REINWARDTIA
A JOURNAL ON TAXONOMIC BOTANY,
PLANT SOCIOLOGY AND ECOLOGY
Vol. 14(2): 249-324, December 23, 2015
Chief Editor
Kartini Kramadibrata (Mycologist, Herbarium Bogoriense, Indonesia)
Editors
Dedy Darnaedi (Taxonomist, Herbarium Bogoriense, Indonesia)
TukirinPartomihardjo (Ecologist, Herbarium Bogoriense, Indonesia)
Joeni Setijo Rahajoe (Ecologist, Herbarium Bogoriense, Indonesia)
Marlina Ardiyani (Taxonomist, Herbarium Bogoriense, Indonesia)
Topik Hidayat (Taxonomist, Indonesia University of Education, Indonesia)
Eizi Suzuki (Ecologist, Kagoshima University, Japan)
Jun Wen (Taxonomist, Smithsonian Natural History Museum, USA)
Managing Editor
Himmah Rustiami (Taxonomist, Herbarium Bogoriense, Indonesia)
Lulut Dwi Sulistyaningsih (Taxonomist, Herbarium Bogoriense, Indonesia)
Secretary
Endang Tri Utami
Layout
Medi Sutiyatno
Illustrators
Subari
Wahyudi Santoso
Anne Kusumawaty
Correspondence on editorial matters and subscriptions for Reinwardtia should be addressed to:
HERBARIUM BOGORIENSE, BOTANY DIVISION,
RESEARCH CENTER FOR BIOLOGY-INDONESIAN INSTITUTE OF SCIENCES
CIBINONG SCIENCE CENTER, JLN. RAYA JAKARTA - BOGOR KM 46,
CIBINONG 16911, P.O. Box 25 CIBINONG
INDONESIA
PHONE (+62) 21 8765066; Fax (+62) 21 8765062
E-MAIL: reinwardtia@mail.lipi.go.id
http://e-journal.biologi.lipi.go.id/index.php/reinwardtia
A
B
C
D
F
E
G
H
Cover images: Zingiber engganoensis Ardiyani. A. Habit B. Leafy shoot and the
inflorescence showing rhizomes, roots and root-tuber C. Leaves D. Ligule and swollen
petiole E. Dissection of inflorescence showing fruit F. Spike and flowers G. Dissection
of flowers and fruits showing bract, bracteole, two lateral staminodes, two petal lobes,
labellum, and the four appendages of the anther H. Flower. Source of materials: E190
(BO). Photo credits: B, C, D by Arief Supnatna. A, E, F, G, H by Marlina Ardiyani.
The Editors would like to thank all reviewers of volume 14(2):
Abdul Latiff Mohamad, Faculty of Science & Technology, Universiti Kebangsaan Malaysia, Malaysia
Abdulrokhman Kartonegoro - Herbarium Bogoriense, Bogor, Indonesia
Agus Susatya - University of Bengkulu, Bengkulu, Indonesia
Axel D. Poulsen - Royal Botanic Garden Edinburgh, Edinburgh, Scotland, UK
Campbell O. Webb - Arnold Arboretum, University of Harvard, USA
Edwino Fernando - Dept. of Forest Biological Sciences, University of the Philippines, Los Baños, Philippines
Fabian Brambach - Dept. of Ecology & Ecosystem Research, Georg August University, Gottingen, Germany
John Mood - Lyon Arboretum, University of Hawaii, USA
Kuswata Kartawinata - Integrative Research Center, The Field Museum, Chicago, USA
Mark Newman - Royal Botanic Garden Edinburgh, Edinburgh, Scotland, UK
Martin Dancak - Faculty of Science, Palacky University, Czech Republic
Mien A. Rifai - Akademi Ilmu Pengetahuan Indonesia (AIPI)
Ridha Mahyuni - Herbarium Bogoriense, Bogor, Indonesia
REINWARDTIA
Vol 14, No 2, pp: 265 – 286
THE VEGETATION OF LAMBUSANGO FOREST, BUTON, INDONESIA
Received April 03, 2015; accepted June 27, 2015
ANDREW POWLING
School of Biological Sciences, University of Portsmouth, Portsmouth, UK. E-mail: andrew.powling@port.ac.uk.
AURORA PHILLIPS
School of Biological Sciences, University of Brighton, Brighton, UK.
ROSIE PRITCHETT
Centre for Biological Sciences, University of Southampton, Highfield Campus, Southampton, UK.
SIMON T. SEGAR
School of Biological Sciences, University of Reading, Reading, UK.
REBECCA WHEELER
School of Biological Sciences, University of Portsmouth, Portsmouth, UK.
ANI MARDIASTUTI
Faculty of Forestry, Bogor Agricultural University, Bogor, Indonesia.
ABSTRACT
POWLING, A., PHILLIPS, A., PRITCHETT, R., SEGAR, S. T., WHEELER, R. & MARDIASTUTI, A. 2015. The
vegetation of Lambusango Forest, Buton, Indonesia. Reinwardtia 14(2): 265 – 286. ― Lambusango Forest is a tropical
rainforest on the island of Buton, which lies close to south east Sulawesi. The forest covers an area of about 95.000 ha,
with different parts of the forest having different levels of conservation protection. It lies on rocks of both calcareous
(limestone) and non-calcareous (sandstone, conglomerate, peridotite and chert) nature, which give rise to soils with
varying pH values, nutrient levels and water-holding capacities. The climate is seasonal, with a dry season of three
months and considerable year-to-year variability due to El Niño and La Niña events. The vegetation on the different
soils and in different habitats has been studied. Over 300 species of vascular plants found in the forest and surrounding
areas are listed, including trees and shrubs, herbs, climbers, epiphytes, ferns and club-mosses. Two genera, Calamus
with 18 species and Ficus with 29 species, are particularly species-rich, apparently due to their ability to occupy numerous edaphic and ecological niches. Species of these two genera are also good colonists and so better able to reach
Buton in the recent past than other species. The plants of the forest indicate that Buton is floristically very similar to
Sulawesi, with at least 83% of the species found in the forest also being known from Sulawesi. Most of the plant families and genera present on Buton are common in SE Asia, indicating colonisation primarily from that continent. Many
fewer families and genera have colonised from the Australasian continent. The conservation of plant diversity is necessary for the forest to continue as a functioning ecosystem, to the benefit of the animals of the forest and also the local
people.
Key words: Buton, Calamus, climate, Ficus, flor istics, Lambusango, soils, Sulawesi.
ABSTRAK
POWLING, A., PHILLIPS, A., PRITCHETT, R., SEGAR, S. T., WHEELER, R. & MARDIASTUTI, A. 2015.
Vegetasi hutan Lambusango, Buton, Indonesia. Reinwardtia 14(2): 265 – 286. ― Hutan Lambusango adalah hutan
hujan tropik di Pulau Buton yang terletak dekat dengan Sulawesi Tenggara. Luas hutan ini adalah 95.000 ha dengan
beberapa bagian hutan yang mempunyai tingkat perlindungan konservasi yang berbeda. Hutan ini berkembang pada
tanah berkapur (limestone) dan non-berkapur (batupasir, konglomerat, peridotit dan rijang) sehingga tanahnya memiliki
berbagai nilai pH, kadar hara dan kapasitas penahan air. Daerahnya beriklim musiman, dengan musim kemarau tiga
bulan dan bervariasi tinggi dari tahun ke tahun karena pengaruh El Niño dan La Niña. Vegetasi pada tanah yang berbeda dan dalam habitat yang berbeda telah dipelajari. Lebih dari 300 jenis tumbuhan yang ditemukan di hutan dan daerah sekitarnya, termasuk pohon-pohon dan semak-semak, herba, liana, epifit, tumbuhan paku dan kerabatnya. Dua marga, Calamus dengan 18 jenis dan Ficus dengan 29 jenis, termasuk jenis yang berlimpah, tampaknya karena kemampuan
mereka untuk menempati berbagai relung ekologis dan edafis. Jenis dari dua marga ini juga merupakan penjelajah yang
baik dan lebih mampu untuk mencapai Buton pada masa lalu dibandingkan jenis lainnya. Tumbuhan di hutan menunjukkan bahwa Buton secara floristik sangat mirip dengan Sulawesi, setidaknya 83% dari jenis yang ditemukan di daerah ini juga terdapat di Sulawesi. Sebagian besar tumbuhan di Buton merupakan jenis umum di Asia Tenggara, sehingga menunjukkan kolonisasi terutama dari benua tersebut. Hanya sedikit suku dan marga yang berasal dari benua Australasia. Konservasi keanekaragaman tumbuhan diperlukan bagi hutan untuk melanjutkan fungsi ekosistem, serta untuk
kepentingan binatang dan juga masyarakat setempat.
Kata kunci: Buton, Calamus, Ficus, floristik, iklim, Lambusango, Sulawesi, tanah.
265
266
REINWARDTIA
INTRODUCTION
The biogeographic region known as Wallacea
includes Sulawesi and its neighbouring islands, the
Moluccas and the Lesser Sunda Islands. The region
has always been separated by deep seawater from
the continental masses of both Asia and Australasia,
so is occupied by organisms which colonised from
one or other of the continents and also by species
that evolved in the region. As a result the forests of
Wallacea are in some ways different from those
known in Asia and Australasia, with different
communities of plants and animals (Primack &
Corlett, 2005). Wallace‟s Line marks the western
edge of Wallacea and the westernmost limit of
organisms which originated on the Australasian
continent.
Buton is an island lying close to South-East
Sulawesi, the biggest island in Wallacea (Fig. 1).
Lambusango Forest is situated in the southern half
of the island (Fig. 2) between 5°S, 122.68°E and
5.5°S, 123.22°E (Widayati & Carlisle, 2012). The
altitude of the forest ranges from sea level to about
[VOL.14
700 m above sea level (asl), so all the area can be
classified as lowland forest. The forest is seasonal
evergreen rain forest with almost continuous canopy
cover. It consists of many different sub-types due to
the varied geology and topology of the area. Some
parts of the forest have steep slopes with thin soils
due to Karst limestone; other parts are flatter with
deeper soils produced from sandstone and ultrabasic
bedrock. Chert ridges and volcanic rocks are very
exposed, with poor and drought-prone soils. Valleys
and their alluvial soils are a major habitat type,
formed by the rivers that flow outwards from the
high ground in the centre of the forest. The forest
covers 95,000 ha, of which 27,000 ha in the west of
the forest is the Lambusango Wildlife Sanctuary, an
area in which no logging, rattan harvesting or
hunting is officially permitted. The remaining
68,000 ha is production forest or limited production
forest with less stringent conservation regulations
(Widayati & Carlisle, 2012). The officially separate
Kakenauwe Nature Reserve, a small (810 ha) spur
of forest immediately adjoining to the north, is here
treated as part of the Lambusango Forest.
Surrounding the forested areas to a width of a few
kilometres is a „non-forest zone‟ which has been
mostly cleared and used for settlements and
agriculture, although some small areas of forest still
exist (Widayati & Carlisle, 2012).
Lambusango Forest is the subject of an ongoing
research effort led by Operation Wallacea and it
was investigated as part of a programme financed
by the World Bank‟s Global Environment Fund
(Purwanto, 2008). The programme ran from 2005 to
2008. It involved the study of biodiversity in
Lambusango Forest by Indonesian and British
scientists, together with conservation agreements
with local villagers whereby they agreed to forest
conservation measures in exchange for investment
in educational and business development, including
ecotourism. Little has been published on the
vegetation of Buton and this paper, part of the
Operation Wallacea research programme, is the first
general description of the vegetation of
Lambusango Forest.
Research Site
Fig. 1. Map of Buton and SE Sulawesi; Lambusango
Forest is shown by the dashed line.
General
Buton is separated from South-East Sulawesi by
a strait 10 km wide with depth less than 100 m. It
has thus been connected by land to Sulawesi at
times of lower sea level during the repeated ice ages
of the Pleistocene Epoch (Voris, 2000). There are
reasons to think that the climate of Sulawesi was
cooler and dryer during the ice ages (De Deckker et
al., 2003; Whitten et al., 2002). Such a climate
2015]
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
267
Fig. 2. South-central Buton showing Lambusango Forest. Dashed line = Production forest; dotted line = Conservation
forest (information from Widayati & Carlisle, 2012); open circles = sites from which the forest was surveyed.
might well have affected the nature and extent of
forest on Sulawesi and Buton.
Following the declaration of conservation zones
in the early 1980s, villages were moved to the
periphery of the forest. Areas formerly inhabited or
cultivated are regenerating but clearly show
evidence of their previous usage. People still exert
an influence on the forest by legal and illegal
logging, legal rattan harvest and hunting. Some encroachment into the forest has occurred due to land
being taken for settlement and conversion to agriculture (Purwanto, 2008).
Geology and Soils
Buton is a fragment of the Australasian continent
that separated from Australia in the late Triassic or
early Jurassic and accreted onto the Eurasian plate
in the early or middle Miocene (Milsom, 2000).
Rocks formed since the Triassic consist of a variety
of marine sedimentary strata, some calcareous and
some non-calcareous in nature. After the middle
Miocene the geological history of Buton is similar
to that of eastern Sulawesi and includes the
emplacement of ophiolite during the mid to late
Miocene. Chalks, marls and reef carbonates were
deposited during the last five million years as Buton
subsided to bathyal depths and later rose again
(Milsom, 2000).
As a result of this history Buton has a wide
variety of rock types which give rise to a range of
soils. The main types of rocks are: i. Limestones
and other calcareous rocks of various ages from late
Triassic to very recent times; ii. Late Cenozoic
sandstones and conglomerates; iii. Mesozoic chert
sediments that now form steep ridges; iv. Ultrabasic
(ultramafic) rocks of the ophiolite body including
peridotites (J. Milsom, pers. comm.) Ultrabasic
rocks have low percentages of silicates and
frequently high levels of minerals that are toxic to
plants. The limestones give rise to calcareous, basic
soils which are often thin and free-draining,
although sometimes containing appreciable clay and
organic components. The sandstones and
conglomerates produce acidic soils which are
generally nutrient poor, whilst the ultrabasic rocks
produce soils, often called ultramafic soils, with a
distinctive vegetation of noticeably small stature. A
minor component of the geology is chert, which
forms very acidic soils with distinctive vegetation.
River valleys contain alluvial deposits that can be
very variable in composition, depending on which
sediments are being washed into the valleys by side
streams. Because of the variety of soils in close
proximity to each other, Lambusango Forest is
expected to have diverse vegetation.
Climate
Buton has a seasonal climate with a dry season
lasting normally from August to October, due to
south easterly winds blowing from the dry continent
of Australia (Whitten et al., 2002). Annual rainfall
has been measured at the coastal town of Kapantori,
on the north east edge of Lambusango Forest, for
the five years from 1997 to 2001. The average
figure was 1967 mm, with a highest figure of 2488
mm in 1999 and a lowest of 941 mm in 1997. The
268
REINWARDTIA
year 1997 was very dry due to a developing El Niño
event, with no rainfall during the months of August,
September and October. The higher parts of
Lambusango Forest almost certainly receive more
rainfall than Kapantori, but no measurements have
been made. The temperatures measured at
Kapantori give a mean daily maximum figure of
26.8°C, with a yearly range from 25.7°C in June to
27.7°C in October and December (Ministry of
Agriculture, Bau Bau, South-East Sulawesi, pers.
comm.).
Two series of annual rainfall measurements exist
for the island of Tobea Besar, which lies close to
the north west coast of Buton, together making a
total of 15 years (1954-1960 and 1975-1982). These
measurements give an average of 1678 mm and the
site normally has its driest month in September or
October (Whitten et al., 2002). Yearly totals for
Tobea Besar (read from the bar chart on page 27 in
Whitten et al., 2002) have been compared with the
yearly cumulative totals of the Oceanic Niño Index
(ONI) produced by the National Oceanic and
Atmospheric Administration, USA, (NOAA, 2014).
The cumulative totals of the ONI give positive
values for El Niño years and negative values for La
Niña years. A significant inverse relationship
between the ONI and yearly rainfall totals was
found (r = -0.641, p=0.010).
Since El Niño and La Niña conditions normally
develop during the second half of the year, the
cumulative ONI values for just the second half were
also tested for correlation with observed rainfall
measurements and a closer correlation was found (r
= − 0.729, p − 0.002). This confirms that in El Niño
years there is below average rainfall in the region
and in La Niña years there is above average rainfall.
As noted above, 1997 was an El Niño year, while
heavy rainfall was experienced on Buton in the
Augusts of 2007 and 2010, both years in which La
Niña conditions developed. Therefore the
vegetation of Lambusango must be able to cope
with very irregular rainfall amounts both during a
year and from year to year.
A yearly average rainfall of approximately 2000
mm or higher with a dry season of about three
months means that Lambusango Forest can be
classified as a seasonal rain forest. It is thus
distinguished from ever-wet forests with no dry
season and monsoon forests with a longer dry
season (Primack & Corlett, 2005).
MATERIALS AND METHODS
Investigation of the soils and the forest was
undertaken during the months of July and August in
the years 2001 to 2012. Soils were tested to
[VOL.14
determine their pH values and also their
conductivities, low values of which indicate low
levels of soluble mineral nutrients. Soil samples
were taken from the top 10 cm of soil, usually from
near paths or transect lines running through the
forest. Two to five sites were sampled for each
underlying rock type, with at least six samples from
each site. The spacing of samples within a site
varied between 30-60 m. A volume of 20 ml of
loose soil was mixed with an equal volume of
deionised water. A Hanna Combo pH and EC meter
was used to make pH and conductivity
measurements. Analysis of Variance, with Tukey‟s
HSD comparisons, was used to determine
significant differences between the means of the
measurements. Analysis of Variance was performed
using the computer program Minitab. Chi-square
tests were done manually.
The forest was explored from sites and camps
within the forest, as shown in Figure 2: Kakenauwe
(limestones), Lapago (limestones, sandstones, alluvial), Anoa (sandstones, ultrabasic), Bala (chert,
conglomerates, alluvial), Wahalaka (limestones)
and Wabalamba (limestones, alluvial). Exploration
was conducted mostly by following pre-existing
paths or transect lines. The transect lines, which had
different orientations in different sites, were 3 km
long and ran straight through the forest, so crossed
many different geological features and habitat
types. Specimens of canopy trees and their
epiphytes were obtained by tree climbers. Plant
parts, mainly leaves, were either: (i) collected and
pressed as herbarium specimens and then sent to
Herbarium Bogoriense for confirmation of
identification, or (ii) photographed and the
photographs compared with specimens in the Royal
Botanic Gardens‟ Herbarium at Kew, London, for
identification to species with the expert assistance
of Herbarium staff.
Information on the identities of many forest trees
was obtained from the listing of Lambusango trees
published on the internet by Widayati et al. (2008).
These authors also collected specimens which were
identified and stored at Herbarium Bogoriense,
Bogor. For the present work nearly all information
on plant species obtained from the various sources
was checked by comparison of photographs with
identified specimens in Kew Herbarium.
RESULTS AND DISCUSSION
Soils
The results of the soil tests are summarised in
Table 1. It was found that soils derived from
calcareous rocks gave relatively high pH values
(mean pH = 6.54), with high values for
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
269
Table 1. pH and conductivity measurements for soils derived from different rock types found at or near the forest sites
Rock Type
Sites
Samples
pH
N
N
Mean
a
Calcareous*
5
66
6.54
Alluvial
2
21
6.43a
b
Non-calcareous*
5
40
5.08
Chert
2
15
4.28b
19
b
Ultrabasic
2
5.03
Conductivity (µS)
SD
Mean
SD
0.61
116.2
a
46.8
0.88
117.1a
50.3
0.63
51.5
b
24.8
0.71
56.7b
30.4
0.37
c
11.9
34.0
*= limestones; **= sandstones, conglomerates; µS = micro Siemens ; SD = standard deviation.
a b c
, , , indicate distinct statistical populations, as determined by Analysis of Variance using Tukey‟s HSD comparison
with a family error rate of p = 0.01.
conductivities also. This suggests that these soils
contain comparatively high levels of mineral
nutrients necessary for plant growth, as might be
expected from the active erosion of surface rocks
creating new soil. Soils in river valleys were mostly
similar to calcareous soils in their pH and
conductivity values, despite the mixed origins of the
sediments forming them. Waters from many rivers
originate from or flow through areas with
calcareous rocks, so it is to be expected that alluvial
soils in the river valleys have some characteristics
of calcareous soils.
The other soils tested were derived from: noncalcareous sandstones, siltstones and conglomerates
(soil mean pH = 5.08); chert (soil mean pH = 4.28);
and ultrabasic rocks such as peridotites (soil mean
pH = 5.03). All these soils gave pH and
conductivity measurements lower than the
calcareous and alluvial soils. This indicates that these soils are strongly leached and have low levels of
mineral nutrients necessary for plant growth.
Analysis of variance using Tukey‟s HSD
comparisons showed that the latter three soil types
are all significantly different, at the 1% level, from
the calcareous and alluvial soils (Table 1). It should
be noted that, although the soils derived from the
ultrabasic rocks have low conductivities, they may
contain levels of certain mineral ions that are toxic
to many plants. Nickel, chromium and manganese
are present at high levels in ultrabasic soils in
Sulawesi (Whitten et al., 2002) and nickel is
produced commercially on Buton.
Vegetation of the Forest
Forest types are divided according to the rock and
soils on which they grow. Two general categories of
calcareous and non-calcareous (acidic) soils are
discussed, but two subcategories of acidic soils,
soils derived from chert and ultrabasic rocks, are
sufficiently different to be described separately.
Other types of forest, namely riverine, coastal and
regenerating secondary, are also described. Two
very species-rich genera, Calamus and Ficus are
described separately.
Over 300 species, consisting of approximately
150 trees, 20 shrubs, over 50 climbing plants and
epiphytes, 40 herbs, 40 ferns and three club mosses
were found growing in the wild in Lambusango Forest, including areas used for human settlements and
agriculture (Table 2). Species planted by people for
food, medicine or decoration are not included. The
list is not complete; many species were seen but not
identified and many more must have remained unseen. Soil preferences for some of the commoner
tree species are shown in Table 3.
Forest on calcareous rocks
Considerable areas of the forest have calcareous
soils formed from limestone. The soils vary from
new, formed recently from the protruding
limestone; to older, with high organic content and in
places regularly waterlogged at times of heavy rain.
Some areas have much limestone rock protruding at
the surface with thin pockets of soil; this soil is
clearly very dry during the dry season, due to the
rapid loss of water through the underlying
limestone. Tree species, considered to be common
based on frequency of encounter, include
Buchanania arborescens and B. sessilifolia,
Canarium asperum, Dillenia serrata, Elaeocarpus
angustifolius, Heritiera trifoliata, Planchonia
valida, Pometia pinnata, Pterocymbium javanicum,
Pterospermum celebicum and P. diversifolium
(Table 2). Many species of figs (genus Ficus) are
present, although most are rare. Among the tallest
trees are Bombax ceiba, Tetrameles nudiflora and
some strangling figs, for example F. glandifera
which can be over 60 m in height.
270
REINWARDTIA
[VOL.14
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area. Families are grouped alphabetically within the Classes listed by Mabberley (2008), Appendix.
No.
1
Class/ Family
Species
Class Lycopodiopsida
Lycopodiaceae
Lycopodiella cernua (L.) Pic.Serm
Huperzia phlegmaria (L.) Rothm.
2
Selaginaceae
Selaginella plana (Desv. ex Poir.) Hieron.
3
Class Psilotopsida
Ophioglossaceae
Ophioglossum pendulum L.
4
Class Marattiopsida
Marattiaceae
Angiopteris evecta (G. Forst.) Hoffm.
5
Class Polypodiopsida
Aspleniaceae
Asplenium longissimum Blume
Asplenium macrophyllum Sw.
Asplenium nidus L.
6
Blechnaceae
Stenochlaena palustris (Burm.) Bedd.
7
Cyatheaceae
Cyathea cf. roroka Hovenkamp
Cyathea cf. elmeri Copel.
Cyathea contaminans Copel.
Cyathea moluccana R. Br. ex Desv.
8
Davalliaceae
Davallia denticulata (Burm.) Mett. ex Kuhn
9
Dennstaedtiaceae
Pteridium aquilinum sensu lato (L.) Kuhn
10 Dryopteridiaceae
Teratophyllum aculeatum (Blume) Mett.
Dicranopteris linearis (Burm. f.) Underw.
11 Gleicheniaceae
Sticherus truncatus (Willd.) Nakai
12 Lindsaeaceae
Lindsaea lucida Blume
13 Lomariopsidaceae
Nephrolepis biserrata (Sw.) Schott
Nephrolepis hirsutula (G. Forst.) C.Presl.
14 Lygodiaceae
Lygodium circinnatum (Burm) Sw.
15 Polypodiaceae
Drynaria quercifolia (L.) J. Sm.
Drynaria sparsisora (Desv.) T. Moore
Microsorum membranifolium (R. Br.) Ching
Microsorum punctatum (L.) Copel.
Phymatosorus scolopendria (Burm. f.) Pic.Serm.
Pyrrosia longifolia (Burm. f.) C.V.Morton
Pyrrosia piloselloides (L.) M. G. Price
16 Pteridaceae
Acrostichum aureum L.
Adiantum malesianum J. Ghatak
Pteris ensiformis Burm.
Pteris moluccana Blume
Pteris tripartita Sw.
Pteris vittata L.
17 Schizaceae
Schizaea dichotoma (L.) Sm.
Schizaea digitata (L.) Sw.
18 Tectariaceae
Pleocnemia irregularis (C. Presl.) Holttum
Pteridrys syrmatica (Willd.) C. Chr. & Ching
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
Species
Stenosemia aurita (Sw.) C. Presl.
Tectaria crenata Cav
19 Thelypteridaceae
Cyclosorus callosus (Blume) Ching
Cyclosorus heterocarpus (Blume) Ching
Cyclosorus subpubescens (Blume) Ching
20 Woodsiaceae
Diplazium sp.
Class Cycadopsida
21 Cycadaceae
Cycas rumphii Miq.
Class Pinopsida
22 Araucariaceae
Agathis dammara (Lamb.) Rich.
23 Gnetaceae
Gnetum gnemon L.
Class Magnoliopsida
Basal Angiosperms
24 Annonaceae
Cananga odorata (Lam.) Hook. f. & Thomson
Polyalthia lateriflora Kurz
25 Hernandiaceae
Hernandia ovigera L.
26 Lauraceae
Alseodaphne borneensis Gamble
Cinnamomum celebicum Miq.
Litsea cordata Hook.f.
27 Myristicaceae
Myristica koordersii Warb.
Myristica malaccensis Hook. f.
28 Piperaceae
Piper abbreviatum Opiz
Piper amboinense C. DC.
Piper betle L.
Piper caninum Blume
Piper cf. bantamense Blume
Piper fragile Benth.
Monocotyledons
29 Amaryllidaceae
Crinum asiaticum L.
30 Araceae
Alocasia cf. balgooyi A. Hay
Amorphophallus sp.
Pothos cylindricus C. Presl.
Pothos scandens L.
Rhaphidophora cf. koordersii Engl.
Rhaphidophora korthalsii Schott
31 Asparagaceae
Dracaena angustifolia (Medik.) Roxb.
32 Cyperaceae
Cyperus kyllingia Endl.
33 Commelinaceae
Commelina diffusa Burm. f.
34 Dioscoreaceae
Dioscorea cf. pyrifolia Kunth
Dioscorea hispida Dennst.
35 Flagellariaceae
Flagellaria indica L.
36 Graminae
Apluda mutica L.
Cenchrus brownii Roem. & Schult.
271
272
REINWARDTIA
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
Species
Coix lacryma-jobi L.
Cynodon dactylon (L.) Pers.
Dactyloctenium aegyptium (L.) Willd.
Digitaria ciliaris (Retz.) Koeler
Eleusine indica (L.) Gaertn.
Eragrostis tenella Roem. & Schult.
Imperata cylindrica (L.) P. Beauv.
Oplismenus compositus (L.) P. Beauv.
Polytrias amaura Kuntze
Saccharum spontaneum L.
Setaria palmifolia Stapf
Sorghum propinquum (Kunth) Hitchc.
37 Marantaceae
Donax canniformis K. Schum.
38 Orchidaceae
Bulbophyllum flabellum-veneris (J. Koenig) Aver.
Calanthe millikenii P. J. Cribb
Trichoglottis geminata J. J. Sm.
39 Palmae*
Areca catechu L.
Arenga pinnata (Wurmb) Merr.
Areca vestiaria Giseke
Calamus koordersianus Becc.
Calamus leiocaulis Becc. ex K. Heyne
Calamus leptostachys Becc. ex K. Heyne
Calamus macrosphaerion Becc.
Calamus minahassae Warb. ex Becc.
Calamus mindorensis Becc.
Calamus ornatus Blume var. ornatus
Calamus pachystachys Warb. ex Becc.
Calamus paucijugus Becc. ex K. Heyne
Calamus pedicellatus Becc. ex K. Heyne
Calamus robinsonianus Becc.
Calamus siphonospathus Mart. var. dransfieldii Baja-Lapis
Calamus suaveolens W.J.Baker & J. Dransf.
Calamus subinermis H.Wendl. ex Becc.
Calamus symphysipus Mart.
Calamus zollingeri Becc.
Calamus L. spp.
Caryota mitis Lour.
Corypha utan Lam.
Daemonorops robusta Warb. ex Becc.
Hydriastele selebica (Becc.) W. J. Baker & Loo
Licuala celebica Miq.
Nypa fruticans Wurmb
Oncosperma horridum (Griff.) Scheff.
[VOL.14
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No. Class/ Family
Species
Pinanga rumphiana (Mart.) J. Dransf. & Govaerts
Saribus rotundifolius (Lam.) Blume
40 Pandanaceae
Freycinetia cf. devriesi Solms
Freycinetia cf. funicularis Merr.
Pandanus cf. borneensis Warb.
Eudicotyledons
41 Acanthaceae
Acanthus ebracteatus Vahl
Andrographis paniculata Nees
42 Achariaceae
Pangium edule Reinw.
43 Anacardiaceae
Buchanania arborescens Blume
Buchanania sessilifolia Blume
Dracontomelon dao (Blanco) Merr. & Rolfe
Koordersiodendron pinnatum Merr.
Semecarpus cf. cuneiformis Blanco
Spondias pinnata (L. f.) Kurz
44 Apocynaceae
Alstonia macrophylla Wall.
Alstonia scholaris (L.) R. Br. var. velutina
45 Araliaceae
Dischidia nummularia R. Br.
Hoya diversifolia Blume
Tabernaemontana macrocarpa Jack
Arthrophyllum sp.
46 Balanophoraceae
Balanophora fungosa J. R. Forst. & G. Forst.
47 Balsaminaceae
Impatiens platypetala Lindl.
48 Burseraceae
Canarium asperum Benth.
Canarium cf. balsamiferum Willd.
49 Calophyllaceae
Calophyllum inophyllum L.
Calophyllum soulattri Burm. ex F. Mull.
50 Capparaceae
Crateva religosa G.Forst.
51 Casuarinaceae
Gymnostoma sumatranum (Jungh. ex deVriese) L.A.S. Johnson
52 Combretaceae
Terminalia copelandii Elmer
53 Compositae
Ageratum conyzoides L.
Blumea balsamifera DC.
Chromolaena odorata (L.) R. M. King & Rob.
Emilia sonchifolia (L.) DC.
Erechtites valerianifolius (Wolf) DC.
Erigeron sumatrensis Retz.
Gynura procumbens Merr.
Pluchea indica (L.) Less.
Synedrella nodiflora Gaertn.
Tridax procumbens L.
Vernonia cinerea (L.) Less.
54 Convolvulaceae
Ipomoea aquatica Forssk.
Ipomoea hederifolia L.
Ipomoea pes-caprae (L.) R. Br.
Merremia peltata Merr.
273
274
REINWARDTIA
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
55 Cucurbitaceae
Species
Gymnopetalum cochinchinense Kurz
Indomelothria W. J. de Wilde & Duyfjes sp.
56 Datiscaceae
Tetrameles nudiflora R. Br.
57 Dilleniaceae
Dillenia serrata Thunb.
58 Ebenaceae
Diospyros aff. lanceifolia Roxb.
Diospyros malabarica Kostel.
59 Elaeocarpaceae
60 Euphorbiaceae
Elaeocarpus angustifolius Blume
Cleistanthus oblongifolius (Roxb.) M. A.
Macaranga cf. grandifolia Merr.
Macaranga gigantea Müll. Arg.
Macaranga tanarius Müll. Arg.
Mallotus floribundus Müll. Arg.
Phyllanthus niruri L.
61 Fagaceae
Castanopsis buruana Miq.
Lithocarpus celebicus Rehder
62 Goodeniaceae
Scaevola sericea Vahl
63 Icacinaceae
Iodes cirrhosa Turcz.
Phytocrene hirsuta Blume
64 Lamiaceae
Clerodendrum paniculatum L.
Hyptis capitata Jacq.
Lantana camara L.
Premna serratifolia L.
Stachytarpheta jamaicensis (L.) Vahl
Vitex cofassus Reinw. ex Blume
Vitex quinata F.N.Williams
65 Lecythidaceae
Barringtonia aff. pendula Kurz
Barringtonia racemosa (L.) Spreng.
Planchonia valida Blume
66 Leguminosae
Albizia lebbeck (L.) Benth.
Clitoria ternatea L.
Cynometra cauliflora L.
Entada Adans. spp.
Erythrina subumbrans Merr.
Erythrina variegata L.
Flemingia strobilifera (L.) W.T.Aiton
Inocarpus fagiferus (Parkinson) Fosberg
Intsia palembanica Miq.
Mimosa pudica L.
Mucuna sp.
Mucuna pruriens (L.) DC.
Parkia sumatrana Miq.
Pterocarpus indicus Willd.
Senna alata (L.) Roxb.
Vigna marina Merr.
[VOL.14
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
67 Loganiaceae
68 Lythraceae
Species
Strychnos axillaris Colebr.
Duabanga moluccana Blume
69 Malvaceae
Bombax ceiba L.
Grewia glabra Blume
Heritiera littoralis Aiton
Heritiera trifoliolata (F.Muell.) Kosterm.
Hibiscus tiliaceus L.
Kleinhovia hospita L.
Microcos paniculata L.
Pterocymbium javanicum R.Br.
Pterospermum celebicum Miq.
Pterospermum diversifolium Blume
Sterculia longifolia Vent.
Sterculia macrophylla Vent.
Urena lobata L.
70 Melastomataceae
Melastoma malabathricum L.
71 Meliaceae
Aglaia odoratissima Blume
Chisocheton kingii Harms
Dysoxylum arborescens Miq.
Xylocarpus granatum Koen.
72 Menispermaceae
Arcangelisia flava Merr.
Pycnarrhena tumefacta Miers
Tinospora crispa (L.) Hook.f. & Thomson
73 Moraceae
Artocarpus elasticus Reinw.
Artocarpus heterophyllus Lam.
Ficus adenosperma Miq.
Ficus benjamina L.
Ficus botryocarpa Miq.
Ficus callophylla Blume
Ficus caulocarpa (Miq.) Miq.
Ficus chrysolepis Miq. subsp. chrysolepis
Ficus congesta Roxb. var. menadana (Miq.) Corner
Ficus cordatula Merr.
Ficus crassiramea (Miq.) Miq. subsp. crassiramea
Ficus disticha Blume subsp. disticha
Ficus drupacea Thunb.
Ficus fistulosa Reinw. ex Blume
Ficus glandifera Summerh.
Ficus gul K.Schum. & Lauterb.
Ficus heteropleura Blume
Ficus hispida L.f.
Ficus hombroniana Corner var. madhucifolia
Ficus lawesii King
Ficus lepicarpa Blume
Ficus magnoliifolia Blume
275
276
REINWARDTIA
[VOL.14
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
Species
Ficus microcarpa L. f.
Ficus nervosa Roth. subsp. pubinervis (Blume) C.C. Berg
Ficus pisifera Wall. ex Voigt
Ficus prasinicarpa Elmer ex C.C. Berg
Ficus racemosa L.
Ficus recurva Blume var. urnigera (Miq.) King
Ficus riedelii Teijsm. ex Miq.
Ficus septica Burm. f.
Ficus sumatrana (Miq.) Miq.
Ficus tinctoria G. Forst. subsp. gibbosa (Blume) Corner
Ficus variegata Blume var. sycomoroides (Miq.) Corner
Ficus variegata Blume var. variegata
Ficus variegata Blume var. viridicarpa Corner
Ficus virens Aiton
74 Myrtaceae
Syzygium zeylanicum (L.) DC.
Syzygium zollingerianum (Miq.) Amshoff
Xanthostemon petiolatus (Valeton) Peter G. Wilson
75 Oleaceae
Chionanthus montanus Blume
76 Oxalidaceae
Averrhoa carambola L.
77 Passifloraceae
Passiflora foetida L.
78 Proteaceae
Macadamia hildebrandtii Steenis
79 Ranunculaceae
Naravelia laurifolia Wall. ex Hook. f. & Thomson
80 Rhizophoraceae
Bruguiera gymnorhiza (L.) Lam.
Rhizophora apiculata Blume
Rhizophora mucronata Lam.
Sonneratia ovata Backer
81 Rubiaceae
Borreria laevicaulis Ridl.
Hydnophytum formicarum Jack
Hymenodictyon horsfieldii Miq.
Ixora sp.
Morinda citrifolia L.
Myrmecodia tuberosa Jack var. bullosa
Myrmeconauclea cf. stipulacea Ridsdale
Nauclea orientalis (L.) L.
Neolamarckia cadamba (Roxb.) Bosser
Neolamarckia macrophylla (Roxb.) Bosser
Neonauclea calycina (Bartl. ex DC.) Merr.
Neonauclea cf. havilandii Koord. ex Ridsdale
Pavetta cf. montana Reinw. ex Blume
82 Sabiaceae
83 Salicaceae
Meliosma sumatrana (Jack) Walp.
Homalium foetidum Benth.
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
277
Table 2. Vascular plant species recorded in Lambusango Forest and the surrounding area (continued)
No.
Class/ Family
84 Sapindaceae
Species
Dimocarpus dentatus Meijer ex Leenh.
Lepisanthes cf. rubiginosa (Roxb.) Leenh.
Lepisanthes tetraphylla Radlk.
Pometia pinnata J. R. Forst. & G. Forst.
Schleichera oleosa (Lour.) Oken
Madhuca betis (Blanco) J.F.Macbr.
Palaquium obovatum (Griff.) Engl.
Palaquium obtusifolium Burck.
Planchonella duclitan (Blanco) Bakh.f.
Planchonella obovata (R.Br.) Pierre
85 Solanaceae
Solanum ferox L.
86 Thymelaeaceae
Phaleria capitata Jack
87 Ulmaceae
Trema orientalis Blume
88 Urticaceae
Dendrocnide oblanceolata (Merr.) Chew
Dendrocnide sinuata (Blume) Chew
Dendrocnide stimulans (L.f.) Chew
Poikilospermum suaveolens (Blume) Merr.
89 Vitaceae
Cissus sp.
Leea aculeata Blume
Leea angulata Korth. ex Miq.
Tetrastigma cf. pedunculare Planch.
Tetrastigma lanceolarium Planch.
Small trees and shrubs commonly found in the
understorey include Caryota mitis, Cleistanthus
oblongifolius, Dysoxylum arborescens, Leea
aculeata and L. angulata, Phaleria capitata and
Pinanga rumphiana. Where water and organic
matter can accumulate a characteristic plant is a
species of Pandanus, possibly P. borneensis,
which forms one of the tallest elements of the understorey when well grown.
Certain species of tree are valued for their timber by local people. The best timber comes from
Vitex cofassus, Intsia palembanica, Pterocarpus
indicus and Madhuca betis. Other species appreciated for their timber include Dracontomelon dao,
Homalium foetidum and various species in family
Sapotaceae including Palaquium obovatum, Planchonella duclitan and Planchonella obovata. A
few species are valued for specialised and ornamental uses, including Pterocarpus indicus and
at least one unidentified Diospyros species. Neolamarckia macrophylla is grown in plantations
as a source of planks for fences to keep pigs out of
fields. Another species appreciated for a particular
use is Tetrameles nudiflora, which is used for dug
-out canoes.
Root-climbing plants are common, among
them two species of Pothos, P. cylindricus and
P. scandens, and two species of Freycinetia,
believed to be F. devriesii and F. funicularis.
Two root-climbing figs, Ficus disticha subsp.
disticha and F. recurva var. urnigera, are also
found. Rhaphidophora korthalsii is a plant of
more open situations. Species that can be
grouped as climbers and lianas include Hoya diversifolia, Dioscorea hispida, Flagellaria indica,
Arcangelisia flava and Poikilospermum suaveolens. The commonest group of climbing plants is
the rattans, which have been described by Powling (2009). As a group these can be dominant in
places where forest is regenerating but also persist at a lower density in areas of mature forest.
Where significant light penetrates under the
normally dense canopy cover various herbaceous
plant species are found. The clubmoss Selaginella plana is widespread and common, while the
fern Lygodium circinnatum is common in places.
Colourful flowers are rare but include Impatiens
platypetala, while Solanum ferox is distinctive.
The root parasite Balanophora fungosa is
relatively common on both basic and acidic soils.
278
Table 3. Tree species present in forest on various soils and of various types.
Soil Type or Forest Type
Tree species
Calcareous
Riverine
Barringtonia racemosa
×
Buchanania arborescens
Castanopsis buruana
Dillenia serrata
Dimocarpus dentatus
Dracontomelum dao
Duabanga moluccana
Heritiera trifoliate
Inocarpus fagiferus
Lithocarpus celebicus
×
×
×
×
×
Secondary
×
×
×
×
×
×
×
×
×
×
×
×
×
Palaquium obovatum
×
Planchonia valida
×
×
Pometia pinnata
×
×
Polyalthia lateriflora
×
×
REINWARDTIA
×
×
×
×
Schleichlera oleosa
Syzygium zeylanicum
×
Terminalia copelandii
×
Tetrameles nudiflora
×
Vitex cofassus
×
×
×
×
×
×
[VOL.14
* = Non-calcareous
Coastal
×
Parkia sumatrana
Xylocarpus granatum
Chert
×
×
Macaranga gigantea
Mallotus floribundus
Xanthostemon petiolatus
Ultrabasic
×
Agathis dammara
Myristica malaccensis
Neolamarckia macrophylla
Non-Calc*
2015]
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
Grasses are all but absent from areas with
substantial canopy cover, but one species found in
clearings is Oplismenus compositus.
Some conspicuous examples of plant-ant
symbioses were observed. Two epiphytic ant
plants of the family Rubiaceae were found,
Hydnophytum formaricum in the forest, and
Myrmecodia tuberosa on trees near the coast. Another species in the genus Rubiaceae, possibly
Myrmeconauclea stipulacea, also has a symbiosis
with ants. This species has ants living in swollen
and hollow sections of its twigs, entering and
exiting through small holes. A common epiphytic
ant plant on trees in more open areas and in
plantations is Dischidia nummularia, while a
damaged specimen of a fern in the ant-sheltering
genus Lecanopteris was once found on the forest
floor after it had fallen. These plants gain
nitrogenous compounds from the excreta and dead
bodies of the ants, and the ants also defend the
plants against herbivores (Primack & Corlett,
2005).
Many epiphytic orchids were seen but only two
were found in flower and therefore identified:
Trichoglottis geminata and Bulbophyllum
flabellum-veneris. Two species of fern are very
common as epiphytes; Asplenium nidus grows in
areas of forest with dense canopy cover, whilst
Drynaria sparsisora tends to occurs where the
canopy is sparser, so light levels are higher and
presumably the humidity lower. Microsorum
punctatum is also a common epiphytic fern, one
that superficially resembles A. nidus. Some
epiphytic species have been observed growing from
the „bird‟s nests‟ of A. nidus, among them the club
moss Huperzia phlegmaria and, once, the fern
Ophioglossum pendulum, but this latter species
appears to be rare. Stag‟s horn ferns (Platycerium
Desv.) have not been observed on Buton. Common
epiphytes on trunks include Pyrrosia longifolia
and Pyrrosia piloselloides. Species of the forest
floor that scramble up tree trunks include Stenochlaena palustris, Teratophyllum aculeatum and
Lygodium circinatum. Phymatosorus scolopendria
is found growing on tree trunks but is also found
on rocks which are exposed and often dry. A species of open areas that can be common in forest
clearings with exposed rock and sparse soil is
Pteris tripartita.
Forest on non-calcareous, acidic, rocks
A small number of tree species are very
characteristic of forests on acidic soils derived
from sandstone, siltstone and conglomerate rocks.
Castanopsis buruana is common in places and its
279
typical form, with many suckers encircling the
main trunk, is a clear marker for sandy, acidic
soils. It is found at altitudes of 350 m and above.
Lithocarpus celebicus is found on similar soils,
although at higher altitudes (500-600 m). It can be
abundant in areas with sandy soil of low pH and
low conductivity. Other species found frequently
on acidic soils include Dillenia serrata and
Dimocarpus dentatus. Some species, such as
Artocarpus elasticus and Barringtonia racemosa,
are like D. serrata in that they are able to grow on
acidic soils but are also found on other soils types
(Table 3). Only one species of conifer was found
in the forest, Agathis dammara (synonym A.
celebica). This was a single well grown individual
at about 400 m asl.
An understorey palm, Licuala celebica, is often
found growing on acidic soils and in places can be
very frequent. Another palm species, Oncosperma
horridum, is less common.
In areas of acidic, nutrient-poor soils the canopy
is usually lower and thinner than on limestone
soils. In these places Drynaria sparsisora is the
most common epiphytic fern, although Asplenium
nidus is still found in darker and damper places,
such as river valleys. A characteristic fern of tree
trunks is Davallia denticulata. The forest floor in
clearings caused by human activities and in other
areas of high light intensity can be dominated by
Dicranopteris linearis, in combination with
Pteridium aquilinum (sensu lato) in places.
Indeed, the presence of D. linearis serves as a
good indicator that the soil is acidic. Steep slopes
with unstable soil can be covered with Sticherus
truncatus. A species of Lindsaea, probably L. lucida, occurs on ground with little cover, often in the
shelter of exposed tree roots. Other forest floor
species on thin acidic soils include Lycopodiella
cernua, Schizaea digitata and S. dichotoma, whilst
on deeper soils Pleocnemia irregularis occurs.
Two species of tree fern were found at altitudes
varying between 400 to 600 m asl., Cyathea contaminans and a species with affinities to C. roroka.
In the west of the forest are steep slopes of
exposed igneous rock (J. Milsom, pers. comm)
with pockets of thin soil. Vegetation is sparse on
these slopes but one tree species is dominant in
very exposed places, Gymnostoma sumatrana,
where it forms apparently permanent stands. This
species gives a completely different, more open,
character to the forest. The palm Hydriastele
selebica grows in more sheltered places on these
slopes, while the rattan Calamus koordersianus
grows widely, including on rocky, exposed terrain.
280
REINWARDTIA
Forest on chert ridges
In the east of the forest are some ridges formed
from chert (J. Milsom, pers. comm.). These can
have steeply sloped or horizontal crests, but
always have thin soils which are very acidic
(Table 1) and must be very drought prone. They
have characteristic vegetation made up of a limited
number of species, in some ways analogous to
heath forest in other parts of SE Asia. Very
common on the higher parts of the steep ridges and
on the horizontal crests are Syzygium zeylanicum,
a small tree with small, hard leaves and
characteristic red, flaking bark; and the palm
Hydriastele selebica. Lower down on these ridges,
where there is more moisture in the soils, are
found two other palms, Areca vestiaria and
Licuala celebica, although these two species are
widespread on acidic soils and not confined to
chert ridge habitats. On the crests of horizontal
ridges are very large specimens of Xanthostemon
petiolatus, which must be able to get their roots
into lower levels of the rocks where there is
sufficient water. These large trees produce
valuable timber but their remote locations save
them from felling, due to the difficulty of carrying
the timber out of the forest. The small tree fern
Cyathea moluccana is common on the very
exposed crests of these ridges, where the soil is
evidently thin and fast drying. Lycopodiella cernua and Schizaea dichotoma are also found on the
ridges.
Forest on ultrabasic (ultramafic) soils
Forest on ultrabasic soil is made up of small
trees, their growth restricted by the general lack of
mineral nutrients and the presence of elements
which inhibit plant growth, such as nickel.
Identifying tree species in these forests presents
problems, due to the stunted growth. There is also
the possibility that some plants present, having
evolved to grow on ultrabasic soils, might be
better considered as new species.
Among the restricted range of tree species
present are some also able to grow on the nutrientpoor sandstone soils, such as Castanopsis buruana
and Syzygium zeylanicum. Some other species present are also found on calcareous soils, for example Syzygium zollingerianum and a species in the
family Rubiaceae believed to be Neonauclea
calycina. Another species present that is very
widespread on different soil types is Dillenia
serrata. The palm Saribus rotundifolius is found
here but not restricted to such areas. A species of
tree fern, probably Cyathea elmeri, appeared to be
limited to ultrabasic soils, while Lycopodiella
[VOL.14
cernua and a species of Nephrolepis, possibly
N. biserrata, are common in places.
River valleys
River valleys can be divided into two general
types: steep-sided valleys with densely vegetated
alluvial soils which must sometimes flood; and
more open valleys with larger rivers which have
areas of incompletely vegetated stone and gravel
deposits. These two types of valley have differing
plant species and will be treated separately.
Due to the abundance of calcium carbonate
containing rocks, water in rivers is often alkaline
(pH > 7.0) even if the rocks in the area it is
flowing through are acidic in nature. This is shown
by the presence of tufa (calcium carbonate) dams
on the river bed.
Steep sided river valleys contain some of the
tallest trees in the forest. These include Terminalia
copelandii, which grows with its roots in the river
water, and Parkia sumatrana, growing on more
acidic alluvial soils in regions of sandstone and
conglomerate rocks. Many species are common on
river alluvium, but not confined to it. These
include Barringtonia racemosa, Calophyllum
soulattri, Crateva religiosa, Hernandia ovigera,
Macaranga gigantea, Myristica malaccensis,
Oncosperma horridum, Pangium edule and
Pometia pinnata. Valleys with dense canopy cover
are also the habitat of large lianas of the genus
Entada, and the epiphytic fern Asplenium nidus is
common as well.
River valleys and areas of wet soil can have
large patches of ground covered by Donax
canniformis, and a species of Alocasia, thought to
be A. balgooyi, also occurs in such habitats. Some
flowering plants of the ground orchid Calanthe
millikenii were seen in 2008, but not in other years.
Among the many ferns found in valleys Angiopteris evecta occurs rarely on the sides of deep river
valleys, whereas Aspidium crenatus occurs on flatter alluvial areas. Tectaria crenata grows as a rheophyte on the rocky beds of rivers, where it is frequently inundated but sometimes common.
The gravel and boulder banks of faster flowing
rivers in open valleys form an unstable substrate
for plant growth, so it is to be expected that many
species found are pioneers in the ecological sense.
Many probably need the open, disturbed habitat
rather than the river water. Small trees that can
establish near the river edge include Duabanga
moluccana and Microcos paniculata, whilst
Terminalia copelandi and Macaranga gigantea are
found in more stable areas. Smaller plants include
Mucuna pruriens and Flemingia
strobilifera.
2015]
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
The grass Saccharum spontaneum forms large
clumps and is probably one of the first species to
colonise bare shingle and gravel. The fern Pteris
moluccana is also found in this habitat. Open,
stony river valleys are possibly the only natural
habitat in the forest that has been colonised by non
-native plants brought to Buton by people. Species
of this type which are present include Lantana
camara, Senna alata and Chromolaena odorata.
These are colonising species that benefit from the
high light levels and lack of established competition.
Mangroves and coastal forest
The main species of mangrove include
Bruguiera gymnorhiza, Rhizophora apiculata, R.
mucronata and Sonneratia ovata. Behind the
mangroves and the Nypa fruticans, in the forest
surrounding tidal creeks, tree species include
Inocarpus fagifer, Schleichera oleosa and Xylocarpus granatum. Also present is Heritiera littoralis, a species which, perhaps surprisingly given
the size of its fruits, occurs in nearby forest on
limestone about 100-200 m asl. The fern
Acrostichum aureum grows in brackish water and
the epiphytic ant-plant Myrmecodia tuberosa was
found in trees, both just behind beaches. Dry,
sandy ground can be covered by Ipomoea
pes-caprae and Vigna marina.
No clear examples of the „Barringtonia‟ or the
„pes caprae‟ formations (Whitten et al., 2002)
were found. This was probably due to few coastal
areas being investigated and also because those
coastal areas that were accessed had been
extensively altered by people. Individual species
characteristic of the formations were found,
however, and examples most probably exist
around the Lambusango Forest. The „pes caprae’
formation is found on the nearby island of Hoga in
the Wakatobi peninsula, off the south east coast of
Buton.
Cleared areas and secondary forests
Around the edge of Lambusango Forest ground
has been cleared at various times for plantations
and arable fields. Many are still in use but some
have been abandoned and are in various stages of
reversion to forest. Within the continuous forest
are many areas where habitations were abandoned
in the late 1970s and early 1980s and which are
now regenerating and can be classified as
secondary forest. The road running around edge of
the forest actually passes through dense forest
between the north of Lambusango and the smaller
Kakenauwe reserve. Various plants of secondary
281
forests grow on these verges, many of which have
been introduced from other places in the tropics,
mainly herbaceous plants (“weeds”), including
many of the members of the families Graminae
and Compositae listed in Table 2. Imperata
cylindrica is a common grass of open, man-made
habitats. Ipomoea aquatica grows in wetter areas,
whilst Ipomoea pes-caprae can appear in more
ruderal, human-dominated habitats.
Two species of fern can be common and even
dominant on cleared and steep soils, particularly
the sides of road cuttings: Nephrolepis hirsutula
and Cyclosorus callosus. Plants of a mid stage of
succession in fields include Blumea balsamifera,
while cleared areas in full sun are often dominated
by the introduced species Lantana camara and
Eupatorium odoratum. These species normally
cover ground densely and must retard the progress
of succession back to forest. Small trees of
regenerating habitats include species of Macaranga and Mallotus, with Neolamarckia macrophylla
and other members of the family Rubiaceae soon
appearing as well.
No examples of deliberate forest regeneration
for nature conservation are known, but some small
-scale plantations of trees, mainly Tectona grandis, Gmelina arborea and Vitex cofassus, have
been started for timber production on abandoned
farm land.
Two species-rich genera: Calamus and Ficus
Table 2 shows that more species were found in
the genera Calamus (16 identified species, also at
least two unidentified species) and Ficus (29
identified species) than in other genera.
Palms, including Calamus, have been covered
in a separate publication (Powling, 2009). Nineteen species of rattan have been found in Lambusango Forest and at least two more (one being
Korthalsia celebica) have been found in northern
Buton. A reason for the co-existence of many Calamus
species is that different species have
adapted to different soil types (Table 4); so they
occupy many different ecological niches, reducing
the
competition between them. Some are colonising species adapted to the high light levels in
areas cleared and abandoned by humans. Such
species include C. zollingeri and C. ornatus. Other species are better able to persist in places with
dense tree cover, where the lack of light prevents
fast-growing pioneer species dominating, for example C. mindorensis and C. leptostachys among
many others.
The most common genus is Ficus, the figs.
Species of fig were found in a wide variety of
282
[VOL.14
REINWARDTIA
Table 4. Palm species present in forest on various soil types.
Tree species
Areca vestiaria
Calamus koordersianus
Calamus leptostachys
Calamus macrosphaerion
Calamus minahassae
Calamus mindorensis
Calamus ornatus
Calamus pedicellatus
Calamus siphonospathus
Calamus suaveolens
Calamus subinermis
Calamus symphysipus
Calamus zollingeri
Caryota mitis
Hydriastele selebica
Oncosperma horridum
Pinanga rumphiana
Saribus rotundifolius
Calcareous
×
×
×
×
Soil Type
Non-Calc*
Ultrabasic
×
×
×
×
×
×
×
×
×
×
Chert
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
×
* = Non-calcareous
habitats, including mature forest, secondary forest,
roadsides, riversides and abandoned fields.
A range of growth forms are present: free-standing
canopy trees (8 species), small trees or shrubs (5
species), hemi-epiphytes (i.e. strangler figs, 14
species) and root-climbers (2 species). Position of
syconia varied in the different species, from
among the foliage (ramiflorus), to on the trunks
(cauliflorous) and in the ground (geocarpic). In all,
29 species were recorded in Lambusango forest
and its environs (roadsides and areas associated
with farming). In addition, two further species
were found on the nearby coral island of Hoga
(Ficus hombroniana and F. prasinicarpa) and
another species (F. lepicarpa) was found in forest
in northern Buton near Ereke. These three species
may also be present in Lambusango. It is probable
that other fig species occur in Lambusango which
were not recorded. In particular, root climbing
epiphytes were probably under-recorded due to the
limited amount of canopy surveying done during
this work. Fig species were particularly diverse in
areas with calcareous soils, presumably due to these soils providing relatively high levels of mineral nutrients. It is known that figs grow best on fertile soils, since their high assimilation rates and
production of latex to protect against
herbivores require such soils (Harrison, 2005). Another
factor relevant to the high diversity of fig species
is the reluctance, for cultural reasons, of local people to cut down fig trees (N. Priston, pers. comm.).
Reasons for such reluctance are discussed by Berg
and Corner (2005).
The different subgenera of Ficus were not
equally represented in the species found. Table 5
shows the subgenera recognised by Berg and
Corner (2005) and their breeding system
(monoecious or dioecious). Also shown are the
numbers of species in the subgenera identified in
Lambusango, the numbers of species known from
Sulawesi (Berg & Corner, 2005) and the
proportion of Lambusango species as a percentage
of Sulawesi species. The percentage figures vary
considerably between subgenera, from 0% for
subgenus Ficus to 67% for subgenus Urostigma.
Although the totals for the numbers of
Lambusango species with the different breeding
systems are similar, the percentage of monoecious
species (58%) is considerably higher than the
percentage of dioecious (26%). If the Ficus species found in northern Buton and the two found on
Hoga (referred to earlier) together with the three
separate varieties of F. variegata are included in
the analysis the percentages show a similar
discrepancy (65% for monoecious and 32% for
dioecious). However, the difference in the
proportions of the two breeding types present in
Lambusango relative to the numbers in Sulawesi is
barely statistically significant (goodness-of-fit
chi-square test with Yates correction factor,
degrees of freedom = 1, P = 0.050). Little of
biological significance can be assumed from this
statistical result since the addition of one more
dioecious species would have given a
non-significant probability value.
The apparent lack of dioecious species in
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
2015]
283
Table 5. Analysis of Ficus subgenera present on Buton and Sulawesi by breeding system.
Subgenus
Ficus
Pharmacosycea
Sycidium
Sycomorus
Sycomorus
Synoecia
Urostigma
Totals
Breeding system*
Dioecious
Monoecious
Dioecious
Monoecious
Dioecious
Dioecious
Monoecious
Monoecious
Dioecious
Buton species
Sulawesi species*
Percentage
0
2
5
1
7
2
12
15
14
9
7
22
1
16
6
18
26
53
0
29
23
100
44
33
67
58
26
*Information from Berg & Corner (2005)
Lambusango may be a result of not finding all the
root climbers in the subgenus Synoecia, due to lack
of work in the canopy. Alternatively, the apparent
excess of monoecious species can be attributed to
the high proportion of species in
subgenus
Urostigma that are present. This might result from
some aspect of their biology other than monoecy,
such as the relatively small sizes of their syconia
making them easily taken by flying
animals
which then disperse the seeds.
DISCUSSION
Many plants of Lambusango Forest belong to
pan-tropical families, including Annonaceae,
Euphorbiaceae, Leguminosae, Meliaceae,
Moraceae, Myristicaceae, Palmae, Rubiaceae and
Sapotaceae. These families are characteristic of SE
Asian forests (Primack & Corlett, 2005). Genera
and species of these families found to the east of
Wallace‟s line are considered to have mostly or
entirely colonised from the west (Primack &
Corlett, 2005). The predominance of Asian
families and genera is due to plants of these taxa
being able to successfully colonise Wallacea
across the relatively narrow water gaps from Asia
and between the islands−tens of km, at least at
times of low sea levels during the Pleistocene ice
ages. Another possible reason for the
predominance has been suggested by Morley
(2003), namely that the south western arm of
Sulawesi was attached to Borneo in the middle
Miocene before moving to its present position.
Flora on the arm would then have been able to
colonise eastwards without having to cross
Wallace‟s Line.
A few genera are considered to have colonised
from the Australasian continent after it started to
collide with the Philippine plate at the OligoceneMiocene boundary (Morley, 2003), e.g. Agathis in
Araucariaceae, Gymnostoma in Casuarinaceae,
Macadamia in Proteaceae and Xanthostemon in
Myrtaceae. These genera are all represented by
one species only. The Australasian continent,
having moved northwards through temperate
latitudes, did not have as many plant species
adapted to tropical climates as Asia and therefore
able to colonise lowland rain forests in Wallacea
and compete with the Asian taxa (Primack &
Corlett, 2005).
Lambusango Forest differs from typical SE
Asian forests in the apparently complete lack of
species of the family Dipterocarpaceae. This is
due to two factors: the low number (six) of
dipterocarp species that are present on Sulawesi
(Keßler et al., 2002) and the seasonal climate of
Buton, which would be unsuitable for the many
dipterocarp species that have evolved for ever-wet
climates.
Of the woody species identified, 174 were
checked against the compilation of trees known on
the island of Sulawesi published by Keßler et al.
(2002). This showed that 73% of the Lambusango
species are also known to occur on Sulawesi.
However, if the known distributions given for
Calamus by Powling (2009) and for Ficus by Berg
and Corner (2005) are used instead, then the figure
becomes 83% of a total number of 176 species
checked. This indicates that Buton can be
considered to be a part of Sulawesi floristically, as
might be expected given the close proximity and
the land connection during the Quaternary ice
ages.
Various factors influence the diversity of plants
in Lambusango. Diversity is increased by the
variable topology, with its flat areas, steep slopes,
ridges and deep river valleys. These features
produce many different habitats and microhabitats
for plants to occupy. Many soil types are present,
with their varying pH values, levels of mineral
nutrients and water contents during the year,
allowing species with different edaphic
requirements to grow. Widayati and Carlisle
284
REINWARDTIA
(2012) showed that tree species diversity in Lambusango increases with slope gradient and altitude,
probably because of lower disturbance by humans
in higher and steeper places, although steep gradients might also increase the variety of habitats
available to trees. The work of Widayati and Carlisle (2012) indicated that soil chemical composition has little effect on tree diversity. The present
work made no quantitative comparisons of overall
diversity on different soil types, but it was observed that figs were very diverse on calcareous
soils. The work showed that some dicotyledonous
tree species and some palms (Tables 3 & 4) are
restricted to certain soil types whilst others are able
to grow on a wide range of soils. It is possible that
some of the widespread species have different
physiological races which are able to grow on
different soil types.
The short dry season on Buton prevents the
growth of some species that require an ever-wet
climate, e.g. some Dipterocarp species. It must also
restrict the number of species adapted to a longer
dry season. However, the thin, dry soils in parts of
Lambusango do permit the existence of some
species adapted to a dryer climate, e.g. Alstonia
angustifolia, Buchanania arborescens and
Cananga odorata, which are characteristic of forests with drier, more seasonal climates than experienced by Buton (Whitten et al., 2002). Another
factor acting to decrease species diversity on Buton
is relative isolation, it being an island off the coast
of an island. It is possible that Sulawesi and Buton
have experienced repeated loss of diversity during the succession of ice ages over the past two
million years, when the climate became colder and
dryer (Whitten et al., 2002; De Deckker et al.,
2003), leaving few refugia for tropical lowland
rainforest species. Local extinctions of
species
may have occurred but recolonisation not subsequently happened.
Some families and genera may be better able to
colonise or recolonise than others, which could
account for some genera being much richer in
species than others. The two species-rich genera
may be examples: Calamus (Palmae) with 18
species and Ficus (Moraceae) with 29 species
(Table 2). Both genera are known to be very
species-rich in SE Asia (Dransfield et al., 2008;
Berg & Corner, 2005), and their fruits are eaten by
frugivorous birds (pigeons, hornbills, etc.) and
fruitbats (Corlett, 1998; Shanahan et al., 2001;
Zona & Henderson, 1989) capable of flying across
the sea from Sulawesi to Buton, carrying seeds in
their guts. Species of other genera might be less
likely to colonise due to their fruits being less
[VOL.14
attractive to birds, on account of colour,
nutritional value or size. However, it could be that
other genera are more species rich than shown in
Table 2, but their species were under-recorded
because they received less attention. Possible
examples include Elaeocarpus, Macaranga and
Syzygium, which are known to be diverse in
Borneo and Sulawesi (de Kok & Utteridge, 2010;
Keßler et al., 2002).
Only one species of conifer was found in the
forest, Agathis dammara (syn. A. celebica). This
was a single well grown individual at about 400 m
asl. Species such as A. dammara, Lithocarpus
celebicus and Castanopsis buruana occur at
altitudes of 400 m to 600 m in Lambusango,
although these species are normally components
of the lower montane rain forest, which is considered to have a lower altitudinal limit of 1200–1500
m (Whitten et al., 2002). However, on isolated,
small mountains the lower limit is 700–900 m
(Grubb, 1971). The reduced heights of vegetation
zones on outlying mountains, compared with main
ranges, is known as the „Massenerhebung‟ effect,
and the low altitudes of the three species mentioned may be an example of this effect. However,
it has been reported that A. dammara occurs naturally at low altitudes in central Sulawesi, but only
at higher altitudes on the peninsulas (Whitten et
al., 2002), which is contrary to expectations if the
Massenerhebung effect applies. Clearly more information on the distribution of A. dammara is
needed.
Many alien species flourish in and near the
settlements surrounding the forest but few have
colonised the intact forest. As noted above, some
are able to colonise unstable shingles in open river
valleys, but not invade the continuous forest. An
introduced species that has become naturalised in
the forest is Areca catechu, which persists at the
sites of deserted settlements. Other non-native
species which show some encroachment on the
forest edges are Muntingia calabura and Psidium
guajava, although there is little sign that they can
naturalise in the continuous forest. Overall there
are very few introduced species in the forest,
despite the presence of many animals capable of
transporting seeds, such as fruit bats, monkeys and
pigs, which feed in cultivated land but spend much
of their time in the forest. The continuous canopy
cover in the forest results in low levels of light on
the forest floor and it seems probable that
competition for light prevents crop and weed
species from establishing. It is noticeable that the
three species mentioned above are plants of the
forest understorey, so adapted to the lack of light
2015]
POWLING et al.: Vegetation of Lambusango forest, Buton, Indonesia
beneath a forest canopy.
The genus Ficus, the figs, was found to be the
most species-rich of the plant genera on Buton.
This high diversity is not a local phenomenon.
There are about 735 fig species worldwide and 367
species in Malesia (Berg & Corner, 2005). Harrison (2005) has observed that Ficus is the most
diverse genus in plant lists for many sites in the
Indo-Pacific and African regions. His
explanation is that the figs display evolutionary flexibility and morphological diversity, and have speciated to occupy many different ecological niches,
some of which are rare. As a result figs can have a
diversity of species in a small forest area, since
they do not compete with each other. The variety
of growth forms and morphologies of the figs on
Buton suggests they do indeed occupy a variety of
ecological niches.
As noted before, fig seeds are efficiently
dispersed over long distances by birds and bats. A
result of this is that a small forest area such as
Lambusango might contain a large proportion of
the fig species found in a much larger
neighbouring area, in this case Sulawesi. In the
present work this was found to be the case, with 29
species out of a Sulawesi total of 78 (Berg &
Corner, 2005), that is 37%, found in Lambusango.
Twelve species in the subgenus of Urostigma were
present (67% of the Sulawesi total), suggesting
species in this subgenus are particularly good
colonisers. In addition, species in this subgenus are
strangler figs, which must be able to survive dry
conditions when they first germinate in the canopy
of a host tree. This ability may allow them to
survive the seasonally dry conditions in some of
the soils of Lambusango.
Figs are pollinated by fig wasps which are often,
but not always, species specific. This is a very
effective mechanism which combines the
advantages of wind pollination, since the wasps
can be carried long distances by winds above the
canopy, with the ability of insects to find rare
individuals of a plant species, by responding to
volatile chemicals released by the plant (Cook &
Rasplus, 2003). Due to this mechanism pollen is
not limited in most fig species in most
environments and species are able to persist at low
densities of individuals. As a result some figs are
able to occupy rare ecological niches which have
low competition due to limited recruitment
opportunities (Harrison, 2005).
Lambusango Forest persists as a functioning
forest ecosystem, with a degree of conservation
protection. From an ecological point of view the
forest needs to maintain a certain level of
285
intactness to function successfully as an
ecosystem. If forest is lost due to human activities
then ecological interactions will start to break
down. If hornbills and pigeons were lost then
rattan fruit would not be successfully dispersed
and there would be less rattan for the local men to
harvest. Hunting of animals and birds would result
in less fig seed being dispersed and fig populations
decreasing, with fewer figs for other forest
animals. Indeed, figs have been shown to be
„keystone‟ species for the forest since so many
animals rely on them as a source of food
throughout the year (Kinnaird et al., 1999), so loss
of figs would cause many further detrimental
changes.
The forest edge is under direct threat from
human activities, such as encroachment by
settlements and loss of peripheral areas to
agriculture (Purwanto, 2008), but all the forest is
affected to a greater or lesser extent by human
actions, since access to the forest is very easy from
the communities around the forest edge. Rattan
canes are extensively collected by local men
(Powling 2009; Widayati et al. 2010). Certain tree
species, of which the most sought is Vitex
cofassus, are felled for their timber. This is much
in demand for local house and boat building,
among other uses. At least one unidentified species of Diospyros, ebony, is taken (often illegally)
because of the commercial value of the wood. It
might be that Agathis dammara was once more
common but has been extensively logged in the
past.
Local people living on the periphery understand
the importance of maintaining the forest as a
reliable source of water for the rice padi fields on
the flood plains of rivers leaving the forest. They
also appreciate the forest as a source of rattan cane
and honey, the two main non timber forest
products. Ecotourism is presently bringing money
to the people living near the forest. Their
co-operation will be needed if the forest is to
survive.
ACKNOWLEDGEMENTS
We thank Operation Wallacea for funding and logistics. We also thank BKSDA South-East Sulawesi for
giving us permission to work in the Lambusango Wildlife Sanctuary and the Kakenauwe Nature Reserve. This
work was done under a research permit issued by the
Indonesian Institute of Sciences (LIPI). Sahimu, Tarahu, Kuseh, Imu and Mesilili, men of Labundo-bundo
village, were essential guides in the forest. We thank
the following people who have helped us with this
work: Nenni Babo, W.J. Baker, D. Blakesley, Tharada
286
REINWARDTIA
Blakesley, Marie Briggs, Eric Clement, J. Dransfield,
Christopher Fraser-Jenkins, Stuart Frost, Graeme Gillespie, Grant Harris, Aelys Humphreys, Rogier de Kok,
Rio Kornel, John Milsom, J.P. Mogea, Patricia Moss,
Titien Ng. Praptosuwiryo, Jean-Yves Rasplus, Alison
Richardson, H. Rustiami, A. Schuiteman, D. Seymour,
Steve Sudworth, Vicky Tough, T. Utteridge, the late
Trevor Walker and Paul Wilkin.
REFERENCES
BERG, C. C. & CORNER, E. J. H. 2005. Flora Malesiana Series 1 (Seed Plants) .17(2). 733 p.
COOK, J. M. & RASPLUS, J. Y. 2003. Mutualists with
attitude: coevolving fig wasps and figs. Trends in
Ecology and Evolution 18: 241‒248.
CORLETT, R. T. 1998. Frugivory and seed dispersal by
vertebrates in the Oriental (Indomalayan) Region.
Biological Reviews 73: 413‒448.
DE DECKKER, P., TAPPER, N. J. & VAN DER
KAARS, S. 2003. The status of the Indo-Pacific
warm pool and adjacent land at the last glacial
maximum. Global & Planetary Change 35: 25‒35.
DE KOK, R. P. J. & UTTERIDGE, T. M. A. 2010. A
Field Guide to the Plants of East Sabah. Richmond,
Kew Publishing.
DRANSFIELD, J., UHL, N. W., ASMUSSENLANGE, C. B., BAKER, W. J., HARLEY, M. H. &
LEWIS, C. E. 2008. Genera Palmarum. Richmond,
Kew Publishing.
GRUBB, P. J. 1971. Interpretation of the
„Massenerhebung‟ effect on tropical mountains.
Nature 229: 44‒45.
HARRISON, R. D. 2005. Figs and the diversity of
tropical rainforests. BioScience 55(12): 1053‒1064.
KEßLER, P. J. A., BOS, M. M., SIERRA DAZA, S. E.
C., KOP, A., WILLEMSE, L. P. M., PITOPANG, R.
& GRADSTEIN, S. R. 2002. Checklist of woody
plants of Sulawesi, Indonesia. Blumea Supplement
14: 1‒160.
KINNAIRD, M. F., O‟BRIEN, T. G. & SURYADI, S.
1999. The importance of figs to Sulawesi‟s imperilled wildlife. Tropical Biodiversity 6: 5‒18.
MABBERLEY, D. J. 2008. Mabberley’s Plant Book
(3rd ed.). Cambridge, Cambridge University Press.
MILSOM, J. 2000. Stratigraphic constraints on suture
[VOL.14
models for eastern Indonesia. Journal of Asian Earth
Sciences 18: 761‒779.
MORLEY, R. J. 2003. Interplate dispersal paths for
megathermal angiosperms. Perspectives in Plant
Ecology, Evolution and Systematics 6: 5‒20.
NATIONAL OCEANIC AND ATMOSPHERIC ADM I NI ST R AT I O N ( U SA) 2 0 1 4 . h t tp : / /
www.cpc.ncep.noaa.gov/products/analysis_monitor
ing/ensostuff/ensoyears.shtml
POWLING, A. 2009. The palms of Buton, Indonesia,
an island in Wallacea. Palms 52: 84‒91.
PRIMACK, R. & CORLETT, R. 2005. Tropical Rain
Forests. Oxford, Blackwell Publishing.
PURWANTO, E. 2008. Lambusango Forest Conservation Project (LFCP), Southeast Sulawesi, Indonesia
– Final Report (December 2008). Bau Bau LFCP.
SHANAHAN, M., HARRISON, R. D., YAMUNA, R.,
BOEN, W. & THORNTON, I. W. B. 2001.
Colonization of an island volcano, Long Island,
Papua New Guinea, and an emergent island,
Motmot, in its caldera lake. V. Colonization by figs
(Ficus spp.), their dispersers and pollinators. Journal
of Bio-geography 28: 1365‒1377.
VORIS, H. K. 2000. Maps of Pleistocene sea levels in
Southeast Asia: shorelines, river systems and time
durations. Journal of Biogeography 27: 1153‒1167.
WHITTEN, T., HENDERSON, G. S. & MUSTAFA,
M. 2002. The Ecology of Sulawesi. Hong Kong,
Periplus Editions (HK) Ltd.
WIDAYATI, A., JONES, S. & CARLISLE, B. 2010.
Accessibility factors and conservation forest
designation affecting rattan cane harvesting in
Lambusango Forest, Buton, Indonesia. Human
Ecology 38: 731‒746.
WIDAYATI, A. & CARLISLE, B. 2012. Impacts of
rattan cane harvesting on vegetation structure and
tree diversity of Conservation Forest in Buton,
Indonesia. Forest Ecology and Management 266:
206‒215.
WIDAYATI, A., CARLISLE, B., & SOEDRADJAT
2008. Trees of Lambusango Forest (ToLF). http://
www.numyspace.co.uk/~unn_szbc1/butontrees/
butontrees.html
ZONA, S. & HENDERSON, A. 1989. A review of
animal-mediated seed dispersal of palms. Selbyana
11: 6‒21.
INSTRUCTION TO AUTHORS
Scope. R einwardtia is a scientific irr egular jour nal on plant taxonomy, plant ecology and ethnobotany
published in December. Manuscript intended for a publication should be written in English.
Titles. Titles should be br ief, infor mative and followed by author ’s name and mailing address in oneparagraphed.
Abstract. English abstr act followed by Indonesian abstr act of not mor e than 250 wor ds. Keywor ds
should be given below each abstract.
Manuscript. Manuscr ipt is or iginal paper and r epr esent an ar ticle which has not been published in any
other journal or proceedings. The manuscript of no more than 36 pages by using Times New Roman 11, MS
Word for Windows of A4 with double spacing, submitted to the editor through
<reinwardtia@mail.lipi.go.id>. New paragraph should be indented in by 5 characters. For the style of
presentation, authors should follow the latest issue of Reinwardtia very closely. Author(s) should send the
preferred running title of the article submitted. Every manuscript will be sent to two blind reviewers.
Identification key. Taxonomic identification key should be pr epar ed using the aligned couplet type.
Nomenclature. Str ict adher ence to the Inter national Code of Botanical Nomenclatur e is obser ved, so
that taxonomic and nomenclatural novelties should be clearly shown. English description for new taxon
proposed should be provided and the herbaria where the type specimens area deposited should be presented.
Name of taxon in taxonomic treatment should be presented in the long form that is name of taxon, author’s
name, year of publication, abbreviated journal or book title, volume, number and page.
Map/line drawing illustration/photograph. Map, line dr awing illustr ation, or photogr aph pr efer ably
should be prepared in landscape presentation to occupy two columns. Illustration must be submitted as
original art accompanying, but separated from the manuscript. The illustration should be saved in JPG or GIF
format at least 350 pixels. Legends or illustration must be submitted separately at the end of the manuscript.
References. Bibliogr aphy, list of liter atur e cited or r efer ences follow the Har var d system as the
following examples.
: KRAENZLIN, F. 1913. Cyrtandraceae novae Philippinenses I. Philipp. J. Sci. 8: 163–179.
MAYER, V., MOLLER, M., PERRET, M. & WEBER, A. 2003. Phylogenetic position and generic
differentiation of Epithemateae (Gesneriaceae) inferred from plastid DNA sequence data. American J.
Bot. 90: 321–329.
Proceedings :TEMU, S. T. 1995. Peranan tumbuhan dan ternak dalam upacara adat “Djoka Dju” pada suku Lio,
Ende, Flores, Nusa Tenggara Timur. In: NASUTION, E. (Ed.). Prosiding Seminar dan Lokakarya
Nasional Etnobotani II. LIPI & Perpustakaan Nasional: 263–268. (In Indonesian).
SIMBOLON, H. & MIRMANTO, E. 2000. Checklist of plant species in the peat swamp forests of
Central Kalimantan, Indonesia. In: IWAKUMA, T. et al. (Eds.) Proceedings of the International
Symposium on: Tropical Peatlands. Pp.179-190.
Book
: RIDLEY, H. N. 1923. Flora of the Malay Peninsula 2. L. Reeve & Co. Ltd, London.
Part of Book : BENTHAM, G. 1876.
Gesneriaceae.
In: BENTHAM, G. & HOOKER, J. D. Genera
plantarum 2. Lovell Reeve & Co., London. Pp. 990–1025.
Thesis
: BAIRD, L. 2002. A Grammar of Kéo: An Austronesian language of East Nusantara.
Australian National University, Canberra. [PhD. Thesis].
Website
: http://www.nationaalherbarium.nl/fmcollectors/k/KostermansAJGH.htm). Accessed 15 February 2012.
Journal
Reinwardtia
Published by Herbarium Bogoriense, Botany Division, Research Center for Biology,
Indonesian Institute of Sciences
Address: Jln. Raya Jakarta-Bogor Km. 46 Cibinong 16911, P.O. Box 25 Cibinong
Telp. (+ 62) 21 8765066; Fax (+62) 21 8765062
E-mail: reinwardtia@mail.lipi.go.id
REINWARDTIA Author Agreement Form
Title of article
:
Name of Author(s) :
I/We hereby declare that:
My/Our manuscript was based on my/our original work.
It was not published or submitted to other journal for publication.
I/we agree to publish my/our manuscript and the copyright of this article is owned by Reinwardtia.
We have obtained written permission from copyright owners for any excerpts from copyrighted
works that are included and have credited the sources in our article.
Author signature (s)
Date
___________________________________________________________________________________________
Name