Dune Management and Reptiles: Implications for Habitat Reconstruction and Conservation Strategies
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Dune Management and Reptiles: Implications for Habitat Reconstruction and Conservation Strategies. Thesis submitted in partial fulfillment of the requirements for the degree of “DOCTOR OF PHILOSOPHY” by Boaz Shacham Submitted to the Senate of Ben-Gurion University of the Negev January, 2010 Beer-Sheva Dune Management and Reptiles: Implications for Habitat Reconstruction and Conservation Strategies. Thesis submitted in partial fulfillment of the requirements for the degree of “DOCTOR OF PHILOSOPHY” by Boaz Shacham Submitted to the Senate of Ben-Gurion University of the Negev Approved by the advisor _______________________ 29-DEC-2010 Approved by the Dean of the Kreitman School of Advanced Graduate Studies ______________ January, 2010 Beer-Sheva 1 Boaz Shacham – PhD dissertation This work was carried out under the supervision of Dr. Amos Bouskila In the Department of Life Sciences Faculty of Natural Sciences Ben-Gurion University of the Negev 2 Boaz Shacham – PhD dissertation Acknowledgements Lack of space prevents me including by name all who deserve my thanks for support, advice and help that bring me to this point. I thank the partners in planning, funding and execution of the Nizzanim project. The staff of the Life Sciences Dept. and the Geography and Environmental Development Dept. at Ben Gurion University of the Negev, for preparations, planning, logistics and execution, for scholarships that gave me some peace of mind. The Jewish National Fund (KAKAL), for starting the journey and initializing management actions. Israel Nature & Parks Authority (INPA) rangers, scientists and permit office staff, and for project funding. Shiqmim Field Study School, my beloved de facto second home, for being our base camp during field sessions. The International Arid Land Consortium (IALC), for initial project funding. The Ministry of Science, for project funding. I thank the wonderful people I was honored to work with and learn from. First and foremost, my supervisor Dr. Amos Bouskila, for being guide and mentor to me since we first met more than 25 years past (we both deny it's been that long). Prof. Pua Bar and Dr. Elli Groner, truly the salt and pepper of our Nizzanim expeditions. My fellow M.Sc. & Ph.D. students at Nizzanim, past and present: Adi Ramot, Constantin Grach, Merav Perry, Ahikam Averbouch, Sharon Renan, Tarin Paz, Yehonathan Rubinstein. Our amazing field technicians: Arnon Tsairi, Yael Bogin-Zilka. Our B.Sc. project students, for leaving their mark in my heart as well as the sand: Arnon Yohanan, Shlomi Aharon, Noa Angel, Eyal Ben David, Na'ama Brener, Irit Messika, Hadas Ast, Ilana Ben David. To our "reservists" who volunteered to supplement or head reptile teams: Amir Arnon, Karmit Arbel, Tamar Amit, Itai Renan, Gal Vine, Eran Banker. My dear friends and partners guiding in the dunes, Avishay Shlomo and Dr. Oded Cohen. Many dozens of students who helped collect data within ecology camps of Life Sciences Dept., Geography & Environmental Development Dept. and Arava Institute for Environmental Studies. My colleagues at the Life Sciences Dept., for hours of fruitful corridor chats, prep and work as teaching assistants in labs and field trips, and forgiveness for my lengthy seminar lectures. The amazing Life Sciences Dept. secretariat, always helpful and smiling even via email: Hanna Hatav, Iris Reuven, Dalit Dahan, Anat Ben Haroush. At INPA, Dr. Yariv Malihi, Dr. Yehoshua Shkedy, Ronen Shavit, Koby Soffer, Itamar Wollis, Shai Cohen. Yair Farjun, man and legend, for nurturing my sand obsession. Elimor Fried, for the continued cooperation of Shiqmim Field School. And my M.Sc. supervisor Prof. Yehudah L. Werner, for supplying (past and present) tools for my evolution as a scientist. And last but not least, I warmly thank my dear family, for continuously accepting me and my accompanying critters. To my mother Nurit for instilling my love for nature and animals from day one, my brothers Lee Or and Uri for forgiving my total lack of interest in soccer, to my beloved Sharon for tolerating weird work hours and dunes wandering into our home, and to our lovely daughter Tamar for giving us a reason to smile every morning. I am lucky to have all of them, including Tamar, as partners in my "hunting expeditions" from time to time. - i - 3 Boaz Shacham – PhD dissertation Dedication I dedicate this work in memory of my late father, Yoram Shacham (Weinraub) ZL, whom I never had the privilege to meet, but as years pass it becomes obvious that an affinity to science and love for nature binds us far and beyond our hereditary connection. And also, to my beloved grandparents, Lena and Moshe Biegeleisen ZL and Donia and Heini Weinraub ZL, and to Moya Shalgi, may they rest in peace, who did not live to see the upcoming piece of familial satisfaction: the kid is about to finish his doctoral degree. - ii - 4 Boaz Shacham – PhD dissertation Table of Contents ABSTRACT …………………………………………………………………………………….. 4 Introduction ……………………………………………………………………………………... 9 Coastal Dune Ecosystems ………………………………………………………................ 9 Coastal Sand Dunes in Israel ……………………………………………………............... 12 Choosing Nizzanim Sands as a Case Study ………………………………………………. 15 The Nizzanim Project ………………………………………………………………….…. 19 Study Questions, Hypotheses & Predictions….………………………………………………… 23 The Research Questions ……………………………………………………...................... 23 Products of This Study ……………………………………………………………………. 23 Hypothesis & Predictions ………………………………………………………………… 24 Methodology ……………………………………………………………………………………. 26 The Study Site ………………………………………………………………………..…… 26 Experimental Design ……………………………………………………........................... 29 Methods for Sampling the Reptiles ………………………………………………………. 35 Data Analysis Techniques ……………………………………………………................... 42 Results I: Reptile Assemblages at Nizzanim Sands …………………………………………….. 45 Species Assemblages on Various Plot Types ……………………………………….…….. 45 Effects of Vegetation Cover on Reptile Species Assemblages ……………………………. 53 Seasonal Effects on Reptile Activity and Abundance …………………………………….. 55 Patterns Seen in Diurnal Lizard Observations ………………………………….................. 59 Results II: Response of Reptiles to Dune Management ………………………………………… 62 Changes in Reptile Assemblages Following Management Actions …………………….… 62 Trajectory of Management Effects Through Time ………………………………………... 70 Effects of Management on Lizard Body Condition ……………………………………… 77 Response of Acanthodactylus to Dune Management ………………………………..…… 85 Discussion and Conclusions ……………………………………………………………………. 92 Reptile Assemblages at Nizzanim Sands ………………………………………………… 92 Reptile Assemblage Response to Manipulation at Nizzanim Sands …………………….. 96 Implications for Management and Monitoring ……………………………….................... 103 Potential Obstacles: How does the Project Measure Up? …………………….................... 111 Summary of Conclusions from Nizzanim Sands …………………………………………. 113 Summary of Recommendations for Monitoring Sand Dune Reptiles ……………………. 114 New Study Questions Arising from this Project ……………………………………..…… 115 References ………………………………………………………………………………………. 117 Appendix I: Restoration Ecology – Definitions, Strategies, Potential Problems, Design ……… 128 Definition of Basic Terms …………………………………………………………….….. 128 Conceptual Framework ……………………………………………………....................... 129 1 Boaz Shacham – PhD dissertation Preliminary Assessment ……………………………………………………...................... 130 Thematic Questions ……………………………………………………............................. 131 Potential Difficulties Interpreting Outcomes of Restoration ……………………………... 131 The Need for Strategic Planning in Restoration ………………………………………..… 131 Appendix II: Herpetofaunal Checklist of Nizzanim Sands and Environs ……………………… 133 Figures and Tables Figure 1: Active dune……………………………………………………………………………………... 27 Figure 2: Semistable dune………………………………………………………………………………… 28 Figure 3: Stabilized dune………………………………………………………………………………….. 29 Table 1: List of the study plots presented in this work……………………………………………………. 30 Figure 4: General map of Nizzanim sands study site……………………………………………………... 31 Figure 5: Bulldozer at work.………..………………………………………….......................................... 33 Figure 6: Plot marking.………………………….…………………………................................................ 33 Figure 7: Debris from removal. ……………………………………………………................................... 34 Figure 8: Aerial photo of manipulated dunes. ……………………………………………………............. 34 Table 2: Characteristics of data obtained by each sampling method……………………………………... 37 Figure 9: Spatial scheme of sampling methods.………….………………………………………….......... 38 Figure 10a: Schematic side view of bucket pitfall trap.….………………………………………………. 39 Figure 10b: Schematic top view of bucket pitfall trap.….………………………………………………... 39 Figure 11: Pitfall configuration and numbering.………….…………………………………………......... 40 Figure 12: Covered pitfall.………………………………….………………….......................................... 40 Figure 13: Pitfall exposure.………………………………….…………………......................................... 41 Figure 14: Track transect.……….……………………………………………............................................ 41 Table 3: Summary of species found.………………….…………………………………........................... 46 Figure 15: Proportional incidence of common species.…….…………………………………………….. 47 Table 4: Average observations and species richness.……….…………………………………………….. 49 Table 5: Average species richness pooled for all methods.……….………………………………………. 50 Figure 16: Average species richness indices.……….…………………………………………….............. 51 Figure 17: