous, biglobular, white, smooth and glabrous, style terete, glabrous, ovary superior, glabrous. Fruit a capsule, valve, dehiscent, ovoid, 5-6 mm long. Seeds 4, late elliptic, 2 mm long, surface pillose at margin and hylum, brown or sometimes black.
Distribution: Lampung and Bangka Belitung.
Habitat and Ecology. Found on roadside, open grassland. Altitude 20-1100 m above sea level.
Note. Widespread in China (Guangdong, Guangxi, Taiwan,Yunnan), India, Myanmar, Laos, Thailand, Vietnam. Malesia: Malaysia, Indonesia (Java, Borneo, Lesser Sunda (Sumbawa), Mollucas, Celebes), Philippines and Australia.
Figure 18 Distribution of Merremia hirta in Sumatra
7. Merremia peltata (L.) Merr.
Merremia peltata (L.) Merr. Interpr. Rumph. Herb. Amb. (1917) 441; id., Enum. Philipp. Fl. Pl. III (1923) 362; Ooststr., Blumea III (1939) 352; Ooststr., FM. I.4.4 (1953) 452.─ Ipomoea peltata Choisy, Mȇm. Soc. Phys. Genȇve VI
(1833) 452.─ Operculina peltata Hall. f. in Engl., Bot. Jahrb. XVI (1893) 549; Backer and Bakhuizen f., FJ. II (1965) 489.─ Convolvulus peltatus Linnȇ, Sp. Pl. (1753) 1194 Type: Indonesia, Java, Blume, C.L. 1505 (Holotype, L!) (by photo).
.─ Merremia nymphaeifolia (Bl.) Hall. f. in Engl., Bot. Jahrb. XLIX.─ Ipomoea nymphaefolia Bl. Bijdr. (1825) 719, non Griseb Type: not seen.
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slender, thin but broader and hairy at the basal at tube, yellow or white, 1.5-1.7 cm; pistill 1, stigma macrostylous, biglobular, white, smooth and glabrous, 2.1 cm long, style terete, glabrous, ovary superior, glabrous. Fruit a capsule, valve, dehiscent, ovoid to conical, 1.6 cm long. Seeds 4, ovoid to conical, 8-9 mm long, surface pillose, black.
Distribution.
Widespread from tropical eastern Africa, Madagascar, Indian Ocean
and Andaman Islands, Thailand, Malaysia, Singapore, Philippines, New Guinea, New Caledonia, Australia (Queensland), Pacific Islands. Naturalized in Indonesia: Sumatra (Nangroe Aceh Darussalam, North Sumatra, West Sumatra, Bengkulu and Lampung), Java, Borneo, Lesser sunda (Flores), Mollucas. In Indonesia (Lampung) it is a invasive alien species.
Habitat and Ecology. Grows in secondary forest or river side at altitude 20-600 above sea level. Lunto/sijunung about 6 km east from sijunjung alt 200-300 m, 21-23 Apr. 2000 no collector 24 (ANDA). LAMPUNG: Sibesi island, 28 Apr. 1921 Dr van leeuwen-reijnvaan 5346 (BO), Bukit Barisan Selatan National Park alt 200-300 m, 29 Maret 2011 Hafni Rahmadani 01 (BIOT); ibid 30 Maret 2011 Hafni Rahmadani
02 (BIOT); ibid 30 Maret 2011 Hafni Rahmadani 03 (BIOT); ibid 31 Maret 2011
Hafni Rahmadani 04 (BIOT), ibid 1 April 2011 Hafni Rahmadani 05 (BIOT). NIAS: 23 July 1991 R Romer Xvii (BO). RIAU ARCHIPELAGO: Anambas Island, Siantan, Terempa alt 75 m, 31 March 1928 Van Steenis 781 (BO).
Figure 19 Merremia peltata (L.) Merr. A. Habit liana; B. blade; C. Flower; D. Corolla limb; F. Pistill; G. Stamen; H. Outer sepal fruit; I. outer sepal flower; J. Inner sepal flower; K. Fruit capsule; L. Seeds.
6 cm
3 cm
2.1 cm cm
1.7 cm
cm 2.4 cm
1.9 cm
1.6 cm 2.1 cm
8 mm
A
B
C D F
G
I J
H
K
L
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Figure 20 Distribution of Merremia peltata in Sumatra
8. Merremia quinquefolia (L.) Hallier f.
Merremia quinquefolia (L.) Hall. f. in Eng., Bot. Jahrb. XVI (1893) 552.─
Ipomoea quinquefolia L., spec. Pl. (1753) 162. Ooststr., Blumea III (1939) 324; Ooststr., FM I.4.4 (1953) 446; Backer and Bakhuizen f., FJ. II (1965) 489 Type:
“Convolvulus quinquefolius glaber Americanus" Plukenet, Phytographia plate 167, fig 6. 1692 (Lectotype designed by D.F. Austin) (not seen)
