Tropical Peat Swamp Forest Silviculture in Central Kalimantan

TECHNICAL PAPERS

Tropical Peat Swamp Forest Silviculture in Central Kalimantan A series of five research papers

Banjarbaru Forestry Research Unit, FORDA and Laura L. B. Graham

Kalimantan Forests and Climate Partnership

TECHNICAL REPORTS

Tropical peat swamp forest silviculture in Central Kalimantan A series of five research papers Banjarbaru Forestry Research Unit, FORDA and Laura L. B. Graham January 2014

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ACKNOWLEDGEMENTS

This report was prepared for the Kalimantan Forests and Climate Partnership (KFCP) by researchers at the Banjarbaru Forestry Research Unit (Litbang Banjarbaru), a regional research unit under the Forestry Research and Development Agency (FORDA) within the Indonesian Ministry of Forestry. Researchers included Rusmana, Dony Rachmanadi, Purwanto Budi Santosa, Tri Wira Yuwati, Pranatasari Dyah Susanti under the supervision of Laura L. B. Graham, Abdi Mahyudi and Grahame Applegate. We wish to thank all team members for their inputs into this report and KFCP for funding the activities. We would like to thank Rachael Diprose for editing this work and KFCP’s communications team (James Maiden and Nanda Aprilia) for their publishing assistance.

This research was carried out in collaboration with the Governments of Australia and Indonesia, but the analysis and findings presented in this paper represent the views of the authors and do not necessarily represent the views of those Governments. Any errors are the authors’ own. The papers in this compendium constitute technical scientific working papers and as such, there is potential for future refinements to accommodate feedback, emerging evidence and new experiences.

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EXECUTIVE SUMMARY

The Kalimantan Forests and Climate Partnership (KFCP) undertook a program of silviculture research for reforestation trials in the KFCP site in Central Kalimantan province. Seedlings were cultivated from seeds or wildlings in local community nurseries and then transplanted and monitored in the field. To increase biodiversity and speed-up natural succession, a wide range of native tropical peat swamp forest tree species were used (following the Framework Species Method). Tropical peat swamp forests (TPSF) house over 500 floral species. There is little species-specific literature published on their silvicultural or ecological traits. To this end, KFCP undertook an extensive grey-literature review and produced ‘A literature review of the ecology and silviculture of tropical peat swamp forest tree species found naturally occurring in Central Kalimantan’. This report, together with discussions with local community members in the KFCP area, facilitated the selection of 46 TPSF tree species native to Central Kalimantan that would become the ‘focal reforestation species’ prioritised in the reforestation trials. There are large gaps in the knowledge on the silvicultural and ecological traits of TPSF species, particularly in the Central Kalimantan peatlands. To address this issue and provide information to KFCP’s reforestation teams, three research activities were commissioned: 1) Two reports documenting the phenology of TPSF tree species in Central and Kalimantan and, more specifically, the focal species mentioned above, based on the analysis of previously collected data, 2) Investigations on the optimal growth conditions and ecological tolerances of the focal species in response to a range of extreme environmental conditions (drought, flooding, light intensity and microbial and nutrient availabilities), 3) Investigations on the extent to which the focal species, post-transplantation, could tolerate the natural environmental conditions of the degraded tropical peatlands in the study site under a range of treatments (weeding, nutrient addition, distance from canal). This report provides the results of the work undertaken to address Research Question 2 above. The research and analysis were undertaken by the Litbang Banjarbaru (Banjarbaru Forestry Research Unit) research team, under the supervision of Laura Graham and Grahame Applegate. The report presents findings on responses of the focal species (between 15–22 species, dependent on specific study) to light intensity, flooding, drought, nutrient and microbial availability. Each of these five environmental conditions are explored separately in five sub-papers presented as a compendium in this report. Paper 1: Light intensity. Twenty species (30 seedlings per species) were cultivated as seedlings, and then grown for four months under three different light intensity treatments: 25 percent, 50 percent and 100 percent light intensity (10 seedlings per treatment). After the four months of treatment, half of the species showed consistently high survival at all light intensity treatments, whilst other species had a lower survival rate at 100 percent light intensity compared with 25 percent or 50 percent light intensity. Nearly all species however, regardless of light intensity, maintained a survival rate of over 80 percent. At 25 percent or 50 percent light intensity, seedlings were able to attain greater height increments. However, no species attained their highest height growth increments at 100 percent light intensity. Over half the species, however, attained their highest stem diameter increments at 100 percent light intensity, with the remaining species showing no difference in stem diameter across the light intensity treatments. The above findings show that all the 21 species chosen have good survival rates for use in silviculture and reforestation activities. The results also indicate that most of the species demonstrate adaptability to light conditions; under low light, species exhibit height growth, whilst under high light species invest in stem strength. Seedling acclimatisation to high light, therefore, should always be carried out for at least two months before seedlings are planted in degraded, open areas of the peatlands. Knowledge of how each species responds to high light intensity can be used to influence selection of planting sites and growth patterns in nurseries (i.e. rapid height growth, or structure-strength investments). A more detailed summary of the tolerance of each species to high light intensity is presented in the analysis. Paper 2: Effects of drought. Twenty-one species (40 seedlings per species) were cultivated as seedlings and then grown under four watering intensity treatments for four months; once per day, twice per week, once

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per week, once per fortnight (10 seedlings per treatment). Based on the recommendations of the available literature, the 21 species were split into two species groups: a pioneer species group and a climax species group. Overall it was found that only a few species could tolerate drought conditions. Survival rates, shoot growth and leaf numbers commonly reduced in response to drought. Root length also reduced or was not affected by drought conditions. This was a surprising finding as this is counter to what the literature would suggest—the roots should grow and forage for water. Instead, the results demonstrate that there was often little difference between watering once per day and twice weekly. Shoot and root biomass generally decreased under drought conditions; there was no consistent trend regarding the shoot-root ratio. In the analysis, it was helpful to delineate the results by species groups as trends were sometimes distinct between them. For example, for the pioneer species group, shoot biomass gradually decreased as watering frequency decreased, which contrasted strongly with the climax species group that had increasing shoot biomass with decreasing watering frequency, except for the once a fortnight treatment. Generally, the pioneer species groups displayed more tolerance to drought than the climax species group. Overall, the results indicate that drought tolerance should be considered when selecting species for reforestation activities. This study has found that several species display good drought tolerance, such as Alstonia spatulata, Knema mandarahan, Licania splendens and Parartocarpus venenosus. Paper 3: Effects of flooding. Seventeen species (40 seedlings per species) were cultivated as seedlings and then grown under four different water-level treatments for four months: half of the roots/polybag submerged, all of the roots/polybag submerged, all of the roots and half the stems submerged, and the entire plant submerged (10 seedlings per treatment). Based on the survival rates and leaf change results, of the 17 species studied, 16 showed at least some adverse reaction to flooding, with only Lophopetalum javanicum maintaining good leaf numbers and survival rates, even during complete submersion for four months. A further five species showed some degree of tolerance, maintaining good survival rates and leaf numbers during partial stem submersion for four months; Alstonia spatulata, Calophyllum sclerophyllum, Dacrydium pectinatum, Disopyros bantamensis, Stemonurus scorpioides. The remaining 11 species, however, showed poor survival and leaf maintenance once some of the stem was submerged. These results indicate the importance of considering flood depths, seedlings transplant size, season, and species-selection when developing a reforestation plan. Paper 4: Macro-nutrients. Nineteen species (50 seedlings per species) were cultivated as seedlings, and then grown under five nutrient addition treatments for four months: control, plus Nitrogen, plus Phosphorus, plus Potassium, plus Calcium and Magnesium (10 seedlings per treatment). Of the 22 TPSF species studied, seven species showed a strong positive response to certain nutrient additions, such as Paratocarpus venenosus’ response to nitrogen additions and Lithocarpus sp.’s response to magnesium-calcium additions. A further three species showed a positive, but less strong response. Only two species, Alstonia spatulata and Aglaia rubignosa showed uniformly positive responses to all nutrient additions. Surprisingly, 12 of the 22 species did not respond or responded negatively to the nutrient additions, indicating these species are highly adaptable to low nutrient environments. The results highlight the importance and usefulness of nutrient additions for seedling survival and growth in TPSF reforestation activities, both during cultivation in the nurseries and after transplanting the seedlings into the forest. However, this study also highlights that nutrient additions are species-specific, differ for different nutrients, and can produce limited results for some species; as such, a blanket-approach in nutrient additions is not advised. Paper 5: Mycorrhizae. Fifteen species (40 seedlings per species) were cultivated as seedlings, and then grown under grown under four mycorrhizal-inoculation treatments for four months; control, inoculated with Glomus clarum, inoculated with Gigaspora decipens and inoculated with Enthrophospora sp (10 seedlings per treatment). Of the fifteen TPSF tree species seedlings studied, only five showed a positive reponse to the mycorrhizal inoculations; Calophyllum sclerophyllum and Parartocarpus venenosus species responded positively to the Enthrophospora sp. inoculum, whilst Lopopethalum javanicum and Syzygium sp. (Mahalilis) responded positively to the Glomus clarum inoculum, and Syzygium sp. (Pakan) responded favourably to both the Enthrophospora sp. and Glomus clarum inoculums. This study also indicated that Gigaspora decipens is not a suitable mycorrhizal inoculum, as none of the 15 species responded strongly to its

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inoculation. These findings can be applied to reforestation activities to increase growth rates of the seedlings both during nursery cultivation, and, potentially, after transplantation. It was surprising, however, that ten of the species showed no positive reaction to the three different mycorrhizae inoculums. The results were inconclusive; a longer period of study might have seen positive reactions to the mycorrhizae inoculums (or not). Further research could have included investigating the seedlings beyond the point of transplantation in the site. More variables could also have been examined such as the dry weights of the shoots and roots, and the nitrogen and phosphorus content of the leaves. This might have revealed other longer-term benefits of mycorrhizae inoculums. Summary findings. The main findings from the five papers are summarised in Table A below. One of the key findings across the five papers is the species-specific nature of tolerance to extreme environmental conditions and the requirements for maximum growth and survival. KFCP and the Banjarbaru Forestry Research Unit invested time and resources in this study to provide much needed, detailed answers to some of the key silvicultural challenges facing tropical peatlands. It is hoped that other rehabilitation projects working in this type of ecosystem apply this species-specific knowledge gained by KFCP to reforestation and rehabilitation programs, and that projects working in other ecosystems adopt a similar investigatory approach to contribute to improved and accurate rehabilitation and reforestation plans.

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No. Species (local name) Drought tolerance Light strategy Flood

tolerance Optimal nutrient

requirements Preferred mycorrhiza Recommended for reforestation?

1 Aglaia rubignosa (Kajalaki) Good All macro nutrients Yes—limited data 2 Alstonia spatulata (Pulai rawa) Good Generalist Acceptable N, Mg, Ca Yes 3 Baccaurea bracteata (Jajantik) Acceptable Generalist Poor Mg, Ca None found In areas without flooding 4 Calophyllum hosei (Bintangur) Poor Shade-tolerant None None found More suited to enrichment planting 5 Calophyllum sclerophyllum (Kapurnaga) Poor Shade-tolerant Acceptable None Enthrophospora sp. More suited to enrichment planting 6 Combretocarpus rotundatus (Perepat) Acceptable None None found Limited data 7 Cotylelobium sp. (Resak) Acceptable Shade-tolerant Poor None None found More suited to enrichment planting 8 Cratoxylum glaucum (Gerunggang) Poor Shade-tolerant None None found More suited to enrichment planting 9 Dacrydium pectinatum (Alau) Acceptable Limited data

10 Disopyros bantamensis (Mahirangan) Acceptable Shade-tolerant Acceptable None None found In shaded areas 11 Knema mandarahan (Mandarahan) Good Generalist Poor P In areas without flooding 12 Koompassia malaccensis Generalist Limited data 13 Licania splendens (Bintan) Good None None found Yes—limited data 14 Lithocarpus sp. (Pampaning) Poor Poor N, Mg, Ca More suited to enrichment planting 15 Lophopetalum javanicum (Perupuk) Poor Shade-tolerant Excellent N, Ka Glomus clarum In flooded areas 16 Mangifera sp. (Mangga-mangga) Good Sun-loving Poor None In areas without flooding 17 Melaleuca leucadendra (Galam) Poor Shade-tolerant N, Ka, Mg, Ca More suited to enrichment planting 18 Palaquium sp. (Nyatoh) Poor Shade-tolerant Poor None None found More suited to enrichment planting 19 Parartocarpus venenosus (Lilin-lilin) Good Generalist Poor N, Ka Enthrophospora sp. In areas without flooding 20 Sandoricum beccanarium (Papung) Acceptable Sun-loving Poor Ka In areas without flooding 21 Shorea sp. (Meranti daun kecil) Poor Shade-tolerant Poor More suited to enrichment planting 22 Shorea sp. (Meranti daun lebar) Poor Generalist None More suited to enrichment planting 23 Stemonurus scorpioides (Medang telur) Poor Sun-loving Acceptable None None found In areas without drought

24 Syzygium sp. 1/pakan (Pakan) No tolerance None Glomus clarum and Enthrophospora sp. Limited data

25 Syzygium sp. (Jambu burung kecil) None found Limited data 26 Syzygium sp. 2 (Mahalilis) Sun-loving Poor None Glomus clarum In areas without flooding

Table A: The key findings of the five presented papers on ecological tolerance to extreme environmental conditions and the conditions required for optimal growth conditions

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CONTENTS ACKNOWLEDGEMENTS ............................................................................................................................. i

EXECUTIVE SUMMARY ............................................................................................................................. ii

CONTENTS .............................................................................................................................................. vi

ACRONYMS ........................................................................................................................................... viii

LIST OF TABLES ....................................................................................................................................... ix

LIST OF FIGURES ....................................................................................................................................... x

BACKGROUND TO THE RESEARCH ............................................................................................................. 1

PAPER 1: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO LIGHT INTENSITY ........................ 2

1.1 Introduction .............................................................................................................................. 2

1.2 Materials and methods ............................................................................................................. 3

1.3 Results and discussion ............................................................................................................... 4

1.3.1 Survival rates ................................................................................................................................ 4

1.3.2 Height ........................................................................................................................................... 6

1.3.3 Stem diameter .............................................................................................................................. 8

1.4 Discussion ............................................................................................................................... 10

1.5 Conclusion .............................................................................................................................. 12

1.6 References .............................................................................................................................. 13

PAPER 2: RESPONSE OF PEAT SWAMP FOREST SPECIES TO DROUGHT .................................................. 15

2.1 Introduction ............................................................................................................................ 15

2.2 Materials and methods ........................................................................................................... 16

2.3 Results .................................................................................................................................... 19

2.3.1 Survival rates ..............................................................................................................................19

2.3.2 Growth—height increment .........................................................................................................21

2.3.3 Growth–stem diameter increment .............................................................................................23

2.3.4 Leaf numbers ..............................................................................................................................25

2.3.5 Root length .................................................................................................................................27

2.3.6 Shoot biomass .............................................................................................................................30

2.3.7 Root biomass ..............................................................................................................................30

2.3.8 Shoot-Root Ratio .........................................................................................................................31

2.4 Discussion and conclusion ....................................................................................................... 33

2.5 References .............................................................................................................................. 36

PAPER 3: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO FLOODING ................................ 37

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3.1 Introduction ............................................................................................................................ 37

3.2 Material and methods ............................................................................................................. 38

3.3 Results, discussion and conclusion ........................................................................................... 39

3.4 References .............................................................................................................................. 45

PAPER 4: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO MACRONUTRIENTS................... 46

4.1 Introduction ............................................................................................................................ 46


4.3 Results and discussion ............................................................................................................. 48

4.4 Conclusion .............................................................................................................................. 62

4.5 References .............................................................................................................................. 63

PAPER 5: RESPONSE OF PEAT SWAMP FOREST TREE SPECIES SEEDLINGS TO MYCORRHIZAL INOCULATIONS ...................................................................................................................................... 64

5.1 Introduction ............................................................................................................................ 64


5.3 Results and discussion ............................................................................................................. 66

5.4 Conclusion .............................................................................................................................. 74

5.5 References .............................................................................................................................. 75

CLOSING REMARKS ................................................................................................................................ 77

RELEVANT LITERATURE .......................................................................................................................... 78

ANNEX 1 ................................................................................................................................................ 79

Photographs of activities .................................................................................................................... 79

ANNEX 2 ................................................................................................................................................ 83

List species found on exploration activities ......................................................................................... 83

ANNEX 3 ................................................................................................................................................ 85

Soil microbes exploration ................................................................................................................... 85

Ectomycorrhizae ........................................................................................................................................85

Endomycorrhizae .......................................................................................................................................87

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ACRONYMS AMF Arbuscular mycorrhizae fungi

ANOVA Analysis of variance

ECM Ectomycorrhiza

EMRP Ex-Mega Rice Project

FORDA Forest Research and Development Agency

KFCP Kalimantan Forests and Climate Partnership

KHDTK Forest for Specific Purposes

PSF Peat swamp forest

REDD+ Reducing Emissions from Deforestation and Forest Degradation

SPSS SPSS predictive analytics software

TPSF Tropical peat swamp forest

VAM Vesicular arbuscular mycorrhiza

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LIST OF TABLES

Table A: The key findings of the five presented papers on ecological tolerance to extreme environmental conditions and the conditions required for optimal growth conditions ...................................................................................... v

Table 1.1: Species used in the light intensity—tropical peat swamp forest tree species seedling tolerance study............. 3

Table 1.2: Optimal light intenisty for high survival ............................................................................................................... 6

Table 1.3: Optimal light intensity for greatest (height) growth increments ......................................................................... 8

Table 1.4: Optimal light intenisty for greatest (stem diameter) growth increments ......................................................... 10

Table 1.5: Summarised optimal light intensities for survival and growth of the 20 studied PSF tree species, with possible light strategies noted ......................................................................................................................................... 11

Table 2.1: Peat swamp forest plant species used in the study of seedlings’ response to drought, with their morphological information at the beginning of the study period ..................................................................... 17

Table 2.2: Grouping of species based on their known growth strategies........................................................................... 18

Table 2.3: Soil chemical content of the growth media in the nursery ................................................................................ 18

Table 2.4: The optimal watering intensity for maximum height increment (cm) of the PSF tree species after four months ........................................................................................................................................................................... 21

Table 2.5: The effect of watering intensity on the height increment (cm) of the PSF tree species after four months ...... 22

Table 2.6: The optimal watering intensity for maximum height increment (cm) and the ability to maintain growth during extreme drought conditions (watering once per week and/or once per fortnight) for the PSF tree species after four months ............................................................................................................................................... 23

Table 2.7: The effect of watering intensity on the basal diameter increments (cm) of PSF tree species after four months ........................................................................................................................................................................... 24

Table 2.8: The optimal watering intensity for maximum stem diameter increment (cm) and the ability to maintain growth during extreme drought conditions for the PSF tree species after four months .................................. 25

Table 2.9: The effect of watering intensity on the final total leaf numbers of PSF tree species after four months........... 26

Table 2.10: The optimal watering intensity for greatest final leaf numbers (cm) and the ability to maintain leaf numbers during extreme drought conditions for the PSF tree species after four months ............................................... 27

Table 2.11: The effect of watering intensity on the root length (cm) of PSF tree species after four months .................... 28

Table 2.12: The watering intensity for maximal root length of the PSF tree species after four months ........................... 29

Table 2.13: The effect of watering intensity on the shoot-root ratio (g) of PSF tree species after four months ............... 32

Table 2.14: The watering intensity for greatest shoot-:root ratio of the PSF tree species after four months ................... 33

Table 2.15: Summary of drought tolerance of the 21 studied PSF tree species, based on morhpological responses to drought conditions ............................................................................................................................................. 35

Table 3.1: Species, plant material source and origin, age at the start of treatment, and seedling average dimensions at the start of the inundation experiment ............................................................................................................. 38

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Table 3.2: Flooding tolerance of 17 TPSF tree species based on the flooding experiment ................................................ 40

Table 4.1: List of species used in the macro-nutrients application study ........................................................................... 47

Table 4.2: Analysis of variance with ortogonal contrast for plant responses to macronutrients ....................................... 48

Table 4.3: Summarised results of the effect of nitrogen (N), phosphorus (P), potassium (Ka), magnesium (Mg) and calcium (Ca) on the height and diameter (Diam.) increments (inc.), the change in leaf number (LN) and the total dry weight (DW) of 22 TPSF tree species seedlings cultivated under nursery conditions ........................ 54

Table 5.1: List of species used in the mycorrhizal inoculation study .................................................................................. 66

Table 5.2: Summarised results of the effect of three mycorrhizal treatments on the height and diameter (Diam.) increments (inc.), the changes in leaf numbers (LN) and the total dry weight (DW) of 15 TPSF tree species seedlings cultivated under nursery conditions .................................................................................................. 68

LIST OF FIGURES

Figure 1.1: Survival rate of PSF tree species seedlings after four months under light intensity treatment ......................... 5

Figure 1.2: Average height of PSF tree species seedlings after four months under light intensity treatment ..................... 7

Figure 1.3: Height increment of PSF tree species seedlings after four months under light intensity treatment ................. 7

Figure 1.4: Average stem diameter of PSF tree species seedlings after four months under light intensity treatment ....... 9

Figure 1.5: Stem diameter increment of PSF tree species seedlings after four months under light intensity treatment .... 9

Figure 2.1: Survival rates of two species group for PSF tree species after four months of watering intensity treatments 20

Figure 2.2: The average of shoot biomass (g) of two groups PSF tree species after four months of watering intensity treatments ......................................................................................................................................................... 30

Figure 2.3: The average of root biomass (g) of two groups PSF tree species after four months watering intensity treatment ........................................................................................................................................................... 31

Figure 3.1: Seedling flood treatment .................................................................................................................................. 39

Figure 3.2: The survival rates of 17 peat swamp forest species based on the inundation experiment ............................. 41

Figure 3.3: The changes in leaf numbers for each treatment in the inundation experiment............................................. 43

Figure 4.1: Changes in height, diameter, number of leaves, and average dry weight of 21 PSF species with macronutrient application ......................................................................................................................................................... 55

Figure 5.1: Changes in height, diameter growth and number of leaves of 15 PSF species, 7 months after three mycorrhizae inoculation treatments ................................................................................................................. 69

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BACKGROUND TO THE RESEARCH

The analysis presented in the following papers presents the research results from the ‘Technical support for the ecological restoration of TPSF, KFCP, Central Kalimantan‘ component of the Silviculture Research and Monitoring program funded by Kalimantan Forests and Climate Partnership (KFCP). The research undertaken in this component consisted of several activities, namely: seed/cutting material sourcing, soil microbes exploration and nursery seedling research.

Seedlings/cutting materials were collected in Block A and E, Mantangai, in the Ex-Mega Rice Project (EMRP) area. More specifically, work was undertaken in the Forest for Specific Purposes (KHDTK) Tumbang Nusa research plots managed by Forest Research and Development Agency (FORDA) at Tumbang Nusa village, Central Kalimantan, in Hampangen forest and Teluk Umpan forest. Investigations of soil microbes were only carried out in Block E of the EMRP. The research in the nursery included seedlings/cuttings preparation and production at the Banjarbaru Forestry Research Unit nursery, South Kalimantan.

The nursery research was the main activity undertaken in this project to determine the response of peat swamp forest (PSF) species under a range of environmental conditions; light intensity, inundation, drought, nutrition supply and soil microbe utilisation. The results of this research are presented in the five sub-papers included in this report. The photos from the research activities are presented in Annex 1. There were 31 PSF species obtained as seedlings or cuttings during the seedling exploration (Annex 2). Only 21 species out of 31 were found in sufficient number to be used in this research.

The exploration of soil microbes were specialised for ectomycorrhiza and arbuscular mycorrhiza species due to the limiting factor of inundation in the degraded peat swamp forest. The mycorrhiza species that were recorded are presented in Annex 3.

The results of this report can be used to support efforts in degraded peat swamp forest rehabilitation. Understanding how each species responds to various environmental conditions provides a basis for species selection for targeted, successful rehabilitation.


PAPER 1: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO LIGHT INTENSITY

Rusmana1, Dony Rachmanadi1, Purwanto Budi Santosa1, Tri Wira Yuwati1 and Laura L. B. Graham2 1Banjarbaru Forestry Research Unit, South Kalimantan (FORDA, Indonesian Ministry of Forestry) 2Kalimantan Forest and Climate Partnership, Palangkaraya, Central Kalimantan

1.1 Introduction

Increased human activity on tropical peatlands has resulted in much of the destruction of the peat swamp forest (PSF) ecosystem. Annual rates of degradation reach up to 1.3 percent (Hooijer et al. 2006) and only 36 percent of tropical peat swamp forest remain currently intact (Posa et al. 2011). In the 1990s, the Mega Rice Project (MRP) in Central Kalimantan opened up one million hectares of peat swamp forest for paddy fields (‘Proyek Pembukaan Lahan Gambut 1 juta hektar’ Anwar 2000). This was achieved by digging drainage canals, which led to drastic changes in the hydrology of the system and resulted in further degradation (fires, flooding, forest loss) (Page et al. 2009). Hoscilo et al. (2011) show that during the 9 years after the project stopped (1996–2005) peatland degradation totaled 72 percent due to fires; whilst no degradation through fires were visible for the 23 years preceding the MRP.

Peatland degradation results in the loss of environmental services such as the hydrological system, the carbon sink, local communities’ livelihoods, and biodiversity (Page et al. 2009; Page et al. 2011). Land degradation has resulted in forest loss and fragmentation, and in the degraded areas the land has mainly become dominated by one-metre high ferns. Under these conditions, light can directly reach the ‘forest’ floor (as the forest canopy is absent) resulting in much higher light intensity than would be experienced under the tropical peat swamp forest canopy.

Light is an important limiting resource for tree seedling establishment, growth, and survival in tropical rainforests (Chazdon et al. 1996). Primary tropical forests are typically comprised of several canopy layers so light conditions within the forest are of heterogeneous but mainly low intensity. In secondary forests more light generally reaches the understorey and the forest floor than in primary forests because logging creates big gaps and more light penetrates. This can trigger seedling and sapling growth (Jans et al. 2004).

Kramer and Kozlowski (1979) also state that light intensity plays a role in the physiological process of photosynthesis. Some plant species require high light intensity, while others require low intensity for optimal growth. The high light intensity found in the degraded peatland area is likely to be one cause of low success rates for reforestation and rehabilitation efforts of peat swamp forests (Graham 2013), with survival rates as low as 0–40 percent for transplanted seedlings younger than one year in age (Rusmana et al. 2005, 2006; Setiyono 2011). Species-specific information regarding the light intensity requirements of peat swamp forest species is still limited, with only a few exceptions, such as ramin (Gonystylus bancanus) (Lazuardi et al. 1996; Jans et al. 2004) and punak (Tetrameristra glabra) (Gavin and Peart 1997).

Determining the species-specific light intensity requirements is an important first step in selecting appropriate species and site-locations for transplanting seedlings in degraded peatlands. Moreover, providing appropriate light intensity can increase productivity during seedling production in nurseries and facilitate optimal species-specific acclimatisation regimes (where shading is gradually removed before seedlings are transplanted).

The objective of this research was to determine the effect of light intensity, through the use of shading nets, on a range of peat swamp forest tree species seedlings during nursery cultivation.


1.2 Materials and methods

After seedlings were collected from various locations, the study was carried out in Banjarbaru Forestry Research Unit greenhouses and seedling nurseries in South Kalimantan. The seed materials collected for this experiment were seeds and wildlings from the Hampangen, Tumbang Nusa, and Mentangai tropical peat swamp forests in Central Kalimantan. At the start of the experiment (August 2011) 20 peat swamp plant species were selected (Table 1.1). Each seed was planted in a separate plastic container/polybag (base diameter 10 cm; height 15 cm), filled with 1:1 (v/v) mixture of top soil and rice husk. Seeds were germinated and maintained in the greenhouse before the experiment began. Wildlings were placed in an area of 1 × 2 m which was covered by plastic films and placed under canopy of tree with light intensity of 60–80 percent. The wildlings were irrigated 3–4 times/day under the plastic cover to maintain a humidity of less than 80 percent and a temperature of 29 °–32 °C.

Table 1.1: Species used in the light intensity—tropical peat swamp forest tree species seedling tolerance study

No Local name Botanical name

1 Resak Cotylelobium sp.

2 Meranti daun kecil Shorea sp.

3 Meranti daun lebar Shorea sp.

4 Kapur naga Calophyllum sclerophyllum

5 Perupuk Lophopetalum javanicum

6 Mahirangan Diospyros bantamensis

7 Bintangur Calophyllum hosei

8 Mandarahan Knema mandarahan

9 Lilin-lilin Parartocarpus venenosus

10 Nyatoh Palaquium sp.

11 Kempas Koompassia malaccensis

12 Galam Melaleuca leucadendra

13 Bintan Licania splendens

14 Pulai Alstonia spatulata

15 Pasir-pasir/Medang telur/tagula Stemonurus scorpioides

16 Mangga-mangga Mangifera sp.

17 Papung Sandoricum beccanarium

18 Syzygium sp. Syzygium sp.

19 Gerunggang Cratoxylum glaucum

20 Jejantik Baccaurea bracteata

Note: K. mandarahan, P. venenosus and L. javanicum were collected as seeds while all other species were from wildlings.

Source: Banjarbaru Forestry Research Unit; funded by KFCP


Sharlon netting was used to control light intensity. Sharlon netting is manufactured from black woven polyethylene with a 50 percent light intensity. The sharlon net was placed over the green house frame, with a floor size of 4 x 25 m. The light intensity of 25 percent was obtained by using two layers of sharlon net, while 50 percent light intensity was obtained by using one layer of sharlon net. For 100 percent light intensity control, the seedlings were placed in the open area of the nursery.

A ‘Randomized Complete Block Design’ experimental design was used for the layout of the 20 peat swamp species across the three light intensity levels, with ten seedling replicates per species per light intensity treatment. The seedling height (from stem base to growing tip), stem diameter (2 cm from soil surface), and the survival rate were observed monthly over four months.

Analysis of variance was carried out across treatments for height, diameter, and survival rate. The data were tested for normal distribution and where this was not upheld, the Kruskal-Wallis signed rank non-parametric test was used. If they results were significantly different, a Tukey post-hoc analysis was conducted. Data that were recorded in the percentage form were transformed using the Arcsine Square Root Transformation for statistical analysis.

1.3 Results and discussion

1.3.1 Survival rates

Ten of the twenty species showed an effect (generally negative) of light intensity at 50 percent or 100 percent, as compared with 25 percent (Figure 1.1 and Table 1.2). Nonetheless, most species attained high survival rates (80 percent–100 percent) across all light intensities, except for the Perupuk, Bintangur and Papung species. This showed that a range of species’ survival rates were affected by light intensity. Planting decisions plans should therefore reflect this variation.


Figure 1.1: Survival rate of PSF tree species seedlings after four months under light intensity treatment

Note: 25 percent—blue, 50 percent—red, 100 percent—green


0

20

40

60

80

100Su

rviv

al ra

te (%

)

Species


Table 1.2: Optimal light intenisty for high survival

No Local name Botanical name Optimal light intensity (%) for survival

1 Resak Cotylelobium sp. 25, 50

2 Meranti daun kecil Shorea sp. 25

3 Meranti daun lebar Shorea sp. 25, 100

4 Kapur naga Calophyllum sclerophyllum 25, 50

5 Perupuk Lophopetalum javanicum 25, 50

6 Mahirangan Diospyros bantamensis 25, 50

7 Bintangur Calophyllum hosei 25, 50

8 Mandarahan Knema mandarahan No difference

9 Lilin-lilin Parartocarpus venenosus No difference

10 Nyatoh Palaquium sp. 25, 50

11 Kempas Koompassia malaccensis No difference

12 Galam Melaleuca leucadendra No difference

13 Bintan Licania splendens No difference

14 Pulai Alstonia spatulata No difference

15 Pasir-pasir/Medang telur/tagula Stemonurus scorpioides No difference

16 Mangga-mangga Mangifera sp. No difference

17 Papung Sandoricum beccanarium 50, 100

18 Syzygium sp. Syzygium sp. No difference

19 Gerunggang Cratoxylum glaucum 25

20 Jejantik Accaurea bracteata No difference


1.3.2 Height

As seedlings were cultivated from a mixture of wildlings and seeds, and had unique growth strategies, the initial and final average heights of the seedlings showed much variation across species (Figure 1.2). Seedlings’ ages were all approximately similar during the study. When comparing within species, but across treatment types, however, the influence of light intensity became apparent. For final heights and height increments attained during the four months study period (Figure 1.2 and Figure 1.3), the majority of the species attained the greatest height increases under the low light intesities, with no species preferring 100 percent light to achieve the highest growth increases (Table 1.3).


Figure 1.2: Average height of PSF tree species seedlings after four months under light intensity treatment



Figure 1.3: Height increment of PSF tree species seedlings after four months under light intensity treatment



0

10

20

30

40

50

60

70He

ight

(cm

)

Species

0

5

10

15

20

25

30

Heig

ht i

ncre

men

t (cm

)

Species


Table 1.3: Optimal light intensity for greatest (height) growth increments

No Local name Botanical name Optimal light

intensity (%) for growth (height)

1 Resak Cotylelobium sp. 25


3 Meranti daun lebar Shorea sp. 50, 100

4 Kapur naga Calophyllum sclerophyllum 25, 50

5 Perupuk Lophopetalum javanicum 25, 50

6 Mahirangan Diospyros bantamensis 25

7 Bintangur Calophyllum hosei 25

8 Mandarahan Knema mandarahan 25

9 Lilin-lilin Parartocarpus venenosus 25

10 Nyatoh Palaquium sp. 25

11 Kempas Koompassia malaccensis 25

12 Galam Melaleuca leucadendra 50

13 Bintan Licania splendens -

14 Pulai Alstonia spatulata 25

15 Pasir-pasir/Medang telur/tagula Stemonurus scorpioides 50

16 Mangga-mangga Mangifera sp. 25, 50

17 Papung Sandoricum beccanarium 25

18 Syzygium sp. Syzygium sp. 25, 50

19 Gerunggang Cratoxylum glaucum 25

20 Jejantik Baccaurea bracteata 25, 50


1.3.3 Stem diameter

Similar to the height measurements, the initial and final average stem diameters of the seedlings showed much variation across species (Figure 1.4, Table 1.4). When comparing within species, and across treatment types, however, the influence of light intensity became apparent. Surprisingly, there were twelve species that showed greater stem diameter increments for the 100 percent light intensity treatment compared with 25 percent light intensity (Figure 1.5 and Table 1.4). This constitutes a reversal of the trend shown for height increments (preferring the low light intensities). Hendromono et al. (2003) show that the stem diameter could influence the stem strength, and that height and diameter ratio should be considered in the seedling production. This may be a strategy employed by the plants, which under high light invest in strength and deposit resources elsewhere, rather than growing tall.


Figure 1.4: Average stem diameter of PSF tree species seedlings after four months under light intensity treatment



Figure 1.5: Stem diameter increment of PSF tree species seedlings after four months under light intensity treatment



0

1

2

3

4

5

6

7

8

9

10St

em d

iam

eter

(mm

)

Species

0

2

4

6

8

10

12

Diam

eter

incr

emen

t (m

m)

Species


Table 1.4: Optimal light intenisty for greatest (stem diameter) growth increments

No Local name Botanical name Optimal light intensity (%)

for growth (stem diameter)

1 Resak Cotylelobium sp. 100


3 Meranti daun lebar Shorea sp. 100

4 Kapur naga Calophyllum sclerophyllum 100

5 Perupuk Lophopetalum javanicum 100

6 Mahirangan Diospyros bantamensis No difference

7 Bintangur Calophyllum hosei No difference

8 Mandarahan Knema mandarahan No difference

9 Lilin-lilin Parartocarpus venenosus 100

10 Nyatoh Palaquium sp. 100

11 Kempas Koompassia malaccensis 100

12 Galam Melaleuca leucadendra 50

13 Bintan Licania splendens -

14 Pulai Alstonia spatulata 100

15 Pasir-pasir/Medang telur/tagula Stemonurus scorpioides 100

16 Mangga-mangga Mangifera sp. No difference

17 Papung Sandoricum beccanarium 100

18 Syzygium sp. Syzygium sp. 50, 100

19 Gerunggang Cratoxylum glaucum No difference

20 Jejantik Baccaurea bracteata No difference


1.4 Discussion

Both survival and growth are important indicators of shade tolerance (Daniels et al. 1979; Lorimer, 1983). The results above indicate that increases in the height of PSF seedlings are commonly less with increasing light levels. However, exposure to full sunlight results in increases to stem diameter. Half the species’ survival rates also responded negatively to high light intensity, but despite this, generally most species still maintained high (over 80 percent) survival rates (Table 1.5).


Table 1.5: Summarised optimal light intensities for survival and growth of the 20 studied PSF tree species, with possible light strategies noted

Botanical name Optimal light

intensity (%) for survival

Optimal light intensity (%) for growth (height)

Optimal light intensity (%) for

growth (stem diameter)

Light strategy

Cotylelobium sp. 25, 50 25 100 Shade-tolerant

Shorea sp. 25 50 100 Shade-tolerant

Shorea sp. 25, 100 50, 100 100 Generalist

Calophyllum sclerophyllum 25, 50 25, 50 100 Shade-tolerant

Lophopetalum javanicum 25, 50 25, 50 100 Shade-tolerant

Diospyros bantamensis 25, 50 25 No difference Shade-tolerant

Calophyllum hosei 25, 50 25 No difference Shade-tolerant

Knema mandarahan No difference 25 No difference Generalist

Parartocarpus venenosus No difference 25 100 Generalist

Palaquium sp. 25, 50 25 100 Shade-tolerant

Koompassia malaccensis No difference 25 100 Generalist

Melaleuca leucadendra No difference 50 50 Shade-tolerant

Licania splendens No difference - - -

Alstonia spatulata No difference 25 100 Generalist

Stemonurus scorpioides No difference 50 100 Sun-loving

Mangifera sp. No difference 25, 50 No difference Sun-loving

Sandoricum beccanarium 50, 100 25 100 Sun-loving

Syzygium sp. No difference 25, 50 50, 100 Sun-loving

Cratoxylum glaucum 25 25 No difference Shade-tolerant

Baccaurea bracteata No difference 25, 50 No difference Generalist


A number of researchers have explored optimum light intensity for some PSF species. Nicholson (1960) found five dipterocarp species - Parashorea malaanonan, Shorea leptoclados, Shorea leprosula, Dryobalanops aromatica and Dipterocarpus stellatus - grew optimally at 87.5 percent relative light intensity. Mori (1980) found Calamus manan (a liana) to grow optimally at 50 percent RLI; and Sasaki and Mori (1981) found Vatica odorata, Dipterocarpus oblongifolia, Shorea assamica and Hopea helferi to grow optimally at 32–53 percent RLIs. Meanwhile, Ashton and de Zoysa (1989) found Shorea trapezifolia to grow optimally in ‘partial sun’; and Zipperlen and Press (1996) found S. leprosula and Drybalanops lanceolata to grow optimally in ‘medium light’. This range of examples highlights two points, also shown by this study: for most species that lack pioneer abilities, 100 percent RLI is too high for optimal growth, but the range of optimal conditions varies greatly from species to species.


In this research, seedlings at the 50 percent light level showed the highest height growth and basal diameter growth. In line with our results, Lee et al. (1996) reported the highest growth rates for ramin seedlings at ≥ 40 percent sunlight. In another study, 15 tropical tree species had the highest growth rates between 50 and 75 percent sunlight, above which growth rates declined (Poorter 1999). Quercus pagoda seedlings had lower leaf mass when grown under 27 percent sunlight compared with full sunlight (Gardiner and Krauss, 2001). An increase in height due to the effect of shade was recorded in species like Ailanthus triphysa (Saju 1992). Sazuki and Jacaline (1986) observed that species like Hopea and Valeria mangachopi grow vigorously out in the open.

Aminuddin (1982) grew seedlings in artificial shadehouses set at at 21, 33, 55, 88, and 100 percent of full sunlight. He observed maximum height and stem diameter in seedlings grown at 33 percent of sunlight. However, these shading trials only spanned the upper end of the range of conditions experienced by seedlings in a natural forest environment (Chazdon et al. 1996). The greatest responses were among plants grown at 40 percent full sunlight. Limited response to varying shade may be characteristic of plants that are tolerant of shade, and intolerant of direct sunlight (Bazzaz and Pickett 1979).

1.5 Conclusion

1) Half of the species showed consistently high survival at all light intensity treatments, whilst others had lower survival rates at 100 percent compared to 25 percent or 50 percent light intensity. Nearly all species, however, regardless of light intensity, maintained over 80 percent survival.

2) 25 percent or 50 percent of light intensitity were optimal to attain high height growth increments, whilst no species attained their highest height growth increments at 100 percent. Over half the species, however, attained their highest stem diameter increments at 100 percent light intensity, with the remaining species showing no difference in stem diameter across the light intensity treatments.

3) The above three findings illustrate that:

a. All the 21 species chosen have good survival rates for use in silviculture and reforestation activities.

b. Most of the species showed adaptability to high light conditions; under low light species invested in height growth, whilst under high light species invested in stem strength.

4) Seedling acclimatisation to high light should always be carried out for at least 2 months before being planted in the degraded, open areas of the peatlands. Knowledge of how each species responds to high light intensity can be used to inform:

a. The selection of planting sites, and

b. Growth patterns in the nursery (i.e rapid height growth, or structure-stength investments).


1.6 References Ashton, P. M. S. and N. D. De Zoysa. 1989. Performance of Shorea trapezifolia (Thwaites) Ashton seedlings growing in

different light regimes. Journal of Tropical Forest Science 1: 356–364.

Anwar, K. 2000. Hambatan lahan gambut rawa untuk pengembangan tanaman pangan dan upaya penanggulangannya. In: Daryono, H.; Sidik, Y.J.; Mile, Y.; Subagyo, E; Hadi, T.S.; Akbar, A. and K. Budiningsih (eds). Prosiding Seminar Pengelolaan Hutan Rawa Gambut dan Ekspose Hasil Penelitian Di Hutan Lahan Basah. Balai Teknologi Reboisasi Banjarbaru. Banjarmasin, 9 March 2000. pp. 144–150.

Aminuddin, H. M. 1982. Light requirements of Dyera costulata seedlings. Malaysian Forester 45: 203–208.

Bazzaz, F. A. and S. T. A. Pickett, 1979. Physiological ecology of tropical succession: a comparative review. Annual Review of Ecology and Systematics 11: 287–310.

Chazdon, R.l., Pearcy R., Lee D., and N. Fetcher. 1996. Photosynthetic responses of tropical plants to contrasting light environments. In Mulkey SS et al. (eds). Tropical Forest Plant Ecophysiology, 5–55. Chapman and Hall, New York.

Gardiner, Es and K. W. Krauss. 2001. Photosynthetic light response of flooded cherrybark oak (Quercus pagoda) seedlings grown in two light regimes. Tree Physiology 21: 1103–1011.

Gavin, D. G. and D. R. Peart. 1997. Spatial structure and regeneration of Tetramerista glabra in peat swamp rain forest in Indonesian Borneo. Plant Ecology 131: 223–231.

Graham, L. L. B. 2013. Restoration from within: An interdisciplinary methodology for tropical peat swamp forest restoration, Indonesia. PhD thesis submitted to the University of Leicester, http://hdl.handle.net/2381/28192.

Hooijer, A., Silvius, M., Wösten, H. and S. Page. 2006. PEAT-CO2, Assessment of CO2 emissions from drained peatlands in SE Asia. Delft Hydraulics report Q3943.

Hoscilo, A., Page, S.E., Tansey, K.J., and J.O. Rieley. 2011. Effect of repeated fires on land-cover change on peatlandin southern Central Kalimantan, Indonesia, from 1973 to 2005. International Journal of Wildland Fire 20: 578–588.

Jans, W., Dibor, L., Kruijt, B., and P. van deer Meer 2004. Leaf properties, photosynthetic rates and growth strategies of ramin. Joint Working Group Malaysia—The Netherlands Sustainable Management of Peat Swamp Forests of Sarawak with Special Reference to Ramin.

Kramer, P.J. and T.T. Kozlowski. 1979. Physiology of Woody Plants, Academic Press Inc. London.

Lazuardi, D. and R. Supriadi. 1996. Hubungan antara ketergantungan air dengan daya hidup tanaman ramin (Gonystylus bancanus) di belukar rawa galam rawa gambut bekas terbakar. Prosiding Ekspose Hasil Penelitian dan Ujicoba Balai Teknologi Reboisasi Banjarbaru, held in Banjarbaru Forestry Institute in 1996.

Lee Dw, Krishnapillay B, Haris M, Marzalina M and Sk. Yap. 1996. Seedling development of Gonystylus bancanus (ramin melawis) in response to light intensity and spectral quality. Journal of Tropical Forest Science 8: 520–531.

Mori, T. 1980. Growth of rotan manau (Calamus manan) seedlings under various light conditions. Malayan Forester 43: 187–192.

Nicholson, D. I. 1960. Light requirements of seedlings of 5 species of Dipterocarpaceae. Malayan Forester 23: 344–356.

Page, S., A. Hoscilo, H. Wosten, J. Jauhiainen, M. Silvius, J. Rieley, H. Ritzema, K. Tansey, L. Graham, H. Vasander, and S. Limin. 2009. Restoration ecology of lowland tropical peatlands in Southeast Asia: Current knowledge and future research directions. Ecosystem 12: 888-905.

Page, S. E., Rieley, J. O. and C. J. Banks. 2011. Global and regional importance of the tropical peatland carbon pool. Global change biology 17: 798-818

Posa, M.R.C., Wijedasa, R.S., and R.T. Corlett. 2011. Biodiversity and Conservation of Tropical Peat Swamp Forests. BioScience, 61, No. 1: 49–57

http://hdl.handle.net/2381/28192


Rusmana. 2005. Laporan ujicoba pembangunan demplot blangeran (Shorea balangeran) di lahan rawa gambut Kalimantan Tengah. Balai Penelitian Kehutanan Banjarbaru. Unpublished.

Rusmana. 2006. Laporan ujicoba pembangunan demplot jelutung (Dyera polyphylla), pulai (Alstonia spatulata) dan katiau (Palaquium sp.) di lahan rawa gambut. Balai Penelitian Kehutanan Banjarbaru. Unpublished.

Saju, P. V. 1992. Effect of shade on the growth of Tectona grandis, Grevilllea robusta and Aillanthus triphysa seedlings. B.Sc. Project work, Kerala Agricultural University, Kerala, India.

Sazuki, T. and D.V. Jacaline. 1986. Response of dipterocarp seedlings to various light conditions under forest canopies. Bulletin of the Forestry and Forest Products Research Institute, Japan (336): 19–34.

Setiyono, 2011. Ujicoba penanaman di lahan rawa gambut terdegradasi di Kalimantan Tengah. Laporan hasil penelitian. Pusat Litbang Konservasi dan Rehabilitasi Lahan. Unpublished.

Zipperlen, S. W. and M. C. Press. 1996. Photosynthesis in relation to growth and seedling ecology of two dipterocarp rain forest tree species. Journal of Ecology 84: 863–876.


PAPER 2: RESPONSE OF PEAT SWAMP FOREST SPECIES TO DROUGHT Dony Rachmanadi1, Pranatasari Dyah Susanti1, Rusmana1, Tri Wira Yuwati1, Purwanto Budi Santosa1, and Laura L.B. Graham2

1Banjarbaru Forestry Research Unit, South Kalimantan (FORDA, Indonesian Ministry of Forestry) 2Kalimantan Forest and Climate Partnership, Palangkaraya, Central Kalimantan

2.1 Introduction

Indonesia is home to 64 percent (about 27 million hectares) of the world’s tropical peat swamp forest. Tropical peat swamp forests (TPSF) in Indonesia have the greatest biodiversity value and the thickest peat in the world and also supply environmental services such as water regulation, carbon pools, unique biodiversity, and others (Barchia 2006; Hadisuparto 2004; McKinnon et al. 1996; Rieley 1992). In the last 10 years, however, forest degradation in Indonesia has reached 1.6–2.0 million hectares per year, or 1.3 percent per year of the total area (Hooijer et al. 2006). Improper forest management, forest conversion, overcutting, illegal logging, land occupation and forest fires have caused forest degradation. The degradation has threatened the peat swamp forest biodiversity and environmental services.

Forest degradation creates land and forest fragmentation resulting in a range of forest conditions from secondary forest to severely degraded, open-condition areas with limited woody vegetation and dominance of understorey vegetations and bushes (Page et al. 2009). This degradation results in changes to the environment, including higher fluctuation of the groundwater table; higher temperature and light intensity; physical, chemical and biological changes to the soil conditions; and increasing competition from undergrowth plants and bushes (Page et al. 2009; Graham 2013). Drought and low water levels occur following the construction of canals in the peat-dome (built to facilitate peat drainage), which are built to drain the peat for various alternative peatland land management practices (Page and Rieley 2005).

Based on the high rates of degradation and the recognised importance of TPSF environmental services, numerous restoration projects are currently underway or being initiated (Page et al. 2009). However, restoration of TPSF is a difficult process that is restricted by several factors. TPSF shows low ability to regenerate, particularly following multiple disturbance events (Hoscilo et al. 2011).

One option is to restore TPSF ecosystem through reforestation: transplanting seedlings into degraded areas where natural regeneration is unlikely (Holl 2012). TPSF tree species are, however, adapted to wet, closed-canopy conditions; very different to the varyingly dry or flooded, high light, and high temperature conditions experienced in degraded peatland areas (Page et al. 2009). There has been little published work that establishes which TPSF tree species can tolerate these degraded condition. Thus, investigating appropriate transplant species is a priority (Page et al. 2009; van Eijk et al. 2009).

One trait that must be understood is the tolerance of these species to drought. Kramer (1983) states that water stress can affect plant growth by modifying species’ anatomy, morphology, physiology and biochemistry. Water stress can decrease the cell development, which leads to decreased plant growth rates, the elongation of the stem, leaf growth, prolonged stomata opening, the thickening of the leaves and increased shoot-to-root ratio.

Water content in plants vary between 70–90 percent depending on the species, the age of the plant and the environment. Water is needed for several functions of plants, namely: 1) as a medium for chemical reactions, 2) as a medium for transportation of organic and inorganic substrates, 3) as a raw material for photosynthesis, the hydrolisis process and other chemical reactions inside the plant cells, 4) as a medium that provides turgor to plant cells, 5) and in transpiration to cool the surface of plants (Cao 2000, Tobin et al.


1999). Without sufficient water supplies, numerous plant functions no longer operate and the plant first suffers dehydration, then desiccation, and eventually mortality.

Given the TPSF restoration activities currently underway, and the lack of knowledge regarding ecological species-characteristics to support these activities, there is at present a gap in research that must be addressed. One important ecological character trait of TPSF tree species that must be better understood is their tolerance to drought. The objective of this research, therefore, was to determine and analyse the tolerance of several TPSF tree species under a range of drought conditions.

2.2 Materials and methods

The study was carried out in the Banjarbaru Forestry Research Unit greenhouse and nursery in South Kalimantan. 21 peat swamp plant species were selected (see rationale for selection in Section 1, and Table 2.1 for the species selected). The species were subsequently split into pioneer and climax species groups based on their known growth strategies (Table 2.2). The seed materials collected for this experiment were seeds and wildlings from TPSF of Hampangen, Tumbang Nusa, Petak Bahandang, in Mantangai (at Central Kalimantan) and Landasan Ulin (at South Kalimantan).

Each seed was planted in a separate plastic container/polybag (base diameter 12 cm; height 17 cm), filled with 1:1 (v/v) mixture of top soil and rice husk. The nutrient status of the growth media is presented in Table 2.1. Seeds were germinated and maintained in the greenhouse before the experiment began. Wildlings were placed in an area of 1 × 2 m, covered by plastic films and placed under the canopy of a tree with light intensity of 60-80 percent. The wildlings were irrigated 3–4 times/day under the plastic cover to maintain a humidity of less 80 percent and temperature 29°–32°C.


Table 2.1: Peat swamp forest plant species used in the study of seedlings’ response to drought, with their morphological information at the beginning of the study period

No Species (Local name)

Plant material

Collection origin

Age (month)

Height (cm)

Diameter (cm)

No. leaves

1 Calophyllum hosei (Bintangur) seedling Tumbang Nusa 5 9–15 0.1–0.3 3–12

2 Stemonurus scorpioides (Medang telur, Pasir-pasir) wildling Mentangai 9 17–23 0.3–0.4 4–9

3 Calophyllum sclerophyllum (Kapur naga) seedling Hampangen 5 17–31 0.2–0.4 3–9

4 Shorea sp. (Meranti daun kecil) wildling Hampangen 5 8–22 0.1–0.2 3–9

5 Shorea sp. (Meranti daun lebar) wildling Hampangen 5 17–

33.5 0.2–0.4 3–9

6 Alstonia spatulata (Pulai rawa) wildling Hampangen 3 23–35 0.3–0.6 4–14

7 Lophopetalum javanicum (Perupuk) wildling Mentangai 9 24–60 0.3–0.6 4–21

8 Disopyros bantamensis (Mahirangan) wildling Tumbang Nusa 9 21–33 0.2–0.5 3–11

9 Knema mandarahan (Mandarahan) seedling Tumbang Nusa 3 7–17 0.1–0.2 3–7

10 Palaquium sp. (Nyatoh) wildling Hampangen 5 12–24.5 0.2–0.4 2–6

11 Melaleuca leucadendra (Galam) wildling Landasan Ulin 5 18–41 0.1–0.2 6–25

12 Cotylelobium sp. (Resak) wildling Mentangai 9 21–53 0.2–0.4 7–52

13 Licania splendens (Bintan) seedling Petak Bahandang 5 11–20 0.1–0.2 2–6

14 Lithocarpus sp. (Pampaning) wilding Tumbang Nusa 6 15–29 0.1–0.3 2–7

15 Parartocarpus venenosus (Lilin-lilin) seedling Tumbang Nusa 6 10–17 0.3–0.4 3–5

16 Mangifera sp. (Mangga-mangga) wilding Hampangen 5 9 -16 0.1–0.2 3–11

17 Sandoricum beccanarium (Papung) wilding Hampangen 5 15-20 0.1–0.4 8 -15

18 Cratoxylum glaucum (Gerunggang) seedling Tumbang Nusa 8 10–15 0.1–0.3 6–10

19 Combretocarpus rotundatus (Perepat) wildling Hampangen 5 16–20 0.2–0.4 7–13

20 Baccaurea bracteata (Jajantik) seedling Hampangen 7 8–13 0.1–0.3 2–4

21 Kajalaki seedling Mentangai 9 9–17 0.2–0.3 2–3



Table 2.2: Grouping of species based on their known growth strategies

No Species characteristic

Pioneer species Climax species

1 Alstonia spatulata Calophyllum hosei

2 Melaleuca leucadendra1 Calophyllum sclerophyllum

3 Cratoxylum glaucum Shorea sp. daun kecil

4 Combretocarpus rotundatus Shorea sp.

5 Stemonurus scorpioides Disopyros bantamensis

6 Lophopetalum javanicum2 Palaquium sp.

7 Knema mandarahan Cotylelobium sp.

8 Licania splendens Parartocarpus venenosus

9 Lithocarpus sp.

10 Mangifera sp.

11 Sandoricum beccanarium

12 Baccaurea bracteata

13 Kajalaki Note: 1) Commonly on sulphate acid soil. 2) Commonly on fresh-water peatswamp forest along river side.


Table 2.3: Soil chemical content of the growth media in the nursery

Sample pH H2O

pH KCl

DHL N C-Org KA Kdd Cadd Mgdd KTK Aldd P Bray 1 Fe

(mS/cm) (%) (cmd(+)/Kg) (ppm P2O5) (ppm)

1. 5.5 4.5 0.084 0.157 2.264 1.94 0.07 0.85 0.397 5 1.02 6.375 37.05

2. 5.5 4.5 0.09 0.086 2.638 1.79 0.073 1.078 0.924 3 0.61 31.199 40.74

3. 5.4 4.5 0.078 0.086 2.271 2.23 0.065 0.876 0.581 5 0.41 30.225 36.15

4. 5.4 4.6 0.084 0.072 1.891 2.18 0.063 0.671 0.281 5.5 0.66 0.755 38.34

5. 5.6 4.6 0.085 0.071 1.738 2.05 0.069 0.608 0.294 5 0.61 0.999 39.8



Seeds were germinated in a germination container. After growing two leaves, the seedlings were transplanted into polybags and grown for approximately one month in the greenhouse. After one month, the seedlings were moved under shading net (paranet) for two months then acclimatised in open area for another two weeks.

Wildlings were packed using newspapers, wet sacks and plastic bags, and carried using boxes. Several leaves were cut to reduce evapotranspiration. A root stimulating hormone Rootone-F was applied before the wildlings were transplanted in polybags. The wildlings were then put inside the simple plastic greenhouse for 1–1.5 months. Gradually, the plastic cover was opened after one month. After that, the wildlings were then moved under a shade net where they remained for approximately three months. The wildlings were then acclimatised in the open area for 1 month.

The design of the research was a Completely Randomized Design with four treatments, namely:

- Treatment 1: watering once a day

- Treatment 2: watering twice a week

- Treatment 3: watering once a week

- Treatment 4: watering once a fortnight

Each treatment was replicated 10 times, except for Kajalaki, which was only replicated 5 times due to limited seedling stocks. Watering was carried out for 10 minutes with approximately 10 litre water applied per watering.

Survival, height growth, diameter growth and changes in leaf numbers were recorded every two weeks. At the end of the experiment, 50 percent of the total remaining seedlings and wildlings were harvested and the root length, shoot-root ratio, and root and shoot biomasses were recorded.

2.3 Results

2.3.1 Survival rates

Seedling survival after four months was calculated for all species and for each of the watering intensity treatments. For all species there was a consistent response of lower survival rates to lower watering intensity. The survival rates between the two species groups were not significantly different with regards to the watering intensity treatments for once a day, twice a week, and once a week. For the once a fortnight watering treatment, however, the climax species showed a lower survival rate than the pioneer species (5 percent and 30 percent respectively). For the once a day and twice a week watering treatments there was no significant difference in the survival rates, with both species groups attaining survival rates of, on average, between 95–100 percent. The survival rates for the once a week and once a fortnight watering intensity treatments were significantly lower, however, for both species groups, as compared with the watering treatment of once a day; with both groups attaining survival rates of, on average, between 5–60 percent (Figure 2.1, Table 2.4). Most species when watered only once a fortnight did not survive. The species that did survive under once a fortnight conditions were L. splendens, A. spatulata, K. mandarahan, Mangifera sp., P. venenosus and Kajalaki (Table 2.4).


Figure 2.1: Survival rates of two species group for PSF tree species after four months of watering intensity treatments

Note: pioneer species group—red; climax species group—blue


0

20

40

60

80

100

Once a day Twice a week Once a week Once a fortnight

Surv

ival

rate

(%)

Watering intensity

Climax species

Pioneer species


Table 2.4: The optimal watering intensity for maximum height increment (cm) of the PSF tree species after four months

Growth strategy Species (local name) Survival tolerance to extreme

drought conditions

Optimal watering for

highest survival rates

Pioneer species

Alstonia spatulata (Pulai rawa) Survived once-fortnightly watering

Every day, or minimal twice

weekly

Baccaurea bracteata (Jajantik) Requires regular watering

Combretocarpus rotundatus (Perepat) Requires regular watering

Cratoxylum glaucum (Gerunggang) Requires regular watering

Kajalaki Survived once-fortnightly watering

Knema mandarahan (Mandarahan) Survived once-fortnightly watering

Licania splendens (Bintan) Survived once-fortnightly watering

Lithocarpus sp. (Pampaning) Requires regular watering

Lophopetalum javanicum (Perupuk) Requires regular watering

Mangifera sp. (Mangga-mangga) Survived once-fortnightly watering

Melaleuca leucadendra (Galam) Requires regular watering

Sandoricum beccanarium (Papung) Requires regular watering

Stemonurus scorpioides (Medang telur, Pasir-pasir) Requires regular watering

Climax species

Calophyllum hosei (Bintangur) Requires regular watering

Calophyllum sclerophyllum (Kapur naga) Requires regular watering

Cotylelobium sp. (Resak) Requires regular watering

Disopyros bantamensis (Mahirangan) Requires regular watering

Palaquium sp. (Nyatoh) Requires regular watering

Parartocarpus venenosus (Lilin-lilin) Survived once-fortnightly watering

Shorea sp. (Meranti daun kecil) Requires regular watering

Shorea sp. (Meranti daun lebar) Requires regular watering


2.3.2 Growth—height increment

The optimal watering regime for all species was once a day, with some species also attaining non-significant different height increments for watering twice, and even once, weekly (Table 2.5). After four months, the greatest height increment was attained by L. javanicum followed by C. glaucum, S. beccanarium, L. splendens and Cotylelobium sp. (9.5 cm, 6.5 cm, 4.9 cm, 5.4 cm and 4.1 cm respectively). The greatest height increments were mainly found in the pioneer species group, with just one species from the climax group (Cotylelobium sp.) attaining greater height increments.


The seedlings growth for the once a week watering treatment was generally very low; ranging from 0.27 cm on S. scorpiodes up to 3.70 cm on L. splendens. Similarly, the seedlings’ height increment for the once a fortnight watering treatment was lower; ranging from 0.1 cm up to 0.22 cm on L. splendens.

Table 2.5: The effect of watering intensity on the height increment (cm) of the PSF tree species after four months

Species

Watering intensity


Average SE Average SE Average SE Average SE

Pioneer species

A. spatulata 3.5 a 0.6 1.9 b 0.3 1.4 b 0.4 0.9 b 0.2

B. bracteata 4.6 a 0.3 4.2 a 0.3 0.4 b 0.2 - -

C. glaucum 6.5 a 2.3 2.2 b 1.6 - - - -

C. rotundatus 1.2 a 0.8 1.6 a 0.8 1.4 a 0.7 0.1 b 0.1

K. mandarahan 2.1 ab 0.4 2.9 a 0.4 1.4 bc 0.2 0.4 c 0.3

Kajalaki 2.0 a 0.4 2.4 a 0.3 0.5 b 0.3 0.2 b 0.1

L. javanicum 9.5 a 1.7 5.4 b 1.2 - - - -

L. splendens 5.4 a 0.5 9.0 a 0.7 3.7 b 0.8 2.2 b 0.8

Lithocarpus sp. 0.9 a 0.3 1.3 ab 0.3 0.3 bc 0.2 - -

M. leucadendra 2.9 a 0.9 3.8 a 0.9 0.4 b 0.3 - -

Mangifera sp. 1.0 a 0.2 1.0 ab 0.2 0.7 ab 0.2 0.4 c 0.1

S. beccanarium 4.9 a 1.1 2.0 b 0.6 1.8 b 0.6 - -

S. scorpioides 2.6 a 0.3 2.1 a 0.2 0.3 b 0.1 - -

Climax species

C. hosei 1.9 a 0.3 1.8 a 0.3 0.3 b 0.3 0.1 b 0.1

C. sclerophyllum 2.4 a 0.3 1.7 ab 0.4 1.0 bc 0.4 - -

Cotylelobium sp. 4.1 a 1.2 5.8 ab 1.3 1.5 bc 0.5 - -

D. bantamensis 1.3 a 0.4 0.9 ab 0.2 0.3 b 0.2 - -

P. venenosus 3.4 a 0.5 4.1 a 0.4 1.4 b 0.2 0.2 b 0.2

Palaquium sp. 1.1 a 0.2 0.8 ab 0.2 0.4 bc 0.2 - -

Shorea sp. (daun lebar) 2.3 a 0.3 2.0 a 0.3 0.2 b 0.1 - -

Shorea sp.(daun kecil) 2.3 a 0.3 1.7 a 0.3 1.5 a 1.3 - - Note: Average value followed by the same letter were not significantly different at the degree of freedom of 5 percent. The statistical analysis was carried out for treatments for each species and not comparing between species; SE = standard error.



Table 2.6: The optimal watering intensity for maximum height increment (cm) and the ability to maintain growth during extreme drought conditions (watering once per week and/or once per fortnight) for the PSF tree species after four months

Growth strategy Species (local name) Optimal watering for greatest

height increments

Maintained height growth during extreme drought

conditions

Pioneer species

Alstonia spatulata (Pulai rawa) Once a day Yes

Baccaurea bracteata (Jajantik) Once a day, twice a week No

Combretocarpus rotundatus (Perepat) Once a day, twice a week, once a week Yes

Cratoxylum glaucum (Gerunggang) Once a day No

Kajalaki Once a day, twice a week Yes

Knema mandarahan (Mandarahan) Once a day, twice a week Yes

Licania splendens (Bintan) Once a day, twice a week Yes

Lithocarpus sp. (Pampaning) Once a day, twice a week No

Lophopetalum javanicum (Perupuk) Once a day No

Mangifera sp. (Mangga-mangga) Once a day, twice a week Yes

Melaleuca leucadendra (Galam) Once a day, twice a week No

Sandoricum beccanarium (Papung) Once a day No

Stemonurus scorpioides (Medang telur, Pasir-pasir) Once a day, twice a week No

Climax species

Calophyllum hosei (Bintangur) Once a day, twice a week Yes

Calophyllum sclerophyllum (Kapur naga) Once a day, twice a week No

Cotylelobium sp. (Resak) Once a day, twice a week No

Disopyros bantamensis (Mahirangan) Once a day, twice a week No

Palaquium sp. (Nyatoh) Once a day, twice a week No

Parartocarpus venenosus (Lilin-lilin) Once a day, twice a week Yes

Shorea sp. (Meranti daun kecil) Once a day, twice a week, once a week No

Shorea sp. (Meranti daun lebar) Once a day, twice a week No


2.3.3 Growth–stem diameter increment

Drought also impacted significantly on the stem basal diameter increments for all species. Many species did not achieve any stem diameter increase during the four month growth period under both once a week and once a fortnight watering treatments (Table 2.7 and 2.8). Under optimal conditions (watering once a day, and for some species, also watering twice-weekly), the highest basal diameter growth was attained by B. bracteata and S. beccanarium for the pioneer species group and C. sclerophyllum and D. bantamensis for the climax species group. Only one species, A. spatulata, showed basal diameter growth under the once a fortnight watering treatment.


Table 2.7: The effect of watering intensity on the basal diameter increments (cm) of PSF tree species after four months

Species

Watering intensity



Pioneer species

A. spatulata 0.04 a 0.01 0.1 ab 0.02 0.02 ab 0.01 0.01 b 0.01

B. bracteata - - 0.7 b 0.1 0.1 a 0.1 - -

C. glaucum 0.6 a 0.2 0.20 b 0.1 - - - -

C. rotundatus 0.1 a 0.1 0.2 a 0.11 - - - -

K. mandarahan 0.02 b 0.01 0.1 a 0.01 - - - -

Kajalaki 0.1 ab 0.02 0.1 a 0.02 - - - -

L. javanicum 0.1 a 0.02 0.1 a 0.02 - - - -

L. splendens - - - - - - - -

Lithocarpus sp. 0.02 a 0.01 0.04 ab 0.01 - - - -

M. leucadendra 0.01 a 0.01 0.02 a 0.01 - - - -

Mangifera sp. - - - - - - - -

S. beccanarium 0.7 a 0.2 0.2 ab 0.1 0.4 ab 0.2 - -

S. scorpioides 0.1 a 0.02 0.04 ab 0.02 - - - -

Climax species

C. hosei 0.03 a 0.01 0.02 ab 0.01 - - - -

C. sclerophyllum 0.03 a 0.02 0.1 ab 0.02 - - - -

Cotylelobium sp. 0.1 a 0.02 0.04 ab 0.01 0.04 ab 0.01 - -


P. venenosus 0.03 b 0.02 0.1 a 0.02 - - - -

Palaquium sp. 0.03 a 0.01 0.04 a 0.02 - - - -

Shorea sp. (daun lebar) 0.1 a 0.02 0.1 a 0.01 - - - -

Shorea sp. (daun kecil) 0.1 a 0.01 0.04 a 0.01 - - - - Note: Average value followed by the same letter were not significantly different at the degree of freedom of 5 percent. The statistical analysis was carried out for treatments for each species and not comparing between species; SE = standard error.



Table 2.8: The optimal watering intensity for maximum stem diameter increment (cm) and the ability to maintain growth during extreme drought conditions for the PSF tree species after four months

Growth strategy Species (local name) Optimal watering for greatest

stem diameter increments

Maintained good stem diameter growth during

extreme drought conditions

Pioneer species

Alstonia spatulata (Pulai rawa) Once a day, twice a week, once a week Yes

Baccaurea bracteata (Jajantik) Twice a week Yes

Combretocarpus rotundatus (Perepat) Once a day, twice a week No


Kajalaki Once a day, twice a week No

Knema mandarahan (Mandarahan) Twice a week No

Licania splendens (Bintan) No increment No



Mangifera sp. (Mangga-mangga) No increment No


Sandoricum beccanarium (Papung) Once a day, twice a week, once a week Yes


Climax species

Calophyllum hosei (Bintangur) Once a day, twice a week No

Calophyllum sclerophyllum (Kapur naga) Once a day, twice a week No

Cotylelobium sp. (Resak) Once a day, twice a week, once a week Yes

Disopyros bantamensis (Mahirangan) Once a day, twice a week Yes

Palaquium sp. (Nyatoh) Once a day, twice a week No

Parartocarpus venenosus (Lilin-lilin) Once a day No

Shorea sp. (Meranti daun kecil) Once a day, twice a week No



2.3.4 Leaf numbers

For the seedling leaf numbers of both the pioneer and climax species groups, there was often a significant effect of watering intensity the from once a week and once a fortnight watering intensity treatments. For these watering treatments, the number of leaves in the pioneer group gradually reduced over the four months, with only four species maintaining their leaf numbers during the once a fortnight watering treatment; C. rotundatus, Mangifera sp., L. splendens, and S. beccanarium. Meanwhile, none of the climax


group species maintained their leaf numbers during extreme drought (either once per week or once per fortnight watering). For the drought conditions of once a week watering, however, more species, including several of the climax species group, maintained their leaves throughout the study period. Watering once a day, or twice a week were generally the optimal watering treatments (Table 2.9 and 2.10).

Table 2.9: The effect of watering intensity on the final total leaf numbers of PSF tree species after four months

Species

Watering intensity



Pioneer species

A. spatulata 1.8 a 0.5 1.2 ab 0.5 - - - -

B. bracteata 2.7 a 0.3 2.4 a 0.4 0.2 b 0.2 - -

C. glaucum 12.0 a 3.8 - - - - - -

C. rotundatus 2.1 a 1.1 0.5 b 0.3 0.4 b 0.3 0.3 b 0.2

K. mandarahan 2.4 a 0.2 2.7 a 0.4 0.9 b 0.3 - -

Kajalaki 1.4 a 0.2 1.8 a 0.2 - - - -

L. javanicum 3.4 a 0.6 0.4 b 0.4 - - - -

L. splendens 2.9 a 0.2 2.8 a 0.4 1.3 b 0.6 0.2 b 0.2

Lithocarpus sp. 1.2 a 0.4 1.4 a 0.3 - - - -

M. leucadendra 5.3 a 1.2 6.2 a 1.7 - - - -

Mangifera sp. 0.8 a 0.3 1.1 a 0.3 0.6 a 0.2 0.5 a 0.2

S. beccanarium 8.3 a 1.5 4.4 ab 1.5 4.2 ab 1.5 0.3 b 0.3

S. scorpioides 1.2 a 0.4 1.4 a 0.3 - - - -

Climax species

C. hosei 4.7 a 0.8 3.2 a 0.4 - - - -

C. sclerophyllum 1.9 a 0.2 1.8 a 0.4 0.2 b 0.1 - -

Cotylelobium sp. 11.4 a 2.3 6.8 a 2.3 0.9 b 0.7 - -


P. venenosus 3.4 a 0.2 3.0 a 0.2 1.3 b 0.4 - -

Palaquium sp. 1.1 a 0.1 0.7 ab 0.2 0.2 bc 0.1 - -

Shorea sp. (daun kecil) 0.9 a 0.3 0.8 ab 0.4 - - - -

Shorea sp. (daun lebar) 1.2 a 0.3 1.1 a 0.2 - - - - Note: Average value followed by the same letter were not significantly different at the degree of freedom of 5 percent. The statistical analysis was carried out for treatments for each species and not comparing between species; SE = standard error.



Table 2.10: The optimal watering intensity for greatest final leaf numbers (cm) and the ability to maintain leaf numbers during extreme drought conditions for the PSF tree species after four months

Growth strategy Species (local name) Optimal watering for greatest final

leaf numbers

Maintained positive leaf numbers during drought conditions

Pioneer species

Alstonia spatulata (Pulai rawa) Once a day No

Baccaurea bracteata (Jajantik) Once a day, twice a week Yes

Combretocarpus rotundatus (Perepat) Once a day, twice a week, once a week Yes


Kajalaki Once a day, twice a week No

Knema mandarahan (Mandarahan) Once a day, twice a week Yes

Licania splendens (Bintan) Once a day, twice a week Yes



Mangifera sp. (Mangga-mangga) Once a day, twice a week Yes


Sandoricum beccanarium (Papung) Once a day Yes


Climax species

Calophyllum hosei (Bintangur) Once a day, twice a week No

Calophyllum sclerophyllum (Kapur naga) Once a day, twice a week Yes

Cotylelobium sp. (Resak) Once a day, twice a week Yes

Disopyros bantamensis (Mahirangan) Once a day, twice a week Yes

Palaquium sp. (Nyatoh) Once a day, twice a week Yes

Parartocarpus venenosus (Lilin-lilin) Once a day, twice a week Yes

Shorea sp. (Meranti daun kecil) Once a day, twice a week, once a week No



2.3.5 Root length

For many species the root length decreased as watering frequency decreased. For several species, however, including the pioneer and climax species, the watering treatment did not significantly affect root length, even under once-fortnightly watering conditions; M. leucadendra, C. rotundatus, Mangifera sp., and Lithocarpus sp. for the pioneer group species; and D. bantamensis, Cotylelobium sp., and Shorea sp. (daun kecil) for the climax group species.


The maximum root length achieved after four months of treatment was attained by Lithocarpus sp. for the pioneer group; with an average of 21.0 cm, and by Shorea sp. (daun lebar) for the climax group; with an average of 21.3 cm.

Table 2.11: The effect of watering intensity on the root length (cm) of PSF tree species after four months

Species

Watering intensity



Pioneer species

A. spatulata 20.5 ab 5.4 25.2 a 2.2 13.3 b 1.3 11.7 b 2.0

B. bracteata 15.3 a 1.9 18.6 a 1.9 10.1 ab 3.5 5.3 b 0.7

C. glaucum 11.3 b 0.8 7.0 a 1.4 5.3 a 2.2 5.4 a 1.2

C. rotundatus 5.3 a 2.1 4.5 a 0.4 3.3 a 1.1 4.3 a 0.6

K. mandarahan 16.8 a 2.0 11.6 ab 3.0 9.8 ab 0.8 5.8 b 0.6

Kajalaki 13.8 a 1.5 12.4 ab 2.1 7.6 ab 1.4 8.4 b 0.5

L. javanicum 20.0 a 1.2 17.3 a 3.6 7.7 b 2.2 12.0 b 3.9

L. splendens 10.8 a 0.3 9.5 ab 0.3 4.2 bc 2.6 2.5 c 0.2

Lithocarpus sp. 21.0 a 4.4 16.5 a 3.6 9.5 a 0.3 11.2 a 0.8

M. leucadendra 6.8 a 0.9 7.4 a 0.7 5.9 a 1.2 6.8 a 0.3


S. beccanarium 15.1 a 2.2 13.5 a 2.4 8.5 ab 1.6 5.2 b 0.8

S. scorpioides 16.8 a 1.5 17.4 a 0.7 8.4 b 1.8 9.2 b 0.9

Climax species

C. hosei 12.8 ab 0.8 16.8 a 2.6 9.4 b 0.7 9.2 b 0.9

C. sclerophyllum 18.4 a 1.5 17.6 ab 2.0 11.4 bc 0.7 9.6 c 1.7

Cotylelobium sp. 16.0 a 2.3 15.6 a 1.2 13.6 a 2.5 8.6 a 1.3

D. bantamensis 14.5 a 2.9 13.5 a 0.9 10.0 a 1.1 10.3 a 1.5

P. venenosus 13.2a 1.2 19.3b 4.3 10.7a 2.3 8.6ab 2.6

Palaquium sp. 10.2 ab 1.3 10.6 b 1.7 5.4 a 0.7 7.8 a 0.6

Shorea sp. (daun kecil) 10.3 a 2.8 8.3 a 1.9 3.7 a 0.5 6.5 a 0.8

Shorea sp. (daun lebar) 21.3 a 3.0 12.5 b 0.8 9.7 b 1.9 8.8 b 1.4 Note: Average value followed by the same letter were not significantly different at the degree of freedom of 5 percent. The statistical analysis was carried out for treatments for each species and not comparing between species; SE = standard error



Table 2.12: The watering intensity for maximal root length of the PSF tree species after four months

Growth strategy Species (local name) Watering treatment for maximal root length

Pioneer species

Alstonia spatulata (Pulai rawa) Once a day, twice a week

Baccaurea bracteata (Jajantik) Once a day, twice a week, once a week

Combretocarpus rotundatus (Perepat) No difference

Cratoxylum glaucum (Gerunggang) Once a day

Kajalaki Once a day, twice a week, once a week

Knema mandarahan (Mandarahan) Once a day, twice a week, once a week

Licania splendens (Bintan) Once a day, twice a week

Lithocarpus sp. (Pampaning) No difference

Lophopetalum javanicum (Perupuk) Once a day, twice a week

Mangifera sp. (Mangga-mangga) No difference

Melaleuca leucadendra (Galam) No difference

Sandoricum beccanarium (Papung) Once a day, twice a week, once a week

Stemonurus scorpioides (Medang telur, Pasir-pasir) Once a day, twice a week

Climax species

Calophyllum hosei (Bintangur) Once a day, twice a week

Calophyllum sclerophyllum (Kapur naga) Once a day, twice a week

Cotylelobium sp. (Resak) No difference

Disopyros bantamensis (Mahirangan) No difference

Palaquium sp. (Nyatoh) Once a day, once a week, once a fortnight

Parartocarpus venenosus (Lilin-lilin) Twice a week

Shorea sp. (Meranti daun kecil) No difference

Shorea sp. (Meranti daun lebar) Once a day



2.3.6 Shoot biomass

For the pioneer species group, shoot biomass gradually decreased as watering frequency decreased, which contrasted strongly with the climax species group that had increasing shoot biomass with decreasing watering frequency, except under the once-fortnightly treatment (Figure 2.2).

Figure 2.2: The average of shoot biomass (g) of two groups PSF tree species after four months of watering intensity treatments



2.3.7 Root biomass

For root biomass, the climax and pioneer species groups both showed the same trend; root biomass increased from the once a day watering treatment to the twice a week treatment and then decreasing after that until the once a fortnight treatment (Figure 2.3). For the climax species group, there was no significant difference in root biomass between the twice a week and the once a week treatment the latter of which attained the greatest root biomass. The once a day and once a fortnight treatments, however, led to significantly lower root biomasses than the two aforementioned watering regimes. For the pioneer species group, the highest root biomass value was also achieved under the twice a week watering treatment, which was significantly greater than the other three watering regimes (no significant difference between these).

0.00

0.50

1.00

1.50

2.00

2.50

3.00

3.50

4.00


Avea

rage

shoo

t bio

mas

s (g)

Watering intensity


Figure 2.3: The average of root biomass (g) of two groups PSF tree species after four months watering intensity treatment



2.3.8 Shoot-Root Ratio

Shoot-root ratio represents the balanced growth of the plant. For both species groups, there was no consistent effect of watering intensity treatment. Almost all species within the pioneer species group showed no significant difference in the shoot-root ratio except for M. leucadendra, L. splendens and S. scorpioides. For the climax species group, the different watering treatments showed no significant difference in the shoot-root ratio for all species.

0.00

0.20

0.40

0.60

0.80

1.00

1.20


Aver

age

root

bio

mas

s (g

)

Watering intensity


Table 2.13: The effect of watering intensity on the shoot-root ratio (g) of PSF tree species after four months

Species

Watering intensity



Pioneer species

A. spatulata 4.6 a 0.8 1.5 a 0.2 2.8 a 1.0 1.1 a 0.1

B. bracteata 3.9 a 0.5 4.2 a 0.5 2.2 a 1.7 0.5 a 0.2

C. glaucum 5.4 a 1.2 4.1 a 0.5 5.8 a 0.7 6.7 a 0.8

C. rotundatus 2.4 a 0.5 1.7 a 0.8 3.9 a 1.1 2.2 a 0.5

K. mandarahan 2.8 a 0.4 2.5 a 0.2 3.1 a 0.4 3.6 a 0.6

Kajalaki 4.5 a 0.9 1.4 a 0.3 1.8 a 0.4 4.9 a 0.8

L. javanicum 4.6 a 1.3 2.5 a 0.4 2.1 a 0.4 2.9 a 1.1

L. splendens 2.6 a 0.6 1.9 a 0.4 2.8 a 0.9 8.9 b 2.4

Lithocarpus sp. 2.3 a 0.4 1.2 a 0.2 2.8 a 0.2 2.1 a 0.2

M. leucadendra 4.5 b 0.4 1.0 a 0.2 2.3 a 0.3 2.4 a 0.4


S. beccanarium 2.8 a 0.6 5.2 a 0.5 2.1 a 0.6 1.8 a 0.3

S. scorpioides 2.4 b 0.2 1.3 a 0.2 2.5 b 0.3 4.5 c 0.3

Climax species

C. hosei 2.2 a 0.3 1.1 a 0.2 3.1 a 0.7 1.5 a 0.2

C. sclerophyllum 2.8 a 0.3 3.1 a 0.8 3.8 a 0.2 5.2 a 0.7

Cotylelobium sp. 8.4 a 1.3 1.2 a 0.2 1.6 a 0.4 7.8 a 1.1

D. bantamensis 4.1 a 0.9 2.6 a 0.5 2.5 a 0.3 2.3 a 0.4

P. venenosus 2.9 a 1.5 1.7 a 0.6 5.5 a 1.5 0.9 a 0.5

Palaquium sp. 3.7 a 0.6 1.9 a 0.3 1.9 a 0.4 3.3 a 0.8

Shorea sp. (daun kecil) 2.9 a 1.5 3.7 a 2.1 3.2 a 0.9 4.5 a 0.5

Shorea sp. (daun lebar) 2.2 ab 0.4 0.9 a 0.3 4.1 b 0.3 3.1 ab 0.8 Note: Average value followed by the same letter were not significantly different at the degree of freedom of 5 percent. The statistical analysis was carried out for treatments for each species and not comparing between species; SE = standard error



Table 2.14: The watering intensity for greatest shoot-:root ratio of the PSF tree species after four months

Growth strategy Species (local name) Greatest shoot-root ratio

Pioneer species

Alstonia spatulata (Pulai rawa) No difference

Baccaurea bracteata (Jajantik) No difference

Combretocarpus rotundatus (Perepat)

Cratoxylum glaucum (Gerunggang) No difference

Kajalaki No difference

Knema mandarahan (Mandarahan) No difference

Licania splendens (Bintan) Once a fortnight

Lithocarpus sp. (Pampaning) No difference

Lophopetalum javanicum (Perupuk) No difference

Mangifera sp. (Mangga-mangga) No difference

Melaleuca leucadendra (Galam) Once a day

Sandoricum beccanarium (Papung) No difference

Stemonurus scorpioides (Medang telur, Pasir-pasir) Once a day, once a week

Climax species

Calophyllum hosei (Bintangur) No difference

Calophyllum sclerophyllum (Kapur naga) No difference

Cotylelobium sp. (Resak) No difference

Disopyros bantamensis (Mahirangan) No difference

Palaquium sp. (Nyatoh) No difference

Parartocarpus venenosus (Lilin-lilin) No difference

Shorea sp. (Meranti daun kecil) No difference

Shorea sp. (Meranti daun lebar) Once a week


2.4 Discussion and conclusion

This research has shown that peat swamp forest plant species can likely survive in dry conditions for 3–4 days. However, intolerance begins from day 7 up to day 14. The conditions would presumably worsen when the rate of evapotranspiration is higher, such as in degraded peat swamp forests (in this research, the evapotranspiration rate for the plants were normal due to the stable temperature of no more than 30oC and humidity levels of 80 percent–90 percent).

Overall, survival rates, shoot growth and leaf numbers commonly reduced in response to drought, as the literature suggests. Root length also reduced or was not affected by drought conditions, which was surprising as this is counter to what the literature would suggest—the roots should grow and forage for water. Shoot and root biomass generally decreased under drought conditions; thus, there was no consistent trend regarding the shoot-root ratio.


In the analysis, it was useful to analyse the data collected by species groups, as trends were sometimes distinct between them. For example, for the pioneer species group, shoot biomass gradually decreased as watering frequency decreased, which contrasted strongly with the climax species group that had increasing shoot biomass with decreasing watering frequency, except for the once a fortnight treatment. Generally, the pioneer species groups displayed more tolerance to drought than the climax species group.

It was found that only a few species could tolerate drought conditions, namely, Pulai, Kajalaki, Mandarahan, Bintan, Mangga-mangga and Lilin-lilin (Table 2.14). Overall, knowledge of drought tolerance should be used when selecting species for reforestation activities as this study has highlighted several species displaying good drought tolerance.

An interesting finding was the slow growth of the Merapat and Gerunggang species and their intolerance to drought. This is contrary to the findings from several other studies (see Page et al. 1999; and Page et al. 2009), that suggest the two species were of the pioneer type that should championed to endure environmental stress compared with others. This effect would be interesting to explore further to determine the photosynthesis ability, leaf water potential and clorophyl reductance (Katahata 2012).


Table 2.15: Summary of drought tolerance of the 21 studied PSF tree species, based on morhpological responses to drought conditions

Growth strategy Species (local name) Drought tolerance

Pioneer species

Alstonia spatulata (Pulai rawa) Good

Baccaurea bracteata (Jajantik) Acceptable

Combretocarpus rotundatus (Perepat) Acceptable

Cratoxylum glaucum (Gerunggang) Poor

Kajalaki Good

Knema mandarahan (Mandarahan) Good

Licania splendens (Bintan) Good

Lithocarpus sp. (Pampaning) Poor

Lophopetalum javanicum (Perupuk) Poor

Mangifera sp. (Mangga-mangga) Good

Melaleuca leucadendra (Galam) Poor

Sandoricum beccanarium (Papung) Acceptable

Stemonurus scorpioides (Medang telur, Pasir-pasir) Poor

Climax species

Calophyllum hosei (Bintangur) Poor

Calophyllum sclerophyllum (Kapur naga) Poor

Cotylelobium sp. (Resak) Acceptable

Disopyros bantamensis (Mahirangan) Acceptable

Palaquium sp. (Nyatoh) Poor

Parartocarpus venenosus (Lilin-lilin) Good

Shorea sp. (Meranti daun kecil) Poor

Shorea sp. (Meranti daun lebar) Poor



2.5 References Barchia, M.F. 2006. Gambut: Agroekosistem dan transformasi karbon. Gadjah Mada, University Press. Yogyakarta.

Bunker, D. E. and W. P . Carson. 2005. Drought stress and tropical forest woody seedlings: Effect on community structure and composition. Journal of Ecology 93: 794–806.

Cao, K. F. 2000. Water relations and gas exchange of tropical saplings during a prolonged drought in a Bornean heath forest, with reference to root architecture. Journal of Tropical Ecology 16: 101-116.

Engelbrecht, B. M. J. and T. A. Kasar. 2003. Comparative drought-resistance of seedlings of 28 species of co-occurring tropical woody plants. Oecologia 136: 383–393.

Graham, L. L. B. 2013. Restoration from Within: An interdisciplinary methodology for tropical peat swamp forest restoration, Indonesia. PhD thesis submitted to the University of Leicester, http://hdl.handle.net/2381/28192.

Hadisuparto, H. 2004. Hutan rawa gambut sebagai sumberdaya alam dan pengawal lingkungan. Makalah Lokakarya Penanganan Kawasan Eks PLG Sejuta Hektar di Kalimantan Tengah. Palangkaraya.

Holl, K. D. (2012) Chapter 9: Restoration of tropical forests. In: van Andel, J. and Aronson, J. (2012) Restoration Ecology: The New Frontier. Second edition. Wiley-Blackwell Press, Oxford, UK.

Hooijer, A., Silvius, M., Wösten, H. and S. Page. 2006. PEAT-CO2, Assessment of CO2 emissions from drained peatlands in SE Asia. Delft Hydraulics report Q3943.

Hoscilo, A., Page, S.E., Tansey, K.J., and J.O. Rieley. 2011. Effect of repeated fires on land-cover change on peatlandin southern Central Kalimantan, Indonesia, from 1973 to 2005. International Journal of Wildland Fire 20: 578–588.

Kramer, P. J. 1983. Plant and soil water relationships (a modern synthesis). McGraw-Hill Publishing, New York, USA.

Katahata, S. 2012. Photoinhibition and photoprotective functions of plants. Materials from field seminars held by Shizuoka University dan Gadjah Mada University in 2012.

McKinnon, K., G.M. Hatta, H. Halim, and A. Mangalik. 1996. The Ecology of Kalimantan (Indonesian Borneo). Periplus Edition. Indonesia–Canada.

Page, S.E., Rieley, J.O., Shotyk, O.W. and D. Weiss. 1999. Interdependence of peat and vegetation in a tropical peat swamp forest. Philosophical Transactions of the Royal Society B: Biological Sciences 354: 1885–1897.

Page, S.E. and J.O. Rieley. 2005. Wise-use of tropical peatlands, Alterra, Wageningen, The Netherlands.

Page, S., A. Hoscilo, H. Wosten, J. Jauhiainen, M. Silvius, J. Rieley, H. Ritzema, K. Tansey, L. Graham, H. Vasander, and S. Limin. 2009. Restoration ecology of lowland tropical peatlands in Southeast Asia: Current knowledge and future research directions. Ecosystem 12: 888–905.

Rieley, J.O. 1992. The ecology of tropical peatswamp forest (A Souteast Asian perspective). In. B.Y. Aminuddin, S.L. Tan, B. Aziz, J. samy, Z. Salmah, H. Siti Petimah and S.T. Choo (eds). Proceedings of the International Symposium on Tropical Peatland. Malaysia. 244–254p.

Tobin, M. F., Lopez, O. R. and T. A. Kursar. 1999. Responses of tropical understorey plants to a severe drought: Tolerance and avoidance of water stress. Biotropica 31: 570–578.

van Eijk, P., Leenman, P., Wibisono, I. T. C. and W. Giesen. 2009. Regeneration and restoration of degraded peat swamp forest in Berbak National Park, Jambi, Sumatra, Indonesia. Malayan Nature Journal 61: 223–241.

http://hdl.handle.net/2381/28192


PAPER 3: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO FLOODING

Purwanto B Santosa1, Tri Wira Yuwati1, Dony Rachmanadi1, Rusmana1, Laura L. B. Graham2

1Banjarbaru Forestry Research Unit, South Kalimantan (FORDA, Indonesian Ministry of Forestry) 2Kalimantan Forest and Climate Partnership, Palangkaraya, Central Kalimantan

3.1 Introduction

The tropical peat swamp forest (TPSF) ecosystem is currently being degraded at a rapid rate (Limin 2000). According Hooijer et al. (2006), 10.6 million hectares (39 percent) of peatland in Southeast Asia were deforested by 2000, with deforestation continuing at an annual rate of 1.5 percent, resulting in approximately 45 percent degradation of all peatland area by 2006 (12.1 million hectares). Boehm et al. (2003) predicted that from the time of publication, the deforestation rate of the TPSF to the south of Palangka Raya would reach 33 percent in 10 years. Page et al. (2009) show that forest cover between 1973 and 2003 reduced by 78 percent in Block C of the Ex-Mega Rice Project (EMRP), caused of fire, exacerbated by drainage and land-use change. Moreover, according to Miettinen and Liew (2010), over 20 percent of TPSF in Sumatera and Kalimantan are categorised as unmanaged degraded landscape, occupied by ferns, shrubs and secondary growth.

This degradation is caused by forest fires, legal and illegal logging, and over-drainage caused by inappropriate land development (Yuwati et al. 2007; Page et al. 2009). Disturbance and degradation further leads to peat subsidence that results in deep and prolonged flooding in the wet season (Page et al. 2009; Hooijer et al. 2012; Woosten et al. 2008). There remains a lack of information regarding the tolerance of TPSF tree species seedlings to a range of flood levels. Such information is crucial in developing appropriate reforestation activities.

In its natural state, TPSF is an ecosystem that is spatially and temporally waterlogged (Nishumua et al. 2007). The TPSF tree species have adapted to this, with seedlings favouring germination on the naturally forming hummocks (Heino et al. 2006), and trees supporting adapted root systems such as pneumatophores and stilt roots (Page and Rieley 2005). Without these adaptations, flooding can cause plant roots to decay, mainly through oxygen deficiency and increased toxic ions in rhizozphere (Yamanoshita et al. 2001). Hapsari and Adi (2010) show that when plant roots are flooded for a short period they face hypoxia (lack of oxygen). Plants that are completely flooded (covering their roots and shoots) must endure completely anoxic conditions (without oxygen). Scott et al. (1989) report that the effect of flooding/inundation is indicated by the yellowing colour of leaves followed by the falling of leaves, stagnant growth and low dry weight of plant tissues. The magnitude, duration and timing of flood events are important factors regulating primary productivity in wetland forests (Megonigal 1992).

Upon degradation, the hummock-hollow topography of TPSF is lost and flooding becomes more severe, resulting in difficult germination and growth for seedlings, inhibiting natural regeneration (Giesen 2004). For successful restoration, and in particular, for reforestation activities to take place, species that can tolerate different degrees of flooding must be explored and used. In this research, the impact of flooding on survival rates and seedlings growth of 17 TPSF tree species under four different flood conditions was investigated.


3.2 Material and methods

The study was carried out in the greenhouses and nurseries of Banjarbaru Forestry Research Unit of South Kalimantan. At the start of the experiment (June 2011), 17 peat swamp plant species were selected (Table 3.1). The seed materials collected for this experiment were seeds and wildlings from TPSF of Hampangen, Tumbang Nusa, and Mentangai, Central Kalimantan. Each seed was planted in a separate plastic container (base diameter 10 cm; height 15 cm), filled with 1:1 (v/v) mixture of top soil and rice husk. Seeds were germinated and maintained in the greenhouse before the experiment began. Wildlings were placed in an area of 1 × 2 m that was covered by plastic films and placed under canopy of tree with a light intensity of 60–80 percent. The wildlings were irrigated 3–4 times/day under the plastic cover to maintain a humidity of less than 80 percent and a temperature of between 29°–32° C.

Table 3.1: Species, plant material source and origin, age at the start of treatment, and seedling average dimensions at the start of the inundation experiment

No. Species

(Local name) Plant

material Place of

collection Age

(months) Height (cm)

Diameter (cm)

No. leaves

1 Alstonia spatulata (Pulai rawa) wildling Hampangen 3 22.5 0.4 8

2 Calophyllum hosei (Bintangur) seedling Tumbang Nusa 3 16.6 0.2 7

3 Calophyllum sclerophyllum (Kapur naga)

seedling Hampangen 5 22.1 0.3 4

4 Cotylelobium sp. (Resak) wildling Mentangai 3 33.3 0.2 22

5 Dacrydium pectinatum (Alau) wildling Hampangen 5 18.8 0.4 13

6 Disopyros bantamensis (Mahirangan)

wildling Tumbang Nusa 9 34.9 0.4 7

7 Knema mandarahan (Mandarahan)

seedling Tumbang Nusa 3 11.3 0.2 3

8 Lithocarpus sp. (Pampaning) wilding Tumbang Nusa 6 22.1 0.25 6

9 Lophopetalum javanicum (Perupuk)

wildling Mentangai 9 44.4 0.5 15

10 Mangifera sp. (Mangga-mangga)

wilding Hampangen 5 14.2 0.2 5

11 Palaquium sp. (Nyatoh) wildling Hampangen 5 21.4 0.3 3

12 Parartocarpus venenosus (Lilin-lilin)

seedling Tumbang Nusa 6 18.7 0.47 9

13 Sandoricum beccanarium (Papung)

wilding Hampangen 5 19.1 0.21 13

14 Shorea sp. (Meranti daun kecil)

wildling Hampangen 5 25.0 0.2 6

15 Stemonurus scorpioides (Medang telur, Pasir-pasir)

wildling Mentangai 9 21.4 0.4 7

16 Syzygium pakan (pakan) wildling Mentangai 9 29.9 0.3 14

17 Syzygium sp. wilding Hampangen 5 15.9 0.2 6



The plants were placed in a transparent glass box filled with water to the specified level depending on the treatment. The seedlings were assigned four flooding conditions: a) half the polybag (part of the roots) flooded; b) all of polybag (all roots) flooded; c) half the seedling height (part of the stem and all of the roots) flooded; and d) Full waterlogging treatment (plants flooded to above top of the seedling) (Figure 3.1). The water level height was controlled everyday. The experimental design was a Randomized Complete Design with four water-level treatments. Each treatment group consisted of 17 species and for each species the treatment was replicated with 10 plants. The number of leaves and survival rates were recorded once every two weeks.


3.3 Results, discussion and conclusion

Based on the survival rates and leaf change results (Figure 3.2 and Figure 3.3), of the 17 species studied, 16 showed at least some adverse reaction to flooding, with only Lophopetalum javanicum maintaining good leaf numbers and survival rates even during complete submersion for four months. A further five species showed some degree of tolerance, maintaining good survival rates and leaf numbers during partial stem submersion for four months: Alstonia spatulata, Calophyllum sclerophyllum, Dacrydium pectinatum, Disopyros bantamensis, and Stemonurus scorpioides. The remaining 11 species, however, showed poor survival rates and leaf maintenance once some of the stem was submerged. These results indicate the importance of considering flood depths, seedlings transplant size, season, and species-selection when developing a reforestation plan (Table 3.2).

A B C D

Figure 3.1: Seedling flood treatment


Table 3.2: Flooding tolerance of 17 TPSF tree species based on the flooding experiment

Species (Local name) Flood tolerance based on survival

Flood tolerance based on leaf change

Overall flood tolerance

Alstonia spatulata (Pulai rawa) Some tolerance Some tolerance Some tolerance

Calophyllum hosei (Bintangur) Poor tolerance Poor tolerance Poor tolerance

Calophyllum sclerophyllum (Kapur naga) Poor tolerance Some tolerance Some tolerance

Cotylelobium sp. (Resak) Poor tolerance Poor tolerance Poor tolerance

Dacrydium pectinatum (Alau) Some tolerance Some tolerance Some tolerance

Disopyros bantamensis (Mahirangan) Some tolerance Some tolerance Some tolerance

Knema mandarahan (Mandarahan) Poor tolerance Poor tolerance Poor tolerance

Lithocarpus sp. (Pampaning) Poor tolerance Poor tolerance Poor tolerance

Lophopetalum javanicum (Perupuk) Excellent tolerance Excellent tolerance Excellent tolerance

Mangifera sp. (Mangga-mangga) Poor tolerance Poor tolerance Poor tolerance

Palaquium sp. (Nyatoh) Poor tolerance Poor tolerance Poor tolerance

Parartocarpus venenosus (Lilin-lilin) Poor tolerance Poor tolerance Poor tolerance

Sandoricum beccanarium (Papung) Poor tolerance Poor tolerance Poor tolerance

Shorea sp. (Meranti daun kecil) Poor tolerance Poor tolerance Poor tolerance

Stemonurus scorpioides (Medang telur, Pasir-pasir)

Some tolerance Some tolerance Some tolerance

Syzygium pakan (Pakan) No tolerance No tolerance No tolerance

Syzygium sp. Poor tolerance Poor tolerance Poor tolerance



Figure 3.2: The survival rates of 17 peat swamp forest species based on the inundation experiment

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Figure 3.3: The changes in leaf numbers for each treatment in the inundation experiment

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3.4 References Boehm, H.D.V., F. Siegert, S.H. Limin and A. Jaya. 2003. Land use change in Central Kalimantan over the period 1991–

2001 including impact of selective and illegal logging, MRP establishment and fires. TROPEAT 2002 Symposium, Bali Kuta 18–19 Sept 2002.

Giesen, W. 2004. Causes of swamp forest degradation in Berbak National Park and recommendations for restoration. Water for Food and Ecosystems Programme Project on: Biodiversity & the EMRP Revised draft Euroconsult Mott MacDonald 45 “Promoting the river basin and ecosystem approach for sustainable management of SE Asian lowland peat swamp forests”. ARCADIS Euroconsult, Arnhem, the Netherlands

Hapsari.R.T and M.M. Adie. 2010. Peluang perakitan dan pengembangan kedelai toleran genangan. Journal Litbang Pertanian 29 (2): 50–57

Heino, I., Lampela, M., Jauhiainen, J. and H. Vasander. 2007. Dynamics of the tropical peat swamp forest floor microtopography. In: H. Hayasaka, and A. Ususp (eds.). Proceedings of International Workshop 'Tropical Rain Forest and Boreal Forest Disturbance and Their Affects on Global Warming', Palangka Raya, Indonesia, 16-18 September, 2006. pp. 108-111.

Hooijer, A., Silvius, M., Wösten, H. and S. Page. 2006. PEAT-CO2, Assessment of CO2 emissions from drained peatlands in SE Asia. Delft Hydraulics report Q3943

Hooijer, A., Page, S., Jauhiainen, J., Lee, W. A., Lu, X. X., Idris, A. and G. Anshari. 2012. Subsidence and carbon loss in drained tropical peatlands. Biogeosciences 9: 1053–1071,

Lenssen, J.P.M., Menting, F.B.J., Van Der Putten, W.H., and C.W.P. Blom. 2000. Vegetative reproduction by species with different adaptations to shallow-flood habitats. New phytol 145:61–70.

Limin, S H., Tampung N. S, Patricia E. P., Untung D., and Layuniyati. 2000. Konsep pemanfaatan hutan rawa gambut di Kalimantan Tengah. Prosiding Seminar Pengelolaan Hutan Rawa Gambut dan Ekspose Hasil Penelitian di Hutan Lahan Basah. Banjarmasin, Maret 2000. Balai Teknologi Reboisasi Banjarbaru. pp9–14

Megonigal, J.P. 1992. Effects of flooding on root and shoot production of bald cypress in large experimental enclosures. Ecology 73(4):1182–1193

Miettinen, J. and S. C. Liew. 2010. Status of Peatland Degradation and Development in Sumatra and Kalimantan. AMBIO 39:394–401

Nishimua T.B.,. Suzuki, E., Kohyama, T., and S. Tzuyuzaki. 2007. Moratlity and growth of trees in peat swamp and heath forest in Central Kalimantran after severe drought. Plant Ecology 188:165–177.

Page, S., Hosciło, A., Wösten, H., Jauhiainen, J., Silvius, M., Rieley, J., Ritzema, H., Tansey, K., Graham, L., Vasander, H., and H. Limin. 2009. Restoration Ecology of Lowland Tropical Peatlands in Southeast Asia: Current Knowledge and Future Research Directions. Ecosystems 12: 888–905

Page, S. E and J. O. Reiley. 2005. Wise-use of tropical peatlands. Alterra Publishing, Wageningen, The Netherlands.

Scott, H.D., De Angulo, J., Daniels, M.B., and L.S. Wood. 1989. Flood duration effect on soybean growth and yield. Agronomy 81:631–636.

Van der Meer. P. J. and B.F. Ibie. 2008. Forestry in the Ex-Mega Rice Project Area in Central Kalimatan. Technical Report Number 6 . Euroconsult Mott MacDonald & Deltares

Wösten, J.H.M., Clymans, E., Page, S.E., Rieley, J. O., and S.H. Limin. 2008. Peat-water interrelations in tropical peatland ecosystems in Southeast Asia. Catena 73:212-224

Yamanoshita.T., Nuyim.T., Tange.T., Kojima K., Yagi.H. and S. Sasaki. 2001. Effect of flooding and adventitious roots formation on height growth of Melalauca cajuputi. Silvicultural research report. The University of Tokyo and Nihon Univesity, Japan. pp.47–55

Yuwati T.W., Santosa. P.B., and B. Hermawan. 2007. Arbuscular Mycorriza Fungi Application for Rehabilitation of Degraded Peat Swamp Forest in Central Kalimantan. Carbon-climate-human interaction on tropical peatland. Proceedings of The International Symposium and Workshop on Tropical Peatland, Yogyakarta, 27–29 August 2007, EU CARBOPEAT and RESTORPEAT Partnership, Gadjah Mada University, Indonesia and University of Leicester, United Kingdom. pp.253–258


PAPER 4: RESPONSE OF PEAT SWAMP FOREST SPECIES SEEDLINGS TO MACRONUTRIENTS

Tri Wira Yuwati1, Dony Rachmanadi1, Purwanto B. Santosa1, Rusmana1 and Laura L.B. Graham2 1Banjarbaru Forestry Research Unit, South Kalimantan (FORDA, Indonesian Ministry of Forestry) 2Kalimantan Forest and Climate Partnership, Palangkaraya, Central Kalimantan

4.1 Introduction

One of the constraints to restoration activities on degraded tropical peat swamp forest (TPSF) is the low nutrient levels that are in available in the peat. Tropical peatlands naturally have very low nutrient inputs and are highly acidic. Limited nutrient availability, essential for TPSF plant survival and growth (Sulistiyanto 2004), cause the slow growth of plants. After TPSF degradation, increased peat decomposition rates can lead to nutrient flushes. However, the loss of the living vegetation means these increased nutrient levels are transient—lost into the waterways or lower peat levels, resulting in lower overall nutrient levels (Page and Rieley 2005). This can lead to low survival rates for TPSF seedlings transplanted into degraded peatland areas.

Essential nutrients are the nutrients needed for plants to grow normally. Arnon (1950) contends that there are three criteria for nutrients to be considered essential: (a) the nutrient is important for the plant to grow normally, (b) a particular nutrient cannot be replaced by other nutrients and (c) the nutrient plays a role inside the plant, not outside. Essential nutrients consist of macro-and micro-nutrients. Macro-nutrients are needed in larger amounts while micro-nutrients are needed only in small amounts. Nitrogen, phosphorus, potassium, calcium and magnesium are the common macronutrients across most plants. Nitrogen is important for vegetative growth in plants. Phosphorus is important in the development of roots and ripening of flowers, fruits and seeds. Potassium is needed during the plant’s physiological processes. Finally, calcium and magnesium improve soil acidity and structure (Kramer and Kozlowski 1979).

Several studies have explored seedlings’ growth response to nutrient additions (Santiago et al. 2012; Newberry et al. 2002; Campo and Vasquez-Yanes 2004; Campo et al. 2012). The addition of both nitrogen (Yap and Moura-Costa 1996, Bungard et al. 2000) and potassium (Gunatilleke et al. 1997) are able to enhance the growth of potted dipterocarps seedlings. Whilst Vincent and Davies (2003) show that below-ground resource availability has a significant effect on the growth performance of dipterocarps seedlings.

In restoration studies, nutrient applications to transplanted seedlings can be used to overcome limited nutrient availability in degraded areas (Holl et al. 2000 2012). However, appropriate nutrient selection and application for particular species is little studied for TPSF. This study explored the response of ten TPSF plant species to the application of macro-nutrients: Nitrogen (N), Phosphorus (P), Potassium (K) and CaMg (Dolomit).


The study was carried out in the greenhouses and nurseries of Banjarbaru Forestry Research Unit of South Kalimantan. At the start of the experiment (June 2011), 22 peat swamp plant species were selected (Table 4.1). The seed materials were seeds and wildlings collected from the TPSF of Central Kalimantan. Each seedling was planted in a separate plastic container (base diameter 10 cm; height 15 cm), and filled with 1:1 (v/v) mixture of soil and rice husk. Seeds were germinated and maintained in the greenhouse before the experiment began. Wildlings were planted into potted compartments and were placed in an area 1 m × 2 m,


covered by plastic films and placed under canopy of tree with a light intensity of 60–80 percent. The wildlings were irrigated 3–4 times/day to maintain humidity under the plastic cover of less than 80 percent and a temperature of between 29°–32° C.

Table 4.1: List of species used in the macro-nutrients application study

No. Species name (Local)

1 Aglaia rubignosa (Kajalaki)

2 Alstonia spatulata (Pulai rawa)

3 Baccaurea bracteata (Jajantik)

4 Calophyllum hosei (Bintangur)


6 Combretocarpus rotundatus (Merapat)

7 Cotylelobium sp. (Resak)

8 Cratoxylum glaucum (Gerunggang)

9 Diospyros bantamensis (Mahirangan)

10 Knema mandarahan (Mandarahan) 11 Licania splendens (Bintan)

12 Lithocarpus sp. (Pampaning)

13 Lophopetalum javanicum (Perupuk)

14 Mangifera sp. (Mangga-mangga)

15 Melaleuca leucadendra (Galam)

16 Palaquium sp. (Nyatoh)

17 Parartocarpus venenosus (Lilin-lilin)

18 Sandoricum beccanarium (Papung)

19 Shorea sp. (Meranti daun lebar)

20 Stemonurus scorpioides (Pasir-pasir)

21 Syzygium sp. 1

22 Syzygium sp.2. (Mahalilis)


Aside from the control, the four treatments included the application of N, P, K, and dolomit. The dosage that was applied were 36.8 mg per polybag, with a twice-weekly application of the nutrients. The experimental design was a Complete Randomized Design, with 10 replications per treatment. The height, diameter and the number of leaves of each seedling was recorded monthly. Before the analysis, the data were checked for normal distribution. If the data were normally distributed, analysis of variance with ortogonal contrast (presented in Table 4.2) was carried out. If the data was not normally distributed, the non-parametric equivilant test was conducted.


Table 4.2: Analysis of variance with ortogonal contrast for plant responses to macronutrients

Treatments Remarks

O (control) Control

O vs N Response to N

(O,N) vs NP Response to P

(O,N,NP) vs NPK Response to K

(O,N,NP) vs NPK (CaMg) Response to CaMg



The height and diameter increments, the change in leaf number and the total dry weights for 22 TPSF tree species seedlings was explored following the additions of N, P, K, Mg and Ca (Table 4.3 and Figure 4.1). Several species showed a positive response to the nutrient additions. Some species showed strong responses, such as Alstonia spatulata (exhibiting greater height, diameter, leaf numbers and dry weight with N, Mg and Ca additions) and Aglaia rubignosa (exhibiting increased leaf numbers and dry weight for all nutrient treatments). Melaleuca leucandendra responded strongly to N additions, achieving increases in height, leaf numbers and dry weight, and to a lesser extent it exhibited positive responses to K and Mg-Ca additions. Similarly, Baccaurea bracteata exhibited greater height, diameter and leaf numbers with Mg-Ca additions. Lophopetalum javanicum also had greater height and leaf numbers after K was added, and saw greater leaf numbers after nitrogen additions. Paratocarpus venenosus showed a strong response to N additions, displaying increased height, diameter and leaf numbers, and to lesser extent, it responded positively to K additions with increased leaf numbers exhibited. After Mg-Ca additions, Lithocarpus sp. responded positively with greater height and diameter increases, and increased stem diameters were exhibited after N additions.

Other species showed a weaker, but also a positive response to the nutrient additions, such as Calophyllum sclerophyllum (greater diameter increases with N and P additions), and Sandoricum beccanarium (increased leaf numbers after K additions). Equally, Knema mandarahan saw increased leaf numbers for P and Mg-Ca additions.

Interestingly, the remaining twelve species showed no significant, and on several occasions, negative responses to the nutrients additions. For example, Palaquium sp. had reduced heights, stem diameters, leaf numbers and total dry weights after all types of nutrients additions, and Cratoxylum glaucum had lower leaf numbers and total dry weights after K and Mg-Ca additions. The lack of responses, and in some cases negative responses, across so many of the species investigated either highlights the degree of adaptation of the tree species to the TPSF ecosystem with its limited nutrient conditions, or questions remain with regards to the volume of nutrients given (could they have reached toxic levels?).

This study therefore shows the important and useful role nutrient additions can play for TPSF reforestation activities, both during cultivation in the nurseries and upon transplanting seedlings into the peatlands. However, this study also highlights that nutrient additions are species-specific, dependent on the nutrient added, and can produce limited results for certain species in the tropical peatland ecosystem; as such, a blanket-approach in nutrient additions is not advised.

KFCP-Sponsored Tropical Peat Swamp Forest Silviculture Research Page 54

Table 4.3: Summarised results of the effect of nitrogen (N), phosphorus (P), potassium (Ka), magnesium (Mg) and calcium (Ca) on the height and diameter (Diam.) increments (inc.), the change in leaf number (LN) and the total dry weight (DW) of 22 TPSF tree species seedlings cultivated under nursery conditions

Species name (Local)

Response to Nitrogen Response to Phosphorus Response to Potassium Response to Mg and Ca Nutrient addition required for optimal growth?

Height inc.

Diam. inc.

Change in LN

Total DW

Height inc.

Diam. inc.

Change in LN

Total DW

Height inc.

Diam. inc.

Change in LN

Total DW

Height inc.

Diam. inc.

Change in LN

Total DW

Aglaia rubignosa (Kajalaki) NS +ve +ve +ve NS NS +ve +ve NS NS +ve +ve NS +ve +ve +ve Yes, all macro nutrients

Alstonia spatulata (Pulai rawa) +ve +ve +ve +ve +ve -ve NS +ve +ve -ve NS NS +ve +ve NS NS N, Mg, Ca

Baccaurea bracteata (Jajantik) NS NS NS NS NS NS NS NS NS NS NS NS +ve +ve +ve NS Mg, Ca

Calophyllum hosei (Bintangur) -ve +ve -ve NS NS NS -ve NS -ve NS -ve NS NS NS -ve NS No

Calophyllum sclerophyllum (Kapur naga) -ve +ve NS -ve NS +ve NS -ve NS NS NS NS NS NS NS NS No

Combretocarpus rotundatus (Merapat) NS NS NS NS NS NS NS -ve NS NS NS -ve No

Cotylelobium sp. (Resak) NS NS -ve NS NS NS -ve NS NS NS -ve NS NS NS -ve NS No

Cratoxylum glaucum (Gerunggang) NS NS NS NS NS NS NS NS NS NS -ve -ve NS NS -ve -ve No

Diospyros bantamensis (Mahirangan) NS NS NS NS NS NS NS NS NS -ve NS NS NS -ve NS No

Knema mandarahan (Mandarahan) NS NS NS NS NS NS +ve NS NS NS NS NS NS -ve +ve NS P

Licania splendens (Bintan) NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS No

Lithocarpus sp. (Pampaning) NS +ve NS NS NS NS -ve NS NS NS NS NS +ve +ve NS NS N, Mg, Ca

Lophopetalum javanicum (Perupuk) NS NS +ve NS NS NS NS NS +ve NS +ve NS NS NS NS NS N, Ka

Mangifera sp. (Mangga-mangga) NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS No

Melaleuca leucadendra (Galam) +ve NS +ve +ve NS NS NS NS NS NS NS +ve NS NS +ve NS N, Ka, Mg, Ca

Palaquium sp. (Nyatoh) NS -ve NS NS NS -ve NS NS -ve -ve -ve -ve -ve -ve -ve -ve No

Parartocarpus venenosus (Lilin-lilin) +ve +ve +ve NS NS NS NS NS NS NS +ve NS NS NS NS NS N, Ka

Sandoricum beccanarium (Papung) NS NS NS NS NS NS NS NS NS NS +ve NS NS NS NS NS Ka

Shorea sp. (Meranti daun lebar) NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS No

Stemonurus scorpioides (Pasir-pasir) NS NS -ve NS NS NS -ve NS NS NS -ve NS NS NS NS NS No

Syzygium sp. 1 NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS NS No

Syzygium sp.2. (Mahalilis) NS -ve NS NS NS NS NS NS NS NS NS NS NS NS NS NS No



Figure 4.1: Changes in height, diameter, number of leaves, and average dry weight of 21 PSF species with macronutrient application Average height increments Average diameter increments Average change in leaf number Average total dry weights

Agla

ia ru

bigi

nosa

(K

ajal

aki)

Alst

onia

spat

ulat

a (P

ulai

raw

a)

cont.

a

a

a a

a

control N NP NPK NPKCaMgTREATMENTS

2,00

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8,00

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ht in

crem

ent (

cm)

]

]]

]

] a

ab

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eter

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emen

t (m

m)

]

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] ]

]

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abab

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ge in

leaf

num

ber

]

]

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]

]

control N NP NPK NPKCaMg

treatm ent

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2,00

tota

l_dr

y w

eigh

t (gr

am)

]

]

]

]]

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a


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ht in

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ent (

cm)

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bc bb

c


treatm ent

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amet

er in

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ent (

mm

)

]

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]

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cbc

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c


treatm ent

0,00

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ge in

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ber

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] ]]

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y w

eigh

t (gr

am)

]

]

]]

]

a

ab

ab

ab

b

b


Average height increments Average diameter increments Average change in leaf number Average total dry weights Ba

ccau

rea

brac

teat

a (J

ajan

tik)

Calo

phyl

lum

hos

ei

(Bin

tang

ur)

Calo

phyl

lum

scle

roph

yllu

m

(Kap

ur n

aga)

a

ab

a

a

b


0,00

10,00

20,00

30,00

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ht in

crem

ent (

cm)

]

]]

]

]

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eter

incr

emen

t (m

m)

]

]

] ]

]

a

ab

ab a

b


TREATMENTS

-5,00

0,00

5,00

10,00

15,00

chan

ge in

leaf

num

ber

]

]

]]

]


treatm ent

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y w

eigh

t (gr

am)

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t (cm

)

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ge in

leaf

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ber ]

]

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] ]


treatm ent

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y w

eigh

t (gr

am)

] ]]

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treatm ent

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ge in

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ber

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y w

eigh

t (gr

am)

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a

a

a

aa

b

b


Average height increments Average diameter increments Average change in leaf number Average total dry weights Co

mbr

etoc

arpu

s rot

unda

tus

(Mer

apat

)

Coty

lelo

bium

sp.

(Res

ak)

Crat

oxyl

um g

lauc

um

(Ger

ungg

ang)

aa

a

a

a


-1,00

0,00

1,00

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ht in

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ent (

cm)

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t (m

m) ]

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ab

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-2,00

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4,00

6,00

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ge in

leaf

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ber

]

]

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am)

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ber

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y w

eigh

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am)

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] ]

a

a

a

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Average height increments Average diameter increments Average change in leaf number Average total dry weights Di

sopy

ros b

anta

men

sis

(Mah

irang

an)

Knem

a m

anda

raha

n (M

anda

raha

n)

Lica

nia

sple

nden

s (B

inta

n)

a aa a

a


treatm ent

0,00

2,50

5,00

7,50

10,00he

ight

inc

rem

ent (

cm)

]

]

]]

]

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a

a

aa


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-0,50

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eter

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t (cm

)

]

]

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]

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a

bab


treatm ent

-4,00

-2,00

0,00

2,00

4,00

chan

ge in

leaf

num

ber

]

]

]

]

]


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0,00

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4,00

6,00

tota

l_dr

y w

eigh

t (gr

am)

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a

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a


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ht i

ncre

men

t (cm

)

]

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] ]

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b


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eter

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)]

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ge in

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ber

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y w

eigh

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am)

] ]

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a


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y w

eigh

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am)

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a

a

aa

a


Average height increments Average diameter increments Average change in leaf number Average total dry weights Li

thoc

arpu

s sp.

(P

ampa

ning

)

Loph

opet

alum

java

nicu

m

(Per

upuk

)

Man

gife

ra sp

. (M

angg

a-m

angg

a)

a

ab

a

ab

b


treatm ent

-2,00

0,00

2,00

4,00

6,00

heig

ht in

crem

ent (

cm)

]

]

]

]

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b

a a

ab


treatm ent

-0,50

0,00

0,50

1,00

diam

eter

incr

emen

t (cm

)

]

]

] ]

]

b

a

aa a


treatm ent

-2,00

0,00

2,00

chan

ge in

leaf

num

ber

]

]

]

]

]


treatm ent

0,00

0,50

1,00

1,50

2,00

tota

l_dr

y w

eigh

t (gr

am)

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] ]

]]

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a

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b

ab


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0,00

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ht in

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ent (

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a

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0,00

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eter

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emen

t (cm

)]

] ]

]

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ab


treatm ent

0,00

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ge in

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ber

]

]

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l_dr

y w

eigh

t (gr

am)

]

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ht in

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ent (

cm)

]

]] ]

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a

a

a

a


0,00

1,00

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eter

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emen

t (m

m)

]

]

]

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]

a

a

a

a

a


-4,00

0,00

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8,00

chan

ge in

leaf

num

ber

]

]

]

]

]


treatm ent

0,00

2,00

4,00

6,00

8,00

tota

l_dr

y w

eigh

t (gr

am)

]

]

]

]

]

aa

a

aa


Average height increments Average diameter increments Average change in leaf number Average total dry weights M

elal

euca

leuc

aden

dra

(Gal

am)

Pala

quiu

m sp

. (N

yato

h)

Para

rtoc

arpu

s ven

enos

us

(Lili

n-lil

in)

a

b

b

b b


treatm ent

-10,00

0,00

10,00

20,00

30,00

heig

ht in

crem

ent (

cm)

]

]] ]

]

a

aa

a

a


treatm ent

-0,50

0,00

0,50

1,00

diam

eter

incr

emen

t (cm

)

]

]

]

]

]

a

bcabc

ab

c


treatm ent

0,00

20,00

40,00

60,00

chan

ge in

leaf

num

ber

]

]

]

]

]


treatm ent

0,00

2,00

4,00

6,00

8,00

tota

l_dr

y w

eigh

t (gr

am)

]

]

]

]

]

a

abab

b

ab

a

aa

b b


-20,00

-10,00

0,00

10,00

heig

ht in

crem

ent (

cm) ]

]]

]

]

a

ab

bc c


-3,00

-2,00

-1,00

0,00

1,00

2,00

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eter

incr

emen

t (m

m) ]

]

]

]

]

aa a

bb


-4,00

-2,00

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chan

ge in

leaf

num

ber

]

]

]

]

]


treatm ent

-2,00

0,00

2,00

4,00

tota

l_dr

y w

eigh

t (gr

am)

]

]]

]

]

a

bb

b

b


0,00

10,00

20,00

heig

ht in

crem

ent (

cm)

]

]]

]

]

a

b

b b b


0,00

2,00

4,00

6,00

diam

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m)

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am)

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Average height increments Average diameter increments Average change in leaf number Average total dry weights Sa

ndor

icum

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cana

rium

(P

apun

g)

Shor

ea sp

. (M

eran

ti da

un le

bar)

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onur

us sc

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ides

(M

edan

g te

lur,

Pasi

r-pa

sir)

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a

a

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ight

incr

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)

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leaf

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15,00

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5,00

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crem

ent (

cm)

]

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1,00

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Average height increments Average diameter increments Average change in leaf number Average total dry weights Sy

zygi

um sp

.1

Syzy

gium

sp.2

(M

ahal

ilis)


4.4 Conclusion

Of the 22 TPSF species studied, seven species showed a strong positive response to certain nutrient additions, such as Paratocarpus venenosus’ response to nitrogen additions and Lithocarpus sp.’s response to Magnesium-calcium additions. A further three species showed a positive, but less strong response. Only two species, Alstonia spatulata and Aglaia rubignosa showed uniformly positive responses to all nutrient additions. Surprisingly, 12 of the 22 species did not show a response, or responded negatively to the nutrient additions, indicating a high adaptation to low nutrient environments. This study therefore highlights the important role of nutrient additions in TPSF reforestation activity, but that a cautious species- and nutrient-specific approach is required.

a

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cm)

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4.5 References Arnon, D. I. 1950. Criteria of essentiality of inorganic nutrients for plants with special reference to molybdenium. Lotsya

3: 31–38.

Bungard, R. A., Press, M. C., and J. D. Scholes. 2000. The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance. Plant Cell Environment 23: 1183-1194

Campo, J. and C. Vasquez-Yanes. 2004. Effects of nutrient limitation on above ground carbon dynamics during tropical dry forest regeneration in Yucatan, Mexico. Ecosystem 7: 311-319.

Campo, J., Solis, E. and J. F. Gallardo. 2012. Effects of fertilization on soil nutrient characteristics and the growth of tree stand in secondary seasonally dry tropical forests in Mexico. Journal of Tropical Forest Science 24: 408-415

Foth, H. D. 1998. Dasar-dasar Ilmu Tanah. Gadjah Mada University Press. Yogyakarta

Gunatilleke, C. V. S., Gunatilleke, I. A. U. N., Gad, P., Burslem, D. F. R. P., Ashton, P. M. S., and P. S. Ashton. 1997. Responses to nutrient addition among seedlings of eight closely related species of Shorea in Sri Lanka. Journal Ecology 85: 301-311

Holl, K. D., Loik, M. E., Lin, E. H. V., and I. A. Samuels. 2000. Tropical montane forest restoration in Costa Rica: Overcoming barriers to dispersal and establishment. Restoration Ecology 8: 339-349

Holl, K. D. 2012. Chapter 9: Restoration of tropical forests. In: van Andel, J. and Aronson, J. Restoration Ecology: The New Frontier. Second edition. Wiley-Blackwell Press, Oxford, UK.

Kramer, P.J. and T.T. Kozlowski.1979. Physiology of Woody Plants. Academic Press Inc. London

Mengel, K and E.A. Kirkby.1978. Principles of Plant Nutrition. International Potash Institute. Switzerland.

Newberry, D. M., Chuyong, G. B., Green, J. J., Songwe, N. C., Tchuenteu, F. and L. Zimmermann. 2002. Does low phosphorus supply limit seedling establishment and tree growth in groves of ectomycorrhizal trees in a central African rainforest? New Phytologist 156: 297-311

Page, S. E and J. O. Reiley. 2005. Wise-use of tropical peatlands. Alterra Publishing, Wageningen, The Netherlands.

Santiago, L. S., Wright, S. J., Harms, K. E., Yavitt, J. B., Korine, C., Garcia, M. N. and B. L. Turner. 2012. Tropical tree seedling growth responses to nitrogen, phosphorus and potassium addition. Journal of Ecology 100: 309-316

Sulistiyanto, Y. 2004. Nutrient dynamics in different sub-types of peat swamp forest in Central Kalimantan, Indonesia. PhD thesis, School of Geography, University of Nottingham, Nottingham.

Tisdale, S.L. and W.L. Nelson. 1956. Soil fertility and Fertilizers. The MacMillan Company. Canada.

Vincent, A. and S. J. Davies. 2003. Effects of nutrient addition, mulching and planting-hole size on early performance of Dryobalanops aromatica and Shorea parvifolia in secondary forest in Sarawak, Malaysia. Forest Ecology and Management 180: 261-271

Yap, S. W., and P. Moura-Costa. 1996. Effects of nitrogen fertilization and soil texture on growth and morphology of Dryobalanops lanceolata seedlings pp.189-196 in Appanah, S., and K.C. Khoo (Eds.) Proceedings of the Fifth Round-Table Conference on Dipterocarps, Dorest Research Institute of Malaysia, November 7-10, Chiang Mai, Thailand

Yuwati, T.W., Susanti, P.D., and B. Hermawan. 2010. Studi Nutrisi Tanaman Meranti Rawa dan Jelutung Rawa. Hasil Penelitian. Balai Penelitian Kehutanan Banjarbaru. Banjarbaru (unpublished)


PAPER 5: RESPONSE OF PEAT SWAMP FOREST TREE SPECIES SEEDLINGS TO MYCORRHIZAL INOCULATIONS

Tri Wira Yuwati1, Laura L.B. Graham2, Dony Rachmanadi1, Purwanto Budi Santosa1 and Rusmana1 1 Banjarbaru Forestry Research Unit, South Kalimantan (FORDA, Indonesian Ministry of Forestry) 2 Kalimantan Forest Climate Partnership

5.1 Introduction

Efforts to restore the degraded tropical peat swamp forest (TPSF) are facing constraints due to, amongst other issues, limited planting techniques (Mulyanto, 2000; Page et al. 2009). Lazuardi and Supriadi (1997) show that the survival rate of TPSF tree species transplanted in Tumbang Nusa, Central Kalimantan was very low (less than 40 percent). Acidic soil is one constraint which causes the slow growth of plants. Environmental manipulations can facilitate seedlings to be transplanted in degraded forest areas (Holl, 2012); one potential for this is the utilisation of soil microbes such as mycorrhiza.

Mycorrhiza are a fungus that form a mutualistic association with plant roots (Harley and Smith 1983). The term symbiosis is used to describe the dependent mutualistic relationship between the fungi that receives the photosynthetic carbon from the host plant, and the host plant that receives soil nutrients, particularly phosphorus, water and protection from root pathogens and toxic elements (Brundrett et al. 1996; Harley and Smith 1983; Dell 2002; Rillig 2004). The two most common forms of mycorrhizal fungi are vesicular arbuscular mycorrhiza (VAM) and ectomycorrhiza (ECM). VAM are part of Zygomycota family which develop arbuscule structures, hyphae and vesicle inside the plant root cortex cells, and ectomycorrhiza (in the Basidiomycetes family) develop a sheath outside the root surface and Hartig net between root cells (Brundrett et al. 1996).

Several studies have shown the importance mycorrhizae in TPSF ecosystems. A preliminary study was conducted to determine the presence of indigenous mycorrhizae in burnt degraded peat swamp forest (Yuwati 2003). The results showed that the spores of Glomus sp. and Gigaspora sp. were present in burnt degraded peat swamp forest of Tumbang Nusa Central Kalimantan Province. Tawaraya et al. (2003) found the presence of mycorrhizal associations on 17 TPSF TREE species, including: Shorea balangeran (Belangeran), Gonystylus bancanus (Ramin), Cratoxylum arborescens (Gerunggang), Calophyllum soulattri (Kapur Naga), Turjaman et al. (2006) show that there was increased growth for Dyera polyphylla and Aquilaria fillaria seedlings under nursery conditions after being inoculated with Glomus clarum and Gigaspora decipiens. Turjaman et al. (2007) also report the positive effect of two arbuscular mycorrhiza species (G. clarum and G. aggregatum) on Ploiarium alternifolium and Calophyllum hosei seedlings under nursery conditions. Alstonia pnematophora and Gonystylus bancanus seedlings increased in height and diameter during 24 weeks of growth in a nursery after being inoculated with VAM (Yuwati 2008; Yuwati et al. 2007).

Despite the potential importance of mycorrhizal fungi in enhancing seedling growth and survival rates (already shown for several TPSF species) and its potential application as a restoration tool in seedling transplants (Turjaman 2007, 2011), there still remains numerous TPSF species for which its relationship is yet to be established. Furthermore, the effect of mycorrhiza in enhancing overall seedling growth is still poorly understood, and as yet, only some have been tested for TPSF tree species such as Dyera polyphylla with Glomus clarum and Gigaspora decipiens and Shorea balangeran with Scleroderma columnare (Graham et al. 2013). Graham et al. (2013) show that mycorrhizal inoculation on S. balangeran and D.polyphylla is recommended due to the higher mycorrhizal colonization level on inoculated seedlings transplanted in the degraded area and the increased nutrient uptake in the transplanted seedlings following mycorrhiza


inoculation. Given the large areas of TPSF which are now degraded and that restoration activities are currently being implemented or continue to be planned, these research gaps need to be addressed.

This report explored the response of single inoculation of several arbuscular mycorrhiza fungi species on 15 peat swamp forest species seedlings under nursery conditions. The hypothesis formulated in this research was that a single inoculation of indigenous arbuscular mycorrhizae fungi (AMF) species would increase the growth of the fifteen TPSF plant species.


The experiment was conducted in the greenhouses of Banjarbaru Forestry Research Unit in South Kalimantan. At the start of the experiment (June 2011), 16 peat swamp plant species were selected (Table 5.1). The seed materials were seeds and wildlings, collected from the TPSF of Central Kalimantan. Seeds of peat swamp forest species were surface-sterilised with H2O2 5 percent for 5 minutes. The seeds were then sown in germination containers. Peat was used as the growth medium for the seedlings. The peat medium was sterilised with autoclave 121o C for 15 minutes. The peat medium was divided into two purposes: a medium for seed germination and a medium for transplanting. After sterilisation, the media were applied in the polybags and the germination containers in the greenhouse.

After 30 days, the germinated seeds were transplanted into polybags. Before sowing the germinated seeds, 10 g of AMF inoculums and 5 alginate beads were applied. The AMF isolates were developed by the Forest Microbiology Lab. The arbuscular mycorrhizal spores that were used were: Glomus clarum, Gigaspora decipiens and Enthrophospora sp. The fungal spores were then mass-produced in zeolite medium (Turjaman et al. 2006). The seedlings were watered daily using tap water. The treatments consisted of a control, and seedlings inoculated with Glomus clarum, seedlings inoculated with Gigaspora decipens and seedlings inoculated with Enthrophospora sp. In total there were four treatments, with 10 replications for each treatment. The height, diameter and number of leaves of the seedlings were measured monthly.

All data obtained (height, diameter and number of leaves) were checked for normality before performing group differences analysis (ANOVA) using SPSS.


Table 5.1: List of species used in the mycorrhizal inoculation study


1 Baccaurea bracteata (Jajantik)

2 Calophyllum hosei (Bintangur)


4 Combretocarpus rotundatus (Merapat)

5 Cotylelobium sp. (Rassak)

6 Cratoxylum glaucum (Geronggang)

7 Diospyros bantamensis (Mahirangan)

8 Jambu burung kecil (Syzygium sp.)

9 Licania splendens (Bintan)

10 Lopopethalum javanicum (Perupuk)

11 Palaquium sp. (Nyatoh)

12 Parartocarpous venenosus (Lilin-lilin)

13 Stemonurus scorpoides (Medang telur)

14 Syzygium pakan (Pakan)

15 Syzygium sp. (Jambu burung kecil)

16 Syzygium sp. (Mahalilis)



Of the fifteen TPSF tree species seedlings studied, only five showed a positive reponse to the mycorrhizal inoculations (Table 5.2, Figure 5.1). Calophyllum sclerophyllum responded strongly to the Enthrophospora sp. inoculum, with significant increases in height and basal diameter observed. Lopopethalum javanicum also showed a strong response, with increased height and leaf numbers exhibited after the application of the Glomus clarum inoculum. Syzygium sp. (Pakan) also responded favourably to both the Enthrophospora sp. and Glomus clarum inoculums, with increased basal diameters displayed for both treatments. Parartocarpus venenosus and Syzygium sp. (Mahalilis) showed less strong responses (increased basal diameter for the Enthrophospora sp. inoculum and increased leaf numbers for the Glomus clarum inoculums respectively). The results also indicated that Gigaspora decipens is not a generalist or a suitable mycorrhizal inoculum, as none of the 15 species responded strongly to its inoculation. These findings can be applied to approaches to reforestation to increase the growth rates of seedlings both during nursery cultivation, and, potentially, after transplantation to degraded areas.

It is surprising, however, that ten of the species showed no positive reaction to the three different mycorrhizae, given that most other studies (for example, Turjaman et al. 2006, 2007; Graham et al. 2013; and Yuwati et al. 2007) showed the strong positive reactions of inoculated seedlings. This might be explained through collecting and analysing data on more variables, such as the dry weights of the shoots and roots, the nitrogen and phosphorus content of the leaves, and the percentage of the inoculum taken up by the roots;


the mycorrhizae may have colonised the roots and served other purposes such protection against drought or disease, as discussed by Graham et al. (2013). A longer period of study, monitoring the seedlings beyond the point of transplantation into degraded TPSF might reveal other, longer-term benefits.


Table 5.2: Summarised results of the effect of three mycorrhizal treatments on the height and diameter (Diam.) increments (inc.), the changes in leaf numbers (LN) and the total dry weight (DW) of 15 TPSF tree species seedlings cultivated under nursery conditions


Response to Glomus clarum

Response to Gigaspora decipens

Response to Enthrophospora sp. Mycorrhizal

inoculation required for optimal growth? Height

inc. Diam.

Inc. Change

in LN Height

inc. Diam.

Inc. Change

in LN Height

inc. Diam.

Inc. Change

in LN 1 Baccaurea bracteata (Jajantik) NS NS NS NS NS NS NS NS NS None found 2 Calophyllum hosei (Bintangur) NS NS NS NS NS NS NS NS NS None found 3 Calophyllum sclerophyllum (Kapur naga) NS NS NS NS NS NS +ve +ve NS Enthrophospora sp. 4 Combretocarpus rotundatus (Merapat) NS -ve NS NS -ve NS NS -ve NS None found 5 Cotylelobium sp. (Rassak) NS NS NS NS NS NS NS NS NS None found 6 Cratoxylum glaucum (Geronggang) NS NS NS NS NS NS NS NS NS None found 7 Diospyros bantamensis (Mahirangan) NS NS NS NS NS NS NS NS NS None found 8 Licania splendens (Bintan) NS NS NS NS NS NS NS NS NS None found

9 Lopopethalum javanicum (Perupuk) +ve NS +ve NS NS NS NS NS +ve Glomus clarum (and

Enthrophospora sp.) 10 Palaquium sp. (Nyatoh) NS NS NS NS NS NS NS NS NS None found 11 Parartocarpous venenosus (Lilin-lilin) NS NS NS NS NS NS NS +ve NS Enthrophospora sp. 12 Stemonurus scorpoides (Medang telur) NS NS NS NS NS NS NS NS NS None found

13 Syzygium pakan (Pakan) NS +ve NS NS NS NS NS +ve NS Glomus clarum and

Enthrophospora sp. 14 Syzygium sp. (Jambu burung kecil) NS NS NS NS NS NS NS NS NS None found 15 Syzygium sp. (Mahalilis) NS NS +ve NS NS NS NS NS NS Glomus clarum



Figure 5.1: Changes in height, diameter growth and number of leaves of 15 PSF species, 7 months after three mycorrhizae inoculation treatments Average height increments Average diameter increments Average change in leaf number

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5.4 Conclusion

Of the fifteen TPSF tree species seedlings studied, only five showed a positive reponse to the mycorrhizal inoculations; Calophyllum sclerophyllum responded positively to the Enthrophospora sp. inoculum, Lopopethalum javanicum to the Glomus clarum inoculum, Syzygium sp. (Pakan) responded favourably to both the Enthrophospora sp. and Glomus clarum inoculums, and Parartocarpus venenosus responded well to the Enthrophospora sp. inoculum, and Syzygium sp. (Mahalilis) to Glomus clarum. The results of this study also indicate that Gigaspora decipens is not a suitable mycorrhizal inoculum, as none of the 15 species responded strongly to its inoculation. These findings can be applied to reforestation activities to increase growth rates of the seedlings both during nursery cultivation, and, potentially, after transplantation in degraded TPSF.

It was surprising, however, that ten of the species showed no positive reaction to the three different mycorrhizae. This might have been explained if a longer period of study had been undertaken; tracking monitoring the seedlings beyond the point of transplantation into the TPSF, and investigating more variables such as the dry weights of the shoots and roots, and the nitrogen and phosphorus content of the leaves. This might reveal other, longer-term benefits.

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5.5 References Brundrett, M., Bougher, N., Dell, B., Grove, T. and N. Malajczuk. 1996. Working with mycorrhiza in forestry and

agriculture, ACIAR, China.

Dell B. 2002. Role of mycorrhizal fungi in ecosystems. CMVJ 1:47–60

Giovannetti M., and B. Mosse. 1980. An evaluation of techniques for measuring vesicular arbuscular mycorrhizal infection in roots. New Phytologist 84, 489–500

Graham L.L.B, Turjaman M, and S.E. Page. 2013. Shorea balangeran and Dyera polyphylla (syn. Dyera lowii) as tropical peat swamp forest restoration transplant species: effects of mycorrhizae and level of disturbance. Wetland Ecology and Management 21 DOI 10.1007/s11273-013-9302-x

Harley, J.L. and S.E. Smith. 1983. Mycorrhizal symbiosis. Academic Press, London.

Holl, K. D. 2012. Restoration of tropical forests. In: van Andel J, Aronson (eds) Restoration ecology: the new frontier, 2nd edn. Wiley-Blackwell, New Jersey.

Lazuardi, D. and R. Supriardi. 1997. Hubungan antara ketergenangan air permukaan dengan daya hidup tanaman ramin pada belukar galam di lahan rawa gambut bekas terbakar. Prosiding Ekspose Hasil Penelitian dan Uji Coba BTR Banjarbaru. Banjarbaru 1997.

Mulyanto, B. 2000. Pendekatan dan Strategi Pemanfaatan Hutan Rawa Gambut Ex-PLG Sejuta Hektar. In: Daryono, H., Jafarsidik, J., Mile, M.Y., Subagyo, E., Hadi, T.S., Akbar, A., Budiningsih, K. (eds.). Prosiding Seminar Pengelolaan Hutan Rawa Gambut dan Ekspose Hasil Penelitian di Hutan Lahan Basah, 1–8, Balai Teknologi Reboisasi Banjarbaru, Kalimantan Selatan.

Page, S. E., Hoscilo, A., Wosten, H., Jauhiainen, J., Ritzema, H., Tansey, K., Silvius, M., Graham, L., Vasander, H., Rieley, J., and S. Limin. 2009. Ecological restoration of lowland tropical peatlands in Southeast Asia- current knowledge and future research directions. Ecosystems 12:288–905

Rillig M. C. 2004. Arbuscular mycorrhizae and terrestrial system processes. Ecology Letters 7:740–754

Tawaraya, K., Takaya, Y., Turjaman, M., Tuah, S.J., Limin, S.H., Tamai, Y., Cha, J.Y., Wagatsuma, T., and M. Osaki. 2003. Arbuscular mycorrhizal colonization of tree species grown in peat swamp forests of Central Kalimantan, Indonesia. Forest Ecology and Management 6258: 1–6.

Tawaraya, K., Takaya, Y., Turjaman, M., Tuah, S.J., Limin, S.H., Tamai, Y., Cha, J.Y., Wagatsuma, T. and M. Osaki. 2003. Arbuscular mycorrhizal colonization of tree species grown in peat swamp forests of Central Kalimantan, Indonesia. Forest Ecology and Management 182: 381–386.

Tawaraya, K., Turjaman, M. and H.A. Ekamawanti. 2007. Effect of Arbuscular Mycorrhiza Colonization on Nitrogen and Phosphorus Uptake and Growth of Aloe vera L. HortScience 42(7): 1737–1739.

Turjaman, M., Tamai, Y., Sagah, H., Santoso, E., OsakI, M. and K. Tawaraya. (2006) Arbuscular mycorrhiza fungi increased early growth of two non timber forest product species Dyera polyphylla and Aquilaria filarial under greenhouse conditions. Mycorrhiza 16:459–464.

Turjaman, M., Tamai, Y., Sitepu, I. R., Santoso, E., Osaki, M. and K. Tawaraya. 2008. Improvement of early growth of tropical peat swamp forest species Ploiarium alternifolium and Calophyllum hosei by two arbuscular mycorrhizal fungi under greenhouse conditions. New Forests 36:1–12.

Turjaman, M., Santoso E., Susanto A., Gaman S., Limin, S.H., Tamai, Y., Osaki, M. and K. Tawaraya. 2011. Ectomycorrhizal fungi promote growth of Shorea balangeran in degraded peat swamp forest. Wetland Ecology and Management 19: 331–339.

Turjaman, M., Y. Tamai, E. Santoso, M. Osaki and K. Tawaraya. 2006. Arbuscular mycorrhizal fungi increased early growth of two nontimber forest product species Dyera polyphylla and Aquilaria filaria under greenhouse conditions. Mycorrhiza 16: 459–464.


Turjaman, M., Santoso, E., Susanto, A., Gaman, S., Limin, S.H., Tamai, Y., Osaki, M. and K. Tawaraya. 2011. Ectomycorrhizal fungi promote growth of Shorea balangeran in degraded peat swamp forest. Wetlands Ecology and Management 19, 331-339.

Vierheilig H., Coughlan A.P., Wyss, U.R.S, and Y. Piche. 1998. Ink and Vinegar: a simple staining technique for arbuscular mycorrhizal fungi. Applied and Environmental Microbiology 64: 5004–5007.

Yuwati, T.W. 2003. Keberadaan mikoriza asli setempat pada hutan rawa gambut pasca kebakaran, Tumbang Nusa, Kalimantan Tengah. Buletin Tekno Hutan Tanaman. Balai Penelitian dan Pengembangan Hutan Tanaman Indonesia Bagian Timur. Banjarbaru. Vol 1 No.1 Oktober 2003.

Yuwati, T.W. 2008. Peningkatan pertumbuhan Pulai Rawa (Alstonia pneumatophora) dengan inokulasi mikoriza dan sterilisasi media. Widya Riset Bulletin Vol. 9 No. 3. LIPI. Cibinong.

Yuwati, T.W., Santosa, P.B., and B. Hermawan. 2007. Arbuscular mycorrhiza fungi application for rehabilitation of degraded peat swamp forest in Central Kalimantan in: Rieley, J.O., Banks,C.J. and B. Radjagukguk. (Eds). 2007. Carbon-climate-human interaction on tropical peat land. Proceedings of The International Symposium and Workshop on Tropical Peatland, Yogyakarta, 27–29 August 2007, EU CARBOPEAT and RESTORPEAT Partnership, Gadjah Mada University, Indonesia and University of Leicester, United Kingdom.

Tawaraya, K., Takaya, Y., Turjaman, M., Tuah, S.J., Limin, S.H., Tamai, Y., Cha, J.Y., Wagatsuma, T. and M. Osaki. 2003. Arbuscular mycorrhizal colonization of tree species grown in peat swamp forests of Central Kalimantan, Indonesia. Forest Ecology and Management 182: 381–386.

Tawaraya, K., Turjaman, M., and H.A. Ekamawanti. 2007. Effect of Arbuscular Mycorrhiza Colonization on Nitrogen and Phosphorus Uptake and Growth of Aloe vera L. HortScience 42(7): 1737–1739.

Turjaman, M. 2007. Improvement of early growth of two tropical peat-swamp forest tree species Ploiarium alternifolium and Calophyllum hosei by two arbuscular mycorrhizal fungi under greenhouse conditions. PhD Thesis Yamagata University. Japan (unpublished).

Turjaman, M., Tamai, Y., Santoso, E., Osaki, M. and K. Tawaraya. 2006. Arbuscular mycorrhizal fungi increased early growth of two nontimber forest product species Dyera polyphylla and Aquilaria filaria under greenhouse conditions. Mycorrhiza 16: 459–464.

Yuwati, T.W. 2003. Keberadaan mikoriza asli setempat pada hutan rawa gambut pasca kebakaran, Tumbang Nusa, Kalimantan Tengah. Buletin Tekno Hutan Tanaman. Balai Penelitian dan Pengembangan Hutan Tanaman Indonesia Bagian Timur. Banjarbaru. Vol 1 No.1 Oktober 2003.

Yuwati, T.W. 2008. Peningkatan pertumbuhan Pulai Rawa (Alstonia pneumatophora) dengan inokulasi mikoriza dan sterilisasi media. Widya Riset Bulletin Vol. 9 No. 3. LIPI. Cibinong.

Yuwati, T.W., Santosa, P.B., and B. Hermawan. 2007. Arbuscular mycorrhiza fungi application for rehabilitation of degraded peat swamp forest in Central Kalimantan in: Rieley, J.O., Banks,C.J. and B. Radjagukguk. (Eds). 2007. Carbon-climate-human interaction on tropical peat land. Proceedings of The International Symposium and Workshop on Tropical Peatland, Yogyakarta, 27-29 August 2007, EU CARBOPEAT and RESTORPEAT Partnership, Gadjah Mada University, Indonesia and University of Leicester, United Kingdom.


CLOSING REMARKS The purpose of this KFCP-funded silviculture study was to explore the ecological tolerances and optimal growth conditions of a range of selected TPSF tree species in order to support both the KFCP trial reforestation rehabilitation activities and wider tropical peatland rehabilitation and reforestation activities. There is little available species-specific, published data on the above topics, and this study has gone far to increase knowledge and provide immediately applicable findings for those working in the field. One of the key messages throughout this report is the species-specific nature of tolerances to extreme environmental conditions and the requirements for maximum growth and survival. KFCP and the Banjarbaru Forestry Research Unit invested time and resources in this study to provide much needed, detailed answers to some of the key silvicultural challenges facing tropical peatlands. It is hoped that other rehabilitation projects working in this type of ecosystem apply this species-specific knowledge gained by KFCP to reforestation and rehabilitation programs, and that projects working in other ecosystems adopt a similar investigatory approach to contribute to improved and accurate rehabilitation and reforestation plans.


RELEVANT LITERATURE Graham, L. L. B. 2009. A literature review of the ecology and silviculture of tropical peat swamp forest tree species found

naturally occurring in Central Kalimantan. IAFCP: Jakarta

Harrison, M. E., Husson, S. J., Zweifel, N., D’Arcy, L. J., Morrogh-Bernard, H. C., Cheyne, S. M., van Noordwijk, M. A. and C. P. van Schaik. 2010. The Fruiting Phenology of Peat-Swamp Forest Tree Species at Sabangau and Tuanan, Central Kalimantan, Indonesia. IAFCP: Jakarta


ANNEX 1

Photographs of activities

Seed material exploration at Mantangai (Block A and E) Seed material exploration at Tumbang Nusa

Seed material exploration at Petak Bahandang Seed material exploration at Hampangen

Nursery activities: seed preparation Nursery activities: wildling preparation


Calophyllum sclerophyllum fruits Knema mandarahan seeds

Lophopethalum javanicum seeds Diospyros bantamensis fruits

Tetramerista glabra fruits Paratocarpus venenosus seeds


Combretocarpus rotundatus seeds Cratoxylon glaucum seeds

Agathis borneensis fruits Sandoricum beccaranium

Syzygium sp.


Research activities: inundation Research activities: nutrient

Research activities: mycorrhiza


ANNEX 2

List species found on exploration activities

No Species/Local name Plant material Collection origin and depth of peat at origin

1 Calophyllum hosei/ bintangur seedling Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

2 Stemonurus scorpioides/ medang telur, pasir-pasir, tagula, tabaras

wildling Mentangai, Central Kalimantan; 1-2 m

3 Calophyllum sclerophyllum / kapur naga seedling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

4 Shorea sp./ meranti daun kecil, meranti bitik

wildling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

5 Shorea sp./ meranti daun lebar, meranti tembaga


6 Alstonia spatulata/ pulai rawa


7 Lopophetalum javanicum/perupuk wildling Mentangai, Central Kalimantan, 1-2 m

8 Disopyros bantamensis / mahirangan wildling Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

9 Knema mandarahan/ mandarahan seedling Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

10 Palaquium sp. / nyatoh wildling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

11 Melaleuca sp. Cf leucadendron / galam wildling Landasan Ulin, South Kalimantan, 1-2 m

12 Cotylelobium sp./ resak wildling Mentangai, Central Kalimantan, 1-2 m

13 Licania splendens/bintan seedling Petak Bahandang, Central Kalimantan, 2-3 m

14 Syzygium sp./ jambu burung wilding Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

15 Lithocarpus sp / pampaning wilding Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

16 Parartocarpus venenosus/ Lilin-lilin

seedling Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

17 Mangifera sp./mangga-mangga wilding Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

18 Sandoricum beccanarium / Papung

wilding Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

19 Cratoxylon glaucum/gerunggang seedling Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m

20 Combretocarpus rotundatus/Perepat wildling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

21 Baccaurea bracteata/Jajantik, rambai hutan

seedling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

22 Kajalaki seedling Mentangai, Central Kalimantan, 1-2 m

23 Nephelium sp. / rambutan hutan seedling Petak Bahandang, Central Kalimantan, 2-3 m

24 Noethopoebe sp. / gemor cutting Tumbang Nusa, Central Kalimantan, Lat.3027’ -30 59 S,1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 4-8 m


25 Syzygium sp. / pakan seedling Mentangai, Central Kalimantan, 1-2 m

26 Dacrydium pectinateum/alau wildling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

27 Syzygium sp/ Syzygium cf, jambu burung daun kecil

wilding Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

28 Agathis borneensis / agathis seedling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

29 Tristaniopsis sp. / pelawan wildling Hampangen, Central Kalimantan, Lat. 0027’ s/d 30 59 S, Long. 1130 2’ 36 ‘- 1140 44’ 00 ‘ E; 1-2 m

30 Leguminosae cf. seedling Mentangai, Central Kalimantan, 1-2 m

31 Koompassia malacensis / kempas seedling Teluk umpan, Central Kalimantan, 3–4 m


ANNEX 3

Soil microbes exploration

Ectomycorrhizae

Mycorrhiza 1. Lactarius sp. Mycorrhiza 2. Calvatia sp.

Mycorrhiza 4. Pholiota sp. Mycorrhiza 5. Suillus sp.


Mycorrhiza 7. Tricholoma sp. Mycorrhiza 7. Tricholoma sp.

Mycorrhiza 3. Calvatia sp. Mycorrhiza 6. Tylopilus sp.


Endomycorrhizae

Association with legume species Association with Syzigium pakan

Association with prupuk Association with tagula/pasir-pasir

88

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Tropical Peat Swamp Forest Silviculture in Central Kalimantan

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