TROPICS Vol. 18 (1) Issued May 31, 2009

Reestablishment of after forest fire in East Kalimantan,

1 1 2 Natsuki M. WATANABE , Eizi SUZUKI , and Herwint SIMBOLON

1Department of Earth and Environmental Sciences, Faculty of Science, Kagoshima University, 1-21-35, Korimoto, Kagoshima 890-0065, Japan 2Research Center for Biology, Indonesian Institute of Science (LIPI), Gedung Botani dan Microbiologi, JL. Raya Jakarta-Bogor Km.46, Cibinong, Java Barat Corresponding author: Natsuki M. Watanabe Present address: Sumitomo Forestry Co., Ltd., Tsukuba Research Institute No. 3-2, Midorigahara, Tsukuba, Ibaraki, 300-2646, Japan TEL: +81-29-847-0153 FAX: +81-29-848-1100 Email: [email protected]

ABSTRACT We quantified the reestablishment of rattans (climbing palms) after severe forest fires INTRODUCTION in 1997–1998 caused by the El Niño Southern During the last few decades, the dry periods associated Oscillation event in Bukit Bangkirai, East with El Niño Southern Oscillation events caused wildfires Kalimantan, Borneo. We established a 1-ha study and damaged the tropical rainforests in Indonesia. The plot in unburned forest (K1) and two 1-ha plots in fires during the 1997–1998 dry period damaged 81,274 burned forest (LD2 and HD2, 200 and 800 m km2 in Kalimantan and 116,984 km2 across Indonesia̶the away from unburned forest, respectively). In 2006 most severely burned area in the world (Tacconi, 2003). the number of species and stem density of rattans The effects of the fires on tree species and tree including seedlings were 16 and 8 species ha–1 and regeneration were investigated in the area. Van 8 8 a n d 2 4 s t e m s h a – 1 i n L D 2 , a n d H D 2 , Nieuwstadt and Sheil (2005) reported a significant respectively. These values were lower than those in negative correlation between tree mortality caused by K1, where 23 species and 3321 stems were fire and bark thickness, which was positively correlated recorded. The dominant species in burned plots with trunk diameter: small trees (<10 cm dbh) showed were Ceratolobus concolor, debilis, and more than 80% mortality and large trees (>70 cm dbh) Plectocomiopsis geminiflora , which were minor showed less than 20% mortality during the 2 years after components in the unburned plot. Rattans likely the surface fire. Species richness and the density of recolonized burned forest sites by transferred seedlings and saplings were significantly lower in burned from neighboring unburned forest by birds and forests than in unburned forests during the 3 years after animals. The distance from unburned forest the fire (Cleary and Priadjati, 2005). In burned forests, appeared to affect the speed of the recovery in the although tree density recovered to prefire levels within 5 burned plots. Although recovery of the stem density –15 years after the fire, species richness remained of rattans was slower than that of trees in burned significantly lower than in unburned forests (Slik et al. plots, it will likely increase gradually because the 2002). Disturbance due to fires has resulted in a reduction number of recruits consistently exceeded mortality of resources for sun bears, causing concerns about during our study period (February 2006 to August the maintenance and recovery of sun bear populations in 2007). However, it is not clear whether species burned forests (Fredriksson et al. 2007). composition and the density in burned forests Few studies, however, have investigated the effect of recover to preburned levels. fire on lianas in Indonesia. One of the most characteristic features of tropical forests is an abundance of lianas, Key words： Bukit Bangkirai, Calamoidae, climbing which contribute to woody species diversity (Putz 1984; palms, dipterocarp forest, lianas, recovery, species Putz and Chai, 1987; Gentry, 1991; Appanah et al. 1993; richness, tropical rain forest, wild fire Nabe-Nielsen 2001; Burnham 2002), forest biomass (Dewalt et al. 2000), and frugivore food sources (Gentry, 14 Natsuki M. WATANABE, Eizi SUZUKI, and Herwint SIMBOLON

1991; Schnitzer and Bongers, 2002). In , rattans (climbing palms) are predominant lianas (Gentry, 1991) and play a particularly important role in the physiognomy of the rain forests (Richards, 1996). is a generic name for 13 genera of climbing palms, with about 600 species, that inhabit the tropics and subtropics of the palaeotropical region (Uhl and Dransfield, 1987). The rattan species diversity is greatest in Southeast Asia (Dransfield and Manokaran, 1994), and Borneo has at least 150 species (Dransfield and Patel, 2005) with additional unidentified species (Dransfield, 1992c). Although rattans are distributed in most forest types (Dransfield, 1992a), the highest species diversity is recorded in mixed dipterocarp forests, which contain a Fig. 1.Map of the study site variety of rattan growth forms from nonclimbers that remain on the forest floor to climbers that reach the canopy (Watanabe and Suzuki, 2008). The canes are the primary or selectively logged mixed dipterocarp forests most important non-timber forest products in Indonesia covered Bukit Bangkirai. The time of selective logging and contribute to the cash income of local people as well within the area was inferred to be the 1970s to 1980s from as global trade (Sastry, 2002). Thus, studies of the the condition of old stumps scattered within unburned reestablishment of rattans after disturbances such as fire forest. During the large-scale forest fires in 1997–1998 in are needed to improve the conservation of biodiversity as Bukit Bangkirai, most forests burned but forests enclosed well as rattan production. by within river systems and nearby the river escaped the The aim of the study was to quantify the fire (Fig. 1). reestablishment of rattans after the severe forest fires in The study was conducted in three permanent plots of 1997–1998 in East Kalimantan, Borneo. We conducted the 100 m × 100 m: K1 in unburned forest (1°1.7’ S, 116°52.2’ investigations in a 1-ha unburned forest plot and two 1-ha E, 110 m a.s.l.), LD2 in burned forest (1°1.6’ S, 116°52.4’ E, plots in burned forests located 200 and 800 m from 80 m a.s.l.), and HD2 in burned forest (1°1.2’S, 116°52.5’E, unburned forest. This study examined rattan species 80 m a.s.l.). LD2 and HD2 are located about 200 and 800 richness, density, stem length increment, and population m from unburned forest, respectively. From the height of dynamics in the three plots. On the basis of a comparison charred parts of standing dead trees, the two burned among the plots, we discuss the reestablishment process plots were inferred to be affected by canopy fire with of rattans after forest fire. high intensity, although LD2 appeared to be less disturbed by fire than HD2. In August 2007, the basal areas and number of tree species (dbh ≥ 4.8 cm) were MATERIALS AND METHODS 30.1, 20.4, and 12.2 m2 ha–1 and 287, 131, and 105 spp. in Study sites K1, LD2, and HD2, respectively (Suzuki E., Simbolon, H., This study was conducted in Bukit Bangkirai, East Oka, N.P., Aiba, S., Ruliyana, S., Watanabe, N.M., and Kalimantan, Borneo, located about 50 km from Shimizu, H. unpublished). The three most dominant tree Balikpapan Airport (Fig. 1). The annual mean species in terms of basal area (dbh ≥ 4.8 cm) were Shorea precipitation from 1971 to 2005 was 2614 mm and the laevis Ridl. (Dipterocarpaceae), Dipterocarpus confertus annual temperature was 26.9 °C (Stasiun Meteologi Klas Sloot. (Dipterocarpaceae), and Madhuca kingiana II Balikpapan 2006). During 1982–1983 and 1997–1998, (Brace) H. J. Lam (Sapotaceae) in K1; Macaranga long dry periods occurred (rainfall <100 mm month–1 over gigantea Muell. Arg. (Euphorbiaceae), Vernonia arborea 3 months; Stasiun Meteologi Klas II Balikpapan 2006) and Buch.-Ham. (Compositae), and Pholidocarpus majadum caused severe forest fires in East Kalimantan. Although Becc. (Palmae) in LD2; and Litsea firma (Bl.) Hook. F. 17% of area of East Kalimantan was damaged by fire in (Lauraceae), M. gigantea, and Schima wallichii (DC.) 1982-1983 (Malingreau et al. 1985), Bukit Bangkirai was korth. (Theaceae) in HD2. Although the fire destroyed not damaged at that time according to the local most trees in the burned plots, several large trees residences. Before the severe forest fires in 1997–1998, survived. Reestablishment of rattans after forest fire 15

Rattan census tied on the petiole base of the top (excluding spear In February 2006 (K1 and LD2) and August 2006 (HD2) leaves) of stems that appeared sound and were less than all rattans, including seedlings rooted within the plots, about 3 m in height. At 6-month intervals from August were tagged and stem height, stem length, location 2006 to August 2007, the stem length increments, the (coordinates), and presence or absence of reproductive length from the colored wire to the base of the youngest organs were recorded. The stem height was measured petiole, were measured. After the measurement, a using a pole for up to 14 m height, and estimated different colored wire was tied on the base of the using a clinometer for plants over 14 m. Only rattans with youngest petiole. At the same time, the number of dead stems at least 20 cm in length could be identified to the stems was counted, and recruits were also tagged and species level because many of those with shorter stems measured as in the first census. showed insufficient characters for identification. Stems up to 1 cm in length were defined as seedlings. Voucher specimens were deposited at Kagoshima University in RESULTS Japan and Herbarium Bogoriense in Indonesia. Species composition, density, and stem height The number of rattan species with stems ≥ 20 cm long Measurement of stem length increment and were 20 (+ 1 variety), 13, and 1 in K1 (unburned), LD2 population dynamics (burned), and HD2 (burned) plots, respectively, at the During the initial rattan census described above, in a half initial census (Table 1), and these numbers remained of K1 (100 m × 50 m) and all of LD2 colored wires were constant during our 1-year observation. Of the 23 species

Table 1. Relative number of rattans (stems ha-1) ≥ 20 cm long in three study plots. Figures in bold type indicate the most dominant species in each plot according to stem density. K1 LD2 HD2 (unburned) (burned) (burned) . caesius Bl. - 6.3 - C. flabellatus Becc. 15.1 - - C. javensis Bl. 0.4 - - C. laevigatus Mart. var. laevigatus 1.6 6.3 - C. laevigatus var. mucronatus (Becc.) J. Dransf. 0.6 - - C. marginatus (Bl.) Mart. 14.4 3.1 - C. nigricans Van. Valk. 5.4 3.1 - C. ornatus Bl. var. ornatus 5.1 3.1 - C. pseudoulur Becc. 4.0 6.3 - C. manan Miq. - 3.1 - Ceratolobus concolor Bl. 5.0 21.9 - dydymophylla Becc. 1.9 - - D. elongata Bl. 3.1 - - D. hystrix (Griff.) Mart. var. hystrix 8.1 - - D. korthalsii Bl. 0.7 - - D. sabut Becc. 8.6 - - D. sp1 - 9.4 - Korthalsia debilis Bl. 6.0 25.0 - K. echinometra Becc. 9.1 3.1 - K. ferox Becc. 0.1 6.3 - K. furtadoana J. Dransf. 1.9 - - K. sp1 0.1 - - Plectocomiopsis geminiflora (Griff.) Becc. 8.4 3.1 100.0 P. mira J. Dransf. 0.4 - - 16 Natsuki M. WATANABE, Eizi SUZUKI, and Herwint SIMBOLON

Table 2. Number of stems and clumps in the three plots. K1 LD2 HD2 (unburned) (burned) (burned) Total number of stems 3321 89 24 Number of stems ≥ 20cm 801 32 6 Number of stems (seedlings) ≤ 1cm 728 14 17 Total number of clumps for all stems 2155 67 19 Number of clumps ≥ 20cm 444 32 1 Number of reproductive stems during census* 29 2 0 *Based on four and three observations during from Feb 2006 to Aug 2007 in K1 and LD2, and from Aug 2006 to Aug 2007 in HD2, respectively.

in total, 1 was found in all three plots, 10 were found only in K1, and 3 were found only in LD2. By adding those cases when we were able to distinguished rattans with stems <20 cm long by leaf characters, the estimated number of species for all sizes were 23, 16, and 8, in K1, LD2, and HD2, respectively. Thus, from one-third to two- thirds of the number of species in K1 became reestablished in the burned plots. The most dominant species, based on the number of stems ≥ 20 cm long, were Calamus flabellatus Becc. and Calamus marginatus (Bl.) Mart. in K1, Korthalsia debilis Bl. and Ceratolobus concolor Bl. in LD2, and Plectocomiopsis geminiflora (Griff.) Becc. in HD2. The most dominant species in the burned plots were also found as minor species in the unburned plot. Two of them, Korthalsia echinometra Becc. and P. geminiflora were confined to canopy gaps within the unburned plot. Contrarily the dominant species in the unburned plot were not found in the burned plots. Stem density and clump density (ha–1) were much lower in the two burned plots than in the unburned plot (Table 2). For all stems including seedlings, at the initial Fig. 2. Frequency distribution of stem height in the census 3321 stems comprising 2155 clumps were found in three plots K1, whereas 89 stems comprising 67 clumps and 24 stems comprising 19 clumps were found in LD2 and HD2. The Recruits and mortality number of stems per clump decreased in the order of K1 The average number of rattan recruits over 6 months was (1.54) > LD2 (1.33) > HD2 (1.26). The cumulative number 105 stems in K1, 26 stems in LD2, and 9 stems in HD2 of reproductive stems recorded during our study period (Fig. 3). The percentage of recruits from seedlings was was 29 in K1, 2 in LD2, and 0 in HD2. higher than that from sprouting in all plots (Test for the The frequency distribution of stem height at the proportion; P <0.05, Fig. 4). The number of recruits initial census of K1 and LD2 plots showed that higher exceeded mortality at every census in LD2 and HD2. The stems occurred at lower frequency (Fig. 2). In K1, rattans average mortality of rattan stems over 6 months was were distributed continuously from the forest floor to the 12.9% in K1, 8.2% in LD2, and 3.6% in HD2 (Fig. 3). The canopy, which was over 20 m high. In LD2, most rattans relatively high mortality in K1 was caused mainly by were shorter than 10 m, and only one stem of Calamus falling trees and debris. manan Miq. reached to the canopy of M. gigantea. In HD2 all rattans were shorter than 1 m, except for stems Stem length increment from a clump of P. geminiflora which reached to over 9 m. Figure 5 shows the mean stem length increments over 6 Reestablishment of rattans after forest fire 17

Fig. 3. Mean recruitment and mortality of rattans within the three plots measured at 6- month intervals. The bars represent standard deviation.

Fig. 4. The percentage of recruitment from seedling in unburned (K1) and burned (LD2 and Fig. 5. Mean stem length increments of dominant HD2) forest plots. The bars represent species measured at 6-month intervals in standard deviation. burned (LD2) and unburned (K1) plots months in K1 and LD2 for the most dominant species: C. flabellatus (K1), C. marginatus (K1), Ce. concolor (LD2), K. DISCUSSION debilis (LD2), and P. geminiflora (HD2). Nearly all the P. Recolonization by rattans after forest fire geminiflora were sampled from K1, and they were all There are three mechanisms for recovery of a distributed in large canopy gaps within the forest. population after disturbance: sprouts from surviving Although there were significant positive correlations (P < organs (Van Nieuwstadt et al. 2001; Cleary and Priadjati 0.05) between the mean stem growth over 6 months and 2005; Simoes and Marques 2007), banks (Quintana- the mean initial stem length, the correlation coefficients Ascencio et al. 1996; Dalling and Denslow 1998), and seed were lower in P. geminiflora (r2 = 0.18) and K. debilis (r2 = rain (Whittaker et al. 1997; Toh et al. 1999; Galindo- 0.40) than in any other species (r2 = 0.69–0.78). The Gonzalez et al. 2000; White et al. 2004). In the burned maximum mean stem length increments of the dominant plots, some climax tree species with long tap roots, such species in the unburned plot were less than 1.5 m, as Fordia splendidissima (Bl. ex Miq.) Buijsen and whereas those in the burned plots were more than 2 m, Cotylelobium melanoxylon (Hook. f.) Pierre regenerated indicating that the dominant rattan species in the burned successfully by sprouting (Oka, N.P., Suzuki E., plots grew more rapidly. Plectocomiopsis geminiflora, the Watanabe, N.M., Tamrin, unpublished). Neary et al. most dominant species in HD2, showed remarkable (1999) reported that soil surface temperature becomes growth: stems a few centimeters long elongated more several hundred degrees centigrade but was only 100 °C than 10-fold over 6 months. at a depth of 16 cm during an experimental combustion. 18 Natsuki M. WATANABE, Eizi SUZUKI, and Herwint SIMBOLON

When soils sampled from our study sites were dispersed in the island and/or between the islands a few experimentally burned for 1 h, the soil temperature was km distant each other by birds and bats (Whittaker and more than 600°C at the soil surface but less than 30°C at Jones, 1994). In Bukit Bangkirai many frugivorous birds 10 cm belowground (Miyazaki T., Mizoguchi, M., and bats exist, such as hornbills and short-nosed fruit Nishimura T., Seki K., Imoto H., Suzuki K., Obuchi A. bats. When we compared the two burned plots, the unpublished). Thus, after the forest fire some trees that number of species and stem density were greater in LD2 lost the aboveground stem could sprout from those parts (200 m away from unburned forest) than in HD2 (800 m under the soil. Although many rattans produce sprouts away). This pattern would be expected, as the number of (sucker shoots) that build up clusters, they never produce visits by flying frugivores would decrease with distance new sprouts on root systems; in all clustering species from the unburned seed source (White et al. 2004). sprouts arise from the base of the stem (Fisher and Dransfield, 1979). Rattans have no tap root and the main Development of rattan communities after forest fire adventitious roots radiate horizontally from the base of At our initial census, it appeared that rattans had just the stem (Dransfield, 1977, 1979), indicating that their begun to colonize the burned forests and the community root systems are distributed within a shallow depth. Thus, recovery process was in its initial stage. The stem rattan individuals in LD2 and HD2 were probably killed densities, including those of seedlings, in the burned by the forest fire, and regeneration by sprouting was plots (89 and 24 stems ha–1 in LD2 and HD2, respectively) impossible after the event. were much smaller than in the unburned plot (3321 stems Recolonization may have occurred through ha–1). For stems longer than 20 cm, the stem density in germination from seed banks that had accumulated the unburned plot (801 stems ha–1) was comparable to the before the fire. Some tree species, especially pioneers of density in other mixed dipterocarp forests in Borneo tropical rain forest, have seeds with life spans of more (600-1000 stems ha-1, Watanabe and Suzuki, 2008), than 1 year as buried seeds (Kanzaki et al. 1997). In whereas the densities in the burned plots (32 and 6 stems contrast, stored seeds of rattans are vulnerable to ha–1 in LD2 and HD2, respectively) were much lower. The dehydration and their life spans are only a few weeks to a tree densities (≥ 10 cm dbh) in once-burned forests in few months (Mori et al. 1980; Ramanayake 1999), even East Kalimantan, located 10–50 km away from our study when stored at 14°C–16°C with a moisture content of 30% sites, reached the prefire level within 5–15 years after the –35% (Mori et al. 1980). No rattans germinated from soils forest fire (Slik et al. 2002). In our study plots, the tree sampled from 0- to 5-cm depth in either unburned or densities (≥4.8 cm) had recovered to 86% and 78% of the burned plots (data not shown). Therefore, rattans prefire levels at LD2 and HD2, respectively, 9 years after probably did not reestablish from seeds buried before the the fire (Suzuki, Simbolon, Oka, Aiba, Ruliyana, fire. Watanabe, and Shimizu unpublished). Although recovery We propose that rattan seeds were transferred from of the stem density of rattans was slower than that of unburned forests into the burned forests by birds and trees in burned plots, it presumably increases gradually, animals. Rattan contain a fleshy fruit wall because the number of recruits consistently exceeded (mesocarp) or a fleshy seed coat (sarcotesta) that attract mortality during our study period. animals (Dransfield 1992a). Frequent seedling The responses of lianas to fire in the Brazilian recruitments were observed in burned plots where a few Amazon can be contrasted with our findings. The stem or no reproductive stems existed, supporting this density of thin lianas dramatically increased after forest hypothesis. Flying frugivores such as birds and bats play fire (Cochrane and Schulze, 1999; Pinard et al. 1999; important roles as seed dispersers for colonization of Gerwing, 2001, 2002). Most lianas that dominate after a degraded tropical forests by plant species (Wunderle, fire are pioneer species, which often inhabit canopy gaps 1997; Toh et al. 1999; Galindo-Gonzalez et al. 2000; White or disturbed areas (Cochrane and Schulze 1999; Pinard et et al. 2004). For instance, in the Krakatau Islands, where al. 1999; Gerwing, 2002), and they may retard forest primary succession began after massive volcanic succession (Pinard et al. 1999). In contrast, rattans eruptions in 1883, two rattan species were found in 1979 produce relatively large fruits with 1-3 seeds, that are and another in 1994 (Whittaker and Jones, 1994; dispersed by frugivores (Dransfield, 1992b); they are Whittaker et al. 1997). The three rattan species were climax species, rather than typical pioneer species that transported by birds to the Krakatau Islands from islands produce fruits with large-number of small seeds that are located more than 19 km across the sea, and seeds were dispersed over long distances by animals or the wind. Reestablishment of rattans after forest fire 19

Rattans climb by hooking their long whip-like climbing forest may be difficult to reestablish in burned forests organ borne on the tip or stems to supports, and their because of the relatively low seed availability. Our stems have high mobility (Putz, 1990). Rattans will not findings also indicate that it will take a longer time for occupy the tree crown of their support over a long period, recovery of burned sites farther from unburned seed because the stems show neither branching (except for 26 sources. The recovery of rattan communities may have Korthalsia species) nor secondary thickening. Therefore, an important effect on those frugivores that rely on rattan rattans are not invasive plants like Amazonian lianas or fruits as their food source. Further research on the vines, which invade disturbed forests such as burned growth and reproductive traits of rattans and their areas and retard forest succession. In another study, we relationship with seed dispersers is needed to understand found a significant correlation between species diversity how to foster the reestablishment of rattans in burned of rattans and trees within a forest (Watanabe and Suzuki, forest. 2008). Recovery of rattan communities after forest fire may accompany that of trees. ACKNOWLEDGEMENT We thank the Indonesian We noted a difference between unburned and Institute of Science (LIPI) for permission to conduct our burned plots in the dominant species (based on number research (permit no. 4049/SU/KS/2006, 4317/SU/ of stems). The most dominant species in K1, C. KS2007) and the staff of Inhutani I for their valuable flabellatus, was not found in the burned plots. Some helps. The study was financially supported by the Global species were minor in K1 but dominated in LD2 and HD2. Environment Research Fund (E-051), the Ministry of the From our results, we cannot explain why the speed of Environment, Japan, and THE SHIKATA MEMORIAL colonization is different among rattan species. Two of the TRUST FOR NATURE CONSERVATION. rattan species that dominated in burned forests were P. geminiflora and K. debilis, which are semelparous plants (i.e., the plant once and then dies) and require REFERENCES light gaps to develop into mature plants (Dransfield, Appanah S., Gentry A.H. & LaFrankie J.V. 1993. Liana 1979). The maximum stem length increments of these diversity and species richness of Malaysian rain species (Fig. 5 d, e) were greater than those of the forests. Journal of Tropical Forest Science 6: 116-123. species dominant in the unburned plot (Fig. 5 a, b), Burnham R.J. 2002. Dominance, diversity and meaning that they had ability to grow more quickly. The distribution of lianas in Yasuni, Ecuador: who is on dominant rattans in the unburned forest are iteroparous top? Journal of Tropical Ecology 18: 845-864. plants (i.e., the plant has the ability to flower continually Cleary D.F.R. & Priadjati A. 2005. Vegetation responses to and flowering is not followed by stem death). In addition, burning in a rain forest in Borneo. Plant Ecology different species should occur in different seral stages, 177: 145-163. such as pioneer stage and late successional stage. Cochrane M.A. & Schulze M. D. 1999. Fire as a recurrent Differences in growth and reproductive traits might be event in tropical forests of the eastern Amazon: relevant to the speed of colonization in burned forests. Effects on forest structure, biomass, and species composition. Biotropica 31: 2-16. Conclusion Dalling J.W. & Denslow J.S. 1998. Soil seed bank Vast areas of East Kalimantan were destroyed by severe composition along a forest chronosequence in forest fires in 1997–1998, but some parts of the forest in seasonally moist tropical forest, Panama. Journal of Bukit Bangkirai surrounded by river systems escaped the Vegetation Science 9: 669-678. fire. Rattans became reestablished in the burned forests 8 Dewalt S.J., Schnitzer S.A. & Denslow J.S. 2000. Density years after the forest fire. Our findings suggested that and diversity of lianas along a chronosequence in a unburned forest plays an important role as a seed source central Panamanian lowland forest. Journal of in the process of reestablishment of rattan communities Tropical Ecology 16: 1-19. in burned forests. In addition, the presence of seed Dransfield J. 1977. Calamus caesius and Calamus dispersers inhabiting the unburned forest is essential for trachycoleus compared. Gardens’ Bulletin, Singapore recovery of the rattan community. At this point, it is 30: 75-78. unclear whether species composition and density of Dransfield J. 1979. A manual of the rattans of the Malay rattans in burned forests will recover to prefire levels. Peninsula. Forest Department Malaysia, Kuala Species with low fertility or at low density in unburned Lumpur, 270pp. 20 Natsuki M. WATANABE, Eizi SUZUKI, and Herwint SIMBOLON

Dransfield J. 1992a. The ecology and natural history of Tropics 7: 9-20. rattans. In: A guide to the cultivation of rattan (eds. Malingreau, J.P., Stephens G. & Fellows L. 1985. Remote Mohd., W. R. W., Dransfield, J. and Manokaran, N.) sensing of forest fires: Kalimantan and North pp. 27-33. Forest Research Institute Malaysia, Kuala Borneo in 1982-83. Ambio 14:314-321. Lumpur. Mori T., Zollfatah A.R. & Tan C.H. 1980. Germination and Dransfield J. 1992b. Morphological considerations: the storage of rotan manau (Calamus manan) seeds. structure of rattans. In: A guide to the cultivation of The Malaysian Forester 43: 44-55. rattan (eds. Mohd, W. R. W., Dransfield, J. and Nabe-Nielsen J. 2001. Diversity and distribution of lianas Manokaran, N.) pp. 11-26. Forest Research Institute in a neotropical rain forest, Yasuni National Park, Malaysia, Kuala Lumpur. Ecuador. Journal of Tropical Ecology 17: 1-19. Dransfield J. 1992c. Rattans in Borneo: botany and Neary D.G., Klopatek C.C., DeBano L.F. & Ffolliott P.F. utilization. In: Proceedings of the International 1999. Fire effects on belowground sustainability: a Conference on Forest Biology and Conservation in review and synthesis. F o r e s t E c o l o g y a n d Borneo, July 30 - August 3, 1990, Kota Kinabalu, Management 122: 51-71. Sabah, Malaysia (eds. Ghazally, I., Murtedza, M. Pinard M.A., Lutz F.E. & Licona J.C. 1999. Tree mortality and Siraj, M.), pp. 22-31. Ya y a s a n S a b a h , and vine proliferation following a wildfire in a Kotakinabalu, Sabah. subhumid tropical forest in eastern Bolivia. Forest Dransfield J. & Manokaran N. eds. 1994. Plant Resources Ecology and Management 116: 247-252. of South-East Asia. No. 6. Rattans. Procea Putz F.E. 1984. The natural history of lianas on Barro Foundation, Bogor, 137pp. Colorado Island, Panama. Ecology 65: 1713-1724. Dransfield J. & Patel M. 2005. Rattans of Borneo: an Putz F.E. 1990. Growth habits and trellis requirements of interactive key. Royal Botanic Gardens, Kew climbing palms (Calamus spp.) in North-eastern (CD-ROM) Queensland. Australian Journal of Botany 38: Fisher J.B. & Dransfield J. 1979. Development of axillary 603-608. and leaf-opposed buds in rattan palms. Annals of Putz F.E. & Chai P. 1987. Ecological studies of lianas in Botany 44: 57-66. Lambir National Park, Sarawak, Malaysia. Journal of Fredriksson G.M., Danielsen L.S. & Swenson J.E. 2007. Ecology 75: 523-531. Impacts of El Nino related drought and forest fires Quintana-Ascencio P.F., Gonzalez-Espinosa M., Ramirez- on sun bear fruit resources in lowland dipterocarp Marcial N., Dominguez-Vazquez G. & Martinezlco forest of East Borneo. Biodiversity and Conservation M. 1996. Soil seed banks and regeneration of 16: 1823-1838. tropical rain forest from milpa fields at the Silva Galindo-Gonzalez J., Guevara S. & Sosa V. J. 2000. Bat- Lacandona, Chiapas, Mexico. Biotropica 28: and bird-generated seed rains at isolated trees in 192-209. pastures in a tropical rainforest. Conservation Ramanayake S. 1999. Viability of excised embryos, shoot Biology 14: 1693-1703. proliferation and in-vitro flowering in a species of Gentry A.H. 1991. The distribution and evolution of rattan Calamus thwaitesii Becc. Jour nal of climbing plants. In: The Biology of Vines (eds. Putz, Horticultural Science and Biotechnology 74: 594-601. F. E. and Mooney, H. A.), pp. 3-49. Cambridge Richards P.W. 1996. The Tropical Rain Forest. 2nd edn. University Press, Cambridge. Cambridge University Press, Cambridge, 575pp. Gerwing J.J. 2001. Testing liana cutting and controlled Sastry C.B. 2002. Rattan in the twenty-first century: an burning as silvicultural treatments for a logged outlook. In: Rattan: Current Research Issues and forest in the eastern Amazon. Journal of Applied Prospects for Conservation and Sustainable Ecology 38: 1264-1276. Development (eds. Dransfield, J., Tesoro, F. O. and Gerwing J.J. 2002. Degradation of forests through Manokaran, N.), pp. 237-244. FAO, Rome. logging and fire in the eastern Brazilian Amazon. Schnitzer S.A. & Bongers F. 2002. The ecology of lianas Forest Ecology and Management 157: 131-141. and their role in forests. Trends in Ecology and Kanzaki M., Yap S.K., Kimura K., Okaichi Y. and Evolution 17: 223-230. Yamakura T. 1997. Survival and Germination of Simoes C.G. & Marques M.C.M. 2007. The role of Buried Seeds of Non-Dipterocarp Species in a sprouts in the restoration of atlantic rainforest in Tropical Rain Forest at Pasoh, West Malaysia. southern Brazil. Restoration Ecology 15: 53-59. Reestablishment of rattans after forest fire 21

Slik J.W.F., Verburg R.W. & Kessler P.J.A. 2002. Effects of fire and selective logging on the tree species composition of lowland dipterocarp forest in East Kalimantan, Indonesia. B i o d i v e r s i t y a n d Conservation 11: 85-98. Stasiun Meteologi Klas II Balikpapan. 2006. Climate and rainfall in Balikpapan. Balikpapan, Indonesia. Tacconi L. 2003. Fires in Indonesia: causes, costs and policy implications. In: CIFOR Occasional Paper No. 38. Center for International Forestry research, Bogor, Indonesia, 21pp. Toh I., Gillespie M. & Lamb D. 1999. The role of isolated trees in facilitating tree seedling recruitment at a degraded sub-tropical rainforest site. Restoration Ecology 7: 288-297. Uhl N.W. & Dransfield J. 1987. Genera Palmarum. Allen Press, Lawrence, KS, USA, 610pp. Van Nieuwstadt M.G.L., Sheil D. & Kartawinata K. 2001. The ecological consequences of logging in the burned forests of East Kalimantan, Indonesia. Conservation Biology 15: 1183-1186. Watanabe N.M. & Suzuki E. 2008. Species diversity, abundance, and vertical size structure of rattans in Borneo and Java. Biodiversity and Conservation 17: 523-538. White E., Tucker N., Meyers N. & Wilson J. 2004. Seed dispersal to revegetated isolated rainforest patches in North Queensland. F o r e s t E c o l o g y a n d Management 192: 409-426. Whittaker R.J. & Jones S.H. 1994. The Role of Frugivorous Bats and Birds in the Rebuilding of a Tropical Forest Ecosystem, Krakatau, Indonesia. Journal of Biogeography 21: 245-258. Whittaker R.J., Jones S.H. & Partomihardjo T. 1997. The rebuilding of an isolated rain forest assemblage: how disharmonic is the flora of Krakatau? Biodiversity and Conservation 6: 1671-1696. Wunderle J.M. 1997. The role of animal seed dispersal in accelerating native forest regeneration on degraded tropical lands. Forest Ecology and Management 99: 223-235.

Received 26th Mar. 2008 Accepted 18th July 2008