The Feeding Biology of Tintinnid Protozoa and Some
Total Page:16
File Type:pdf, Size:1020Kb
THE FEEDING BIOLOGY OF TINTINNID PROTOZOA AND SOME OTHER INSHORE MICROZOOPLANKTON by David John Blackbourn A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in the Department of Zoology and Institute of Oceanography We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA September, 1974 > In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of ^OoCoG Y The University of British Columbia Vancouver 8, Canada . Date i ABSTRACT Tintinnids are among the largest and most abundant of the marine cillate microzooplankton but there is very little published information on their feeding rates and abilities. The feeding of Tintinnopsis subacuta (and to a lesser extent, that of 12 other species) was investigated with three methods 1) direct observation 2) counts of accumulated food cells and 3) Coulter Counts of the particles in the experimental medium. There was reasonable quali• tative agreement between the results obtained by the three methods but quantitative agreement was poor. Many of the results showed no signi• ficant differences due to very great variability in the results for a single tintinnid species within and between experiments. Much of this variability may be due to the methods used but it also reflects the vari• ability of tintinnids in natural populations. A wide variety of items was eaten by tintinnids, including smaller tintinnids; and the maximum food size can be related to tintinnid cell volume over a wide range but is dissimilar in tintinnid species of similar cell size. Several tintinnid species showed differential pre• dation on various types of laboratory phytoplankton. This differential predation was based upon the ability of the predator to handle prey, or on prey size or prey type depending upon the particular tintinnid species. 'Negative' selection of some types of laboratory phytoplankton in mixed- prey samples was also shown for some tintinnid species, particularly Tintinnopsis subacuta on members of the Cryptophyceae. Feeding rates measured with the accumulation method were equivalent ii to 0.65% ml/hr/tintinnid for T_. subacuta and usually much less. Feeding rates for this species measured with the Coulter Counter technique ranged from 0.33 to 3.8% ml/hr/tintinnid. Very little feeding was observed directly but feeding rates estimated with this method were somewhat higher than those estimated for the same species from accumulation experiments. Tintinnids apparently both consumed, and caused the production of particles during experiments. Correlations between feeding rate and 9 other experimental variables were such that it would be impossible to predict the feeding rate of a tintinnid species using only the size dis• tribution of avaifeble particulate biomass of less than 20 um diameter. There were large differences between the apparent feeding rate asymptotes of T_. subacuta and _S_. ventricosa as measured with the Coulter Counter and the accumulation method. The latter method gave lower asymptotes than did the former. Ivlev electivity indices for T_. subacuta were most consistently positive in those middle Coulter size classes which also showed the greatest growth in controls. Increased temperature had little effect on the rate of food accumu• lation by four tintinnid species, but there was some evidence of a faster rate of disappearance of ingested food at very high temperatures. The relationship between the gain of new food and the loss of old food in in• dividual T. subacuta and Stenosomella ventricosa was highly variable and may strongly reflect the physiological history of the cell. The rate of gain of new food may be largely independent of the amount of old food in a tintinnid, but the average rate of loss of old food is faster in cells given new food than in starved cells. iii It was shown that natural concentrations of T_. sub acuta can apparently control the growth of natural populations of phytoplankton of less than 20 um dia. in under 24 hours. From a comparison with some other types of microzooplankton it was concluded that the larger species of tintinnid could probably have a potentially predominant effect upon the highly pro• ductive phytoplankton of less than 10 um diameter in English Bay and other coastal localities. iv TABLE OF CONTENTS Page ABSTRACT i TABLE OF CONTENTS iv LIST OF TABLES vi LIST OF FIGURES ix ACKNOWLEDGEMENTS xi 1. INTRODUCTION 1 2. TINTINNID BIOLOGY . 5 3. MATERIALS AND METHODS a) Sampling and initial treatment 32 b) Experimental methods (i) General comments 35 (ii) Counts of accumulated food 38 (iii) Observations of feeding behaviour 39 (iv) Coulter Counter experiments 41 Glossary 49 4. RESULTS AND DISCUSSION a) Accumulation experiments (i) Qualitative results 51 (ii) Quantitative results 60 b) Observations of tintinnid motions and feeding behaviour 117 c) The effect of microzooplankton on natural and laboratory phytoplankton populations (Coulter Counter experiments) 127 5. GENERAL DISCUSSION 163 Table of Contents (Cont'd) Page 6. SUMMARY 179 REFERENCES 182 APPENDICES 187 vi LIST OF TABLES Page TABLE 1, List of tintinnid species and their measurements. 52 TABLE 2. Food eaten by microzooplankton. 56 TABLE 3. Eutintinnus tubulosus feeding on Isochrysis 62 galbana at two concentrations. TABLE 4. Eutintinnus tubulosus feeding on Monochrysis- 62 lutheri at two concentrations. TABLE 5. Eutintinnus tubulosus feeding on Monochrysis 64 lutheri at three concentrations. TABLE 6. Tintinnopsis parvula feeding on Monochrysis 64 lutheri at three concentrations. TABLE 7. Tintinnopsis subacuta feeding on Dunaliella 66 tertiolecta at three temperatures and three food levels. TABLE 8. Tintinnopsis parvula and Tintinnopsis 68 cylindrica feeding on Monochrysis lutheri in dim light and in darkness. TABLE 9. Eutintinnus tubulosus and Helicostomella 69 kiliensis feeding on Monochrysis lutheri at three concentrations TABLE 10. Various tintinnid species feeding on 'new' 71 and 'old' cultures of Dunaliella tertiolecta at four concentrations. TABLE 11. Tintinnopsis parvula feeding on Isoselmis ssp. 73 and Monochrysis lutheri. TABLE 12. Tintinnopsis subacuta feeding on Etitreptiella 73 sp. and Isochrysis galbana. List of Tables (cont'd) Tintinnopsis subacuta feeding on Eutreptiella sp.,Isochrysis galbana and Dunaliella tertiolecta. Tintinnopsis subacuta and Tintinnidium mucicola feeding on Eutreptiella sp. and Isoselmis sp. Various tintinnid species feeding on Monochrysis lutheri and Dunaliella tertiolecta. Tintinnopsis subacuta and Stenosomella ventricosa feeding on Eutreptiella sp., Monochrysis lutheri and Isoselmis sp. singly and in combination. Tintinnopsis subacuta (etc.) starved for various periods in filtered seawater feeding on Dunaliella tertiolecta at unknown, but dense, concentrations. Tintinnopsis subacuta and other predators starved for various periods and feeding on Eutreptiella sp. Loss rate of Stenosomella nivalis at two levels of dilution of medium with filtered seawater. Change of food contents of -T-intinnidium mucicola with time at four levels of dilution o of medium with filtered seawater. Tintinnopsis subacuta, T_. parvula, _T. rapa and Tintinnidium mucicola feeding on new food - Monochrysis lutheri and Cryptomonas sp., and loss rate of old food of various types at four temperatures. viii List of Tables (cont'd) Page TABLE 22. Feeding and loss rates of Tintinnopsis 99 subacuta; losing Monochrysis lutheri and Plagioselmis sp. and either starved or gaining Eutreptiella sp. and Isoselmis sp. TABLE 23. Accumulation and loss rates of Tintinnopsis 101 cylindrica, Helicostomella kiliensis, Tintinnidium mucicola and Eutintinnus latus, feeding oh Monochrysis lutheri or Isoselmis sp., or starved; and losing M. lutheri, Isoselmis sp. or Dunaliella tertiolecta. TABLE 23A. Summary of accumulation experiments with 103A Tintinnopsis subacuta. TABLE 24. The relationship between tintinnid cell 115 length and number of accumulated food items in two species taken from different experiments. TABLE 25. The effect of immobilization by sonication 121 on the successful ingestion of algal flagel• lates by the tintinnid, Eutintinnus latus. TABLE 26. Observed contact rates of various tintinnid 124 species on natural and laboratory food items. TABLE 27. Multiple correlation coefficients from Coulter 129 Counter experiments. TABLE 28. Microzooplankton lower threshold feeding values 136 and regression coefficients of Logmean E (variable 3) when food consumption rate (variable 1) is zero. TABLE 29. Results of Coulter Counter experiments with 159 Synchaeta littoralis and Synchaeta sp. eating Dunaliella tertiolecta. TABLE 30. Approximate relative sizes and feeding rates 177 of various types of marine microzooplankton. ix LIST OF FIGURES Page FIGURE 1. Diagram of Favella sp. (modified from 7 Campbell, 1927). FIGURE 2. Possible theoretical relationships be- 15 tween tintinnid lorica length and frequency. FIGURE 3. Lorica length-frequency data for Tintinnopsis 18 subacuta from three successive field samples. FIGURE 4. Seasonal abundance and lorica lengths of 100