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Effect on the Feeding Behaviour of White-Bellied Sunbirds, Cinnyris Talatala (Nectariniidae)

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Effect on the Feeding Behaviour of White-Bellied Sunbirds, Cinnyris Talatala (Nectariniidae) The feeding response of white-bellied sunbirds (Cinnyris (Nectarinia) talatala) to sugar concentration and viscosity of artificial nectar Eckart Stolle By Carolina D. C. Leseigneur Supervised by Prof. S. W. Nicolson Submitted in partial fulfilment of the requirements for the degree MSc Zoology in the Faculty of Natural and Agricultural Science University of Pretoria Pretoria February 2008 © University of Pretoria MSc Candidate Miss Carolina D. C Leseigneur Department of Zoology & Entomology University of Pretoria Pretoria 0002 South Africa [email protected] [email protected] Student no.: 20077582 Carolina Leseigneur One must ask children and birds how cherries and strawberries taste. - J. W. Von Goethe Supervisor Prof. S. W. Nicolson Department of Zoology & Entomology University of Pretoria Pretoria 0002 South Africa [email protected] 2 Acknowledgements I would firstly like to thank my supervisor, Prof. Sue W. Nicolson, without her guidance and support this thesis would certainly have taken me a lot longer to put together. I have enjoyed learning from you and your sincerity and encouragement was much appreciated. Next I would like to thank all my colleagues and friends, who gave me some valuable comments in the preparation of this thesis, and who were a wonderful support group throughout this time. A special word of thanks goes to Luke Verburgt, for all his assistance and statistical help in the work of the third chapter of this thesis. Building circuit boards, soldering cables and programming computers late into the night with a beer was both enjoyable and productive. My fellow “sunbirders”, Craig Symes, Cromwell Purchase and Angela Köhler, and also Dr. Hannelie Human and Dr. Christian Pirk, your inputs were very helpful, and the general conversations to take our minds off work were very fun. To my special friends in and outside the department, Anne Dolinschek, Rosie van Zyl, Lia Rotherham, Katarina Medger, René Wolmerans, Michelle Fourie, Tracy Shaw, Eckart Stolle, Mike Ellis, Cornel Du Toit and Bernard Coetzee, thanks for being there when I just needed to get away from it all. Your kindness, love, laughter and general enthusiasm has been great to share in. Special thanks to Eckart Stolle for the cover photo of the white- bellied sunbird. Most importantly, I would like to thank my family (Mom, Dad, Ivan and our feathered and four-footed family members). Your love, support and helping me stay sane during these years has been fantastic. Never saying no to anything I asked of you and encouraging me throughout is truly appreciated. No bird soars too high if he soars with his own wings. - William Blake 3 General Abstract Plant nectar is a simple food and is easily digested by many different species of pollinators. Many compounds make up the composition of floral nectars, but the most abundant are sugars, generally dominated by sucrose and the hexoses, glucose and fructose. Nectar sugars have been measured for many plant species visited by hummingbirds, sunbirds and other passerines, revealing a range of concentrations. The nectars of passerine- pollinated flowers are generally dilute compared to those of bee-pollinated flowers. The question why bird nectars are so dilute has been addressed in many studies. Many hypotheses have been proposed, among them the relationship between viscosity and drinking by birds. The viscosity of sugar solutions increases exponentially with increasing concentration, and capillarity is inversely proportional to viscosity. Nectarivorous birds imbibe nectar by capillarity, and high sugar concentrations could impose constraints on their feeding efficiency. Feeding in nectarivorous birds, especially hummingbirds, has been mostly devoted to assessing sugar type preferences. However, concentration preferences have received less attention, and the effect of viscosity on feeding has not been examined separately from sugar concentration for any bird species. Do nectarivorous birds show a preference for specific concentrations at a broad and a fine scale of difference, given a specific sugar type? Does viscosity impose a feeding limitation on nectarivorous birds? Does it affect their feeding behaviour? Sunbirds and other nectar-feeding birds can choose amongst various flowering plant species at any one time. Their feeding responses may have important consequences on pollination ecology. In this study, concentration preferences of white-bellied sunbirds were examined using paired solutions of either sucrose or equicaloric 1:1 mixtures of glucose and fructose, at a 4 both a broad and a fine scale of difference between pairs over the concentration range of 0.25 to 2.5 M. I hypothesized that sunbirds would prefer concentrations of 1 M and higher on sucrose solutions, while preferring concentrations less than 1 M on hexose solutions. On both sugar types at the broad scale, the higher concentration was significantly preferred up to 1 M, suggesting a preference for 1 M sugar solutions. At a finer scale, white-bellied sunbirds were able to discriminate 0.03 and 0.05 M (1 and 2% w/w) concentration differences between sucrose and hexose solutions respectively. This discrimination is similar to that reported at low concentrations for other passerine nectar-feeders, and at higher concentrations for hummingbirds. To determine if high viscosity nectars limit the sugar intake of avian nectar consumers, white-bellied sunbirds were exposed to three different test series of sucrose solutions: control series (CS, pure sucrose 0.25 – 2.5 M), constant viscosity series (CVS, 0.25 – 0.7 M with increased viscosity equivalent to that of 1 M sucrose) and constant concentration series (CCS, 1 M with increased viscosities equivalent to that of 1.5, 2 and 2.5 M sucrose). Viscosities were artificially altered with Tylose ®. The sunbirds had reduced intake rates and gained less energy on more viscous sucrose solutions. Also, sunbirds did not alter their feeding behaviour (feeding frequency, feeding duration, total feeding duration and feeding interval) in any significant way when feeding on more viscous sucrose solutions. This lack of change in feeding behaviour led to lower sugar intake rates and sugar consumption. These results suggest that sunbirds suffer a preingestional limitation when consuming nectars with viscosities higher than those due to sugar concentration alone. 5 TABLE OF CONTENTS Acknowledgements................................................................................................................ 3 General Abstract ................................................................................................................... 4 Disclaimer............................................................................................................................... 7 Chapter 1................................................................................................................................8 General Introduction..................................................................................................................... 8 The white-bellied sunbird........................................................................................................................... 8 The nectars of bird-pollinated flowers ....................................................................................................... 9 Sugar type preferences of bird pollinators............................................................................................... 10 Concentration preferences of bird pollinators......................................................................................... 13 Discrimination abilities of nectarivorous birds ....................................................................................... 15 Viscosity of sugar solutions and effects on preingestional processing..................................................... 16 Aim of study.............................................................................................................................................. 19 Chapter 2.............................................................................................................................. 29 Nectar concentration preferences of the white-bellied sunbird, Cinnyris talatala (Nectariniidae).............................................................................................................................. 29 ABSTRACT............................................................................................................................................ 30 Introduction............................................................................................................................................ 31 Materials & Methods ............................................................................................................................. 35 Results ..................................................................................................................................................... 39 Discussion................................................................................................................................................ 48 Chapter 3.............................................................................................................................. 59 The viscosity of artificial nectar: effect on the feeding behaviour of white-bellied sunbirds, Cinnyris talatala (Nectariniidae) ................................................................................................
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