Growth and Acclimation Responses of Dipterocarp Seedlings to Logging Disturbance
Total Page:16
File Type:pdf, Size:1020Kb
Growth and acclimation responses of dipterocarp seedlings to logging disturbance Michael J. Clearwater A thesis submitted for the degree of Doctor of Philosophy Institute of Ecology and Resource Management University of Edinburgh August 1997 Declaration This thesis has been composed by myself from the results of my own work, except where stated otherwise, and has not been submitted for any other application for a degree. 11 Abstract The growth and acclimation response of wild dipterocarp seedlings to environmental change caused by logging disturbance was examined in lowland dipterocarp forest in Central Kalimantan (Indonesian Borneo). The aim was to identify environmental and physiological limitations to natural regeneration after logging. The microclimate of logged forest was compared with undisturbed forest. Seedling growth and mortality after logging was described at the population level, and the physiological causes of growth limitation and mortality after logging were explored. A system for estimating daily below canopy PPFD (photosynthetic photon flux density) from hemispherical photographs was developed and calibrated against direct measurements of PPFD. Predictions from photographs were repeatable, accurate (r 2 > 0.95) and linearly related to measured PPFD over at least the range 5 - 50 % of above canopy daily PPFD. Photographic estimates of the diffuse proportion and above canopy PPFD were independent of canopy cover and representative of measured above canopy values over a range of seasonal conditions, thus supporting the validity of the model. The most important effect of logging on the seedling microclimate will be the increased exposure of seedling leaves to direct sunlight. Air temperature and the vapour pressure deficit of the air were influenced more by season and site location than by canopy opening. Low wind speed was an important feature which will decouple seedling leaves from ambient conditions. It was hypothesised that low boundary layer conductance and high leaf temperatures during periods of direct sunlight would have important effects on dipterocarp seedling regeneration in logging gaps. Logging increased patchiness and removed seedlings from disturbed areas where they were most required. Seedling mortality was lower than in undisturbed forest. Wild light hardwood seedlings (Shorea johorensis, S. leprosula, S. parvifolia, S. ovalis) reached maximum growth rates at moderate PPFDs (5 - 10 mol m 2 day'), and in disturbed soil growth declined with further increases in PPFD. Canopy opening after conventional logging often exceeds that required for maximum seedling release. Seedling nutrient status, mycorrhizal infection and the capacity for photosynthetic acclimation were investigated as potential causes of limitation to growth in open sites. Seedlings of S. leprosula and S. parvifolia were tolerant of low nutrient availability when grown in high light under controlled conditions. Mycorrhizal infection was increased when nutrient availability was low and seedlings were growing fast in high light. Both species exhibited significant architectural and photosynthetic plasticity. S. johorensis seedlings were found to be capable of acclimation to extreme post logging conditions, including changes in leaf nutrient partitioning, but the respiratory cost of acclimation to high leaf temperatures and PPFD may contribute to limitation of seedling growth. Water availability and the response of stomata to humidity were not important limitations during the wet season. Variation in the acclimation response was attributed to soil conditions and seedling nutrient status. It was concluded that the most important limitation to natural regeneration after logging was the direct physical impact on seedling populations and the residual stand at the time of logging. Limitations imposed by microclimate and seedling physiological characteristics will be less important if logging impacts are reduced. 111 Acknowledgements First of all, I would like to thank my supervisor, Paul van Gardingen, who gave me the opportunity to work on the Indonesian project and provided lots of support along the way. Thanks also to my second supervisor, John Grace, who was always ready with useful ideas. I am deeply grateful to the many Indonesian people who allowed me the privilege of working in their country and helped me adjust to their culture and forests. In particular the Indonesian Institute of Science (LIPI) and the Forest Research Institute, Samarinda (BPK) gave permission to work at Wanariset Sangai and to export plant material. Riskan Effendi, Wisnu Rusmantoro, Thomas Nifinluri, Ismanto, Merang, Tanyit Hendrik, and Yan provided vital logistical and scientific support. Chay Asdak assisted with collection of meteorological data. Bidu and the rest of the pondok/camp 48 crew taught me much about life in Kalimantan and provided essential help and entertainment in the field. Simon Armstrong and Peter Wilkie were there in the early days when the dried fish and super mie got a bit too much. The staff of PT Kayu Mas were always helpful when needed. Runci Susilawaty, Yuni Norma Idha, Merty Ilona and Khairul Basyar were collaborators for the work in Chapter 6 and endured through the falling trees, leeches and mud. This research forms a component of the Indonesia UK Tropical Forest Management Programme, funded by DFID and the Indonesian Ministry of Forestry, and I am grateful to Alistair Fraser, David Taylor, Richard Scott, Robert De Kock and other project staff for their support and guidance. In the UK, thanks to Bob Munro and Kevin Ingleby of JTE who counted and identified the mycorrhizas. Steve Long kindly loaned the spectroradiometer. Plant material was imported into Scotland with permission from the Scottish Agricultural Science Agency (Licence No. IMP/SCEIREAI2/1995). None of this would have been possible without excellent support from the Association of Commonwealth Universities in the UK and the Vice Chancellors Committee in New Zealand in the form of a Commonwealth Scholarship. Linda Sharp and Shiela Wilson could always be relied upon. Andy Grey, Derek Scott, Steve Scott, Dave MacKenzie, Bob Astles and Malcolm Ritchie provided essential technical support. Yadvinder Malhi explained the principles of boundary layer meteorology and suggested the possibility of a sea breeze front. Thanks to all my fellow students here in Edinburgh who made life in the department at least as entertaining as that in the jungle. The cries of Patrick Meir and Yadvinder could always be heard from down the corridor, and the menagerie of the first and second floors was always open for a visit. Thanks to James Irvine who always knew the answer to a problem but never believed it. Allen Young helped me right from the beginning and provided lots of stimulating debate as we had one more for the road. Thanks to Mark Rayment for being the second fastest thesis writer in the office. Thanks also to my flatmates for putting up with me over the last few months, and to Julie and Rob for endless dinners, tarantulas, distraction and encouragement. Last but not least, thanks (again) to my parents and my sister Sue, who have always supported me in everything I do. iv Table of Contents Abstract iii Acknowledgements iv List of Abbreviations and Symbols ix 1. Introduction i 1.1 Achieving sustainable forest management 1 1.2 The Dipterocarpaceae 1 1.3 Silvicultural systems and sustainable management 3 1.4 Defining the problem 5 1.5 Dipterocarp seedlings: general characteristics and physiological factors potentially limiting to regeneration 6 Light 7 Nutrients 7 Mycorrhizal associations 8 Soil water and drought 8 Acclimation potential 9 1.6 Study site io 1.7 Thesis Aim: 12 1.8 Thesis structure, objectives and hypotheses 12 2. Calibration of hemispherical photography and estimates of the diffuse component for predicting below canopy light climate in Bornean tropical rain forest 14 2.1 Introduction 14 2.2 Methods 17 PPFD measurements 17 Hemispherical photography 19 2.3 Results 23 Global site factor vs measured PPFD 23 V Daily variation iflfd and Qa - measured and modelled values 26 Seasonal variation infd and Qa 28 Combined results 30 2.4 Discussion 31 Conclusions 34 3. Effects of logging and drought on the understorey microclimate of lowland dipterocarp rain forest 36 3.1 Introduction 36 3.2 Methods 39 Site description 39 Meteorological measurements 39 Sunshine hours 40 Soil surface temperatures 41 3.3 Results 42 Logged versus undisturbed forest 42 Daily versus seasonal variation 44 Daily time courses 45 Variation in cloudiness and sunshine hours 48 Soil temperatures 49 3.4 Discussion 51 Effects of logging 51 Seasonal changes in forest microclimate 54 Nocturnal wind - a sea breeze front? 55 Conclusions 56 4. Population dynamics and growth of dipterocarp seedlings after logging in Central Kalimantan, Indonesia 57 4.1 Introduction 57 4.2 Methods 59 Site description 59 Seedling tagging and measurement 60 Seedling light climate 61 An alternative approach - light climate estimates for individual seedlings 62 4.3 Results 63 Logging damage and seedling densities 63 Population structure 65 Mortality 68 vi Growth 69 4.4 Discussion 75 Spatial heterogeneity after logging 75 Growth response to light 75 Mortality 77 Interspecific comparisons 78 Conclusions 79 Effects of light and nutrient supply on the growth, mycorrhizal infection and photosynthesis of Shorea seedlings 80 5.1 Introduction 80 5.2 Methods 83 Plant material 83 Experimental treatments 84 Seedling growth