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Palaeogeography, Palaeoclimatology, Palaeoecology 171 (2001) 213228

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A Late Pleistocene and Holocene pollen and charcoal record from peat swamp forest, Lake Sentarum Wildlife Reserve, West Kalimantan, Indonesia
Gusti Anshari*, A. Peter Kershaw, Sander van der Kaars
Centre for Palynology and Palaeoecology, School of Geography and Environmental Science, Monash University, Monash, Vic. 3800, Australia Received 20 October 1999; received in revised form 20 April 2000; accepted for publication 25 September 2000

Abstract A preliminary palynological record is presented from a peat swamp forest in the Upper Kapuas River Basin in West Kalimantan. Although the record is discontinuous, it provides a picture of changing vegetation, riverine and swamp environments, and climate through parts of at least the last 30,000 years. During the Late Pleistocene, the composition of both riverine and swamp forests was different to that of the Holocene. Temperatures were cooler, especially during the Last Glacial Maximum, as indicated by the presence of components of submontane and montane vegetation. The presence of charcoal demonstrates that the tropical lowland forests experienced re through the whole of the recorded period. However, charcoal values generally rise through the period suggesting increased human impact through time. Highest burning levels and forest indicators of disturbance indicate most intensive human impact within the last 1400 years. q 2001 Elsevier Science B.V. All rights reserved.
Keywords: Quaternary; Indonesia; Palynology; Peatland forest; Climate change; Biomass burning

1. Introduction Late Quaternary records of environmental change in the Indonesian region, derived from pollen analysis, have largely focused on montane environments, which have been sensitive to changes in temperature but have revealed little information regarding changes in precipitation, a critical variable within this region (Flenley, 1998). Some attention has been paid to coastal swamp forests (Anderson and Muller, 1975; Caratini and Tissot, 1988; Morley, 1981) (see Fig. 1) but records have been limited to the Holocene period
* Corresponding author. Tel.: 161-3-9905-2929; fax.: 161-39905-2948. E-mail address: gusti.anshari@arts.monash.edu.au (G. Anshari).

and local inuences of sea level variation and vegetation succession have masked regional environmental changes. More recently, lowland records have been constructed from discrete basins within lowland areas (Hope and Tulip, 1994; Van der Kaars et al., 2001) but these sites are also near coastal in their location. A study has been initiated in the Lake Sentarum Wildlife Reserve of West Kalimantan, which is situated in the central part of the island of Borneo (Fig. 1), primarily to provide an inland, and more regional, record of environmental change. The area was selected also because it possesses some unique characteristics and a knowledge of the history may contribute to the future management of what is a very sensitive system. Lake Sentarum is an extensive

0031-0182/01/$ - see front matter q 2001 Elsevier Science B.V. All rights reserved. PII: S 0031-018 2(01)00246-2

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Fig. 1. Location of study area within the Indonesian region and with respect to previous palynological studies at Berakas and Marudi (Anderson and Muller, 1975), Sebangau (Morley, 1981) and Mahakam Delta (Caratini and Tissot, 1988).

uviolacustrine basin, much of which is ooded during wet seasons when incoming rivers ll some 80 lakes. These lakes partially or totally drain during dry periods. Interuve areas largely support a range of swamp forests that are inundated to varying degrees. Due to this environmental variability, exacerbated by oSouthern Oscilthe strong inuence that the El Nin lation exerts on the hydrology and other environmental components, the system is delicately balanced. The stability of the system is further inuenced by the large number of landuses, despite the reserve status of the basin. The local Dayak population, mainly the Iban tribe, traditionally practice shifting cultivation in the dry land areas and wet rice paddy cultivation in swamp environments, while the Malays engage in intensive commercial shing within the lakes. Further disturbance to forests results from local timber usage, commercial logging by private companies, and forest re. This paper presents results of the pollen and charcoal analysis of one core (HN3) extracted from a peat

swamp forest in the vicinity of Lake Pemerak, a small lake near the Tawang river (see Fig. 2). The core was taken in late August 1997, during an extremely dry o. At this time, most lakes period attributed to El Nin in the reserve were totally dry while peat swamp forest appeared to maintain relatively high water tables. The signicance of ENSO and human inuences appeared very evident at the time of eldwork as the exceptionally dry conditions restricted access into the lakes any further than Lake Pemerak, while a hazardous plane landing at Pontianak airport and subsequent, continuous haze paid testimony to the opportunistic burning practised throughout the island as a result of the severe drought conditions. 2. The study area Lake Sentarum Wildlife Reserve is located in the upper Kapuas River basin, some 700 km east of Pontianak, the capital of West Kalimantan Province

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Fig. 2. Location of the study site.

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(Fig. 2). The reserve, which lies virtually on the equator at an altitude of between 35 and 50 m asl, contains some 80 seasonal and shallow lakes that can be regarded as temporary oodplain extensions of the Kapuas River. These lakes occupy 27% of the total 80,000 hectares of the reserve. The remaining area consists mainly of seasonally or permanently ooded swamp forests (Giesen, 1996). The geology of the reserve is poorly known. The sediments are mainly clays in the lake basins and peats and peat soils in the terrestrial lowlands with sandstone forming the surrounding hills. The low nutrient status of parent soils materials has been considered to be responsible for the stunted and pole-like nature of the vegetation around the lakes (Giesen, 1996). However, structural features of the vegetation may also be a result of regular oods that produce an inundated environment. The upper Kapuas basin experiences a wet tropical climate with average annual rainfall estimated to be over 4000 mm. Rainfall exhibits some seasonality with a weak monsoon inuence extending from August to May. The wettest months are November and April while the driest months, which still average at least 100 mm of rainfall, are June to August (MacKinnon et al., 1996). However, Borneo regularly oSouthern Oscillafaces seasonal droughts. El Nin tion (ENSO) is being increasingly recognised as a major inuence on interannual rainfall variability and consequently on lake level uctuations. The few temperature measurements available suggest a range of between 30 and 368C during the day and between 23 and 298C at night throughout the year. Several types of forest dominate the region. Tropical lowland forests grow in the hills, tall swamp forests are common on peat soils while stunted and dwarf swamp forests are found in riparian habitats. Floating grass mats occur within the lakes. Agricultural elds, mainly swiddens, are common in upland sites, usually close to human settlements. Plant diversity is not very high, at least in comparison to other lowland tropical forests in Borneo. The common plant families found in the reserve are Dipterocarpaceae (40 species), Euphorbiaceae (36 species), Rubiaceae (35 species), Myrtaceae (26 species), Fabaceae (21 species), Lauraceae (20 species), Melastomaceae (20 species), Guttiferae (19 species), Moraceae (14 species), and Arecaceae (14 species). About three-

quarters (73%) of these families are trees and shrubs (Giesen, 1996). The upper Kapuas basin is surrounded on its northern, eastern and southern boundaries by chains of low mountain ranges. These chains mark the boundaries between Sarawak and West Kalimantan in the north and West and Central Kalimantan in the south (Fig. 2). Occasional peaks, rising to above 2000 m asl., support both lower and upper montane vegetation and these occurrences are between 100 and 200 km from the study site. Some information on the nature of vegetation associated with the highest of these mountains, Bukit Raya (Fig. 2), is provided in Nooteboom (1987) although detailed description is restricted to lowland forest in the foothills. An altitudinal transect of surface pollen samples from above 1670 m revealed very high percentages of the oaks (Lithocarpus/Castanopsis and or Quercus), indicative of lower montane forest, to about 2000 m, and signicant percentages of Ericaceae (including Vaccinium and Rhododendron types), well represented in upper montane forest, above this altitude (Maloney, 1987). The southern conifers, Dacrydium, Podocarpus, Dacrycarpus and Phyllocladus were also recorded. Of these taxa, only Lithocarpus/Castanopsis and Dacrydium have been detected growing in the Lake Sentarum area. The surface samples also showed high levels of ferns and Casuarinaceae, and Myrtaceae above 2000 m. 3. The study site Lake Pemerak represents the ooded area of a small river that originates in the surrounding hills and disgorges into the Tawang River (Fig. 2). At the time of eldwork, the basin was dry except for a reduced meandering river channel. The basin was sparsely vegetated with low growing grasses and sedges, except around the margins that supported a riparian tree cover of Barringtonia agutanguia. Tree cover was dense at the very margins but became progressively lower and more open into the lake basin. The lake margin was dened by a terrace that became more pronounced downstream from about 1 m adjacent to the coring site to about 10 m near the conuence with the Tawang River. Rainforest surrounds the lake and in places, like the

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Fig. 3. Pollen diagram from Pemerak peat Core HN3.

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Fig. 3. (continued)

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Fig. 3. (continued)

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Fig. 3. (continued)

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Table 1 Radiocarbon dates from Pemerak peat Core HN3. Calibrated ages were calculated using CALIB 4.2 (Stuiver and Reimer, 1993), calibrated age in years B.P. and two sigma range are given Sample ID OZE-133 Wk-6278 OZE-134 Wk-6275 OZE-135 Wk-6277 OZE-136 Wk-5779 Depth (cm) 1415 4142 6061 6768 7172 91.592.5 9495 104124 Radiocarbon age (years B.P.) 265 ^ 35 1366 ^ 72 2290 ^ 50 3117 ^ 57 13070 ^ 70 16840 ^ 120 28600 ^ 250 28780 ^ 100 Calibrated age (years B.P.) 300 (430153) 1290 (14071171) 3075 (32402888) 3355 (34673170) 15713 (1619014652) 20058 (2075819393) d 13C () 230.0 229.9 230.0 229.4 229.5 229.5 230.0 230.1 ^ 0.2 Remarks AMS date AMS date AMS date AMS date AMS date AMS date AMS date Conventional date

coring area, occurs on peat that extends to the lake edge.

4. Field and laboratory methods The core, 124 cm in length, was collected with a modied Livingstone sampler within the swamp forest about 100 m from the lake. It was wrapped in plastic and protected in PVC tubing. In the laboratory 1 cm slices were taken at 4 cm intervals along the core length. From each slice an uncontaminated cm 3 was extracted for pollen analysis while the remainder of the sediment was oven dried for 24 h at 1008C to allow a measure of the water content, and dried sediment ignited in a furnace at 5508C to remove the organic content and provide an estimate of the inorganic matter content. A basal sample was taken originally for conventional radiocarbon dating to get some general estimate of the age of the sequence. Subsequently, additional samples were submitted for accelerator mass spectrometry radiocarbon dating to age major changes in the constructed pollen record. These samples were pretreated in a similar manner to those for pollen analysis in an attempt to exclude possible contamination from penetrating roots and mobile humic acids. Determinations were made at the University of Waikato Laboratory in New Zealand and the Australian Nuclear Science and Technology Organisation (ANSTO). A standard methodology was followed to concentrate the pollen (Lowe et al., 1996). Initially, each sample was disaggregated in 10% Na-pyrophosphate for 24 h before sieving through a 180 mm mesh to

remove larger fragments. The residue was then ltered through a 7 mm sieve to remove ne particles before the dissolution of cellulose substances by a 10 min acetolysis. Remaining organics were then separated from mineral matter using the heavy liquid, sodium polytungstate, and the residue washed in acetic acid, distilled water and 100% ethanol before being mounted on microscope slides in glycerol. Pollen counts were undertaken on an Olympus BH-2 microscope at a magnication of 750. Initial identications were veried using a 100 oil immersion objective, giving a magnication of 1875. A total of 200 pollen grains per sample were counted where possible but, due to low pollen concentrations, only about 100 pollen grains were counted in some samples. Identication was assisted by comparison with reference slides in the collection of the Centre of Palynology and Palaeoecology, School of Geography and Environmental Science, Monash University, and published pollen oras especially Huang (1972) and Tissot et al. (1994). Most taxa could only be identied to genus level while a proportion of pollen types, between about 10 and 20% in individual samples, could not be identied. All black, opaque, angular particles, greater than or equal to 10m maximum diameter, were counted as charcoal particles along three transects from each pollen slide.

5. Stratigraphy and dating of Core HN3 The core can be divided into three sections on the physical nature of the peat sediment. The upper section from 0 to 66 cm depth is brous (bris) peat

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with wood remains, the middle section from 66 to 95 cm is humied and compact sapris peat, while the bottom section is sub-brous peat (Fig. 3). The base of this peat deposit was not reached. The Munsell colours of these sections overlap, from dull yellowish brown (10Y R4/3) to brown (10YR4/4). Moisture content averages 78% of wet weight with the top section slightly wetter than the lower sections while the inorganic residue after ignition is extremely low, averaging only 3%. Results of radiocarbon dating are presented in Table 1. At a depth between 104 and 124 cm, the conventional radiocarbon age is 28,780 ^ 10 years B.P. The seven AMS dates are younger and all dates conform to the stratigraphic sequence. 6. The pollen diagrams All pollen and charcoal data, along with features of the core stratigraphy and radiocarbon dates, are shown in Fig. 3ad. Fig. 3a provides a summary of stratigraphic and pollen data with the sums of pteridophytes and indeterminate or unidentied taxa percentaged against the sum of trees and shrubs plus herbs. In Fig. 3bd, total woody plant pollen provides the sum on which percentages for all taxa from each stratigraphic level are based. Herbaceous taxa, represented only by Poaceae and Cyperaceae, and pteridophyte taxa are excluded from the sum. Woody plant taxa, which are predominantly trees, are divided into broad ecological groups although, as indicated by the categories, lowland peatland/riparian and lowland peatland/dryland, there is substantial overlap between groups as a great deal of ecological diversity is contained within many represented taxa. In addition, the identication status of taxa is variable. The sufx `comp.' indicates that the pollen grain compares well with the given taxon although this does not exclude the possibility that it could be derived from another, closely related, taxon whose pollen is not represented in the reference collection. The sufx `type' relates more generally to a group of taxa of which the most likely one is named. Pollen morphological sufxes are added to taxa either to indicate a particular type within that taxon or where, in the case of `monolote' and `trilete', the pteridophyte spores have either lost their exosporia preventing more rened identication

or to fern taxa whose taxonomic status is unknown. Concentrations of total pollen and charcoal particles per cm 3 and charcoal values expressed as percentages of the pollen sum, i.e. charcoal/pollen ratios, are included in Fig. 3d. The diagrams were constructed using the program TILIA 2.0 B4 (Grimm 1991) and zoned with the aid of a stratigraphically constrained classication subroutine (CONISS) contained within the TILIA program. All taxa were used as input into CONISS. The CONISS dendrogram is included on Fig. 3a and allowed the recognition of 6 zones. Zones B3-B1 show high levels of internal spectrum similarity except for the topmost sample of zone B1 (the surface sample). As the difference between this sample and others in B1 is the result largely of a single high value for Gymonstoma sumatrana, it is considered that this is insufcient to warrant a separate zone allocation. Internal variation between samples is greater in the lower zones of B6B4 but, again, much of this variation is caused by occasional high taxon values which do not alter the general composition of zones. 6.1. Zone B6 (120110 cm): .ca. 30,000 years B.P. This basal zone has consistent representation of the riparian tree Gluta renghas and relatively high values for Ilex, palms, Gonystylus bancanus, Meliaceae/ Sapotaceae and Rosaceae within the peat forest component. The three latter taxa, together with the palm taxon with small echinae (spines) are also characteristic of this zone. Pteridophyte representation is high with signicant contributions from Polypodiaceae type and from both Monolete and Trilete spores. Pollen and charcoal concentrations are low but the charcoal/pollen ratio is notable. 6.2. Zone B5 (11094 cm): ca. 30,00028,000 years B.P. This zone shows similar values for riparian taxa including relatively high Gluta renghas percentages to those in zone B6, but no other consistently high woody plant percentages apart from Palaquium Type 1, whose values rise abruptly at the beginning of the zone and Ilex whose percentages are maintained from zone B6. There are notable percentages of Sterculiaceae, Sterculiaceae (Reevesia comp.) and Tecoma comp. in one or more samples. The Pteridophyte

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percentages are very similar to those in the basal zone but concentrations are much higher. Charcoal values are, on average, the lowest for the whole record. 6.3. Zone B4 (9466 cm): ca. 17,00013,000 years B.P. This zone shows strong pollen domination by Gluta renghas and Palaquium Type 1. There is highest representation for the diagram of Sterculiaceae (Reevesia comp.), Quercus, Longetia and the Podocarpaceae, including Dacrycarpus, Dacrydium, Phyllocladus which are all restricted to the zone, and Podocarpus, as well as a peak towards the top of the zone in Gymnostoma. Pteridophytes also have their highest percentages, largely due to very high values of Trilete spores and additional signicant presence, except for the top two samples, of Lycopodium cernuum and L. phlegmaria. Pollen and charcoal concentrations achieve high levels and there is a strong increase in the charcoal/pollen ratio. 6.4. Zone B3 (6640 cm): ca. 30001350 years B.P. There are major changes at or close to the zone B43 boundary with sustained sharp declines in Gluta renghas and Palaquium Type 1 and the pteridophytes Monolete and Trilete spores as well as Polypodiaceae type, sustained increases in Calophyllum and Combretocarpus rotundus and generally higher values for the riparian taxa Anacardiaceae type, Baccaurea type and Carallia, and for the herbs Poaceae and Cyperaceae. The riverine taxon Barringtonia occurs for the rst time.The zone is characterised by high values for Planchonella type and Nephelium. This and the previous zones are the only ones containing more than the occasional value of Ericaceae (probably derived from Rhododendron or Vaccinium). Pollen concentrations are slightly reduced, those of charcoal are variable but include highest values for the diagram, while the charcoal/pollen ratio is similar to that in the previous zone. 6.5. Zone B2 (4018 cm): ca. 1350300 years B.P. There is little consistent change in the woody plant spectra from the previous zone, apart from a strong increase in Pandanus, with the only other major features being a further decrease in pteridophytes,

assisted by the near elimination of Lycopodium species, and higher values of herbaceous taxa, particularly Poaceae. Pollen and charcoal concentrations are low but there are high values in the charcoal/ pollen ratio. 6.6. Zone B1 (180 cm): ca. 3000 years B.P. The base of the zone is marked by the rst presence and substantial representation of Palaquium type II while Gymnostoma increases substantially through the zone. Rutaceae type and Trema, both of which are rst recorded in the top sample of zone B2, are well represented. The herbs Cyperaceae and Poaceae maintain signicant representation but values do not achieve those recorded in the previous zone. Pollen concentrations continue to be low but there is a marked increase in charcoal concentrations and attainment of highest average charcoal/pollen ratios for the record. 7. Vegetation and environmental reconstruction The topmost samples of the core provide some basis for interpretation of the record in that they can be compared directly with the present day landscape. The extremely high proportion of rainforest taxa, most of which can be found in peat swamp forests, reects well the local vegetation and suggests that most pollen may be derived from trees growing in close proximity to the core site. Unfortunately there are few vegetation data to allow a more detailed assessment of pollen/ vegetation relationships. The vegetation of nearby Lake Pemarak is probably reected in the values for Poaceae, Cyperaceae and Barringtonia, in particular, while the charcoal level would be reecting the degree of burning that was noted to have taken place both within the forest and on the dry lake surface where there was sufcient fuel to burn, during the present dry period. Disturbance to the forest itself is best indicated by the signicant values for indicator taxa such as Trema, Terminalia and Pandanus. In reconstructing the history of the site, it must be stressed that it is likely that only short periods of time within the last 30,000 years or so are represented. Estimates of average rates of tropical peat accumulation from elsewhere in Indonesia suggest values of between 0.3 and 13.3 mm/yr (Neuzil, 1997) and

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even using the lowest gure, an accumulation of some 10 m rather than 1.24 m would be expected if accumulation had been continuous and regular. Furthermore, even in peats which have achieved a thickness of over 10 m in the last few thousand years, recent dating has indicated major phases of non-deposition (Page et al., 1999). The radiocarbon dates provide some basis for assessment of the timing and length of non-depositional periods although contamination is always a possibility in discontinuous sequences. The basal date, taken from bulk sediment, is likely to be the most suspect as a large component of tropical peatlands are composed of penetrated roots (Brady, 1997) which will date younger than the sediment matrix. The AMS dates should be less inuenced by rootlet contamination although the prepared samples contained resistant plant material other than pollen that may have derived from decomposed root material. The earliest period, indicated by zone B1, may date to around or beyond 30,000 years B.P. The subbrous nature of the peat, together with substantial variation in composition of pollen spectra and the low pollen concentration values, suggest that accumulation was fairly rapid and that it covers a short period of time. The composition of the swamp forest was clearly different to that of today, suggesting perhaps different climatic conditions. But what these conditions were is difcult to determine. The relatively high values for ferns could suggest higher effective precipitation than today and more frequent ooding on the basis that ferns are not a major component of any recognised peat swamp forest and that fern spores, being well dispersed by water, could have derived from dry land rainforest in the upper catchment. There is no evidence of vegetation types like those of Lake Pemerak within the region today and it is possible that the lake did not exist, at least in its present form, and that the area was more generally ooded rather than the water being concentrated within the lake area. Evidence for peat sediments including tree stumps exposed in the margins of the small stream meandering across the dry bed of Lake Pemerak suggest that, at this time, the present lake area may have been part of the peat swamp forest system. However, the presence of Gluta renghas does indicate that some form of river system did exist. In addition, the postulation of more general

inundation appears to be contradicted by the consistent presence in the pollen samples of Gonystylus bancanus which is a peat swamp forest tree that is not tolerant of water logging (Soerianegara and Lemmens, 1994). Under these circumstances, the possibility that the ferns were local, either growing within the immediate peat forest or within the `lake' system, has to be considered. Although ferns are not very evident today, they may well have been during the last glacial period when temperatures were possibly lower. The occurrence of Podocarpus, and Quercus, are the only other possible indicators of reduced temperatures and perhaps suggest regional expansion of montane and submontane forest in the highlands, but are insufcient to indicate a local presence and therefore any substantial temperature lowering in the lowlands. Gonystylus bancanus virtually disappears during the period represented by zone B5, eliminating but not clarifying the fern problem. This decline was probably a regional feature as today it is a relatively rare species. Despite the decline of G. bancanus and other marked changes in the composition of the peat swamp forest at the zone B6-5 boundary, there is little change in riparian vegetation or differences in hydrology between the two periods. The consistent presence of Ericaceae, in addition to Quercus, Podocarpus, and the maintenance of relatively high pteridophyte percentages, provides greater condence in the postulation of temperatures lower than today. As the two basal dates are so similar, it cannot be determined whether or not there is a time gap between the periods represented by zones B6 and B5. The dates bracketing the zone B54/B4 boundary of around 29,000 and 17,000 years B.P. indicate a long period of non-deposition, or erosion of accumulated sediment, that included the earlier part of the Last Glacial Maximum. The date of the end of the period represented by zone B4 is less certain. Judging by the date of 3117 years B.P., the zone could extend well into the Holocene. However, the date of 13,070, only 4 cm below the 3117 years B.P. date, makes it likely that most of the zone falls into the Last Glacial Maximum and Late Glacial. Support for a Last Glacial Maximum date is provided by highest representation of the montane taxa Podocarpus, Phyllocadus and Quercus, together with some representation of Ericaceae, the only representation in the diagram of

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Dacrycarpus and Dacrydium and, possibly, the very high values for pteridophyte spores. It is surprising, however, that the montane conifers are best represented towards the end of the period, which dates less certainly to the Last Glacial Maximum than the early part of the period. Either the dates are not totally reliable or it took montane taxa some time to arrive within the pollen catchment area of the site from the mountains, some 100200 km away. It appears that at least part of the Last Glacial Maximum experienced sufciently wet conditions to allow peat development although precipitation levels lower than those of the other peat-forming periods might be inferred from the decomposed nature of the peat during the period represented by zone B4, as well as the fact that temperatures are likely to have been lower than today. Drier conditions might also be suggested from the relatively high representation of species of Lycopodium that are indicative of open canopied vegetation and increased charcoal levels that indicate greater re frequency or effectiveness than previously. The zone B4/B3 boundary marks the largest changes in the pollen and sedimentary record and supports the evidence for a major time gap, possibly extending from about 13,000 to 3000 years B.P. It may be surprising that the late Holocene period, rather than the early-mid Holocene, is represented considering that the bulk of peat in other parts of Kalimantan was formed during the earlier period (Neuzil, 1997; Page et al., 1999). However, a second phase of peat growth is evident within the last 2000 years in other areas and the early growth phase was probably more related to sea level rise and the subsequent attainment of hydrological equilibrium than any climate change. Consequently it could be that, regionally, effective precipitation has been higher in the late Holocene than it was in the early Holocene. Alternatively some change in the hydrology of the system may have prompted renewed peat growth in the Late Holocene. In comparison with the late Pleistocene, riparian components of the vegetation were very different with a mix of taxa including Barringtonia replacing Gluta renghas A. more open environment is indicated by increasing representation of Poaceae and Cyperaceae, as well as some maintenance of Lycopodium values. One explanation for these features is increased

environmental variability, which initiated or caused further development of the discrete lake system which dries periodically. The destruction of Gluta communities may have been a cause or effect of this variability, the latter possibly the result of human activity. This tree species would have been vulnerable due to its high timber quality and location in relation to more open environments and shing grounds. As this riparian environment was opened up, sustained disturbance and perhaps increased variability would have resulted in the replacement of Gluta by a variety of other colonists that formed a more open and stunted vegetation. The sharp reduction in fern spores is consistent with greater channelling of water within the lake basin and consequently less ooding of the peatlands, although fern reduction is likely to have been predominantly a response to increased temperatures from the Pleistocene to Holocene, indicated by the disappearance of montane elements and increased representation of the lowland dipterocarps. Charcoal levels were as high if not higher than during the Last Glacial Maximum and a further increase in burning is indicated, most likely the result of environmental variability induced by climate, people, or a combination of the two inuences. There is little change in the riparian environment through zones B2 and B1 where peat accumulation is likely to have been continuous. Higher values for grasses may indicate more frequent lake drying although the lack of any similar response in Cyperaceae suggests that the grasses may be reecting an opening up of terrestrial and peatland forests. Increased representation of other opportunistic taxa such as Terminalia, Trema and Pandanus, as well as highest charcoal/pollen ratios for the whole recorded period, provide supporting evidence for increased forest disturbance. It is likely that these changes reect increased human impact on the landscape although ENSO variability would most likely have facilitated high disturbance levels. 8. General discussion and conclusions This study has temporally extended the late Quaternary record of peat swamp forest environments by some 20,000 years and provided the rst evidence for the nature of these environments during the last

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glacial period. The record demonstrates substantial vegetation and environmental change that can be linked largely to regional inuences rather than autogenic successional processes that appear to dominate other more coastal peat forest sequences from the region. In line with other studies on the history of tropical peatlands, the record is discontinuous and it is possible that gaps are caused by periods of erosion or wastage as well as non-deposition. Based on the radiocarbon dates and changes in pollen spectra, major gaps incorporated the major transitions into the Last Glacial Maximum (from about 28,000 to 17,000 years B.P.) and between the Pleistocene and Holocene (between about 13,000 and 3000 years B.P.) with the possibility of one at or beyond 30,000 years B.P. Underlying causes of times of peat growth and disconformity are difcult to determine, particularly as they do not conform with regionally established patterns of climate change (Van der Kaars et al., 2000; Van der Kaars et al., 2001). It is tempting to equate times of peat accumulation with highest effective precipitation but some inconsistency is suggested by the formation of distinctly different peat types (i.e. bris and humic peat) within the sequence. The fact that there is no evidence of peat accumulation during the early to mid Holocene, regionally the period of highest recorded rainfall within the last 30,000 years, and a time of major peat accumulation elsewhere in Indonesia, could indicate that peat accumulation was restricted to times of lower effective rainfall. Such a situation could be explained by high water discharge during wetter periods ooding the peats and inhibiting sediment accumulation or causing erosion of accumulated peat. This scenario does not, however, explain the lack of sediment accumulation during the early part of the Last Glacial Maximum that experienced regionally similar conditions to the later part of this period. It is also possible that in such a hydrologically complex system, times of growth and stability vary substantially over the peatland and that there is no clear regional climatic signal discernable from the pattern of peat accumulation at any one spot. Palynologically, the glacial phases represented are characterised by high values of Gluta renghas which probably dominated closed, riparian forest adjacent to the peat forest, while the peat forest component itself is dominated by pollen of a species of Palaquium

although the peat forest assemblages are diverse. High percentages of fern spores were most likely derived from locally growing species and, together with the presence of conifers, Ericaceae, and Quercus, indicate the local presence of a montane element in the forest vegetation. The data add weight to the proposition of Colinvaux (2001) that equatorial rainforests, during the height of the last glacial period, had no modern analogue but were composed of a mix of lowland and montane taxa. Major changes to the vegetation are indicated midway through the record although the abrupt nature of the change is likely to be a product of an inferred gap between the Last Glacial Maximum and late Holocene. By at least 3000 years B.P. the stable closed riverine forest dominated by Gluta had been replaced by a more open and stunted riverine vegetation indicating higher levels of disturbance and most likely the development of the present dened lake system. It is difcult to allocate a cause for these changes, especially as they most likely occurred during the gap phase, but both the development of ENSO climatic variability, which would have stressed the preexisting stable riverine forest, and physical destruction by people could have contributed. Major Holocene peat forest taxa include Calophyllum, Combretocarpus and Hopea that are common in other Holocene peat sequences in Borneo (Anderson and Muller, 1975; Morley 1981). Unlike the riparian forest, there is no apparent change in the diversity of the peat swamp forest and unlike the other Holocene sequences, there is no clear evidence of successional vegetation changes. However, human impact is evident, particularly with increases in disturbance taxa, both woody plants and grasses, especially since about 1400 years B.P. The presence of at least some charcoal in all samples demonstrates that re has been a component of these humid tropical environments through the last 30,000 years. However, the evidence for the presence of people within inland Borneo from at least 35,000 years ago (Flood, 1995) prevents a determination of whether lowland rainforests did burn under natural conditions. There is a general increase in the charcoal/pollen ratio throughout the record suggesting that the inuence of people has increased through time although the beginning of a sustained rise during the Last Glacial Maximum may, as previously implied,

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have been, at least partially, a result of lower effective precipitation. As with the pollen evidence, greatest human impact is witnessed within the last 1400 years. The study has provided a general picture of environmental change within the region. It has also raised some interesting questions relating to environmental stability, which can be addressed in the immediate future for the Lake Pemerak area by examination of additional cores taken from peat swamp forest and also from the present lake basin. Future research should be directed towards other parts of Lake Sentarum to examine the regional signicance of the patterns of change recorded, towards the extension of records to provide baseline data for assessment of the initial impact of people on the landscape, and towards ne resolution studies of the last few thousands years in an attempt to decouple the inuence of people and climate variability on the development of this unique landscape. Acknowledgements We thank Ir. Adi Susminanto, MSc., the Head of Sub Balai Konservasi Sumber Daya Alam Kalimantan Barat, for allowing us to do this important research in the Lake Sentarum Wildlife Reserve Region; Yefri Dahrin, Ismail, Ruslan Kabulman, Pak Tangkong, Darwis, Adi and other friends in the village of Nanga Pemerak for accommodation and great assistance with eld work; Gary Swinton for preparing the nal gures; Rona Dennis for contributing a base map of Danau Sentarum Wildlife Reserve; and Raymonde Bonnelle and Azmi Mohd Yakzan for very valuable comments on the manuscript. Gusti Anshari has been supported by an AusAid scholarship, while nancial support was provided by an ARC large grant to Peter Kershaw, Nigel Tapper and Paul Bishop and an Australian Institute of Nuclear Science and Technology (AINSE) grant to Peter Kershaw and Gusti Anshari. References
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