M I N G and Swimming Behaviour of Bythotmphes Cedemtmemi
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Mingand swimming behaviour of Bythotmphes cedemtmemi Schoedler James Robert Muirhead A thesis submitted in conformity with the requirernents for the degree of Master of Science Graduate Department of Zoology University of Toronto O Copyright by James Robert Muirhead, 1999 National Library Bibliothèque nationale 1+1 ,cana, du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. NO Wdtingtm OnawaON KlAûN4 OPEawaON KIAM Canada Canada The autbor has granted a non- L'auteur a accordé une licence non exclusive Licence allowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, distribute or sell reproduire, prêter, distribuer ou copies of this thesis in microfonn, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/nIm, de reproduction sur papier ou sur format électronique. The author retains ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantial extracts fiom it Ni la thèse ni des extraits substantiels may be printed or otherwise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Feeding and swimming behaviour of Bythoîrephes cedersimemi Schoedler Master of Science, 1999 James Robert Muirhead Graduate Department of Zoology University of Toronto Abstract Predatory zooplankton detect prey with mechanoreception, visual cues or chemoreception. The objectives were to quantify swimming behaviour of Bytbnephes cederstroemi under the influence of prey, prey kairomones, and light and to mode1 Bythotrephes' encounter rate with prcy f?om Harp Lake. 1 used video capture and motion tracking to rneasure Bythrrephes swimming behaviour with Daphnia, Daphnia kairomnes and Polyphemus as prey. 1 also measured Bythotrephes' reaction distance to prey under different light levels. Bythoirephes swam fastest and in a more directeci fashion in the presence of light and prey than in treatments with Light or prey only. Also, Bythotrephes' reaction distance increased in higher light levels and thus is likely to use vision as a detection rnechanism Results fkom the encounter mode1 show that small, slow-moving prey faced the greatest risk fkom Byrhomephes. Acknowledgements 1 sincerely thank my supervisior, Dr. W. Gary Sprules, for his advice and support when 1 rnost needed it, and for his encouragement throughout. I also thank my CO-supervisor, Dr. Rob Baker, for helpful advice on animai behaviour. Special thanks go to Dr. Charles Rarncharan who provided me with the motion-tracking and analysis software he had personnaiiy written. Additional thanks go to Dr, Hank Vandcrploeg at the Great Lakes Environmental Research Labs in Ann Arbr, Michigan for the use of his video setup for critical stages of my experiments; Dr. Nom Yan and Bob Guard at the Ministry of Energy and the Environment (Dorset) for providing equipment and data on a moment's notice and Cristina Dumitru and Giliian Morgan for providing Bythutrephes abundance and diet preference data. Thanks also go to Sam Genova for his help with field collections of Bythotrephes. The easy-going, fnendly atmosphere and personal support in the lab would not be possible without the presence of Dr. Stuart Whipple (Uncle Stu) and Agnes Blukacz (Rzapka). Stuart: We taiked about issues arising from my thesis into the wee hours of the rnorning more often than 1 can remember. Agnes: 1 depended the rnost on Agnes while we were coliecting zooplankton fiom Harp Lake. I don't think either one of us is going to forget the trouble we went through to bring biick îive Bythotrephes. To Stuart and Agnes, 1 thank for completing experiments critical to my thesis; without their help I probably would not be able to finish. Findy, 1 thank my famiiy Keith, Ga& Jackie and Karen who were always in my thoughts while 1 was away working on my Master's; I dedicate my thesis to them iii Table of Contents Page Abstract ii Acknowledgements iii List of Tables v List of Figures vi List of Appendices vii Generai Introduction 1 Chapter 1 3 Inmductio n 4 Materials and Methods 7 Results 20 Discussion 30 References 38 Chapter 2 42 Introduction 43 Materials and Methods 45 Results 58 Discussion 76 References 83 General Discussion and Conclusions 86 References 88 Appendices 90 References 96 Page Table 1. Second-order mean vector r and swimming velocity 23 with bootstrapped standard emrs. Table 2. MANOVA summary tables for Daphnia kairomone and Light experiments. Table 3. Mean of bootstrapped Sb for second-order angles, eigenvalues of ma& T and probabilities. Table 1. Average prey swimming speeds. Table 2. Prey risk factor (percentage in Bythotrephes diet) of species and lifestages. Table 9. Interpretation of spherical distribution of directions based 94 on eigenvalues and eigenveçtors of matix T. Page Figure 1. Video setup Figure 2. Frequency histogram of swUnming behaviours 12 Figure 3. Second-order analysis of angles 18 Figure 4. Bythotrephes swïmming tracks under different treatrnents 21-22 Figure 5. Prey versus light interaction plot 27 Figure 1. Reaction distance Nming chamber 46 Figure 2. Rdonsphere for Byrhomphes 50 Figure 3. Circular distributions of Bythtrephes apparent reaction 59 distances under different light levels. Figure 4. Relationship between Light level and reaction distance 61 for a third instar Bythotrephes. Figure 5. Enco unter rate, to ta1 Bythotrephes-calanoid CO pepodid 62-64 encounters and prey risk. Figure 6. Average encounter rates for Byrhotrephes and associated 66-68 prey risk at 0-10 m depth over 24 hows. Figure 7. Encounter rate and prey risk wnsitivity responses to a 20% 70 change in surîàce Light levels. Figure 8. Encounter rate and prey risk sensitivity responses to a 20% 71 change in Byrhotrephes swimming velocit y. Figure 9. Encounter rate and prey risk sensitivity responses to a 20% 73 change in Dophnia swimming velocity. Figure 10. Mean number and standard error of encounters of cyclopoid 74 nauplii for a single third instar Bythotrephes. Figure 11. Mean number and standard emr of total cyclopoid naupüi 75 and Bythotrephes encounters. Page Appendix 1. Description of motion aacking program 90 Appendix 11. Circulat and spherical gcarnctry and analysis 91 Appendix III. Effect of sample rate on Bythoîrephes swimming 95 parame ters. vii General Introduction One of the rnost ofien asked and pressing questions about invading species is what are some of the fùnctional characteristics of the species that permit a successful invasion. From individual case studies and literature surveys, many ecologists have proposed that successful invading species typically have r-selected traits, high dispersal rates. high genetic variabity. phenotypic plasticity, and polyphagy (Lodge 1993; Johnson and Carlton 1996; Rejmanek and Richardson 19%). Although many successful invading species share cornmon traits, it is not possible to predict with certainty the invasive potencial of individual species in target comrnunities (Burke and Grime 19%). Predictive studies, according to Lodge (1993), require focused, quantitative studies on pYtrular invaders and comrnunity characteristics. Bythoîrephes cederstroerni Schoedler (Onychopoda, Cercopagidae) is a large-bodied carnivorous hshwater moplankter with an elongated caudal spine that invaded the Great Lakes in the early-1980's kom Lake Lagoda, CIS (Bur et aL 1986; Evans 1988; Sprules et aL 1990) and has since spread to several inland lakes (Hall and Yan 1997). Bythotrephes' invasion success can be partly attributed to adaptations such as the barbed caudal spine which provides some protection against gape-lunited small fish (Branstrator and Lehrnan 1996). fast pmhenogenic reproduction and the production of resting eggs which serves as a source for restockïng the population after the winter (Rivier 1998). Adaptations for a predatory lifestyle include a large, media1 compound eye, fast swimming, reduced carapace and dedicated thoracic limbs used in prey capture. Despite the interest shown in Bythotrephes as an invasive species. there has been litt le work on Bythotrep hes' predatory be haviour. Predatory zooplankton typicaily search for prey by detecting hydrornechanical signals produced by their prey's swimming (Kerfoot 1978; Rice 1988). In addition, iike the predatory zooplankton Mysis relicfa (Ramcharan and Sprules 1986), Leprodora kindrii (Wolken and Gallik 1965) and Polyphemus pediculus (Odselius and Niisson 1983), Bythoh-ephes' compound eye is likely able to form images, in Chapter 1,I examine how Bythotrephes detects prey by rneaswing changes in Bythotrephes swimming be haviour under various experimental conditions. By using vision, Bythotrephes is able to detect prey at a greater distance in the presence of visible light than by mec hanoreception or direct enco unter and thus increase its foraging efficiency. In Chapter 2,I measure the distance at whic h Bythotrephes reacts to prey under different light Ievels. 1 then model the encounter rate of Byrhotrephes with dflerent prey fÏom Harp Lake based on this reaction distance and swimming speeds fiom Chapter 1. B y incorporating a diunial light cycle and Lght attenuation in the water column into the model, 1 am able to identify spatial and temporal patterns of maximum risk faced by prey as well as qualify the relative rïsk among zooplankton species in Harp Lake. Predation pressures exened by invading species such