MaiaYali Nature Jor~mal.

2000, 54·4, 271 - 288

A Study oftbe Vegetati.on of the Forests in the Lower Kinabatang.an Region, Sabab, Malaysia

RAMESH BOONRATANA'

Abstracl: A study of the vegetation was carried out at two sites. Sukau and Abai, in. the Lower Kinabatangan regj-Oll in eastern Sabah, Sukau had a bi.gher tree densily and was more species-rich than Abai. Young leaf production was similar at both sites, but Abai had a higher rate of flower and frui [ production.

INTRODUCTION

The Lower Kinabatangan region (Fig 1) is mostly under forest on flat land that has been SUbjected to different degrees of disturbance. Open water, hills, villages and plantations are sparsely scattered throughout (payne 1989). This paper aims to describe the structure and composition of the forests at Sukau and Abai, in the

Figure t. Map of Sa bah.

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I p, O. Box 54, Chiang Mai University, Chiang Mai 50202, Thailand,

271

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lower region of the Kinabatangan River (Fig 2). This was a major component of a two-year study on the ecology and behaviour of Nasalis larvatus in the region, described in Boonratana (J 993).

The .Klnabatangan River

The Kinabatangan is Sabah's largest river, 560km long and with a catchment area of 16,800km1 (Scott 1989). Its main source of water is run-off from mountains in the south-eastern interior. The lower Kinabatangan River meanders through a large flat floodplain, much of which is subjected to seasonal flooding, resulting in lowstature forest with little timber of commercial value. The Kinabatangan floodplain, measuring approximately 280;000ha, is the largest and most important wetland in Sabah (Scott] 989). The river empties into a large delta, which is brackish and tidal. This deltaic area has the largest intact area of mangrove in the state, covering approximately 40,500ba (Scott 1989). Tidal range in the lower Kinabatangan River averages I.! m, but the river level can rise Sm overnight if there is widespread rain upriver. There are many ox-bow lakes between the middle reaches of the river and the coastal plains, which are at various stages of infilling (Boonratana L993).

Soilsand Geology

The soil in the flood-prone area is alluvium which lies on top of sedimentary rocks, mainly sandstones with some mudstones and limestones. Raised alluvial terraces and plains are found along the banks of the main river and its larger tributaries. Behind and scattered among these terraces and plains is low-lying freshwater swamp forest. The alluvial soils are only thousands of years old, which is quite recent in geological terms. Scattered throughout the region are small but steep, hard sandstone hills, which protrude from massive layers beneath. The remainder of the Kinabatangan landscape is made up of soft standstones and mudstones. An intere ling feature of the floodplain is the limestone outcrops. The sandstone and limestone hills were formed during the Miocene, between 26 and 7 million years ago (Payne 1989).

Climate

The area has a humid tropical climate with an average annual rainfall of about 2600mm and mean diurnal temperatures ranging from 22°C to 32°C (Scott 1989). Mean temperatures do not vary much; the mean monthly minimum for both 1990 and 1991 was 23.7°C, whereas the mean monthly maximum for 1990 was 32.9°C, and for 199\, 33°C (Boonratana 1993). The total rainfall for 1990 was 1816mm and for 1991, 2975mm (Fig 3). It rained for 159 days in 1990, and for 183 days in

. 199 J . This means that it rained on average once in two days. There were two dry months in ] 990, February and April, with less than 50mm of rain, whereas in 1991 there was only one dry month, March (Boonratana 1993).

272

Figure 2. Natural features of Sukau and Abai regions.

~ Mangrove and mps forest

~

Mam river and tributary

R. River
L. Lake
0 5 10
I
Kilometres Hills and outcrop. above 70m elevation

* Sukau Village

D Fresn.W.3ter. swamps and : riverine rarest

Lowland dlperocarp fO\'<'51 dod dry land below 70m elevauon

[c:v(;1 Ox-bow lakes

273

Figure 3. Mean monthly rainfall for 1990-199] at Sandakan Airport (c. 40km from Abai and 50km from Sukau).

800
~ 600
E
E
'-'
ro 400
'+-
C
m
rr: 200
0 JFMAMJJASONDJFMAMJJASOND Months

Logging History

The lower Kinabatangan was subjected to cotnmercial logging from the early 195'05 until the present (Payne 1989). Virtually all of the forests of the Kinabatangan area have been logged at least once. In the early 19805, logging of mangroves in eastern Sabah was widespread, mainly for woodchips, It was banned in early 1987 (Bennett 1991), More than 60,000ha of the lowland rainforest in the flood-free zone of the lower Kinabatangan region is being replaced by cocoa and oil-palm plantations (Boonratana 1993; Boonratana and Sharma ]994).

METHODS

Fieldwork was conducted from January 1990 to December 1991 inclusive. At the Sukau study area, Zm-wide straightline transects were placed on both sides of the Menanggul River at the Ist, 3rd, 5th, 7th and 9th kilometres, measured from the river mouth. All trees of at least 30cm girth at breast height (g.b.h.) located within the transects were measured, tagged and identified. A 30cm g.b.h. was used to aUow comparison with other plots in Sabah (Davies 1984). Girths of buttressed tree were measured immediately above the buttresses (Bennettt 1983; Davies 1984). Trees that were partIy within the transects were included if at least 50% of their bases were within the strip. The basal area of each tree was calculated from the girth at breast height. The basal area is a good indicator of tree size and foliage biomass (Gross and net primary production and growth parameters 198 I). The first 50 trees on each side of the river of every transect were used for phenology" making a total of 500 trees (Boonratana 1993).

274

"

At the Abai study area, straight line transects were similarly placed on both sides of the Merah River, at the 3rd, 6th and 9th kilometres, measured from the: river mouth. The Ise and 5th kilometres were not selected because the ISland 5th transects had few or no standing trees. Being equidistant from the 3rd and 9th kilometre, the 6th kilometre was selected instead of the 5th and 7th kilometres.

Every month from March 1990 to December 1991 the transects were examined for the presence of mature and young leaves, ripe and unripe fruits, and flowers. Young leaves were defined as the new leaves that had yet to attain full size. were sometimes stm unfurled, and whose colouration was from pinkish-red to light green. Phenology was monitored as the percentage of trees and not the percentage of plant parts per tree. This was becanse many of the trees were covered with Hanas and creepers, making it impossible to determine the abundance of the different plant parts per tree.

RESULTS

Sukau Study Area

At Sukau, the principal vegetation types in the flood-prone areas were riverine forest and freshwater swamp forest. There was also some open reed swamp. In the flood-free zone, there were remnants of pristine lowland dipterocarp forest, loggedover swamp forest and burnt lowland dipterocarp forest, and cocoa and oil-palm p lantati ons.

Structure. The 2m- wide transects at Sukau covering a total area of 1.96ha, contained 1378 trees. Thus, the density of trees of more than or equal to 30cm g.b.h. was 703 trees/ha. Most of the trees in the transects at Sukau had girths of less than 140cm, with the maximum recorded girth being 447cm (Fig 4). The mean and median girths of trees in the transects were 65Acm and 49.0cm respectively. The total basal area for trees with more than or equal to 30cm g.b.h. was 67.77m2 or 34.58m2Iha. The maximum basal area recorded for anyone of the trees in the transects was 15,896cm2, and the mean. basal area of an trees was 492cm2 (Boonratana 1993).

Composition. The 1378 trees in the transects at Sukau comprised 109 species from 37 families. The species, number of stems and basal area of each are listed in Appendix I. To assess species richness, the rate at which the number of tree species increased with increasing tree sample ize was plotted (Fig 5). Although the curve was close to being level by the end, there was still a tendency for the number of species to increase as the total sample size increased.

Euphorbiaceae was the most abundant tree family in terms of the number of stems at Sukau and had the greatest basal area (Table 1). Although Dipterocarpaceae ranked seventh in terms of abundance, it had the next greatest basal area. Tree families that were also abundant in tenus of number of stems included CJusiaceae Myrtaceae Rubiaceae, Lauraceae and Dilleniaceae.

The most common tree species in the transects, Syzygium bankense (Myrtaceae) made up 8.8% of the total number of trees of more than or equal to 30cm g.b.h.

275

Figure 4. Frequency distribution of girths at breast height of trees in the transects at SUKaU (n = 1378).

% trees

100~------------------------------------------------'

10

0.1

Girth at breast height (em)

FigureS. Number of tree species against the number of trees sampled in tile transects at Sukau (n = 1378).

No. of species

,.20 r------------------------~

O~----~._------._----_,,_----_.r_----~------_.------~

o

200

400

600

800

1000

1200

1400

No. of trees

276

Table t. Abundance and basal area of tree families in botanical transects at Sukau.

Family % Total Trees % Total Basal Area Mean Basal Area (ern')
Alangiaceae 0.3 0.09 1.54.72
Anacardiaceae 4.1 2.80 333.49
Annonaceae 0..9 1.24 643.93
Apocynaceae 0.1 0.04 152.19
Bornbacaceae 0.1 0 . .14 477.49
Burseraceae 3.7 3.27 434.94
Combretaceae 0.6 1.20 1014.75
Datiscaceae 0.4 0.27 307.10
Dijleni aceae 7.8 2.83 180.63
Dipterocarpaceae 6.7 10.21 742.19
Bbenaceae 2.8 2046 427.84
Elaeccarpaceae 0.9 0.76 397.08
Euphorbi aceae 13,4 14.32 527.57
fagaceae 0.1 0.,03 231.98
Placourtiaceae 15 2.16 732.21
Guttiferae 9 . .0 8.38 457.97
Jrvingiaceae 0.2 0.045 1021.80
Lauraceae 8.2 8.35 500.84
Lecythidaceae 2.2 0..99 215.36
Leguminosae 1.6 2.04 626.28
Melastomataceae 4 .. 0 6.42 791.63
MeLiaceae 0.8 0.59 361.93
Moraceae 1.3 2.76 1040.67
Myristicaceae 1.0 0.73 350..94
Myrtaceae 8.8 8.90 498.62
Olaeaceae 0.1 O.!4 980.19
Polygalaceae 1.3 1.20 451.10
Rhi zophoraceae 0 . .1 0.,04 l51.79
Rosaceae 0.3 0..23 386.72
Rubiaceae 8.5 9.07 520.68
Rutaceae 0.4 0.Q7 99.57
Sapindaceae 0.3 0.10 164.00
Sapctaceae 1.3 0.50 [87.81
Sonneratiaceae '0.4 0.69 783.90
Sterculiaceae 2.2 2.55 556.75
Tiliaceae 2,[ 1.71 398,96
Verbenaceae 2.3 2.27 495.77 277

(Table 2). The next most common tree species was Dillenia grandifolia (Dilleniaceae), followed by Ludekia bernardoi (Rubiaceae), Litsea elliptica (Lauraceae) and Mallotus muticus (Euphorbiaceae). In terms of basal area, however, M. muticus ranked the highest at 10.95%, followed by S. bankense at 8.9%.

Table 2 .. Abundance and basal area of the IS most common tree species at Sukau,
Family Species No. of % Total Basal Area % Total
Trees, Trees (em') Basal Area
Myrtaceae Syzygium bankense 122 8.9 60313.75 8.90
Dilleniaceae Dillenia grandifolia 98 7.1 15680.12 2.31
Rubiaceae Ludekia bernardoi 90 6.5 36811.30 5,43
Lauraceae Litsea elliptica 81 5.9 28877.22 4.26
Euphorbiaceae Madotus muticus 76 5.5 74230.78 10.95
Anacardi aceae Buchanania insignis 56 4.1 18801.98 2./7
Melastornataceae Ptemandra galeata 55 4.0 43539.54 6.42
Burseraceae Conarium apertum 51 3.7 22182.11 3.27
Guttiferae Calophyllum borneense 50 3.6 14084.78 2.08
Guttiferae Cratoxylum sumatranum 47 3.4 35801.92 S.28
Euphorbiaceae Glochidion borneense 39 2.8 13417.40 1.98
Ebenaceae Diospyros pendula 38 2.8 16593.65 2,45
Dipterocarpaceae Vatica oblongifolia 38 2.8 12973.86 1.91
Lecythidaceae Barringtonia lanceolata 31 2.3 6676.22 0.99
Rubiaceae Neolamarckia cadamba 27 2.0 24542.31 3.62
Total 899 65.3 424503.87 62.72 Phenology. The production of young leaves was high throughout most of the study period, being exceptionally high from October 1990 to January 1991, and from March to December 199] (Fig 6). These peaks coincided with wet periods, except March 199 J, which just followed a wet period .. Leaf production was moderate from March to May 1990, just after a dry period. There was, however, no correlation between young leaf production and rainfall (rs ;:::; 0.163, n = 22, p > 0.05). The minimum recorded leaf production during the study period was 58% of trees in May 1990, just after a dry period.

There was a distinct peak in flower production in April 1990, when 20.6% of trees in the transects bore flowers, coinciding with a dry period (Fig 7). There were two peaks in flower production for 1991, in April (16.4%), just after a dry period, and in October (16.8%), when rainfall was moderate. No correlation existed between rainfall and flower production (rs ;:::; - 0,170, n = 22, p > 0.05). During the study period, flower production was lowest in December 1990, when only 1.4% of trees bore flowers.

278

Figure 6. Young leaf production at Sukau (n = 500) and Abai (0 = 300).

100
90
so
70
C 60
ill
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Months
.Sukau OAbai I "Figure 7. Plower and fruit production at Sukau (n = 500) and Abai In = 300),

45 -.--- -- - - .. -- -- •. -- - •. -- -- -- , .. ------ .. ----- -- ---.-

40~----------------------------------~------or~nr_----~

35+---------------------------------~--------~r-r-------~ 30 +----------------~--------~__i t-- I--:-----;t--

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Ell Sukau fruits Ii!! Abai fruits

There was a distinct fruiting season from June to September 1990, with a peak in June (14.2% of all trees). This was during a wet period. There were two fruiting seasons in ] 991, during a wet and a dry period, from January to April, and during a wet season, from July to September. There was no correlation between fruit production and rainfall (r s = 0.1] 6, n = 22, P > 0.05). The peak in fruit production for 1991 was in August, when 12% of trees bore fruits. The lowest fruit production was in March] 990, when only 1.4% trees bore fruits.

Abai Study Area.

The principal vegetation at Abai is mangrove forest" with extensive stands of Nypa fruticans (Palmae) at the inland edge and the upper tidallimit of the estuaries. Upriver from this, beyond the influence of seawater, riverine forest and lowland

279

· .

swamp forest dominate in the seasonally flood-prone ZOne. Transitional forest exists where mangrove forest intergrades to riverine forest. There are also some forests on steep hills and flat ground. The trees at Abai are highly clumped, set between stands of pure N. fruticans. The tree species and their proportions in the botanical transects at Abai are given in Appendix If.

Structure. The number of trees in the transects totalled 300, covering an area of 2906m1 and having a total basal area of 12.0Sm1 .. Thus, the density of trees with more than or equal to 30cm g.b.h. was estimated at 1034.5 trees/ha, and basal area was estimated at 41.64m2lha. Most of the trees in the transects at Abai had girths of less than 120cm, with the maximum girth being 408cm (Fig 8). Mean and median girths of trees in the transects were 59 and 47cm respectively. The maximum and mean basal area of tree in the transects were 13.243cm2 and 402.6cm2 respectively (Boonratana, 1993).

Composition. The botanical transects at Abai contained 300 trees. comprising a total of 42 species belonging to 26 families. The rate at which the number of tree species increased with increasing tree sample size was plotted to assess species richness (Fig 9). From the figure, very little increase in the number of tree species is expected.

Melastomataceae was the most abundant tree family at Abai, followed by Myrtaceae, Elaeocarpaceae, Verbenaceae, Anacardiaceae and Rhlzophoraceae (Table 3). Although trees belonging to the family Rhizophoraceae were not very abundant, it had the greatest basal area of all families (J 9.5] % ofthe total). Moraceae also was not abundant, but had the next greatest basal area.

Figure 8. Frequency distribution of girths at breast height of trees in the transects at Abai (n = 300).

% trees

30 60 70 90 110 130150 1701·90'210 230250270.290 310 330360 370390 410

Girth at breast height (em)

280

Figure 9. Number of tree species against the number of trees sampled m the transects at Abai (n = 300).

No. of species

50.-----------------------------------------------~

40

30

20

10

O~-----_.----_,~----._----_.------r_----._----~

o

50

100

150

200,

.250

300

350

No. of trees

The most common tree species at Abai was Pternandra galeata (Melastomataceae), which made up 14.0% of the total trees (Table 4). This was followed by Syzygium bankense (Myrtaceae), Elaeocarpus sp. (Elaeocarpaceae), Yitex pinnata (Verbenaceae), and Buchanania insignis (Anacardiaceae),

Phenology. Young leafproduction was high from December 1990 to January 1991. and from March to December 1991 (Fig 6). These peaks coincided with periods of high rainfall, except in March when it was dry. April 1990 was the month with the lowest leaf production, when only 43.3% of trees in the transects produced young leaves. This coincided with the dry period for that year. There was a positive correlation between young leaf production and rainfall (rs "" 0.491, n = 21, P < 0.05), implying the importance of rainfall for leaf production.

In 1990, flower production was highest in October, during a wet period, when 29.3% of trees in the transects bore flowers (Fig 7). There were two flowering seasons in 1991, from March to May and from July to September. December 1990 had the lowest flower production, when only 8.4% oftrees in the transects flowered, while September 1991 had the highest at 41 % of trees .. No correlation, however, existed between flower production and rainfall erg = - 0.183, n = 21, P > 0.05), implying that rainfall did not influence flower production.

Fruit production was high from December 1990 to February 1991, in April 1991,. and from August to October 1991.llighest fruit production for 1990 was in December, at 28.4%, and for 1991 it was in January and April, both at 29.7%. The lowest fruit production was in April 1990 at 6%, coinciding with a dry period. Peak fruit production was during the wet periods. A significant positi ve correlation existed between fruit production and rainfall (rs = 0.513, n = 21, P < 0.05), implying that rainfall strongly influenced fruit production.

281

'Table 3. Abundance and basal area of tree families in botanical transects at Abai,

Family % Tree Sterns % Tree Basal Area Mean Basal Area (crnZ)
Anacardiaceae 7.7 5.48 287.58
Annonaceae 3 . .0 2.99 400.62
Apocynaceae 3..3 1.50 181.00
Burseraceae 4 . .0 3.16 318.06
Cornbretaceae 0.7 2.01 1212.17
Dilleniaceae 3.0 0.79 106.46
Oi pterocarpaceae 2.3 2.66 458.13
Ebenaceae 2.7 0.85 127.75
Elaeocarpaceae 8 .. 3 6.90 333.23
Buphorbi aeeae 3.3 2.03 272.17
Flacourtiaceae 1.0 0.76 305.62
Guttiferae 4.3' 1.90 176.56
Lauraceae 1.3 1.23 372.71
Lecythidaceae 0.3 0 . .09 108.91
Leguminosae 0.3 0.10 121.0.0
Melastomataceae 14.0 8 . .05 231.50
Moraceae 2.7 14.71 222.0.57
Myrtaceae L1.3 5.99 212.92
Polygalaceae 4 . .0 3.61 363.66
Rhizophoraceae 6.3 19.86 1262.33
Rubiaceae 1.7 3.91 943.12
Rutaceae 0.3 0.29 336.12
Sapotaceae 1.7 0.66 132.92
Sterculiaceae 2.3 4.34 748.78
Thymelaeaceae 1.7 0.89 215.24
Verbenaceae 8.3 5.26 254.08 Table 4. Abundance and basal area of the 15 most common tree species at Abai,

Family Species No, of % Total Basal Area % Total
Trees Trees (em') Basal Area
Melastornataceae Pternandra galeata 42 14 . .0 9723.22 8.05
Myrtaceae Syrygium bankense 34 11.3 7239.14 5.99
EJ aeocarpaceae Etaeocarpus sp. 25 8.3 8330.86 6.90
Verbenaeeae Vitex pinnata 24 8.0 5880..19 4.87
Anacardiaceae Buchanania insignis 22 7.4 4752.0.9 3.93
Rhizophoraceae Brugulera sexangula 15 5.0 2.3558.60. 19.51
Burseraceae Canarium apertum 12 4.0. 3816 . .70. 3 . .16
Polygalaceae Xanthophyllum rufum 12 4.0 4364.96 3.6,1
Apocynaceae Alstonia angustifolia ]0 3.3 1810.02 1.50.
Dilleniaceae Dillenia excelsa 9 3.0 58.15 0.79
Annonaceae Polyalthia glauca 9 3.0 3605.57 2.99
Moraceae Ficus condensa 8 2.3 17189.84 14.23
Buphorbiaceae Baccaurea pubera 6 2.0 1627.68 1.35
Guttiferae Calophyllum borneense 6 2.0. 855.3 1 0..71
Ebenaceae Diospyros pendula 6 2.0 797.93 0..66
Total 240 79.6 945.0.9.16 78.25
282 DISCUSSION: COMPARISON BETWEEN SUKAU AND ABAI

The forest at Sukau was more species-rich than that at Abai. The Shannon-Weiner diversity index (H') at the species level was calculated for each study area. This index showed that the forest at Sukau (H' :::; 3.72) was more diverse than that at Abai (H' :::; 3.13), although this may be due to the larger sample size at Sukau,

Twenty-five families were common to both areas. These made up 98.3% of the total trees in the transects at Abai and 90.2% of those in Sukau. The basal area of species common to both areas was 99.1 % of the total for Abai, and 94.8% of the total for Sukau. Melastomataceae (14%) had the highest percentage of trees in the transects at Abai, whereas at Sukau it was Euphorbiaceae (13.4%). The highest percentage in terms of basalarea at Abai was Rhizophoraceae (19.9%), while at Sukau it was Eupborbiaceae (14.3%).

When the 15 most common species at Sukauand at Abai were compared, only six species were common to both areas. They were Syzygium bankense (Myrtaceae), Buchanania insignis (Anacardiaceae), Pternandra galeata (Melastomataceae), Canarium apertum (Burseraceae), Calophyllum borneense (Clusiaceae) and Diospyros pendula (Ebenaceae). At Abai, the most common tree in the transects was Pternandra galeata (14%), but the tree with the highest percentage of total basal area was Bruguiera sexangula (19.5%). At Sukau the most common species was Syzygium bankense (8.8%) •. and the tree with the highest percentage basal area was Mallotus muticus (11 %). These differences between sites can be attributed to their distances from the sea. Sukau was much further inland and not affected by seawater, whereas Abai was closer to the coast and inundated by seawater. The straight line transects at Sukau and Abai were primarily located in riverine and mangrove forests, respectively.

A comparison of the phenological patterns at Abai and Sukau indicates that there was no significant difference in young leaf production between the two sites (Mann-Whitney U = 203.5, "1 :::; 21, 02 :::; 22, p > 0.05). More flowers were produced at Abai than. Sukau during most months and the trend was significant (U :::; 38, n 1 :::; 21, 02 == 22. P < 0.05). The trend throughout the year, however, was similar to that at Sukau, In October 1990, however, there was a peak in flower production at Abai, but not at Sukau.

Except in March 1990, the trend in fruit production was similar at both sites, although production was generally less at Sukau. This difference was significant (U = 65.5, n 1 = 21, n2 :::; 22, p < 0.05). At Abaiin 1991, there was more growth in March. April, May. August and September. Sukau had more growth in April and October 1991. Fruit production at Sukau and Abai was much higher than at Sepilok (Davies 1984) and Kuala Lompat (Bennett (983), which might partly explain the high primate density and diversity in the Lower Kinabatangan (Boonratana 1993; Boonratana and Sharma 1994).

CONCLUSION

The Lower Kinabatangan region does not have much pristine vegetation, but it has a relatively large tract of riverine, freshwater and mangrove forest, supporting a

283

high density and diversity of wildlife. The forest at Sukau, which was predominantly riverine, had a higher tree density and was more species-rich than at Abai, which was predominantly mangrove. At Sukau,if the sample size in the transects was increased, the number of species recorded was also expected to increase, There was no significant difference in young leaf production at Sukau and Abai, but flower and fruit production were higher at Abai. The differences in tree density and composition between sites were most likely due to the influence of seawater at the Abai study area.

ACKNOWLEDGEMENTS

I acknowledge Datuk Wilfred Lingharn, former Permanent Secretary, Ministry of Tourism and Bavironmental Development, and Me Mahedi Andau Director, Sabah Wildlife Department for permission to conduct this study and for support. Trees in the transects were identified by Me Paul Langgi of the Sabah Wildlife Department. Mr Dionysius S. Sharma of World Wide Fund for Nature (Malaysia) provided excellent field assistance. Thts study was supervised by Dr Elizabeth L. Bennett and Prof Warren Y Brockelman, and funded by the Wildlife Conservation Society.

REFERENCES

Bennett, E.L. 1983. The banded langur: ecology of a colobine in We~t Malaysian rainforest Unpublished doctoral dissertation, Cambridge University.

Bennett, EL. 1991. Diurnal Primates. Pp, 150--172 in: R. Kiew (ed.) The State of Nature Conservation in Malaysia. United Selangor Press Sdn. Bhd., Kuala Lumpur.

Bocnratana, R. 1993. The ecology and behaviour of the proboscis monkey (Nasalis larvams) in the Lower Kmabatangan, Sabah, Unpublished doctoral dissertation, Mahidol University.

Boonratana, R. and D .. S. Sharma .. 1994. Conservation of proboscis monkeys in the lower Kinabatangan, Sabah, Pp, 93-JOO in: B. Thierry. R. J. Anderson. J. 1. Roeder and N. Herrenschmidt (eds.), Current Primatology, Volume I: Ecology and Evolution. Strasbourg: Universite Louis Pasteur.

Davies, A.G, 1984 .. An ecological study of the red leaf monkey tPresbytis rubicunda; in ttLe dipteroearp forest of northern Borneo, Unpublished doctoral dissertation, Cambridge University.

Gross and net primary production and growth parameters. 198 L Pp, 233-248 in: Tropical Forest Ecosystems. UNESCO, Paris.

Payne. 1. 1989. A tourism feasibility study for the proposed Kinabatangan Wildlife Sanctuary. Kuala Lumpur, WWF Malaysia.

Scott, D.A., ed. 1989. A Directory of Asian Wetlands, ruCN, Gland/Cambridge.

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APPENDIX L Tree speciesand their proportions in the botanical transects at Sukau (n = 1378).
Family Species No. of % Total % Total
Trees Trees BasalArea
Alangiaceae Alangiun: ebenaceum 4 0.3 0 .. 09
Anacardiaceae Buchanania insignis 56 4.1 2.77
Anacardiaceae Dracontomelon dao 1 0.1 0.03
Annonaceae Cananga odorata 3 0.2 0.77
Annonaceae Phaeanthus crassipetalus 3 0.2 0.14
Annonaceae Polyalthia glauca 7 0.5 0.33
Apocynaceae Alstonia angustifolia 0.1 0.02
Apocynaceae Alstonia angustiloba 0.1 0.02
Bombacaceae Coelostegia griffithii 0.1 0.04
Bombacaceae Durio grandiflorus I 0.1 0.12
Burseraceae Conarium apertum 51 3.7 3.27
Chrysobalanaceae AtwlG racemosa 2 0.[ 0.15
Chrysobalanaceae Parinari oblongifolia 2 0.1 0.08
Combretaceae Terminaiia foetidissima 8 0.6 1.20
Datiscaceae Octomeles sumatrana 6 0.5 0.27
Dilleniaceae Dillenia excelsa I 0.1 0.35
Dilleniaceae Dillenia grandifolia 98 7.1 2.30
Dilleniaceae Dillenia indica 7 0.5 0.18
Dipterocarpaceae Dipterocarpus applanatus 4 0.3 0.46
Dipterocarpaceae Dipterocarpus caudiferus 7 0.5 0.94
Di pterocarpaceae Diprerocarpus stellatus 0.1 0.Q3
Diprerocarpaceae Dryobalanops lanceolata 2 0.1 0.28
Di prerocarpaceae Hopea nervosa 2 0.1 0.63
Di pterocarpaceae Parashorea malaanonan I 0.1 0.03
Di pterocarpaceae Parashorea tomentella 2 0.1 0.04
Diplerocarpaceae Shorea beccariana 7 0.5 0.24
Dipterocarpaceae Shorea faguetina 0.1 0.01
Di pterocarpaceae Shorea falciferoides 2 0.1 0.04
Di pterocarpaceae Shorea inappendiculata I 0.1 0.08
Dipterocarpaoeae Shorea leprosula 8 0.6 1.64
Dipterocarpaceae Shorea macroptera 4 0.3 0.64
Dipterocarpaceae Shorea platyclados 1 0.1 0.16
Dipterocarpaceae Shorea scrobiculata 2 0.1 0.08
Diprerocarpaceae Shorea smithiana 2 0.1 2.40
Di pterocarpaceae Shorea xanthophylla 1 0.1 0.05
Dipterocarpaceae Yatica umbonata ssp. acrocarpa 7 0.5 0.56
Dipterocarpaceae Vatica oblongifolia 38 2.8 1.91
Ebenaceae Diospyros pendula 38 2.8 2.45
Elaeocarpaceae Elaeocarpus sp. 13 0.9 0.76
Euphorbiaceae Baccaurea bracteatd" 7 0.5 0.18
Euphorbiaceae Baecaurea parviflora 27 1.9 0.49
Euphorbiaceae Baccaurea pubera 2 0.1 0.05
Euphorbiaceae Erismanthus sp, 2 0.[ 0.02
285 Family Species No. of % Total % Total
Trees Trees Basal Area
Euphorbiaccae Glochidion borneensc 39 2.8 1.98
Euphorbi aceae Koilodepas laevigatum 2 0..1 0.0.4
Euphorbiaceae Macaranga coni/era 18 1.3 0..38
Euphorbiaceae Macaranga hypoleuca 11 0..8 0..23
Euphorbiaceae Mallows muticus 76 5.5 10.95
Fagaceae Luhocarpus lucidus I C.I 0.0.3
Fl acou rti aceae H omalium foetidum 6 0..4 0..46
Flacourtiaceae Hydnocarpus woodli 14 1.0 1.70.
Guttiferae Cratoxylum. sumatranum 47 3.4 5.28
Guttiferae Cratoxylum sp. 1 0. . .1 0..0.4
Guttiferae Calophyllum borneense 51 3.7 2.0.8
Guttiferae Calopkyllum canum 12 0..9 0..57
Gurtiferae Garcinia bancana 12 0..9 0..39
Guttiferae Garcinia beccarii I 0..1 0..0.2
lrvinginceae lrvingia malayana 3 0..2 0..45
Lauraceae A lseodaphne bancana 0..1 o..Q3
Lauraceae Beilschmiedia micrantha 0..1 o.oi
Lauraceae Dehaasia cUlleatCl D.] 0..12
Lauraceae Eus ide mxy ton zwageri 10. 0.8 2.86
Lauraceae Litsea firma 15 1.1 0.83
Lauraceae Litsea elliptica 8] 5.9 4.26
Lauraceae Nothaphoebe kingiana 3 0.2 0.21
Lauraceae Phoebe elliptica 0.] 0.Q3
Lecyt hidaceae Barringtonia lanceolata 31 2 .. 3 0.99
Legurninosae Albiria chinensis 0.1 0.0.4
Legurninosae Albisia pedicel/ala 0.,1 0.D2.
Leguminosae Fordia cf. gibbsiae 5 0.4 0.16
Leguminosae Intsia palembanica 0.1 0.42
Legurninosae Koompassia excelsa 0.) 0..24
Legumi nnsae Parkia speciosa 0.1 0.l4
Leguminosae Sindora irpicina 5 0.4 0..1.4
Legumi.nosae Spatholobus hirsutus I 0.1 0.02
Legurni nosae Sympetalandra borneensis 6 0.4 0..56
Lythraceae Duabanga moluccana 6 0.4 0.69
Melastomataceae Pternandra galeata 55 4.0 6.42
Meliaceae Aglaia rubiginosa 6 0..4 0,14
Meliaceae AgJaia malaccensis 3 0..2 0.15
Meliaceae Chisocheton pentandrus 2 0.1 0.30
Moraceae Artocarpus dadah 0.1 0.D2
Moraceae Artocarpus elasticus I 0,1 0.04
Moraceae Artocarpus tamaran 4 0.3 0.09
Moraceae Ficus condensa l2 0..9 2.60
Myristicaceae Knema latifolia 14 1.0 0..73
Myrtaceae Syzygium bankense 122 8.9 0..14 286

Family Species No. of % Total % Total
Trees Trees Basal Area
Olacaceae Scorodocarpus bomeensis I 0.1 0.14
Polygalaceae Xanthophyllum rafum 18 1.3 1.20
Rhizophoraceae Carallia brachiata 2 0.1 0.04
Rubiaceae Neolamarckia cadamba 27 2.0 3.62
Rubiaceae Ludekia bernardo! 90 6.5 5.43
Ruraceae Meliocope confusa 5 0.4 0.07
Sapindaceae Nephelium cuspidatum 1 0.1 0.01
Sapindaceae Paranephelium xestophyllum 3' 0.2 0.09
Sapotaceae . Palaquium rostratum 6 05 0.11
Sapotaceae Payena Lucida 12 0.9 0.39
Sterculiaceae Heritiera borneensis 3 0.2 0.09
Sterculiaceae Heritiera elata 2 0.1 0.03
Sterculiaceae Heritiera littoralis 1 0.1 0.21
Sterculiaceae Pterospermum elongatum 25 1.8 2.21
Tiliaceae Microcos antidesmifolia 21 1.6 0.99
Tiliaceae Pentace adenophora 1 0.1 0.04
l'iliaceae Pen/ace bomeensis 3 0.2 0.24
Tiliaceae Pentace erectinervia 4 0.3 0.44
Verbenceae Teijsmanniodendron bo goriense 2 0.1 0.13
Verbenceae Teijsmanniodendron sympliciodes 10 0.7 0.25
Verbenceae V!tex pinnata 20 1.5 1.89 287

APPENDIXll. Tree species and their proportions in the botanical transects at Abai (n = 300).
Family Species No .. of % Total % Total
Trees Trees Basal Area
Anacardiaceae Buchananla ills ignis 22 7.4 3.93
Anacardiaceae Mangifera pajang I 0..3 L55
Annonaceae Polyalthia glauca 9 3.0 2.99
Apocynaceae Alstonia angustifolia 10 3.3 1.50
Burseraceae Conarium apertum 12 4.0 3.16
Combreraceae Terminalia foetidissima 2 0.7 2.0.1
Dilleniaceae Dillenia excelsa 9 3.0. 0.79
Dipterocarpaceae Shorea leprosula 3 1.0. 0..68
D ipterocarpaeeae Vatica oblongifolia 3 1.0 0.21
Dipterocarpaceae Varica rassak 0..3 0..13
Ebenaceae Diospyros evena 2 0..7 0..19
Ebenaceae Diospyros pendula 6 2.0 0..66
Elaeocarpaceae Elaeocarpus sp. 25 8.3 6.90
Euphorbiaceae Baccaurea pubera 6 2.0. 1.35
Euphorbiaceae Glochidion bomeense 2 0..7 0.58
Euphorbiaceae Macaranga conifera 2 0..7 0..10
Flacourtiaceae Homalium [oetidum 3 1.0. 0..76
Guttiferae Calophyllum bomeense 6 2.0 0.71
Guttiferae Calophyllum canum 4 1.3 0.85
Guttiferae Calophyllum sp. 2 0..7 0.21
Guttiferae Garcinia bane alia 0..3 0.13
Lauraceae Litsea elliptia 4 1.3 1.23
Lecythidaceae Barringtonia lanceolata 0..3 0..09
Leguminosae Sindora irpicina 0.3 0.10
Melastornataceae Pternandra galeata 42 14.0. 8.05
Moraceae Artocarpus dadah 1 ·0..3 0.48
Moraceae Ficus condensa 7 2.3 14.23
Myrtaceae SYl.ygium bankense 34 11.3 5.99
Polygalaceae Xanthopltyllum rufum 12 4.0 3.61
Rhlzophoraceae Bruguiera sexangula 16 5.3 19.51.
Rhizophoraceae Ceriops tagal 3 1.0 0.35
Rubiaceae Ludekia bernardoi 5 1.7 3.91
Rutaceae Melicope confusa 1 0.3 0.29
Sapotaceae Madhuca motleyana 0.3 0.08
Sapotaceae Madhuca burckiana 0.3 0.'12
Sapotaceae Palaquium rostratum 2 0.7 0.35
Sapotaceae Pouteria obovata 1 0.3 0.11
Sterculiaceae Heritieria littoralis 7 2.3 4.34
Thyrnelaeaceae Gonystylus keithii 5 1.7 0.89
Verbenceae Teijsmanniodendron symp~ociodes 1 0.3 0.39
Verbenceae Vifex pill/lata 24 8.0 4.87 288