Platalina Genovensium, Pollinate the Colurnnar Cactus
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UNIVERSITY OF MIAMI BAT AND HUMMINGBIRD POLLINATION OF TWO SPECIES OF COLUMNAR CACTI: EFFECTS ON FRUIT PRODUCTION AND POLLEN DISPERSAL by CATHERINETERESASAHLEY A DISSERTATION Submitted to the Faculty ofthe University ofMiami in partial fulfuillment of the requirements for the degree ofDoctor ofPhilosophy Coral Gables, Florida May, 1995 ~ UNlVERSITY OF MIAMI A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy Bat and hummingbird pollination of two species of columnar cacti: Effects on fruit production and pollen dispersal Catherine Teresa Sahley Approved: -,.. 7f~~Hfh ~@ Dr. Tarek M. Khalil Professor of Biology Dean of the Graduate School Chairperson of the Dissertation Cornmittee (J\~l \ - \ Dr\J(eith Wáddington Dr. David Janos <:»: Professor ofBiology Associate Professor ofBiology ~c.~ Dr. Carol Horvitz qf.' j~mes riarnnck Associate Professor ofBiology Professor ofBotany •••• ~j :;-:tf~~,-. ¡,;.~-, SAHLEY,CATHERINETERESA (Ph.D., Biology) Bat and hummingbird pollination oftwo species of columnar cacti: Effects on fruit production and pollen dispersal. (May 1995) Abstract of a doctoral dissertation at the University ofMiami. Dissertation supervised by Professor Theodore H. Fleming No. of pages in text: 115 Two species of hummingbirds, Patagona ~ and Rhodopis vesper, and one species ofnectar-feeding bat, Platalina genovensium, pollinate the colurnnar cactus Weberbauerocereus weberbaueri in southwestern Peru. Initial fruit-set frorn pollinator exclusion experiments conducted in 1991 and 1993, just prior to and after the El Niño event of 1991-92, revealed that bats were the most important pollinators in 1991, but that hurnmingbirds were most important in 1993. In both years, however, autogamy and lepidopteran larval infestation of fruits reduced differences in mature fruit praduction among pollinator exclusion treatments so that differences among treatments in mature fruit-set were not statistically significant. Platalina ~novensium populationsdecreased at the study site between 1.991and 1993. Because the data indicate that bats are specializing on W. weberbaueri flowers and fruits for food, I hypothesize that bats emigrated from Arequipa during the drought because cactus flower and fruit production declined during this time. Persistence of small bat populations in Arequipa may be threatened because bats are harvested for use in folkloric medicinal remedies. In Sonora, Mexico, one species of bat, Leptonycteris curasoae, and one species of hummingbird, Cynanthis lasirostris, were the most important pollinators of the organ pipe cactus, Stenocereus thruberi. Bats and hummingbirds deposited large quantities of pollen on stigmas per flower visit. Pollinator exclusion experiments indicated that the praportion ...•. ~.•. ~....~--. --~~- of fruits produced by hummingbird pollination was constant while that by bats was greater in 1993 than in 1992. Pateroity exclusion analysis ofbat-and hummingbird-pollinated seeds indicated that most pollen probably carne from within 75 m although substantial amounts (5-11 %) carne from beyond 125 and 150 meters respectively, for the two ~ thurberi populations studied. To rny father, Theodore, who instilled in us a sense of adventure and respect for nature and rny rnother, Teresa, who raised us so that we would carry a little bit ofPeru in our hearts. iii Acknowledgments 1 would like to thank the members of my dissertation committee, Drs. Carol Horvitz, David Janos, James Harnrick, Keith Waddington and Theodore Fleming for their constructive comments regarding my research. Their editing of this manuscript has improved it significantly, and for this 1 am most grateful. 1 would like to give special thanks to my dissertation advisor, Theodore Fleming, for introducing me to the fascinating world ofbats, cacti, and deserts. Dr. Stephen Buchmann assisted my analysis ofpollen deposition at the Carl Hayden Agricultural Research Station in Tucson, Arizona. Dr. Jim Harnrick allowed me to work in his genetics laboratory at the University of Georgia. My research in Peru would not have been possible without the assistance and friendship of several people. Victor Pacheco of the University of San Marcos Natural History Museum in Lima first suggested that 1 visit Arequipa. Once in Arequipa, Professor Percy Jimenez ofthe Universidad Nacional de San Agustin gave me important logistic advice and introduced me to Luis Baraybar, a biology student who would become my field assistant. Luis provided excellent full time assistance during the course of my field research. His knowledge regarding plants and animal s at the study site as well as his dry sense ofhumor contributed greatly to this study. Joachim Ugarte and Jose E stefanero , also biology students, assisted me in the field at the very outset ofthe study, when help was most needed. Eleana Linares Perea offered me advice on where to find columnar cacti in Arequipa, and patiently taught me how to identify many plant species at the study site. Horacio Zeballos provided me with important data on rainfall for Arequipa. While 1 was away from my study site and in Arequipa, many people made me feel iv as ifl was at home. Blenda Femenias and Pablo Vera de la Cruz offered me not only their friendshíp, but gave me an archeological and anthropological perspective of Arequipa and its surroundings. Yolanda Hurtado and her family treated me as one of their own. I thank Luis Baraybar' s family: Mr. and Mrs. Baraybar, his two sisters, Marta and Mayu, and their assorted pets, for good food, drink, and laughter on weekends. During my stay in Mexico, I was fortunate to be able stay at the Prescott School's field station at Bahía Kino. Tad Pfister moved out of his house and into a very small, uncornfortable trailer so that I could stay in cornfort at his home. I am most grateful to Tad for introducing me to the Sea of Cortez, where I finally got to see great whales. Vinicio Soza, although busy with his own field work, never hesitated to lend me a helping hand ifl needed one (and I often did). Whíle I learned the techniques of electrophoresis at the "Hamrick lab", John Nason, Victoria Aspit, Elizabeth Stacey, and Mary Hagin made me feel welcome and were extremely patient with the many questions I had regarding the alien world of isozymes and gels. I willleave the University ofMiarni with many fond memories. My experience as a graduate student here was enhanced by the opportunity to interact with many people too numerous to name here. In particular, I would like to give special thanks to Mindy Nelson, Karen Lips, Devon Graham, Maureen Donelly, and Roger Gunther, who at various times throughout my graduate career have sat patiently through practice talks and read many drafts of proposals and manuscripts. Dr. Jay Savage not only taught me about biogeography, but also about how to relax and have a good time when a day' s work was done. Dave Janos, in addition to editing my manuscripts, has offered me advice and discussion on many aspects regarding my career interests. I feel very fortunate to have v had the opportunity to participate in his tropical biology field course, otherwise known as "the hike". It added a bit of adventure to rny life, to say the least. Finally, 1 rnust gratefully acknowledge the best support group one could ever hope for: rny farnily. My graduate education has been a farnily affair in every sense ofthe word. During rny graduate educatíon, rny parents Theodore and Teresa, rny sister Carol, rny brother Teddy, and rny aunt Mery, have offered me unfailing support and love. Financial support carne frorn the New York Zoological Society/The Wildlife Conservation Society, the World Wildlife Fund-Garden Club of America Scholarship in Tropical Botany, Bat Conservation Internatíonal, the University ofMiarni, and Theodore and Teresa Sahley. vi TABLE OF CONTENTS CHAPTER 1: Introduction........................................................................... 1 CHAPTER 2: The natural history and population status ofthe nectar feeding bat, Platalina genovensium (PhylIostomidae: Glossophaginae) in southwestern Peru 12 CHAPTER 3: PolIination biology and breeding system of a columnar cactus in southwestern Peru 39 CHAPTER 4: Vertebrate poIlination of the organ pipe cactus, Stenocereus thurberi: Effects on polIen deposition, polIen dispersal, and fruit production 78 vii CHAPTERONE INTRODUCTION The Cactaceae comprise a diverse and ecologically important New World plant family. Approximately 1500 species and 100 genera are distributed throughout all major arid and semi-arid ecosystems of North, Central, and South America (Barthlott & Hunt 1993, Gibson & NobeI1986). Despite their prominence in New World habitats, astonishingly little is known about the basic ecology of these plants. Cacti probably playa critical role in maintaining biodiversity of desert habitats because they provide food and shelter for a variety of desert animals including insects, lizards, birds, and mammals. Because of their tolerance to prolonged dry spells, some species of cacti probably are the only plants to consistently provide nectar, pollen and fruit for animal s during droughts; thus, they may be considered "keystone species" in New World desert habitats ( sensu Terborgh 1986, Gilbert 1980). Humans harvest cacti and/or cactus products for a variety ofpurposes. Cactus fruit (especially that ofthe genus Opuntia) is commercially valuable in both North and South America, and many indigenous communities harvest fruits of other species for local consumption