RESPONSE OF MADAGASCAR’S ENDEMIC CARNIVORES TO FRAGMENTATION, HUNTING, AND EXOTIC CARNIVORES ACROSS THE MASOSALA-MAKIRA LANDSCAPE. Zachary J. Farris Dissertation submitted to the faculty of the Virginia Polytechnic Institute and State University in partial fulfillment of the requirements for the degree of Doctor of Philosophy In Fish & Wildlife Sarah M. Karpanty, Co-Chair Marcella J. Kelly, Co-Chair Dean F. Stauffer Steig E. Johnson November 24, 2014 Blacksburg, VA Keywords: behavior, bushmeat, camera trap, community ecology, co-occurrence, demographics, lemur biology, occupancy, poaching, temporal activity Copyright © 2014 Zach J. Farris RESPONSE OF MADAGASCAR’S ENDEMIC CARNIVORES TO FRAGMENTATION, HUNTING, AND EXOTIC CARNIVORES ACROSS THE MASOSALA-MAKIRA LANDSCAPE. Zach J. Farris ABSTRACT The carnivores of Madagascar are likely the least studied of the world’s carnivores, thus little is known about threats to their persistence. I provide the first long-term assessment of Madagascar’s rainforest carnivore community, including: 1) how multiple forms of habitat degradation (i.e., fragmentation, exotic carnivores, human encroachment, and hunting) affect native and exotic carnivore occupancy; 2) how native and exotic carnivore temporal activity overlap and how body size and niche explain these patterns; 3) how native and exotic carnivores spatially co-occur across the landscape and which variables explain these relationships; and 4) how native and exotic carnivores and humans co-occur with lemurs across Madagascar’s largest protected landscape: the Masoala-Makira landscape. From 2008 to 2013 I photographically sampled carnivores and conducted line- transect surveys of lemurs at seven study sites with varying degrees of degradation and human encroachment, including repeat surveys of two sites. As degradation increased, exotic carnivores showed increases in activity and occupancy while endemic carnivore, small mammal, and lemur occupancy and/or activity decreased. Wild/feral cats (Felis sp.) and dogs (Canis familiaris) had higher occupancy (0.37 ± SE 0.08 and 0.61 ± SE 0.07, respectively) than half of the endemic carnivore species across the landscape. Additionally, exotic carnivores had both direct and indirect negative effects on native carnivore occupancy. For example, spotted fanaloka (Fossa fossana) occupancy (0.70 ± SE 0.07) was negatively impacted by both wild/feral cat (beta = -2.65) and Indian civets (beta = -1.20). My results revealed intense pressure from hunting (ex. n = 31 fosa Cryptoprocta ferox consumed per year from 2005-2011 across four villages), including evidence that hunters target intact forest where native carnivore and lemur occupancy and/or activity are highest. I found evidence of high temporal overlap between native and exotic carnivores (ex. temporal overlap between brown-tail vontsira Salanoia concolor and dogs is 0.88), including fosa (Cryptoprocta ferox) avoiding dogs and humans across all seasons. However, I found no evidence of body size or correlates of ecological niche explaining temporal overlap among carnivores. Estimates of spatial co-occurrence among native and exotic carnivores in rainforest habitat revealed strong evidence that native and exotic carnivores occur together less often than expected and that exotic carnivores may be replacing native carnivores in forests close to human settlements. For example, falanouc show a strong increase in occupancy when dogs are absent (0.69 ± SE 0.11) compared to when they are present (0.23 ± SE 0.05). Finally, the two-species interaction occupancy models for carnivores and lemurs, revealed a higher number of interactions among species across contiguous forest where carnivore and lemur occupancy were highest. These various anthropogenic pressures and their effects on carnivore and lemur populations, particularly increases in exotic carnivores and hunting, have wide-ranging, global implications and demand effective management plans to target the influx of exotic carnivores and unsustainable hunting affecting carnivore and primate populations across Madagascar and worldwide. iii Dedicated to Alisha, aka “Skinny love” Oliver, “Tiny prancer” & & Fletcher “Stinky Pete” Nothing will ever compare to the great joy of being a part of your lives. Your humor, excitement, and outright craziness made this experience a blast! Thanks for being part of this crazy ride. iv ACKNOWLEDGEMENTS First and foremost, I am truly grateful to my wife Alisha and two boys, Oliver and Fletcher. Your amazing support and ridiculous humor helped me stay sane. I would follow you three anywhere! I am thankful to my advisers Sarah Karpanty and Marcella Kelly for this amazing opportunity. From your last minute edits, to your miraculous and mysterious sources of funding, to your encouraging and kind emails after a tough day, you both went above and beyond your role as an adviser. I consider you both as great mentors, colleagues, and friends. I am thankful to my additional committee members, Dean Stauffer and Steig Johnson. Dean, you indeed live by the motto that has accompanied your emails all these years, “Integrity is when what you think, say, and do are all in alignment.” Steig, you played a big role in shaping my career early on as a researcher in Madagascar. I am grateful that you were able to serve as one of my committee members. I am thankful to all the members of the Whapa lab (L. Rich, A. Hillborne, M. Kane. K. Thapa, D. Montague, D. Morin, E. Poor, C. Wultsch, C. Tredick, M. St. Germain, Sunarto, C. Rowe, C. Satter, B. Augustine, B. Mesa, and C. Proctor), the Karpanty lab (K. Davis, T. and K. Geider, A. Murphy, M. Wura, B. Gerber, M. Kotschwar, K. Ray, and M. Hillman), and Fraser lab (S. Ritter, A. DeRose-Wilson, D. Catlin, J. Cohen, J. Felio, D. Fraser, K. Hunt, M. Friedrich, and J. Fraser) for their friendship, v encouragement, humor, and support these many years. I am also thankful to all the remaining staff, faculty, and students of Virginia Tech FiW for their role they have played over the last five years to help me get to this point. I am grateful to Tom (“Tboy”) who played a massive role in making this experience really enjoyable for me and my family (not so much my kids though as you have scarred them for life). The truly immense data set collected from 2008 to 2013 in Madagascar would not have been possible without the assistance of countless organizations and their supporting staff. I am indebted to Chris Holmes and his family for providing this opportunity, for offering his unending support, and for looking out for my family while in Madagascar. The numerous researchers and staff of the Wildlife Conservation Society Madagascar Program are the reason all these data and subsequent results are coming to light. Felix Ratelolahy is a fearless researcher and true friend. Vonjy Andrianjakarivelo, I am thankful for your extensive insight and impressive knowledge of the Masoala-Makira landscape and its diverse wildlife. To the remaining staff and researchers of WCS Madagascar your fingerprints are all over these vitally important results on the worlds least known and most threatened carnivores. Your tireless work to conserve Madagascar’s biodiversity is truly inspiring and does not go unnoticed. I am truly thankful to Donah, Marka’Helin, and Wilson for their amazing work ethic and their fabulous sense of humor all those days and nights as we trekked through impenetrable rainforest and suffered through some of the hardest conditions imaginable. I am also thankful to the numerous Malagasy assistants that played a significant role in this vi project (C.B. Beandraina, B.A. Salofo, R.C. Christian, Didice, B. Papin, Rabeson, Tobey, Cressent, J. Fernando, and Sassid). I am grateful to each student who sacrificed their lives, their budgets, and their sanity to travel to Madagascar to assist on this project (Andi Evans, Toby Nowlan, Kat Miles, Hunter Doughty, Kate Galbreath, Chelsea Miller, Joe Larson, and Hanna Davis) Further, without the countless Virginia Tech undergraduate volunteers who helped enter the plethora of photos and ID carnivores I would have been here another 10 years. I am thankful for the opportunity to collaborate with Chris Golden and his well-qualified team, as well as with Brian Gerber. I would like to acknowledge my numerous funders including: Cleveland Metroparks Zoo, European Association for Zoos and Aquariums, Idea Wild, National Geographic Society- Waitts grant (#W96-10), Peoples Trust for Endangered Species, Virginia Tech Chapter of Sigma Xi, Virginia Tech McGinnis fellowship and Mosby fellowship, as well as logistical and financial support from the Wildlife Conservation Society Madagascar Program. In addition, I acknowledge the funders would supported the bushmeat surveys by Chris Golden and his team, including: Margot Marsh Biodiversity Fund (grant #023815), Mohammed bin Zayed Species Conservation Fund (grant #1025935), National Geographic Society Conservation Trust (grant #C135-08), National Science Foundation Doctoral Dissertation Improvement grant (#1011714) and Coupled Human Natural Systems grant (NSF-GEO 1115057). I would not be where I am today without my family. Jeff, Josh, Jonathan, and Jonah, your continued friendship means the world to me. I could not envision a more ridiculous or vii rewarding way to be brought up. To my grandparents, you paved the way allowing me to have this opportunity. I am so happy to continue following in your footsteps. Becca and Judge, thank you both for your friendship and your valuable insight. Mom, thank you for your unwavering love and support. Your amazing sacrifices you have made and continue to make to ensure our family (and other random families in need) are taken care of will never be fully realized. Your faith continues to amaze me. Dad, I cannot thank you enough for the passion you gave me for the natural world. Clearly I would not be where I am without all your insane, backwoods family camping and floating trips. I have always loved and will continue to love all the time we have been able to share exploring and hiking together.