A TAXONOMIC REVISION
OF THE GENUS
POPOWIA
Endlicher
(ANNONACEAE) IN MALESIA
By
SOETJIPTO MOELJONO
POST GRADUATE SCHOOL
BOGOR AGRICULTURAL UNIVERSITY
B O G O R
STATEMENT OF RESEARCH ORIGINALITY
AND INFORMATION SOURCE
This is to verify that my dissertation entitled “A Revision of the of Genus
Popowia Endlicher (Annonaceae) in Malesia” is my own original work under
supervision committee and has never been submitted in any form to any institution before. All of the incorporated data and information source or citations from other litterature which have been published or not are valid and stated clearly in the text and listed in the references.
Bogor, April 2009
ABSTRACT
Soetjipto Moeljono, Edi Guhardja, J.P. Mogea, Soedarsono Riswan and P.J.A. Kessler, 2009. A Revision of The of Genus Popowia Endlicher (Annonaceae) in Malesia. Description of the genus followed by key identification to the species, description of the species, information regarding the habitat, geographical distribution, anatomy, and on morphological notes is presented. There are 24 species and one variety of Popowia are found in Malesia i.e. Popowia pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. fusca King, P. velutina King, P. perakensis King, P. tomentosa Maingay ex
Hook.f., P. tomentosa var. crinita J. Sinclair, P. bancana Scheffer, P. hirta
Miquel, P. cuspidata Miquel, P. cordata Moeljono, P. odoardi Diels, P.
lanceolata Merrill, P. beccariiScheffer, P. clavataDiels, P. cyanocarpaLauterb.
and K. Schum., P. novoguineensisMiquel, P. papuana Scheffer, P. schefferiana
Diels, P. pachypetalaDiels,P. polytrica Diels, P. platyphyllaDiels, P. parvifolia
Scheffer, P. nooteboomii Moeljono, andP. hentyii Moeljono. One species from Java (P. cordata Moeljono) and two species (P. nooteboomii Moeljono, and P.
hentyii Moeljono) from Papuasia are described and illustrated for the first time.
Thirteen species were found in Papuasia. No species is found in Lesser Sunda Islands. Orophea minahassae Boerlage was placed as a synonym of Popowia pisocarpa(Blume) Endl.
ABSTRAK
Soetjipto Moeljono, Edi Guhardja, J.P. Mogea, Soedarsono Riswan dan P.J.A. Kessler, 2009. Revisi Marga Popowia Endl. (Annonaceae) di Malesia. Pertelaan marga dan jenis disertai dengan kunci identifikasi jenis, keterangan mengenai habitat, persebaran geografi, anatomi dan keterangan morfologi disajikan. Di Malesia ada 24 jenis Popowia dan satu varietas: Popowia
pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. fusca King, P.
velutina King, P. perakensis King, P. tomentosa Maingay ex Hook.f., P. tomentosa var. crinita J. Sinclair, P. bancana Scheffer, P. hirta Miquel, P.
cuspidata Miquel, P. cordata Moeljono, P. odoardi Diels, P. lanceolata Merrill, P. beccarii Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P.
novoguineensis Miquel, P. papuana Scheffer, P. schefferiana Diels, P. pachypetala Diels,P. polytrica Diels,P. platyphyllaDiels,P. parvifolia Scheffer,
P. nooteboomii Moeljono, dan P. hentyii Moeljono. Satu jenis (P. cordata
Moeljono) dari Jawa dan dua jenis (P. nooteboomii Moeljono dan P. hentyii Moeljono) dari Papua dipertelakan dan dilengkapi dengan gambar untuk yang pertama kali. Tiga belas jenis ditemukan di Papua. Tidak satu jenispun dijumpai di kepulauan Sunda kecil. Orophea minahassae Boerlage sebagai sinonim
Popowia pisocarpa(Blume) Endl.
SUMMARY
SOETJIPTO MOELJONO. A Revision of the Genus Popowia Endl. (Annonaceae) in Malesia; advised by: Edi Guhardja, J. P. Mogea, Soedarsono Riswan and P.J.A. Kessler.
This dissertation presents a taxonomic revision ofPopowia Endl. in Malesia. It is based on an examination of 574 collection numbers which are preserved in the Herbarium Bogoriense, Bogor ( BO); Herbarium Manokwariense, The State University of Papua, Manokwari (MAN); and National Herbarium of the Netherlands, Leiden Branch( L). The Leiden collections were loaned to Bogor for this project.
Method used follows the normal practice in herbarium taxonomy. Morphological characters, which have been used as the diagnostic taxonomic features, are used in evaluating the species of Popowia Endl. Vegetative morphology: leaf shape, leaf base, leaf texture, number of main nerves, and the petiole length; and reproductive morphology: pedicel length, sepal shape, petal shape, and fruit size, are the main characters used for determinity the species in the genus.
There are twenty four species and one variety ofPopowia Endl. in Malesia, but absent in the Lesser Sunda Islands (Nusa Tenggara). In Malay Peninsula seven species have been identified including: Popowia pisocarpa (Blume) Endl., P.
pauciflora Maingay ex Hook. f., P. fusca King, P. velutina King, P. perakensis
King, P. hirta Miquel, andP. tomentosa Maingay ex Hook. f.; and one variety:P.
tomentosa Maingay ex Hook.f. var. crinita Sinclair. In Sumatra seven species:P.
bancana Scheffer, P. fusca King, P. hirta Miquel,P. pisocarpa (Blume) Endl.,P. velutina King, and P. tomentosa Maingay ex Hook. f.; and one variety: P. tomentosa Maingay ex Hook.f. var. crinita Sinclair. In Java three species: P.
pisocarpa (Blume) Endl., P. cordata Moeljono and P. cuspidata Miquel. In
Borneo six species: P. bancana Scheffer, P. pisocarpa (Blume) Endl., P. fusca
species: P. pisocarpa (Blume) Endl. and P. lanceolata Merrill. In the Moluccas three species: P. pisocarpa (Blume) Endl., and P. schefferiana Diels Thirteen species are known from New Guinea: Popowia pisocarpa (Blume) Endl., P. beccarii Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensis Miquel, P. phacypetala Diels, P. papuana Scheffer, P. parvifolia
Scheffer, P. platyphylla Diels, P. schefferiana Diels, P. polytrica Diels, P. hentyii Moeljono,P. nooteboomii Moeljono.
The vegetative and generative characters upon which the taxonomic revision are based will be discussed. First, this includes detail studies on the leaf surface using the microscope and detail measurement of the leaf morphology and anatomy. The flowers were selected and studied in detail. Botanical descriptions are included of all species with notes on their habitats and a list of their vernacular names. An identification key is provided. All examined collections are listed in Appendix 1. The collections are from Malesia.
COPYRIGHT © 2009,
BOGOR AGRICULTURAL UNIVERSITY
ALL RIGHT RESERVED
1. It is prohibited to cite all or part of this dissertation without referring to and mentioning the source.
a. Citation only permitted for the sake of education, research scientific writing, report writing or reviewing scientific problems. b. Citation does not inflict the name and honour of Bogor
Agricultural University.
A TAXONOMIC REVISION
OF THE GENUS
POPOWIA
Endlicher
(ANNONACEAE) IN MALESIA
By
SOETJIPTO MOELJONO
A dissertation submitted to fulfill one of the requirements for the Doctor degree at the Study Program of Biology,
Graduate School, Bogor Agriculture University
DEPARTMENT OF BIOLOGY
POST GRADUATE SCHOOL
THE BOGOR AGRICULTURAL UNIVERSITY
B O G O R
External Reviewer in Closed Examination : Dr. Ir. Tatik Chikmawati, M.Si.
External Reviewer in Open Examination : 1. Kuswata Kartawinata, Ph.D.
T i t t l e : A Taxonomic Revision of the Genus Popowia Endlicher (Annonaceae) in Malesia.
N a m e : Soetjipto Moeljono Reg. No. : 975073
Study Program : Biology
Approved by :
1. The Advisory Committee
( Prof. Dr. Ir. Edi Guhardja, M.Sc.) Chairman
(Prof. Dr. J. P. Mogea) (Soedarsono Riswan, M.Sc., Ph.D.) (Dr. P.J.A. Kessler) Member Member Member
2. Head of Study Program 3. Dean of the Graduate School
(Dr. Ir. Dedi Duryadi S., DEA) (Prof. Dr. Ir. Khairil Anwar Notodiputro, MS.)
ACKNOWLEDGEMENTS
During the course of this study there were many people who helped me in many ways; to all those who are rendered thanks which individually I give my sincere appreciation for their helps. First I wish to acknowledge support of finance accepted from DEPDIKNAS - DIKTI ( BPPS), which enabled me to study in Bogor Agriculture University.
To my advisors, Professor. Dr. Ir. Edi Guhardja, Msc., Dr. Soedarsono Riswan APU, Professor. Dr. J.P. Mogea, and Dr. P.J.A. Kessler, I am grateful for their generosity in allowing me to carry out this study. To them I also indebted for their consistent encouragement, guidance, positive attitude and the various kinds of support.
I must express my sincere thank to Head of the Herbarium Bogoriense, the Development Center for the Biology - Institute of Science of Indonesia in Bogor, not only for providing space available for my use during this study, but also for generously placing the facilities and services of their staff at my disposal.
I would like to thank to Director of the Rijksherbarium, Professor. Dr. Pieter Baas, who kindly permitted specimens to be sent on loan to Bogor for my research project.
CURRICULUM VITAE
The author ( Soetjipto Moeljono) was born on 23 th of December 1964 in Surabaya (East Java Province). He is the seventh child from his parents Mr Dasoeki and Mrs Moerah who were passed away. He entered Elementary school in 1972 at Sekolah Dasar Negeri I, Biak and finished in 1977. He started Junior high school at Sekolah Menengah Pertama Negeri I, Biak in 1977/1978, and continued his study in Senior High school at Sekolah Menengah Atas YPK, Biak in 1981/1982.
After graduated from Senior High school in 1984, he was accepted as a student in the Faculty of Agriculture, Cenderawasih University, Manokwari, Irian Jaya, with majoring in Forest Management under Department of Forestry. He finished his study (S-1) in the same university in 1989. In the year 1994, he entered Post Graduate Program in Bogor Agriculture University leading for Master of Science degree in Biology with majoring in Plant Taxonomy. He received his Master of Science degree from Bogor Agriculture University in January 11, 1997. In 1997, he entered the Post Graduate School for Doctor of Philosophy degree in Biology with majoring in Plant Taxonomy in the same university. He received a scholarship from Government of Indonesia through a Ministry of National Education.
In 1990, he started his work as a lecturer in the same university where he serves up to present. His main responsibility is as researcher and curator in Herbarium Manokwari, The State University of Papua, Manokwari.
CONTENTS
Pages
STATEMENT OF RESEARCH ORIGINALITY AND
INFORMATION SOURCE ... i
ABSTRACT ... ii
ABSTRAK ... iii
SUMMARY ... iv
COPYRIGHT © 2009, BOGOR AGRICULTURAL UNIVERSITY ALL RIGHT RESERVED ... vi
ACKNOWLEDGEMENTS ... x
CURRICULUM VITAE ... xi
CONTENTS ... xiii
LIST OF TABLES... xv
LIST OF FIGURES ... xvi
LIST OF APPENDICES ... xviii
I. INTRODUCTION ... 1
II. LITERATURE REVIEW ... 3
2.1. Malesia Region... 3
2.2. History of Classification ofPopowia Endl. ... 5
2.3. History of Research onPopowia Endl.in Malesia ... 10
2.4. Distribution and Ecology ... 12
III. RESEARCH METHODOLOGY... 13
3.1. Materials ... 13
3.2. Methods ... 13
IV. RESULT AND DISCUSSION ... 15
4.1. Vegetative Morphology... 15
4.2. Generative Morphology ... 18
4.3. Leaf Anatomy ... 20
4.5. Distribution and Ecology ofPopowia Endl. ……… 29
4.6. Taxonomy ... 34
Key Identification to the Species ofPopowia Endl. in Malesia ... 35
1.Popowia pisocarpa (Blume) Endl. ... 37
2.P. pauciflora Maingay ex Hook.f. ... 44
3.P. fusca King ... 47
4.P. velutinaKing... 51
5.P. tomentosa Maingay ex Hook.f. ... 54
6.P. tomentosa var.crinita Sinclair ... 58
7.P. hirta Miquel ... 60
8.P. cyanocarpa Lauterb. and K. Schum. ... 63
9.P. novoguineensis Miquel ... 65
10.P. clavata Diels ... 67
11.P. schefferianaDiels ... 69
12.P. polytricha Diels ... 71
13.P. platyphylla Diels . ... 73
14.P. nooteboomii Moeljono ... 75
15.P. bancana Scheffer. ... 78
16.P. hentyii Moeljono . ... 81
17.P. perakensis King ... 84
18.P. beccarii Scheffer ... 87
19.P. cordataMoeljono ... 89
20.P.odoardii Diels ... 92
21.P. cuspidata Miquel ... 95
22.P. pachypetala Diels ... 97
23.P. lanceolata Merrill ... 99
24.P. parvifolia Scheffer ... 101
25.P. papuana Diels ... 103
V. CONCLUSION ... 105
REFERENCES ... 107
LIST OF TABLES
Pages
Characters and Character States Used in the Phylogenetic Analysis
of the Malesian Taxa ofPopowia Endl. ………..……….. 25
Data Matrix of Morphological Characters of Popowia Endl.
used for Phylogenetic Analysis ……… 27
LIST OF FIGURES
Pages
1. Malesia and its divisions as described by Van Steenis ... 5
2. Wallace’s line ... 5
3. Tribe Mitrephoreae ... 9
4. Place where measurements of plant parts were taken for the species description . ... 14
5. Portion of a cross-section of the leaf ofPopowia Endl... 21
6. Epidermal cells and crystals on the lower leaf surface ... 23
7. Epidermal cells and stomata on the lower leaf surface. ... 24
8. Cladogram ofPopowia Endl. In Malesia ... 28
9. Density map ofPopowia Endl. in Malesia ... 33
10. Distribution ofPopowia pisocarpa (Blume) Endl. ... 42
11.Popowia pisocarpa (Blume) Endl... 43
12. Distribution ofP. paucifloraMaingay ex Hook.fi. ... 45
13.P. paucifloraMaingay ex Hook.fi. ... 46
14. Distribution ofP. fusca King... 49
15.P. fusca King ... 50
16. Distribution ofP. velutinaKing ... 52
17.P. velutinaKing ... 53
18. Distribution ofP. tomentosa Maingay ex Hook.fi. ... 56
19.P. tomentosa Maingay ex Hook.fi. ... 57
20. Distribution ofP. tomentosa var.crinita J. Sinclair ... 59
21. Distribution of P. hirtaMiquel ... 62
22. Distribution ofP. P. cyanocarpaLauterb. and K. Schum. ... 64
23. Distribution ofP novoguineensisMiquel... 66
24. Distribution ofP. clavataDiels ... 68
25. Distribution ofP. schefferiana Diels ... 70
26. Distribution ofP. P.. polytrica Diels ... 72
27. Distribution of P. platyphyllaDiels ... 74
29.P. nooteboomii Moeljono ... 77
30. Distribution of P. bancana Scheffer ... 80
31. Distribution ofP. hentyii Moeljono ... 82
32.P. hentyii Moeljono ... 83
33. Distribution of P. perakensisKing ... 85
34.P. perakensisKing ... 86
35. Distribution ofP. beccariiScheffer ... 88
36. Distribution ofP. cordataMoeljono ... 90
37.P.cordata Moeljono ... 91
38. Distribution ofP. odoardii Diels ... 94
39. Distribution ofP. cuspidataMiquel ... 96
40. Distribution ofP. pachypetalaDiels... 98
41. Distribution ofP. lanceolataMerrill ... 100
42. Distribution ofP. parvifoliaScheffer ... 102
LIST OF APPENDICES
Pages
I. INTRODUCTION
The Annonaceae is a pantropical family with about 130 genera and 2.300 species. The family is known in Indonesia under the name ”Sirsak-sirsakan”. It is confined mostly to humid tropical lowland forests. They are abundant common in Asia, Africa, Central and South America and few in Australia. Two related genera Asimina and Deeringothamnus occur in warm temperate North-East America. In Malesia Annonaceae is abundant in the low altitude and becomes scarce at about above 600 meters elevation (Sinclair 1955 and Kessler 1993). 47 out of 130 genera of the family occur in Malesia, including genusPopowia.
The genus Popowia was firstly published by Stephano Endlicher in the
Genera Plantarum secundum Ordines Naturales page 831 in the year of 1839. In
1842 in Walp. Repert. 1, page 74, he published Popowia pisocarpa to revise
Guatteria pisocarpa Blume in 1825, which is written in Bijdragen tot de Flora van Nederlandsch Indie page 21 and Sprengel also published this name in
Systema Vegetabilium Vol. IV Part II, page 214.28 in 1827 as a basionym. Other species of Popowia were published by Miquel (1865) namely, P. affinis (Sumatra),P. cuspidata (Java), P hirta (Sumatra),P. novoguineensis (New
Guinea) and P. rufula (Borneo). In 1855, Hooker and Thomson published
Popowia ramosissima (Malay Peninsula). Later in 1872, they published again
seven species of Popowia namely P. beddomeana, P. foetida, P. helferi, P.
nervifolia, P. pauciflora, P. ramosissima, and P. tomentosa. In 1885, Scheffer
published four species of Popowia namely, P. bancana (Sumatra), P. beccarii
(New Guinea), P. papuana (New Guinea) and P. parvifolia (New Guinea). In 1893, King published twelve species from Malay Peninsula namely,P. foetida,P.
fusca, P. helferi, P. hookeri, P. kurzii, P. nervifolia, P. nitida, P. pauciflora, P.
perakensis, P. ramosissima, P. tomentosa andP. velutina. In 1901, Lauterback &
K. Schumman describedP. cyanocarpa (New Guinea). In 1908, Merrill described
P. polyandra (Philippine) and in 1921, he described P. lanceolata (Philippine).
New Guinea. In 1929, he describedPopowia polytricha (New Guinea). Finally, in 1931 he describedP. odoardi (Borneo). In 1913, Ridley published five species of
Popowia namely,P hirta, P. novoguineensis P. pisocarpa, P. ramosissima andP. tomentosa. In 1922, he published ten species of Popowia namely, P. foetida, P. fusca, P. nervifolia, P. nervosa, P. pauciflora, P. perakensis, P. pumila, P. ramosissima, P. tomentosa and P. velutina. Later in 1926, he published againP.
rufescent. Finally, in 1955, Sinclair published six species and one variety of
Popowia from Malay Peninsula namely, P. fusca, P. pauciflora,P. perakensis, P. pisocarpa, P. tomentosa, P. tomentosa var. crinita and P. velutina, and removed
Popowia affinis,P. ramosissima, P. rufescens and P. rufula to P.pisocarpa andP.
foetida to Neouvaria foetida, Popowia nervifolia to Trivalvaria macrophylla,
Popowia nervosa toTrivalvaria nervosa, Popowia pumila toTrivalvaria pumila.
II. LITERATURES REVIEW
2.1. Malesia Region
Malesia is the floristic region that comprises the following political entities: Malaysia, Singapore, Indonesia, East Timor (previously Portuguese Timor), the Philippines, Brunei Darussalam, and Papua New Guinea. The concept and demarcation of this floristic region was firstly proposed by Van Steenis in the late 1940’s, under the name “Malaysia” (Van Steenis 1950). To avoid confusion with the State of Malaysia (granted independence from the British in 1953), the name Malesia was introduced to replace the previous name (Whitmore 1975).
In the original demarcation Van Steenis (1950) did not include the islands surrounding the mainland of New Guinea, namely the Bismarck, Solomon, and New Hebrides Archipelagos (Figure 1). He explained that these islands were excluded mainly because knowledge of their floras was limited. However, he suggested that the floras on these islands were mainly derived from Malesia. Furthermore, he noted that the demarcation of Malesia without these groups of islands was artificial, thus the exclusion of the islands was basically for practical reasons only (Van Steenis 1950). Van Steenis concluded that the Indo-Asiatic flora ends rather abruptly in the mainland of New Guinea. The studies of Van Balgooy (1971) support the suggestion by Van Steenis that the floras of the Bismarck, Solomon and New Hebrides Archipelagos indeed share many similarities with the Malesian flora rather than with the Australian flora.
consisting of the Philippines, Celebes, Moluccas and the Lesser Sunda Islands (Nusa Tenggara). Later Dickerson (1925; 1928) proposed the name Wallacea for this “region in between”.
Combining Wallace’s line, at least between Borneo and Celebes, with his own studies on plant generic distribution in the Malay Archipelago, Van Steenis (1950; 1954; 1979; Van Balgooy 1987) proposed three areas, known to him as “demarcation knots”, which are not crossed by many genera in either direction. He thus divided Malesia into three divisions, West Malesia (including the Malay Peninsula, Sumatra, Borneo, and the Philippines), East Malesia (including Celebes, Moluccas, and New Guinea with its nearby islands), and South Malesia (including Java and the Lesser Sunda Islands).
Fig. 1. Malesia and its divisions as described by Van Steenis (1950)
Fig. 2. Wallace’s line.
2.2. History of Classification ofPopowia Endl.
The Annonaceae is a pantropical family with about 128 genera and ca. 2.300 spp. (Sinclair 1955, Keng 1978 and Kessler 1993). All members of the Annonaceae are trees, shrubs, or climbers. Among the Asiatic and African members of the family, the climbing habit is frequent. According to Sinclair (1955) the Annonaceae are readly distinguished from other families by twigs alone, which often have lozenge-shaped striations resembling a sort
B
A
of trellis work and the fruit is of considerable importance in separation of genera but not species.
Although the Annonaceae as a family are well characterized, subdivision has always been problematic. This is reflected by the different proposals that have been made in recent systematic treatments ( Sinclair 1955; Fries 1959; Hutchinson 1964; Walker 1971; and Koek-Noorman et al. 1990). These classifications are far from being comparable with each other, although all contain valuable elements.
All systematists, maintain a fundamental subdivision of the family into Annonoideae and Monodoroideae. This is based on the peculiar gynoecium structure of Monodora and Isolana (paracarpous or monocarps with laminal placentation. The classification of the group of Annonaceae according to Sinclair 1955 is divided into 2 subfamily and 6 tribes as follow:
KEY TO THE GROUPS OF ANNONACEAE
A. Carpels free or if united forming a many-celled syncarp; stigma erect. Subfamily (1) ANNONOIDEAE
B. Carpel free or if slightly so, then always quite free in fruit.
C. Sepal imbricate or valvate. Petals 2-seriate, one or both series Imbricate in bud. Stamens many. Closely packed, their anther cells concealed at the top by the connective. Indumentum of leaves often stellate or lepidote, seeds many to few.
Tribe (1) UVARIAE
C. Sepals valvate. Petals al valvate either in two distinct series or rarely in one series, usually 6, rarely 4, more rarely 3, in the latter case either the inner or the outer series missing.
concave at the base only, the inner flat, terete, or filiform, or concave or not at the base. Stamens many, connectives flat-topped or convex, rarely apiculate, the anther cells concealed by the connective.
Tribe (2) UNONEAE
D. Petals sub-equal or more often the inner shorter, the outer orbicular,ovate, ooblong but usually lanceolate, spreading, often triquetrous, the inner nearly always triquetrous, not often spreading but remaining erect and touching each other, smaller, than the outer. Stamen concealed by the connectives.
Tribe (3) XYLOPIEAE
D. Petals, one or both sets but usually the inner saccate or at least broad at base, the outer small and like the sepals, free, the inner larger, free or less often tending to adhere to some extent by the margin (both sets sub- equal in Alphonsea). Stamens few, loosely imbricate, anther cells not concealed by the flat-topped, rounded or pointed connectives.
Tribe (4) MILIUSEAE
D. Petals, inner larger or smaller than the outer, less often sub-equal, usually dissimiliar, concave, connivent, arching over the sexual organs and forming a dome, if free their edges at first united for a short time, clawed, the claw often long and narrow or vestigial or absent. Stamens many, flat-topped or convex, the connectives hiding the anther cells.
Tribe (5) MITREPHOREAE
B. Carpels united into a fleshy mass, especially in fruit.
A. Carpels united into a 1- celled ovary with parietal placentas, stigmas radiating ( No examples in Malesia )
Subfamily (2) MONODOROIDEAE
TRIBE MITREPHOREAE
According to Sinclair (1955) that tribe Mitrephoreae came from the Unoneae. They are characterised by the development of a mitriform corolla. Fig.3. shows the development and differentiation of such a corolla. Pseudu-varia seem to be the most advance where the inner petals are well
differentiated into claw and blade and the blades remain united to form a dome. Sepals valvate. Petals valvate, inner larger or small than outer, less often sub-equal, usually dissimilar, concave, connivent, arching over the sexual organs and forming a dome, if free, their edges at first united for a short time, clawed, the claw often long and narrow or vestigial or absent.
Stamens 20 - 30, flat-topped or convex (uvaroid).
KEY TO THE GENERA OF THE TRIBE MITREPHOREAE
a. Flowers axillary
b. Flowers unisexual ... Pseuduvaria b. Flowers hermaphrodite
c. Petals sub-equal; stellate hairs present on leaves and young twigs
... Neo-Uvaria c. Inner petals much small than outer; no stellate hairs ... Goniothalamus a. Flower extra-axillary
d. Inner petals shorter than outer or equal to them
e. Climbers. Inner petals not clawed ... Oxymitra e. Trees. Inner petals with long narrow claws 1 mm broad or less
... Mitrephora d. Inner petals longer than outer with a vestigial claw, the edges at first
Fig. 3. Tribe Mitrephoreae. Tree based on development and differentiation of corolla (Sinclair 1955).
Outer petals slightly clawed or not. Inner larger, with very narrow claws, the blades united forming dome
Outer petals slightly or not clawed, spreading. Inner slightly shorter, with very narrow claws, erect, edges adhering, later free.
Outer petals slightly or not spreading. Inner smaller, clawed, erect, connivent, edges adhering
Outer petals spreading. Inner much smaller, not clowed, connivent, erect, their edges adhering.
Outer petals spreading. Inner longer with incurved apices and rudimentary claw, some-what connivent and erect, united by edges at first, later free.
The genus Popowia name can be found in Genera Plantarum
secundum Ordines Naturales page 831 in 1839 by Stephano Endlicher,
which he named asPopowia pisocarpa. It was based on the nameGuatteria pisocarpa Blume inBijdragen tot de Flora van Nederlandsch Indie page 21
in 1825 and in Systema Vegetabilium Vol. IV Part II, page 214.28 is basonym. Guatteria pisocarpa transferred to Popowia pisocarpa in Walp. Repert. 1 (1842) 74 by Endlicher and Bocagea pisocarpa Bl. in Flora of Java. Anon (1830) p.90, Tab. 45 is synonim ofP. pisocarpa. The reason for these changes will be found in the systematic section i.e. position of inflorescence or single flower leaf-opposite, aestivation of corolla valvate, inner petals longer than outer ones, shape of petals clawed, coherent of petals with reproductive organs, numbers of carpels 2- 20, numbers of ovule per capel 2 – 5 and placentation of ovule lateral.
2.3. History of Research on Popowia Endl. in Malesia
The first species currently placed in the genus Popowia wasPopowia pisocarpa which described by Ejusd asGuatteria pisocarpa from G. Parang,
Tjiandjoer (Cianjur), West Java. Guatteria pisocarpa transferred to
Popowia pisocarpa in Walp. Repert. 1 (1842) 74 by Endlicher; Bocagea pisocarpa Bl. is synonim of P. pisocarpa. Other species of Popowia were
published by Hooker and Thomson (1855) published Popowia ramosissima
(Malay Peninsula) in Flora Indica Vol.1 page 105. Miquel (1865) published in Annales Musei Botanici Lugduno Batavi Volume 2, page 20 – 22 namely, P. affinis and P hirta (Sumatra), P. rufula (Borneo), P. cuspidate (Java) and P. novoguineensis (New Guinea). In 1872 Hooker and Thomson published again seven species from Malay Peninsula namely, P.
ramosissima, P. beddomeana, P. herferi, P. pauciflora, P. foetida, P. nervifolia andP. tomentosa inThe Flora of British India Vol. 1. page 68-70.
In 1885, Scheffer published four species of Popowia namely, P. bancana (Bangka), P. papuana, P. beccarii and P. parvifolia (New Guinea) in
Annales du Jardin Botanique de Buitenzorg Vol. 2, page 15 - 17. In 1893,
P. ramosissima, P. nitida, P. herferi, P. foetida,P. perakensis, P. fusca, P.
velutina, P. tomentosa P. nervifolia, P. kurzii and P. hookeri in Journal Asiatic Society Bengalis Part 2 Natural History Vol. 61, page 92 -98. In
1901, Lauterback & K. Schumman describedP. cyanocarpa (New Guinea) in Nachr Flora Deutsch Sudsee page 317. In 1908, Merrill described P. polyandra (Philippines) in The Philippine Journal of Science Vol. 3, page
224. In 1921, he described P. lanceolata (Philippines) in The Philippine Journal of Science Vol 17, page 252. Later in 1923, he transferred P. polyandra from Philippines to P. pisocarpa in An Enumeration of Philippine Flowering Plants Vol. 2, page 164. In 1912, Diels describedP.
schefferiana andP. pachypetala (New Guinea) in Botanische Jahrbucher fur
Systematik, Pflan-zengeschichte und Pflanzengeographie page135 -138. In
1915, he again described P. clavata and P. platyphylla (New Guinea) in
Botanische Jahrbucher fur Systematik, Pflanzengeschichte und Pflanzengeographie page181-182. In 1929, he describedPopowia polytricha
(New Guinea) inJournal of The Arnold Arboretum. Vol. X : 76. In 1931 he described P. odoardi (Borneo) in Notizblatt des Koniglich Botanischen
Gartens und Museum zu Berlin Vol. XI, page 82. In 1913, Ridley published
five species of Popowia (P. ramosissima, Popowia pisocarpa, P hirta, P. tomentosa and P. novoguineensis) inSarawak Musei Journal Vol.1, Part 3,
page 87. In 1922, he published ten species of Popowia from Malay Peninsula namely, P. ramosissima, P. pauciflora, P. foetida, P. perakensis, P. fusca, P. velutina, P. nervifolia, P. nervosa, P. pumila and P. tomentosa
in The flora of the Malay Peninsula Vol.1, page 74 - 79. Later in 1926, he published again P. rufescens (Siberut) in Royal Botanic Garden Kew Bullentin of Miscellaneous Information Vol. 14, page 59. Finally, in 1955, Sinclair published six species and one variety of Popowia from Malay Peninsula namely P. pisocarpa, P. pauciflora, P. fusca, P. velutina, P.
perakensis, P. tomentosa and P. tomentosa var. crinita; and Popowia ramosissima, P. rufula, P. affinis and P. rufescens transferred to P.
macrophylla, P. nervosa to Trivalvaria nervosa, P. pumila to Trivalvaria
pumila in The Gardens Bulletin Singapore Vol. XIV, Part 2, page 149 – 516.
2.4. Distribution and Ecology
The Annonaceae are confined mostly to moist tropical lowland forest. They are more plentiful in the Old World than in the New. They are not found in Europe. Only the two related genera Asimina and Deeringo-thamnus occur in warm temperate eastern North America. Asimina triloba
III. RESEARCH METHODOLOGY
3.1. Materials
This study was based mainly on herbarium materials from herbaria listed below. Five hundred and seventy four specimens were examined during the course of this study. Herbaria which I personally visited, and from which most of the material studied came, are indicated with an asterisk (*).
BO : Herbarium Bogoriense, Bogor. *
MAN : Herbarium Manokwariense, The State University of Papua,
Manokwari, Papua*
L : Rijksherbarium, Leiden, The Netherland (on loan to Bogor).
3.2. Methods
In general the methods in this study follow the normal practice of herbarium taxonomy (Rifai, 1976 and de Vogel, 1987).
Lauterb. and K. Schum., P. novoguineensisMiquel, P. papuanaScheffer, P.
schefferiana Diels, P. pachypetala Diels, P. polytrica Diels, P. platyphylla
Diels, P. parvifolia Scheffer P. cordata Moeljono, and P. nooteboomii
[image:33.612.94.496.175.690.2]Moeljono, P. hentyii Moeljono) were not included because of incomplete material available.
Fig. 4. Place where measurements of plant parts were taken for the species description (Popowia hentyii Moeljono) p.82.
A. Leaf, lower surface. B. Flower bud. C. Outer and inner petals D. Flower, petals removed. E. Stamen F. Gynoecium. G. Fruit
E
A B
C
D
IV. RESULT AND DISCUSSION
This present research was based on herbarium materials. Based on the available specimens, twenty four species and one varieties ofPopowia Endl. are recognized in Malesia. There are Popowia pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. fusca King, P. velutina King, P.
perakensisKing, P. tomentosa Maingay ex Hook.f., P. bancana Scheffer, P.
hirtaMiquel, P. cuspidataMiquel, P. odoardi Diels, P. lanceolataMerrill, P.
beccariiScheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensis Miquel, P. papuana Scheffer, P. schefferiana Diels, P.
pachypetala Diels, P. polytrica Diels, P. platyphylla Diels, P. parvifolia
Scheffer, P. cordata Moeljono, P. nooteboomii Moeljono, P. hentyii Moeljono, andP. tomentosa var.crinita J. Sinclair.
The general morphological characters which have been used as diagnostic taxonomic feature were found usefull in evaluating the relationship of species ofPopowiaEndl. The vegetative morphology i.e leaf shape, leaf base, leaf texture, number of main nerves, and petiole length; the floristic morphology i.e pedicel length, sepal shape, petal shape, and fruits size; are the main characters used for determinating the species in the genus
Popowia Endl. (Blume, 1830; Diels, 1912; Sinclair, 1955 and Kessler,
1993).
4.1. Vegetative Morphology 4.1.1. Habit.
Popowia beccariiScheffer,P. cuspidataMiquel, P. pachypetala
Diels, P. cordata Moeljono, and P. nooteboomii Moeljono are shrub.
Popowia pisocarpa (Blume) Endl., P. fusca King, P. perakensis
King, P. hirta Miquel, P. odoardi Diels, P. clavata Diels, P. cuspidata Miquel P. platyphylla Diels, P. Schefferiana Diels, P. novoguineensis Miquel, and P. papuana Scheffer are small tree. P.
pauciflora Maingay ex Hook.f., P. velutina King, P. tomentosa
Maingay ex Hook.f., P. tomentosa var. crinita J. Sinclair, P. lanceolata Merrill, P. bancana Scheffer, P. cyanocarpa Lauterb. and
K. Schum., P. parvifolia Scheffer, P. hentyii Moeljono, and P.
polytrica Diels are tree.
4.1.2. Leaves
Leaves of Popowia Endl. are simple, alternate, membranous characters or coriaceous, granular or subgranular in texture, glabrous, pubescent or tomentose. The leaves of most Popowia
Endl. species vary in size and shape. Popowia pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. fuscaKing, P. velutina
King, P. perakensis King, P. tomentosa Maingay ex Hook.f., P. tomentosa var. crinita J. Sinclair,P. hirtaMiquel,P. odoardi Diels, P. beccarii Scheffer, P. cyanocarpa Lauterb. and K. Schum., P.
novoguineensis Miquel, P. platyphylla Diels, P. Schefferiana Diels,
P. papuanaScheffer, P. cordata Moeljono, P. nootebomii Moeljono,
and P. polytrica Diels are elliptic – oval or oblong. P. cuspidata
Miquel, P. bancana Scheffer, P. clavataDiels, P. pachyptala Diels, and P. hentyii Moeljono are ovate. P. lanceolata Merrill, P. papuanaScheffer, andP. parvifoliaScheffer are lanceolate.
All Malesian species have the upper leaf surface glabrous or glossy, except in P. tomentosa Maingay ex Hook.f., P. tomentosa
var. crinita J. Sinclair and P. hirta Miquel are tomentose – puberulent. The lower leaf surface are pubescent in Popowia
perakensis King, P. tomentosa Maingay ex Hook.f., P. lanceolata
Merrill, P. clavata Diels, and P. polytrica Diels. The lower leaf surface are pilose in P. pauciflora Maingay ex Hook.f., P. beccarii
Scheffer, P. parvifolia Scheffer, P. platyphylla Diels, and P. hentyii
Moeljono. The lower leaf surface are glabrous inP. hirtaMiquel, P. cuspidata Miquel, P. cordata Moeljono, P. bancana Scheffer, P.
odoardi Diels, P. novoguineensis Miquel, and P. nooteboomii
Moeljono.
The leaves are membranaceous inP. pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. perakensisKing, P. tomentosa
Maingay ex Hook.f., P. tomentosa var. crinita J. Sinclair, P.
cuspidata Miquel, P. odoardi Diels, P. beccarii Scheffer, P.
pachypetala Diels, P. cordata Moeljono, and P. hentyii Moeljono.
Chartaceous or coriaceous leaves were found in P. fusca King, P.
velutina King, P. lanceolata Merrill, P. hirta Miquel, P. bancana
Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensis Miquel, P. platyphylla Diels, P. schefferiana
Diels,P. nooteboomii Moeljono, andP. polytrica Diels.
Acuminate leaf apex can be found in P. pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. perakensis King, P. tomentosa var. crinita J. Sinclair, P. lanceolata Merrill, P. hirta
Miquel, P. cuspidata Miquel, P. odoardi Diels, P. clavata Diels, P. novoguineensisMiquel, P. pachyptala Diels, P. platyphyllaDiels, P. parvifolia Scheffer, P. hentyii Moeljono and P. nootebomii
Moeljono.
Acuter or obtuse leaf apex is observed in P. fusca King, P. velutina King, P. tomentosa Maingay ex Hook.f., P. bancana
Scheffer, P. cordata Moeljono, P. beccariiScheffer, andP. papuana Scheffer. The rounded leaf base can be found in Popowia pisocarpa
(Blume) Endl., P. fusca King, P. velutina King, and P. tomentosa
Maingay ex Hook.f. The cordate leaf base can be found in P.
pauciflora Maingay ex Hook.f., P. perakensis King, P. lanceolata
Merrill, P. hirtaMiquel, P. cuspidataMiquel, P. bancana Scheffer, P. clavata Diels, P. novoguineensisMiquel, P. papuana Scheffer, P. parvifolia Scheffer, P. pachyptala Diels, P. platyphylla Diels, P. nootebomii Moeljono, andP. hentyii Moeljono.
The number of main nerves is 5 - 10 pairs in Popowia
pisocarpa (Blume) Endl., P. pauciflora Maingay ex Hook.f., P. fusca King, P. velutina King, P. perakensis King, P. bancana
Scheffer, P. cuspidata Miquel, P. lanceolata Merrill, P. beccarii
Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum.,
P. novoguineensis Miquel, P. papuana Scheffer, P. schefferiana
Diels, P. pachypetalaDiels, P. polytrica Diels,P. platyphyllaDiels,
P. parvifoliaScheffer, and P. nooteboomii Moeljono, andP. hentyii
Moeljono. The number of main nerves is 11 – 18 pairs in P.
tomentosa Maingay ex Hook.f., P. tomentosa var.crinita J. Sinclair,
P. hirtaMiquel, P. cordata Moeljono, andP. odoardi Diels.
The leaf petiole is 1 - 2 mm long in Popowia lanceolata Merrill, P. bancana Scheffer, P. beccarii Scheffer, P. parvifolia
Scheffer, P. platyphylla Diels, P. papuana Scheffer, P. polytrica
Diels, P. cordata Moeljono, and P. nootebomii Moeljono. The leaf petiole is 3 – 5 mm long in Popowia pisocarpa (Blume) Endl., P.
pauciflora Maingay ex Hook.f., P. fusca King, P. velutina King, P. perakensisKing, P. tomentosa Maingay ex Hook.f., P. hirtaMiquel, P. cuspidata Miquel, P. odoardi Diels, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensis Miquel, P. Schefferiana Diels, P. pachypetala Diels, P. hentyii Moeljono, and
P. tomentosa var.crinita J. Sinclair.
4.2. Generative Morphology
4.2.1. Sepal.
Popowia Endl. has three sepals which are valvate and small
size. The shape and size of sepals of each species is very characteristic. In most species the sepal are different in shape. The sepal shape is triangular – ovate. It can be found in Popowia pisocarpa (Blume) Endl., P. perakensis King, and P. platyphylla
Diels. Ovate sepals can be found in P. pauciflora Maingay ex Hook.f., P. fusca King, P. hirta Miquel, P. cuspidata Miquel, P. cordata Moeljono,P. pachypetala Diels, andP. parvifolia Scheffer.
Broadly cordate sepals can only be found in P. bancana Scheffer. The outside surface of the sepals is tomentose in P. pisocarpa (Blume) Endl., P. fusca King, P. perakensis King, P. tomentosa Maingay ex Hook.f., and P. bancana Scheffer. It is pilosa in P. pachypetala Diels, P. parvifolia Scheffer, and P. platyphylla Diels.
It is pubescent or hairy inP. lanceolataMerrill and P. hirtaMiquel. All Malesian species have the inside surface of the sepals glabrous. Most of Malesian species have an acute sepal apex, except in P.
fuscaKing which has obtuse sepals.
4.2.2. Petal
Popowia Endl. has six petals, 3 at the outer, and 3 at the inner.
Petals are sub-orbicular – broad ovate, valvate, coriaceous, and thick. Outer petals small, spreading, slightly larger than the sepals; inner petals larger than outer, concave inside, erect, their edges at first adhering, later free, their apices often incurved, base shortly clawed, sometimes the outer and inner petals united at the base forming a sympetalous corolla. The size of petals of each species is very characteristic. The outside surface of the petals is pubescent in
Popowia pisocarpa (Blume) Endl., P. pauciflora Maingay ex
Hook.f., and P. lanceolata Merrill. It is pilosa – villous in P. pachypetala Diels, P. parvifolia Scheffer, and P. platyphylla Diels.
of most of the Malesian species has glabrous inside surface, except inP. platyphylla which is pilose.
4.2.3. Stamen
Commonly cuneate with broad, flat-topped or slightly concave connectives, number of stamen 10 - 18 in two rows.
4.2.4. Ovary
Ovaries 3 - 12, ovoid to oblong; style very short; stigma sub-capitate or wedge-shape, grooved on the top and down the inner side.
4.2.5. Ripe Carpels
Globose or ovoid, sub-sessile or stalked. Seeds 1 – 4, rugose and with a circumferential ridge.
4.3. Leaf Anatomy
1 2
3 4
[image:40.612.98.484.75.641.2]5 6
Fig. 5. Portion of a cross-section of the leaf ofPopowia Endl.
1. Popowia velutinaKing 2.P. cyanocarpaLauterb. and K. Schum 3. P. bancana Scheffer 4. P. nooteboomii Moeljono
5. P. hirtaMiquel 6. P. tomentosa var.crinita J. Sinclair. Epidermis
Epidermis Epidermis
Spongy Spongy Spongy
Palisade Palisade Palisade
Epidermis
7 8
9 10
[image:41.612.95.507.99.675.2]11 12
Fig 5. continued
7.Popowia pisocarpa (Blume) Endl 8. P. perakensisKing 9.P. kostermansii Moeljono 10.P. cuspidataMiquel 11.P. odoardi Diels 12.P. tomentosa Maingay ex Hook.f.,
A. Epidermis B. Palisade C. Spongy A
A
A B
B
B C
C
C
A B
[image:42.612.121.499.76.410.2]C D
Fig.6. Epidermal cells and crystals on the lower leaf surface A. Popowia tomentosa B. P. novoguinensis . C.P bancana D. P. tomentosavar. crinita
A B
C D
E F
[image:43.612.108.503.75.658.2]G
Fig. 7. Epidermal cells and stomata on the lower leaf surface
A. Popowia tomentosa B.P. novoguinensis C.P hirta. D.P clavata . E.P bancana F. P. tomentosavar. crinita G.P. velutina
1
1
1
1 2
2
2
2
4.4. Relationships Species ofPopowia Endl.
A cladistic analysis of Popowia Endl. was undertaken based on 34 morphological characters (Table 1). Mitrephora maingayi Hook.f
was chosen as an outgroup because it has the closest relationship to
[image:44.612.92.509.239.632.2]Popowia Endl. Mitrephora Hook. f and Popowia Endl.have extra axillary flower (Sinclair, 1955).
Table 1. Characters and Character States Used in the Phylogenetic Analysis of the Malesian Taxa ofPopowia Endl.
Characters Characters state
1 : Habit 0 = shrub
1 = tree 2 : Petioles length 0 = < 4 mm
1 = > 4 mm 3 : Petioles surface 0 = glabrous
1 = pubescent or tomentose 4 : Leaves length 0 = < 10 cm
1 = > 10 cm 5 : Leaves wide 0 = <4 cm
1 = > 4 cm 6 : Leaves shape 0 = lanceolate
1 = ellipt or ovate 7 : Leaves texture 0 = membranaceous
1 = chartaceous or coriaceous 8 : Apex of leaves 0 = acute
1 = acuminate 9 : Base of leaves 0 = oblique
1 = unequal
10 : Leaves base shape
0 = acute or cuneate
1 = rounded
2 = cordate or sub cordate
11 : Upper leaves surface 0 = glabrous
1 = pubescent tomentose 12 : Lower leaves surface 0 = glabrous
1 = pubescent tomentose 13 : Midrib surface 0 = glabrous
1 = pubescent tomentose 14 : Number of nerves 0 = < 10 pairs
1 = > 10 pairs 15 : Reticulation 0 = not visible
1 = visible 16 : Secunder nerves 0 = faint
Table 1. continued
Characters Characters state
17 : Pedicel length 1 = < 10 mm
0 = >10 mm
18 : Pedicel surface 0 = strigose or hairy
1 = pubescent or tomentose 19 : Sepal shape 0 = triangular
1 = ovate
2 = cordate or sub cordate 20 : Sepal length 0 = < 4 mm
1 = > 4 mm
21 : Sepal apex 0 = acute or sub acute
1 = obtuse 22 : Outer surface of sepal 0 = hairy
1 = pubescent or tomentose 23 : Petal shape 0 = orbicular or sub orbicular
1 = ovate or elliptic 24 : Outer surface of petal 0 = pubescent or tomentose
1 = villous
25 : Inner surface of petal 0 = pubescent or tomentose 1 = villous
26 : Outer petal length 0 = < 4 mm
1 = > 4 mm 27 : Inner petal length 0 = < 5 mm
1 = > 5 mm 28 : Number of ovary 0 = < 8
1 = > 8
29 : Ovary surface 0 = glabrous
1 = pubescent – tomentose
30 : Ripe carpel length 0 = < 15 mm 1 = > 15 mm
31 : Ripe carpel shape 0 = globose
1 = ovoid 32 : Ripe carpel surface 0 = glabrous
1 = pubescent tomentose 33 : Stalk length 0 = < 10 mm
1 = > 10 mm 34 : Number of seed 0 = < 1
1 = > 1
Data matrix of the characters which has been used for the analysis shown in table 2.; when a characters state can not be defined it was noted as “ ? “.
(Fig. 5). The characters, steps, consistency index and resistency index for the morphological characters shown in Appendix 2.
Table 2. Data Matrix of Morphological Characters of Popowia Endl. used for Phylogenetic Analysis.
Characters
Species 1 2 3 4 5 6 7 8 9 1 0 11
M. maingayi 1 1 1 1 1 1 1 0 1 0 0
Popowia pisocarpa 1 1 1 0 1 1 0 1 1 0 0
P. pauciflora 1 1 1 1 1 1 0 1 0 0 0
P. fusca 1 1 1 1 1 1 1 0 1 1 1
P. velutina 1 0 1 0 0 1 1 0 1 1 0
P. perakensis 0 1 1 1 1 1 0 1 1 0 0
P. tomentosa 1 0 1 1 1 1 0 0 1 1 1
P. tomentosa var. crinita 1 0 1 1 1 1 0 1 1 2 1
P. odoardi 0 1 1 1 1 1 0 1 1 1 0
Table 2. continued
Characters
Species 12 13 14 15 16 17 18 19 20 21 22
M. maingayi 1 1 1 1 1 0 1 0 0 0 2
Popowia pisocarpa 1 1 0 0 1 1 1 1 0 0 2
P. pauciflora 1 1 0 0 ? 1 0 0 0 ? 1
P. fusca 1 1 0 1 1 0 1 0 0 1 2
P. velutina 1 1 0 1 ? 1 1 0 ? 0 2
P. perakensis 1 1 0 1 1 1 0 0 0 0 2
P. tomentosa 1 1 1 1 1 1 1 0 1 0 2
P. tomentosa var. crinita 1 1 1 1 1 1 1 0 1 0 2
P. odoardi 0 0 1 1 1 0 1 0 0 0 2
Table 2. continued
Characters
Species 23 24 25 26 27 28 29 30 31 32 33 34
M. maingayi 0 1 0 0 1 ? 2 1 ? 1 1 ?
Popowia pisocarpa 2 0 0 0 0 0 2 0 0 1 0 0
P. pauciflora 0 0 0 0 0 0 1 ? 0 0 ? 0
P. fusca 1 1 1 0 0 0 2 1 0 1 0 ?
P. velutina 0 1 1 ? ? ? ? 0 1 1 0 ?
P. perakensis 0 1 1 ? ? 1 0 0 1 0 1 1
P. tomentosa 1 1 1 1 1 1 2 1 0 1 1 1
P. tomentosa var. crinita 1 1 1 1 1 1 2 1 0 1 1 1
1>0 1>0 1>0 1>0 8 14 27 30
0>1 1>0 1>0 10 28 33
0>1 1>0 8 10
1>0 1>0 2>1 2>0 2>1 1>0 2 10 11 20 23 26
0>1 1>2 8 10 1>0 1>0 1>0 1>0 1>0
1 18 29 31 32
1>0 1>0 1>0 1>0 2 4 5 31 1 12 13 14 17
7 17 25
0>1 1>0 0>1 0>1 0>1 11 17 21 23 30 0>1 1>0 1>0
1>0 0>1 4 19
0>2 1>0 1>0 23 24 25
1>0 1>0 1>0 1>0 1>0
1>0 1>0 2>1 2>0 2>1 1>0 1>0 1>0 1>0
10 15 34
9 18 22 23 29 32
Mitrephora maingayi
Popowia pisocarpa
P. pauciflora P. odoardi P. fusca P. velutina P. perakensis P. tomentosa
[image:47.792.64.696.131.499.2]P. tomentosavar.crinita
Within the cladogram three groups of branching are shown. The
first branch is Mitrephora maingayi (outgroup) separated from
Popowia Endl. By a plesiomorph character of the outgroup (leaves texture is coriaceous, pedicel length is 2 cm and inner surface of
petal is pubescent. The second branch is separated by apex of
leaves (character 8), Number of nerves (character 14), inner petal
length (character 27) and ripe carpel length (character 30). The other
Popowia is separated by petioles length (character 2), leaves base shape (character 10), upper leaves surface (character 11), sepal
length (character 20), petal shape (character 23) and outer petal
length (character 26). The last group finished in three sister group,
they are P. pisocarpa and P. pauciflora, they are joined by leaves base shape (character 10), reticulation (character 15) and number of
seed (character 34); P. fusca andP. velutina, are joined by apex of leaves (character 8), and leaves base shape (character 10); the sister
group is closely relate with P. tomentosa and P. tomentosa var.
crinita is separated by apex of leaves (character 8), and leaves base shape (character 10).
4.5. Distribution and Ecology ofPopowia
The Annonaceae is a pantropical family with about 130 genera and 2.300 species. It is confined mostly to humid tropical lowland forests. They are abundant common in Asia, Africa, Central and South America and few in Australia. The genusPopowia formerly recorded from Asia and Africa, about 90 species and majority from Tropical Africa. The remainder, Madagascar, S.India, Burma, Siam, Malaya, Indo-China, Malay island to Australia.
borders, road sides, waste land, and the genus has a wide altitudinal range, from sea level up to the montain area at 0–600 meters altitude.
In the Malesian region, only a few species have a wide distribution, namelyPopowia pisocarpa (Blume) Endl. Most species have rather limited range of distribution such asP. fusca King ( Malay Peninsula, Sumatra, and Borneo), P. hirta Miquel (Malay Peninsula, Sumatra, and Borneo), P.
tomentosa Maingay ex Hook.f. (Malay Peninsula, Sumatra, and Borneo),P. tomentosa var. crinita J. Sinclair (Malay Peninsula, Sumatra, and Borneo),
P. velutina King (Malay Peninsula, and Sumatra), P. bancana Scheffer
(Sumatra, and Borneo); P. schefferiana Diels (Moluccas and Papuasia). Some species are only known from a single locality of the type collection (P.
pauciflora Maingay ex Hook.f. and P. perakensis King from Malay
Peninsula; P. odoardi Diels from Borneo; P. cuspidata Miquel and P. cordata Moeljono from Java; P. lanceolata Merrill from Philippines, P.
beccariiScheffer, P. clavataDiels, P. cyanocarpa Lauterb. and K. Schum.,
P. novoguineensis Miquel, P. papuana Scheffer, P. pachypetala Diels, P. polytrica Diels,P. platyphylla Diels, P. parvifoliaScheffer,P. nooteboomii
Moeljono, andP. hentyii Moeljono from Papuasia. (Table 3).
In the present study, seven species and one varieties are recorded in Malay Peninsula (Popowia pisocarpa (Blume) Endl., P. pauciflora
Maingay ex Hook.f., P. fusca King, P. hirta Miquel, P. velutina King, P.
perakensis King, P. tomentosa Maingay ex Hook.f., and P. tomentosa var.
crinita J. Sinclair). In Sumatra, six species and one varieties were recorded
(P. pisocarpa (Blume) Endl., P. fusca King, P. hirta Miquel, P. velutina
King, P. bancana Scheffer, P. tomentosa Maingay ex Hook.f., and P. tomentosa var. crinita J. Sinclair). In Java, three species were recorded (P. pisocarpa (Blume) Endl., P. cuspidata Miquel, and P. cordata Moeljono).
In Borneo, six species and one varieties were recorded (P. pisocarpa (Blume) Endl., P. odoardi Diels, P. fusca King, P. hirtaMiquel,P. bancana
pisocarpa (Blume) Endl.). In Moluccas, two species were recorded (P.
pisocarpa (Blume) Endl.,and P. schefferiana Diels). In Papuasia, thirteen
species were recorded (P. pisocarpa (Blume) Endl., P. beccarii Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensis
Miquel, P. papuana Scheffer, P. schefferiana Diels, P. pachypetala Diels,
P. polytrica Diels, P. platyphylla Diels, P. parvifolia Scheffer, P.
nooteboomii Moeljono, and P. hentyii Moeljono (Fig. 9). Papuasia is the
richest in species number of Popowia in the Malesian region, with 13 species recorded. This area has the highest number of endemic species, with eleven species (P. beccarii Scheffer, P. clavata Diels, P. cyanocarpa Lauterb. and K. Schum., P. novoguineensisMiquel, P. papuanaScheffer, P.
pachypetala Diels, P. polytrica Diels, P. platyphylla Diels, P. parvifolia
Scheffer, P. nooteboomii Moeljono, and P. hentyii Moeljono) recognized being endemic (Fig. 9). In contrast, Philippines, Celebes, and Moluccas are the poorest in species number recorded from these areas, but no single species in Lesser Sunda Island. The distribution area of Popowia in Malesia can be grouped into three subregions: Western Malesia, Central Malesia and Eastern Malesia.
a. Western Malesia: Malay Peninsula, Sumatra, Borneo, and Java. There are 12 species occuring in this subregion, two species are considered to be endemic in Malay Peninsula, two species in Java and one species in Borneo.
b. Central Malesia: Philippine, Celebes, Moluccas and the Lesser Sunda Islands. In this area occur 3 species, one of which endemic in the Philippine.
c. East Malesia: New Guinea. There are 13 species, eleven of which are considered to be endemic in this area.
Tabel 3. Distribution ofPopowia Endl. in Malesia
Species M S J B P C L N G
01.P. pisocarpa 1
02.P. fusca
03.P. hirta
04.P. tomentosa 05.P. tomentosa
var.crinita 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 2 2
06.P. velutina 0 0 0 0 0 0 3
07.P.pauciflora
08.P.perakensis
* * 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 4
09.P.bancana
10.P.odoardi
0 0 0 0 0 * 0 0 0 0 0 0 0 0 5 5
11.P.cuspidata 12. P.cordata
0 0 0 0 * * 0 0 0 0 0 0 0 0 0 0 6 6
13.P.lanceolata 0 0 0 0 * 0 0 0 7
14.P.beccarii
15.P.clavata
16.P.cyanocarpa 17.P.novoguineensis
18.P.pachypetala
19.P.papuana 20.P.parvifolia
21.P.platyphylla
22.P.polytricha
23.P.hentyii
24.P.nooteboomii
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 * * * * * * * * * * * 8 8 8 8 8 8 8 8 8 8 8
25.P.schefferiana 0 0 0 0 0 0 9
Endemic 2 0 2 1 1 0 0 11
Non endemic 6 7 1 6 1 1 2 2
4.6. Taxonomy
Popowia
Endl.
Popowia Endl. in Walp. Repert. 1 (1842) 74; Sinclair, J. 1955. Gard. Bull. Sing. Vol. 14 Part 2 : 149 - 516.
Type Species :Popowia pisocarpa (Blume) Endlicher.
Shrub or small trees. Leaves granular or subgranular in texture, glabrous, pubescent or tomentose. Flower rather small, single or in fascicle, opposite the leaves. Sepal 3, valvate. Petal 6, valvate, thick; outer small, spreading, slightly larger than the sepals; inner large than the outer, concave inside, somewhat connivent and erect, their edges at first adhering, later free, their apices often incurved, base shortly clawed, sometimes the outer and inner united at the base forming a sympetalous corolla. Stamens cuneate with broad, flat-topped or slightly concave connectives, number of stamens 10 – 18. Ovaries 3 - 12, ovoid to oblong; style very short; stigma sub-capitate or wedge-shape, grooved on the top and down the inner side.
Ripe carpels globose or ovoid, sub-sessile or stalked. Seed 1- 4, rugose or
pitted and with a circumferential ridge.
Distribution : About 90 species, majority from Tropical Africa. The remainder, Madagascar, S.India, Burma, Siam, Malaya, Indo-China, Malay island to Australia. There are twenty four species and one varieties of
PopowiaEndl. are recognized in Malesia, but this genus is absent in Lesser
KEY IDENTIFICATION TO THE SPECIES OFPOPOWIA ENDL. IN MALESIA
1. Leaves elliptic or oblong – ovate.
2. Reticulation not visible in adult leaves.
3. Inner petal 2 mm long, fruit has one seed …………..…… P. pisocarpa 3. Inner petal 5 mm long, fruit has more than one seed…….P. pauciflora 2. Reticulation visible in adult leaves.
4. The upper surface leaf pubescent – tomentose
5. Leaves chartaceous or coriaceous, nerves up to 10 pairs
6. Leaves oval, pubescent beneath with thickened pubescent edge; nerves 9 - 10 pairs; ripe carpel globose …..……….… P. fusca 6. Leaves less oval, pubescent to tomentose; nerves 6 – 7 pairs;
ripe carpel elliptic – ovoid ………..… P.velutina 5. Leaves membraneous, nerves more 12 pairs
7. Apex acute or shortly acuminate, nerves 12 pairs, tomentum of leaf short and dense with hairs 1 mm long ….. P. tomentosa 7. Apex caudate – acuminate, nerves 16 – 18 pairs, tomentum of
leaf less dense, hair 2 mm long ……..P. tomentosa var.crinita 4. The upper surface of leaf glabrous - glossy
8. Leaves chartaceous or coriaceous
9. Petiole more than 2 mm long, ripe carpel globose.
10. Leaves base obtuse – cordate, nerves 11 pairs or more, ripe carpel elliptic ………..…. ………. P. hirta 10. Leaves base acute – cunneate, nerves up to 9 pairs, ripr
carpel globose.
11. Midrib on lower surface hairy
12. Leaves base cunneate and slight rounded, scalariform …………..…..……….P. cyanocarpa 12. Leaves base acute, reticulate …. P. novoguineensis 11. Midrib on lower surface glabrous
13. Lower surface glabrous, reticulation fine, pedicel 5 mm long ………... P. schefferiana 9. Petiole less than 3 mm long
14. The lower surface of leaves pilose or pubescent
15. Peduncle 5 - 7 mm long ……….…...P. polytrica 15. Peduncle short 2 - 3 mm long ………… P.platyphylla 14. The lower surface of leaves glabrous
16. Pedicel up to 12 mm long ... P. nooteboomii 16. Pedicel 1,5 mm long ………….….……… P. bancana 8. Leaves membranous
17. The lower surface of leaves pubescent – tomentose
18. Petiole 5 – 7 mm long, nerves 6–7 pairs ……..…P. hentyii 18. Petiole 3- 4 mm long, nerves 10 pairs ……. P. perakensis 17. The lower surface of leaves glabrous
19 Petiole less 2 mm long, base obtuse or cordate
20. Obtuse leaf base, nerves 8 – 11 pairs ….. P. beccarii 20. Cordate leaf base, nerves 13 – 14 pairs .... P. cordata 19. Petiole 3 mm long or more, base cunneate
21. Midrib on lower surface hairy, pedicel 2 – 5 mm long, ripe carpel pubescent …………P. odoardi 21. Midrib on lower surface glabrous, pedicel up to 10
mm long, ripe carpel glabrous
22. Trees, ripe carpel ellipsoid ……P. cuspidata 22. Shrub, ripe carpel globose, glabrous………
………..…………P. pachypetala 1. Leaves lanceolate
23. Petiole more than 2 mm long ………..……...… P. lanceolata 23. Petiole less 1 mm long.
1. Popowia pisocarpa(Blume) Endlicher - Fig.11 (p.43)
Popowia pisocarpa(Blume) Endlicherin Walp. Repert. 1 (1842) 74; Boerlage
in Icon. Bogor 1 Fasc. 2 (1899) 144; Koorder et Val. in Meded. Lands Plantent. 61 (1903) 339; Ridleyin Sarawak Mus. Journ. 1/3 (1913) 87; Merrill Enum. Phil. Fl. Plants 2 (1923) 164; Craib, Fl. Siam. Enum. 1 (1925) 47; Ast
in Suppl. Fl. Gen. L’Indo-Chine 1 (1938) 90. Sinclairin Gard. Bull. Singapore XIV (1955) 468 - 472. Bocagea pisocarpa Blume. - Type: Fl. Jav. Anon. (1830) 90, T.45. Basonym :Guatteria pisocarpa Blume in Bijdr. I (1825) p. 21.
Popowia ramosissima (Wallich) Hooker, f. et Th. Fl. Ind. 1 (1855) 105 et in Fl. Br. Ind. 1 (1872) 68; King, Mat. F.M.P. Vol. 1 No. 4 (1892) 341 et in Ann. Roy. Bot. Gard. Calc. 4 (1893) 117 Pl. 159 B; Ridley, F.M.P. 1 (1922) 75.
Guatteria ramosissima Wall. Cat. (1832-47) Nos. 7274 and 8006, nomen nudum.Type: Wall. Cat. (1832-47) Nos. 7274 and 8006
P. rufula Miquelin Ann. Mus. Bot. Lugd. Bat. 2 (1865) 20. Type: Borneo, South Borneo, Mt. Pamatton and Sakoembang, Korthals s.n. n.v. (not seen in BO and L)
P. affinis Miquelin Ann. Mus. Bot. Lugd. Bat. 2 (1865) 20. Type: Sumatra, West Sumatra,Korthals s.n. n.v. (not seen in BO and L)
P. polyandra Merrill in Philp. Journ. Sc. Bot. 3 (1908) 224. B. polyandra Presl, Rel. Haenk. 2 (1835) 77. Type: Philippines. Luzon, Sorsogon (Prague),
Haenke s.n. n.v.
P. rufescens Ridley in Kew Bull. (1926) 59. Type: Sumatra, West Sumatra, Mentawai Islands, Siberut, 1924, SFN 14607Boden Kloss (BO!).
Orophea minahassae Boerlage in Koorder Meded. Lands Plantentuin 19 (1898) 336. Type : Celebes, Kajoewatoe, Alt. 50 m. asl.,Koorders 16026 (BO!, L!) and 16028 (BO!, L!).
Shrub or small tree 3 - 7 m high with spreading, slender. Young twigs rufous -pubescent, the older dark brown, glabrous, striate and with numerous leaf scars.
Petioles 2 - 5 mm long, pubescent. Leaves, 5.
