Ecology and Behavior of Two Sympatric Chameleon Species on Bioko Island, Equatorial Guinea Elliott Chiu Dr
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Ecology and Behavior of Two Sympatric Chameleon Species on Bioko Island, Equatorial Guinea. A Thesis Submitted to the Faculty of Drexel University by Elliott Chiu in partial fulfillment of the requirements for the degree of Master of Science June 2013 ©Copyright 2013 Elliott Chiu. All Rights Reserved i ACKNOWLEDGEMENTS First and foremost, I would like to thank my parents Serena and Edward, my brother Alvin, and my sister Melissa. Without them, I would not be here; they supported me emotionally and financially and I can truly say that this project is as much theirs as it is mine. I would like to acknowledge Dr. Tom Butynski and Dr. Torsten Wronski for their help in developing my study during the early stages of my thesis. Originally my project focused on rare occurrences between primate and duiker feeding habits, but you made me realize that projects that I envisioned would be difficult to finish even for a PhD and helped me develop a suitable Master’s thesis. I thank all of my friends who despite my constant weariness, never gave up on me and were a welcome respite from the long hours in the lab. Long absences can strain many relationships, but I have been lucky to know all of you. Despite many rejected offers, you never abandoned me and that is the test of a true friendship. Of course, much of the brunt of the fieldwork was shared by many dedicated volunteers. The students of the Drexel Study Abroad program on Bioko (including Evan Neyland, Kadeem Gilbert, Ryan Quinn, Liz Long, Beth Darby, and Baltasar), Nabil Nasseri, Andrew Fertig, Djibrilla Ousman, and Maya Lipschutz were instrumental in the overall collection of my chameleons. Reed Power provided incredibly useful habitat and density information to me when I was unable to return the field in early 2013. I would be remiss to forget my three amazing lab volunteers, Ellen Wildner, Lexi Khan, and Halle Choi, who spent many hours looking through dissecting scopes and ii separating out thousands of tiny invertebrates. Not only that, you remained patient during the entire bomb calorimetry fiasco. Special mention should be given to the graduate students of the Hearn lab Drew Cronin, Jake Owens, and Pat McLaughlin who shared many experiences with me regarding working in the tropics and never let bureaucratic nonsense impede on my progress. Additionally, I’d like to thank Drew Cronin and Steve Hromada for supplying me with the beautiful maps that have been generated for my thesis. Funders for my project were greatly appreciated. As mentioned before, my parents were my biggest financial support, but the Bioko Biodiversity Protection Program paid for many of my expenses in country and provided me with hot food and a “comfortable” bed/tent to sleep in. The Garden Club of America has been extremely generous in providing funds for the acquisition of many of my research supplies. Lastly, I would like to thank ExxonMobil for their support in conservation efforts on Bioko Island. Finally, I would like to thank the members of my thesis committee. Dr. Gail Hearn has been a great mentor and advisor for me, helping me with developing the sense to work in science abroad. I would be nowhere without the help of Dr. Mike O’Connor who taught me how to approach and answer scientific questions with statistics. Dr. Krystal Tolley advised me in all things chameleon, and was an irreplaceable member of my committee. Dr. Dan Duran provided me with much needed help in collecting and identifying the insects that were used in my study. And then the most important of all is Dr. Shaya Honarvar, who sat with me through many drafts of my thesis and personally devoted time and energy into seeing me succeed. iii TABLE OF CONTENTS LIST OF TABLES……………………………………………………………………....iv LIST OF FIGURES……………………………………………………………………...v ABSTRACT…………………………………………………………………………….vii 1. CHAPTER 1: General Introduction………………………………………………….1 2. CHAPTER 2: A life history of two of Bioko Island chameleons, the Fea’s chameleon (Trioceros feae) and the spectral pygmy chameleon (Rhampholeon spectrum) Introduction…………………………………………………………………….…6 Methods…………………………………………………………………………..10 Results…………………………………………………………………………....13 Discussion………………………………………………………………………..17 3. CHAPTER 3: Differential feeding strategy adaptations of Trioceros feae and Rhampholeon spectrum in Bioko Island, Equatorial Guinea Introduction……………………………………………………………………..30 Methods…………………………………………………………………………34 Results…………………………………………………………………………..37 Discussion………………………………………………………………………40 4. CHAPTER 4: Conservation implications of diet partitioning in response to differential nutritional requirements in the Fea’s chameleon (Trioceros feae) and the African leaf chameleon (Rhampholeon spectrum) on Bioko Island Introduction……………………………………………………………….……53 Methods…………………………………………………………………….…..57 Results……………………………………………………………………….…60 Discussion……………………………………………………………………...63 5. CHAPTER 5: Notes on the natural history of Bioko Island chameleons………….74 6. CHAPTER 6: General Conclusions………………………………………………..88 LIST OF REFERENCES……………………………………………………………....90 iv LIST OF TABLES 1. Stomach contents of gastric lavage were dissected and identified to the level of order. The number of individual prey items is listed with the number of stomach pellets in which the individual prey was collected in parentheses………………………………22 2. Intraspecific behavioral data: Total distance traveled (TDT), average speed (AS), moving speed (MS), percent time spent moving (PTM), and feeding attempts (FA) did not differ significantly with the change among sexes within species……………..….45 3. Interspecific behavioral data: Total distance traveled (TDT), average speed (AS), and moving speed (MS) were insignificantly variable between species. Percent time spent moving (PTM) and feeding attempts (FA) differed significantly between species….46 4. Binomial models determining variables affecting activity levels: The full model included five variables (sex, species, time, temperature, and the interaction between species and sex). Subsequent models removed one sequential variable until significant changes occurred with the removal of species and time………………………….…..47 5. Available invertebrates collected along each trail are categorized by the method by which they were collected. Percent (%) composition displays what percentage of the total available invertebrate selection was made up of each taxon………………..…..67 6. Stomach contents retrieved via gastric lavage are displayed for each species along with the trail on which they were collected. Numbers of prey items are reported along with the number of chameleons from which those individuals were collected, demarcated by the parentheses…………………………….……………………..…...68 7. Electivity indices (E*) are reported for T. feae along both trails. Invertebrate taxa are first blocked by prey hardness and then by evasiveness of individual prey taxa…….69 8. Electivity indices (E*) are reported for R. spectrum along both trails. Invertebrate taxa are first blocked by prey hardness and then by evasiveness of individual prey taxa…70 9. Combined electivity indices (E*) are reported for T. feae and R. spectrum. The Nature trail and the Cascades trail were grouped to produce one E* value. Invertebrate taxa are first blocked by prey hardness and then by evasiveness of individual prey taxa…71 10. Biometric measurements that were used for each captured chameleon in 2011…....78 11. Additional biometric measurements were used for each captured chameleon in 2012 (Hopkins & Tolley, 2011)…………………………………………………….………79 v LIST OF FIGURES 1. Four species of chameleons on Bioko Island. A) Rhampholeon spectrum perched upon tree fern branches undergoing ecdysis. Its skin is shed in one solid piece instead of flaking off in small pieces. B) Trioceros cristatus photographed on the 2008 National Geographic Expedition on Bioko Island. Photo credit: Christian Ziegler ILCP Bioko RAVE. C) Trioceros oweni is a mid-sized chameleon that was originally described from the island but its presence is not disputed. D) Trioceros feae is a mid-sized chameleon endemic to sub-montane forests of Bioko Island……………………...…..23 2. Distribution maps of R. spectrum, T. cristatus, T. feae, and T. oweni in Africa…..…..24 3. General map of study sites: The Nature trail and Cascades Trail are marked in respect to the Moka Wildlife Center (MWC) located just outside of the village of Moka. …..25 4. Density of chameleons were estimated on each trail. The Nature trail is classified as a more disturbed habitat compared to the Cascades trail…………………………….….26 5. Photos of the understory of A) Cascades trail and B) the Nature. The understory has been cleared of much of the flat broad leaf vegetation in the Nature trail whereas that secondary growth is still present along the Cascades trail………………………….….27 6. Sleeping perches utilized by T. feae during the 2011 field season. The majority of sleeping perches utilized were exotic species. …………………………………...……28 7. A sub adult male T. feae and adult male R. spectrum are shown here shortly after being aroused by observers in the forest. A) The T. feae is utilizing the small graspable twigs and branches to perch upon whereas B) R. spectrum use flat broad leaves to rest upon. They typically are not found to grasp small perches…………………………………..29 8. Differences in total distance traveled is plotted with respect to sex within each species as a result of direct 12-hour observation cycles (n=5). There are no statistical differences between species or sex……………………………………………….……48 9. Differences in average speed is plotted with