Dipterocarpaceae: Forest Fires and Forest Recovery
Aldrianto Priadjati Aldrianto Priadjati Dipterocarpaceae: Forest Fires and Forest Recovery Thesis Wageningen University, the Netherlands – With ref. – With summaries in English, Dutch and Bahasa Indonesia
Keywords: forest fires, forest recovery, Dipterocarpaceae, Shorea leprosula, population phenotype, soil, mycorrhizae, ecophysiology, light intensity, Kalimantan, Indonesia.
ISSN 1566-6522 ISBN 90-5808-753-0 (thesis version) 90-5113-062-7 (Tropenbos version)
© 2002 MoF – Tropenbos Kalimantan Programme
The opinions expressed in this publication are those of the author(s) and do not necessarily reflect the views of Tropenbos International
No part of this publication, apart from bibliographic data and brief quotations in critical reviews, may be reproduced, re-recorded or published in any form including print photocopy, microfilm, electromagnetic record without the prior written permission.
Cover photo: Forest Fires 1997-98 in the Wanariset Samboja, Kalimantan, Indonesia (W.T.M Smits) Printed by: Ponsen & Looijen, Wageningen, the Netherlands. DIPTEROCARPACEAE: FOREST FIRES AND FOREST RECOVERY
Aldrianto Priadjati
PROEFSCHRIFT ter verkrijging van de graad van doctor op gezag van de Rector Magnificus van Wageningen Universiteit, Prof.dr.ir. L. Speelman, in het openbaar te verdedigen op dinsdag 26 november 2002 des namiddags te 16.00 uur in de Aula, Wageningen Promotoren: Prof.dr.ir. R.A.A. Oldeman, Hoogleraar in de bosteelt en bosoecologie, Wageningen Universiteit, Nederland
Prof.dr.ir. Joedoro Soedarsono, Hoogleraar in de microbiologie, Gadjah Mada University, Yogyakarta, Indonesia
Prof.dr. Steph B.J. Menken Hoogleraar in de evolutionaire biologie Universiteit van Amsterdam, Nederland
Promotiecommissie: Prof.dr. A.M. Cleef, Wageningen Universiteit, Nederland Prof.dr.ir. A.F. van der Wal, Wageningen Universiteit, Nederland Prof.dr. R.F. Hoekstra, Wageningen Universiteit, Nederland Dr.ir. Irsyal Yasman, PT Inhutani I, Jakarta, Indonesia Dipterocarpaceae: Forest Fires and Forest Recovery
Aldrianto Priadjati
The Tropenbos Foundation Wageningen, the Netherlands 2002 Tropenbos-Kalimantan Series The Tropenbos-Kalimantan Series presents the results of studies and research activities related to sustainable use and conservation of forest resources in Indonesia. The multi-disciplinary MoF-Tropenbos Kalimantan programme operates within the framework of the international programme of the Tropenbos Foundation. Executing Indonesian agency is the Forest Research Institute, Samarinda, governed by the Forestry Research and Development Agency (FORDA). Other Indonesian executing agencies include the Association of Indonesian Concession Holders (APHI) and PT INHUTANI I.
Wageningen University, the Ministry of Forestry, Indonesia Netherlands !#
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General Introduction 1.3.2 The taxonomical characteristics of Shorea leprosula Miq. Seedling (Fig. 1.5A) A S. leprosula seedling has an epigeal germination, cotyledons equal and spreading vertically with first leaves opposite (Foxworthy, 1932). The seedlings are very easy to recognize because they have domatia at the back of the leaf along the main nerve but these will become less abundant when the seedling reaches pole size. Furthermore, Meijer and Wood (1964), Yasman (1994), Newman et al. (2000) and Sidiyasa (2001) describe the seedling of S. leprosula as follows: Main stem, twigs and petioles glabrous or with soft hairs; Stipules 0.5 by 0.1 to 0.2 mm, oblong, reddish green, c. 4-nerved, stellate pubescent, persistent; petiole 0.7 cm; Leaves 4 to 5.6 by 9 to 12.7 cm, oblong or oblong lanceolate including 0.7 cm long gradually tapering acumen, glabrous above except for minute stellate hairs on midrib and nerve bases, 12 to15 nerves, glabrous below except for fine, small stellate hairs on the midrib, on upper surface margin of midrib swollen by the domatia, center sunken, below elevated and densely longitudinally channeled; base rounded or almost truncate; nerves sharply turned up at the margin; petiole 0.3 in. long; Domatia present on lower surface, arranged continuously along the main nerve (midrib) or sometimes up to half the base of the secondary nerve when young. 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All individual plants were sampled within their respective subplots, and 200 butterflies were sampled per total plot. Standardising sample size is preferable for mobile animals because sampling regimes based on a standard amount of sampling days can introduce an unacceptable bias when weather conditions are adverse. Relative density in this case is the frequency of encounter (number of individuals per species divided by sampling time). The area sampled is the same for all plots. The frequency of encounter can be used to estimate the density based on the total length of the transect sampled, crypsis and effective search width (MacKinnon et al., 1996) all of which are standardised here so that only the actual frequency is given. The collection was discontinued under adverse weather conditions (rain). 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Data points are daily measurements over a 30-day period. Humidity and temperature follow a standard linear model. Light follows an exponential model. Both temperature and humidity show no significant pattern at 07.00 hrs and a maximum variation between burned and unburned areas at 12.00 hrs. Average maximum and minimum are calculated at the extremes of the 200 m transect. Maximum average humidity is 91.07% (± 1.47%) and minimum average humidity is 74.88% (±1.47%) which is a difference of 16.18% between burned and unburned forest. The maximum average temperature is 30.85°C (± 0.30) and the minimum average temperature is 29.39°C (± 0.30) which is a difference of 1.46°C between burned and unburned forest. Light is even more extreme and ranges from 183.428 Lux in unburned forest to 2170.16 Lux in burned forest, which is more than a ten-fold increase in light intensity. 26 7KH,QLWLDO,PSDWRI)RUHVW)LUHVRQ3ODQWDQG%XWWHUIO\&RPPXQLWLHVLQWKH6XQJDL:DLQ)RUHVW 7DEOH 5HVSRQVHVRIYDULRXVVHHGOLQJDQGVDSOLQJYDULDEOHVWRIRUHVWEXUQLQJ &DWHJRU\ 7UDQVH!W 5 6LJ ,QWHVW 6( 7VWDW 6ORSH 6( 7VWDW HVW 6DSOLQJ 'HQVLW\ 7RWDO %XUQHG 1V 8QEXUQHG 1V 'LDPHWHU 7RWDO %XUQHG 1V 8QEXUQHG 1V *URZWK G 7RWDO 1V %XUQHG 1V 8QEXUQHG 1V *URZWK K 7RWDO 1V %XUQHG 1V 8QEXUQHG 1V +HLJKW 7RWDO %XUQHG 1V 8QEXUQHG 1V 6SH!LHV 7RWDO %XUQHG 1V 8QEXUQHG 1V 6HHGOLQJ 'HQVLW\ 7RWDO %XUQHG 1V 8QEXUQHG 1V *URZWK K 7RWDO %XUQHG 1V 8QEXUQHG 1V +HLJKW 7RWDO %XUQHG 8QEXUQHG 1V 6SH!LHV 7RWDO %XUQHG 1V 8QEXUQHG 1V 6SH!LHV &OLPD[ 7RWDO %XUQHG 1V 8QEXUQHG 1V 3LRQHHU 7RWDO %XUQHG 1V 8QEXUQHG 1RWH 'LDPHWHU KHLJKW %RWK WKH \LQWHU!HSW DQG VORSH !RHIIL!LHQW DUH JLYHQ LQ!OXGLQJ D WWHVW WR VHH LI HLWKHU VLJQLIL!DQWO\ GLIIHU IURP ]HUR $OO UHJUHVVLRQV DUH EDVHG RQ VWDQGDUG OLQHDU PRGHOV RYHU WKH WRWDOWUDQVH!WOHQJWKDQGLQEXUQHGDQGXQEXUQHGIRUHVWVHSDUDWHO\7KHODWWHUWHVWVIRUDQ\HGJH HIIH!W1RWHWKDWDOPRVWDOOYDULDWLRQLVEHWZHHQEXUQHGDQGXQEXUQHGIRUHVW7KHUHLVYHU\OLWWOH YDULDWLRQZLWKLQHLWKHUEXUQHGRUXQEXUQHGIRUHVW7KHRQO\YDULDEOHWRVKRZDVLJQLIL!DQWHGJH HIIH!WLVWKHVHHGOLQJKHLJKWZKL!KKDVDVLJQLIL!DQWO\QHJDWLYHUHVSRQVHLQEXUQHGIRUHVW Dipterocarpaceae: Forest Fires and Forest Recovery a. b. burned unburned burned unburned burned unburned burned unburned Figure 2.3 Distance from the edge versus height (cm) and growth of seedlings and saplings. Data points are measurements of individual plants in 2 x 2 m subplots (seedlings) and 5 x 5 m subplots (saplings). Seedling height (a) is significantly greater in unburned than in burned forest. Seedling growth however, is significantly greater in the burned forest. Those data points greater than 100 in plot (b) all belong to either Mallotus mollissimus or Mallotus paniculatus (Euphorbiaceae). Sapling height (c) is significantly greater in burned than in unburned forest. Compared to unburned forest all saplings shorter than 5 m died. In the burned area there is a patch of unburned forest (Plot 4, subplots 7 and 9) that can be seen in plot (c). There is no significant difference between the growth of saplings in burned and unburned forest. It is evident, however, that there was very little growth for saplings outside the unburned patch in the burned forest. 2.3.2 Species richness and density The species richness (Ancova) and density (Anova) of seedlings is significantly lower in burned than in unburned forest (Table 2.3). In butterflies, only the tree-und functional group has significantly fewer species in burned forest than in unburned forest. However, there is much stronger variation in pooled functional group density with significantly higher densities of mix-gap, mix-und, shrub-und and tree-und butterflies in unburned forest. The mix-edge functional group has a significantly 28 7KH,QLWLDO,PSDWRI)RUHVW)LUHVRQ3ODQWDQG%XWWHUIO\&RPPXQLWLHVLQWKH6XQJDL:DLQ)RUHVW KLJKHUGHQVLW\LQEXUQHGIRUHVW7KLVLQGL!DWHVWKDWGHVSLWHDVSLOORYHUDVLQGL!DWHGE\ WKH SDX!LW\ RI VLJQLIL!DQW YDULDWLRQ EHWZHHQ ERWK IRUHVW VWDWHV WKHUH LV VWLOO D VLJQLIL!DQW UHVWUX!WXULQJ RI WKH EXUQHG IRUHVW !RPPXQLW\ UHODWLYH WR WKH XQEXUQHG IRUHVW 7DEOH 5HVXOWVRIWZRZD\$QRYD¶VDQGWZRZD\$Q!RYD¶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ipterocarpaceae: Forest Fires and Forest Recovery Figure 2.4 Biplot of the butterfly community from the correspondence analysis of all butterfly species (n > 5). The largest amount of variation (16.48%) in the butterfly community is between burned and unburned states along CA-1. Species to the extreme left of the biplot (low CA-values) are strongly associated with unburned forest, i.e. Arhopala pseudomuta and Arhopala norda. Species to the extreme right of the biplot are associated with burned forest, i.e. Appias paulina and Eurema lacteola (Appendix 2). Figure 2.5 Biplot of the seedling plant community from the correspondence analysis. As in the butterfly community the largest amount of variation (23.34%) is between burned and unburned sites along CA-1. Plant species to the extreme right are strongly associated with unburned forest i.e. Aglaia simplicifolia (Meliaceae) and Dipterocarpus convertus (Dipterocarpaceae). Plant species to the extreme left are strongly associated with burned forest, i.e. Dillenia sp. (Dilleniaceae) and Macaranga gigantea (Euphorbiaceae). 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The Initial Impact of Forest Fires on Plant and Butterfly Communities in the Sungai Wain Forest significant preference for unburned forest. The seeds of understory tree Fordia splendidissima were clearly the most successful of the pre-fire trees although it also showed a significant preference for unburned forest. It is interesting (see Figure 2.8) that many typical forest butterflies such as J. pura, various Tanaecia species and Erites elegans (not shown) exhibited a rather abnormal behaviour by their often pronounced presence in severely perturbed, burned areas. Normally these species are confined to the forest. Figure 2.8 Distribution of the four most abundant butterfly and seedling species in the Sungai Wain forest. All species except T. clathrata show a significant preference for either burned or unburned forest. The preference of germination for seedlings is marked with almost no germination of rainforest trees such as S. laevis and M. lowii in burned forest and no germination of the pioneer M. gigantea in unburned forest. With the seedling species the third column indicates the number of saplings. Saplings indicate the prefire forest community and most of them died in the burned forest. The pioneer M. gigantea, for example, was absent from the prefire forest as indicated by the complete lack of saplings. The butterfly species O. medus is also a typical pioneer associated with secondary vegetation. 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The Impact of Environmental Conditions on the Growth of S. leprosula: the Greenhouse Studies but reduces the number of unwanted contaminant organisms, i.e. pasteurisation. The experimental result hence is only considered as far as the statistical outcomes are highly significant, because a significance at lower levels may be a false significance, due to ‘environmental noise’, i.e. a variation in environmental factors that is random in the context of the experiment. In the light of laboratory science, such an experimental set-up is unsatisfactory. As a field experiment, it has the drawback of many artificial structures and factors. Still, it is preferred in the present book to remain closer to natural conditions than to perfect laboratory regulation of experimental conditions. Our experiments, therefore, have the virtue of streamlining mere empirical data from nursery practices while at the same time the results are directly relevant to silvicultural practices. This viewpoint will be evaluated in the discussion at the end of this chapter. 4.2 EXPERIMENT 1: THE INFLUENCE OF SOIL TYPES AND ORIGINAL POPULATION ON Shorea leprosula. Assessment of factors buffering stress in S. leprosula; origin of the plant material versus soil type. 4.2.1 Location and time of the experiment The experiment was carried out in a greenhouse at the Wanariset Samboja research station (0°59’48” latitude South and 116°58’37” longitude East), located some 38 kilometers northeast of Balikpapan in Samboja village, Kutai district, East Kalimantan (Fig. 4.1). The experiment started in April 2000 by planting S. leprosula wildlings and ended after 20 months in December 2001. 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