AVAILABLE ENERGY AND SPECIES DIVERSITY: THEORY AND EXPERIMENTS WITH BEES (COMMUNITIES, FLOWERS, FORAGING). Item Type text; Dissertation-Reproduction (electronic) Authors WRIGHT, DAVID HAMILTON. Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 26/09/2021 08:46:34 Link to Item http://hdl.handle.net/10150/187808 INFORMATION TO USERS This reproduction was made from a copy of a document sent to us for microfilming. While the most advanced technology has been used to photograph and reproduce this document, the quality of the reproduction is heavily dependent upon the quality of the material submitted. 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MI48106 8500480 Wright, David Hamilton AVAILABLE ENERGY AND SPECIES DIVERSITY: THEORY AND EXPERIMENTS WITH BEES The University of Arizona PH.D. 1984 University Microfilms International 300 N. Zeeb Road, Ann Arbor, MI48106 AVAILABLE ENERGY AND SPECIES DIVERSITY: THEORY AND EXPERIMENTS WITH BEES by David Hamilton Wright A Dissertation Submitted to the Faculty·of the DEPARTMENT OF ECOLOGY AND EVOLUTIONARY BIOLOGY In Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY In the Graduate College THE UNIVERSITY OF ARIZONA 1 9 8 ~ THE UNIVERSITY OF ARIZONA GRADUATE COLLEGE As members of the Final Examination Co~nittee, we certify that we have read the dissertation prepared by ~p~a~v~j~d~H~awm~l~'lwt~Q~n~W~r~j~g~h~t _______________________ entitled Available Energy and Species Diversity: Theory and Experiments with Bees and recommend that it be accepted as fulfilling the dissertation requirement for the Degree of ___D~o~c~t~o~r~o~f~Ph~l~'l~o~s~o~p~h~y ________________ ~ ____________ __ i/fod1 Date 7 20 ,?y Date A:r-£ } ?-o ~~ \et&' L.( Date <..52:..~ ____ ..-/ Date / /~~2ht-~ ~t:';d7:b~ Date Final approval and acceptance of this dissertation is contingent upon the candidate's submission of the final copy of the dissertation to the Graduate College. I hereby certify that I have read this dissertation prepared under my direction and recommend that it be accepted as fulfilling the dissertation requirement. //D,i:ifserta tion Director Date t~ STATEMENT BY AUTHOR This dissertation has been submitted in partial fulfillment of requirements for an advanced degree at The University of Arizona, and is deposited in the University Library to be made available to borrowers under the rules of the Library. Brief quotations from this dissertation are allowable without special permission, provided that accurate acknowledgement of the source is made. Requests for permission for extended quotation from or reproduction of this manuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in his or her judgement the proposed use of the material is in the interests of scholarship. In all other instances, however, permission must be obtained from the author. SIGNED, /kM~ ~ To my parents iii ACKNOWLEDGEMENTS In the course of the work represented by this dissertation, I have been provided with a wide variety of kinds of essential assistance from a large number of people, to whom I would like to express my deep appreciation: James H. Brown, Astrid Kodric-Brown, Michael L. Rosenzweig, Robert L. Smith, David Vleck, Dav:ld W. Inouye, George E. Bohart, William A. Calder III, Svata M. Louda, Dorothy Johnson, John C. Johnson Jr., Michael Zimmerman, Kathryn L. Iverson, Mary Price, Nikolas M. Waser, Shari Uhl, Michael A. Bowers, L. Susan Anderson, Brian A. Maurer, Erik Van Hoogenstyn, Mari Jensen, Eric Larsen, Thomas Gibson, David F. Gori, Richard H. Thomas, Mike Michael, John Pleasants, and Bernd Heinrich. The staff and researchers of the Rocky Mountain Biological Laboratory were very helpful and made field work pleasant and productive. Many thanks to all. This research was financed in part by grants from Sigma Xi, The Scientific Research Society, and from the University of Arizona Graduate Student Program Development Fund. I am grateful for this support. iv TABLE OF CONTENTS Page ACKNOWLEDGEMENTS ••••••••••••••••••••••••••••••••••••••• iv LIST OF TABLES •• ....................................... viii LIS T 0 F FIG U RES. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •• ix A B S T RA CT. • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • x GENERAL INTRODUCTION................................... 1 CHAPTER 1 • SPECIES-ENERGY THEORY: AN EXTENSION OF SPECIES-AREA THEORy •••••••••••••• 4 Summary •••••• • •••••••••••••••••••••••• 8 ••••••••••• 4 Introduction. · ................ ......... 5 The meaning of available energy in species- energy theory ••••••••••••••••••••••••••••••• 8 The effect of available energy in the equili- brium model, and species-energy curves •••••••••••• 9 Testing species-energy theory ••••• ................ 16 Discussion ••• ·................................... 24 Species occurrence and abundance. ............ 24 Patterns in species diversity •••••••••••••••• 26 General theories of diversity •••••••••••••••• 31 v vi TABLE OF CONTENTS, continued CHAPTER 2. AVAILABLE NECTAR AND BUMBLEBEE FORAGING PROFITABILITy ••••••••••••••••••••••••••• 33 Summary •••••• · ................................... 33 Introduction. · ................. .............. 34 Materials and methods. · ........ 36 Study sites •••••• · .......................... 36 Nectar availability. ......................... 36 Observation colonies. · .................... 37 Foraging profitability. • ••••••••• 0 ••••••••••• 39 Results ..•......•........ .. .. .. .. .. .. 41 Nectar availability. · .................... 41 Foraging profitability....................... 44 Discussion •••••••••••••••••• · .................... 54 CHAPTER 3. PATCH DYNAMICS OF A FORAGING ASSEMBLAGE OF BEES.· •••••••••••••••••••••••••••••••• 63 Summary •••••• ·................................... 63 Introduction. · ................................... 64 The arrival-departure ra te mode 1. •••••••••• It •••••• 67 Population s iz e ••• . .. .. 67 Species occurrence and number of species present •••••••••• · ...................... 70 Materials and methods. · .......................... 71 vii TABLE OF CONTENTS, Chapter 3 continued Results........................................... 74 Effect of enrichment on assemblages visiting !.El.!. 0 b .!J!~ and f.!.~~~!.~. • . • • • • • . ..• . • • • • . • • • • 74 Effect of differentially available resources on response to enrichment: Experiments with !'££!!.l~:!!!. and .Q~.!..E.!:!1!!..!J!:!!!. • • • • • • • • • • • • • • • • • • • • • • 81 Me c han ism s a f f e c tin g de par t u r era t e.. ••• • • ••• 85 Discussion......... •••••••••••••• •••••••••••• 91 Species-level heterogeneity in resource availability and use.................... 91 Island analogies.......................... 92 The species-level approach to community composition ••... 0 •••••••••••••••••••••••• 94 Energy and community structure............... 95 CONCLUSION. •• •• • • • • • • • • • • • • • •• • • • • • • • • • • • • • • • •• • • • • • • • • 97 APPENDIX A: Bombus appositus nectar loads ••••••••••••• 104 APPENDIX B: Bombus appositus foraging trip durations ..........•.•.•..•..... 106 REFERENCES ••••••••••••••••••••••••••••••••••••••••••••• 113 LIST OF TABLES Page Table 1.1: Is1ands, island characteristics, and species numbers of angiosperms and land and freshwater birds ••••••••••••••••••••••• 18 Table 1.2: Rarity and patchy distribution of zooplankton................................ 30 Table 2.1: 1982 Flower density changes and utiliza- tion by Bombus appositus