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Holm Jenniferallison 2012.Pdf i ii Copyright 2012 © By: Jennifer A. Holm All rights reserved i Abstract Because of human pressures, the need to understand and predict the long-term dynamics and development of subtropical dry forests is urgent. The new gap model ZELIG-TROP was developed, parameterized, and successfully modeled the forest dynamics of the Puerto Rico subtropical dry forest. Along with new forest parameterization, additions to ZELIG-TROP increased the level of detail in gap models by including a new natural disturbance routine and by tracking litterfall, leaf production, coarse woody debris, and carbon components at the individual tree level. This research developed a valuable tool to understand the process of rehabilitating dry tropical forests, understand the unique aspects and predict the future direction of subtropical dry forests. In addition to validating a new model, simulating the succession of secondary forests on abandoned fields was achieved, showing that secondary forests have a delayed response to recover and reach a mature forest status (100+ years). Realistic hurricane simulations were implemented within the modeled Puerto Rican subtropical dry forest from ZELIG-TROP, for the first time. A major gap in the literature is to compare storm effects on vegetation from storms varying in intensity, duration, and frequency that are forecasted in climate change scenarios. Increasing hurricane intensity of severe storms did not create a large shift in the average aboveground biomass, net primary productivity (NPP), annual litterfall, leaf production, coarse woody debris, or annual autotrophic live carbon accumulation (AALCA) from that of historical hurricane regime (control). During both control and increased hurricane ii intensity scenarios, AALCA was negative indicating a carbon transfer from the live forest to the surrounding ecosystem. Increasing the frequency of hurricanes had a much stronger effect on forest dynamics. In comparison to the control scenario, increased hurricane frequency decreased the aboveground biomass by 5%-40%, increased NPP by 32%-50%, increased leaf production, decreased litterfall and coarse woody debris, and AALCA became positive indicating a carbon sink into the forest over 600 years. Overall community composition does not substantially change from the control treatment when there is an increase in hurricane intensity or frequency, but species diversity was the highest during all treatments which increased the frequency of hurricanes. iii TABLE OF CONTENTS ABSTRACT……………………………………………………………………………I TABLE OF CONTENTS………………………………………………………………III ACKNOWLEDGMENTS……………………………………………………………..VI LIST OF FIGURES……………………………………………………………………VII LIST OF TABLES……………………………………………………………………..IX LIST OF PICTURES…………………………………………………………………..X LIST OF APPENDICES……………………………………………………………….XI CHAPTER 1 INTRODUCTION TO THE DISSERTATION AND BACKGROUND………………………………………………………………………1 1.1 Introduction……………………………………………………………………....1 1.1.1 Field Site Description…………………………………………………….4 1.1.2 Guanica Forest History…………………………………………………..7 1.1.3 Field Data Collection……………………………………………………..9 1.1.4 Background and Introduction to Hurricane Disturbances………………16 1.2 Model Background……………………………………………………………….17 1.3 Dissertation Objectives and Questions…………………………………………..19 1.3.1 Overall Project Questions……………………………………………….20 1.3.2 Specific Objectives……………………………………………………….20 1.3.3 Composition of Dissertation……………………………………………..22 1.4 Literature cited…………………………………………………………………...32 1.5 Appendix…………………………………………………………………………38 1.5.1 Growth Sub-routine……………………………………………………...39 1.5.2 Regeneration Sub-routine………………………………………………..44 1.5.3 Mortality Sub-routine…………………………………………………….45 CHAPTER 2 MODEL DEVELOPMENT, VALIDATION, AND APPLICATION TO SUCCESSION OF SECONDARY SUBTROPICAL DRY FORESTS IN PUERTO RICO………………………………………………………………………..50 2.1 Introduction………………………………………………………………………50 2.2 Methods…………………………………………………………………………..54 2.2.1 Description of ZELIG-TROP…………………………………………….54 2.2.2 Site Description…………………………………………………………..55 2.2.3 Data Collection/Model Parameterization………………………………..56 2.2.4 ZELIG-TROP model modifications………………………………………60 2.2.4.1 Height-diameter relationship and PET modifications……………….…..61 2.2.4.2 Basal-sprouting…………………………………………………………..62 2.2.4.3 Mortality estimation……………………………………………………...63 2.2.4.4 Aboveground biomass and crown architecture………………………….64 iv 2.2.5 Validation Methods………………………………………………………65 2.2.6 Model Testing: Transition of Abandoned Fields to Secondary Forest…..67 2.3 Results……………………………………………………………………………70 2.3.1 Validation Results………………………………………………………..70 2.3.2 Total Basal Area and Biomass Analysis………………………………....71 2.3.3 Species Composition……………………………………………………..72 2.3.4 Density Analysis…………………………………………………….........75 2.3.5 Percent error testing using actual data……………………………….....77 2.3.6 Model testing: transition of abandoned fields to secondary forests……..79 2.4 Discussion………………………………………………………………………..84 2.4.1 Model validation…………………………………………………………84 2.4.2 Need to incorporate disturbance into tropical gap modeling……………86 2.4.3 Relation to Previous Gap Models in the Tropics………………………...87 2.4.4 Model Testing: Transition of abandoned fields to secondary forests……89 2.5 Conclusions………………………………………………………………………92 2.6 Literature cited…………………………………………………………………...93 CHAPTER 3 SIMULATING HURRICANE DISTURBANCES IN CLIMATE CHANGE SCENARIOS: BIOMASS AND NET PRIMARY PRODUCTIVITY CHANGES FOR A SUBTROPICAL DRY FOREST………………………………102 3.1 Introduction…………………………………………………………………….102 3.1.1 Hurricanes in Puerto Rico……………………………………………..104 3.1.2 Hurricane modeling……………………………………………………105 3.1.3 Changes in forest biomass and productivity with disturbances……….107 3.2 Methods………………………………………………………………………...109 3.2.1 ZELIG-TROP model……………………………………………………109 3.2.2 ZELIG-TROP and Stand Visualization System…………………………110 3.2.3 Hurricane data collection and modeling……………………………….111 3.2.4 Hurricane disturbance simulations and modifications…………………116 3.2.5 Net primary productivity model additions……………………………...119 3.3 Results…………………………………………………………………………..120 3.3.1 ZELIG-TROP with hurricanes and SVS results………………………...120 3.3.2 Disturbance simulations………………………………………………..122 3.3.3 Effects of hurricane treatments in climate change scenarios…………..125 3.3.4 Effects of disturbance on net primary productivity (NPP)……………..128 3.4 Discussion………………………………………………………………………132 3.4.1 Hurricane treatments…………………………………………………...132 3.4.2 Effects of disturbance on net primary productivity……………………..136 3.4.3 Simulation modeling in comparison to field studies……………………138 3.4.4 Strategies for disturbance management and monitoring………………140 3.5 Conclusions…………………………………………………………………….142 3.6 Literature cited………………………………………………………………….143 3.7 Appendix……………………………………………………………………….152 v CHAPTER 4 ANNUAL CARBON PRODUCTION AND FUNCTIONALITY OF A SUBTROPICAL DRY FOREST AFTER HURRICANE DISTURBANCES…….155 4.1 Introduction…………………………………………………………………..…155 4.1.1 Components to annual autotrophic live carbon accumulation (AALCA)………………………………………………………………..158 4.1.2 Disturbance, forest productivity, and net carbon accumulation……….161 4.2 Methods………………………………………………………………………...165 4.2.1 ZELIG-TROP model and hurricane modeling…………………………165 4.2.2 Annual autotrophic live carbon accumulation (AALCA) model additions………………………………………………………………...166 4.3 Results…………………………………………………………………………..172 4.3.1 Litterfall and hurricanes in subtropical dry forests…………………….172 4.3.2 Leaf production and hurricanes in subtropical dry forests…………….176 4.3.3 Effects of disturbance on annual autotrophic live carbon accumulation (AALCA)………………………………………………………………..178 4.3.4 Relationship between CWD and litterfall………………………………183 4.4 Discussion………………………………………………………………………186 4.4.1 Importance of litterfall, leaf production, and woodfall measurements...186 4.4.2 Modeling annual carbon production with disturbances………………..188 4.4.3 Previous methods of determining AALCA components………………...190 4.4.4 Advantages to annual carbon production calculation………………….192 4.4.5 Management purposes and final conclusions…………………………..194 4.5 Literature cited………………………………………………………………….195 4.6 Appendix………………………………………………………………………..206 CHAPTER 5 SPECIES DIVERSITY ASSESSMENT WITH VARYING HURRICANE DISTURBANCES…………………………………………….....…..208 5.1 Introduction……………………………………………………………………..208 5.1.1 Hurricane disturbances and species diversity………………………….210 5.1.2 Intermediate disturbance hypothesis…………………………………...213 5.2 Methods…………………………………………………………………………214 5.2.1 ZELIG-TROP model and hurricane modeling………………………….214 5.2.2 Species diversity assessment……………………………………………216 5.3 Results…………………………………………………………………………..219 5.3.1 Species composition shifts in response to increased hurricanes……….220 5.3.2 Successional traits in response to increased hurricanes…………….…224 5.3.3 Diversity analysis with hurricane disturbances……………………...…228 5.4 Discussion………………………………………………………………………231 5.5 Literature cited………………………………………………………………….233 5.6 Appendix………………………………………………………………………..238 vi ACKNOWLEDGMENTS I gratefully thank my supervisory committee, Dr. Howard Epstein, Dr. Bob Davis, Dr. Gerard Learmonth, Dr. Guy Larocque, and most importantly my committee chair and advisor, Dr. Herman H. Shugart for their time and effort. I owe a large amount of gratitude to Dr. Hank Shugart who picked me up as a graduate student and graciously gave me much assistance, guidance, and support while completing my research. This work was made possible by the Environmental Sciences Department at University of Virginia (Charlottesville, USA) through
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