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The Life Cycle of the Pteropod Limacina Helicina in Rivers Inlet (British Columbia, Canada)

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The Life Cycle of the Pteropod Limacina Helicina in Rivers Inlet (British Columbia, Canada) The Life Cycle of the pteropod Limacina helicina in Rivers Inlet (British Columbia, Canada) by Kang Wang B.Sc., The University of British Columbia, 2009 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF Master of Science in The Faculty of Graduate and Postdoctoral Studies (Oceanography) THE UNIVERSITY OF BRITISH COLUMBIA (Vancouver) April 2014 c Kang Wang, 2014 Abstract The life cycle of Limacina helicina has been continuously debated within the literature. We believe the current lack of consensus regarding fundamental aspects of its life cycle (e.g. seasonal times of spawning, seasonal development of the population size structure, as well as the life cycle longevity) is primarily due to using datasets of low temporal resolution. Using fort-nightly data, two population cohorts were identified using the mixdist statistical package and tracked for more than 400 days, throughout 2008 to 2010. From this, a life cycle longevity of 1.2{1.5 years was estimated for L. helicina in Rivers Inlet. Throughout the seasons, the population size struc- ture showed a continually high presence of the smaller size-groups suggesting continuous spawning, however, based on total densities of > 600 ind.m−3, the late spring was put forward as the period of peak spawning. Continuous spawning was confirmed with the use of daily data. Identifi- cation of a summer peak spawning established late spring and summer as two periods of enhanced spawning, although continuous spawning occurred through- out the season (in a limited fashion). Short-term periods of significant growth were observed prior to peak spawning in late spring and summer. This was not directly coupled with chlorophyll concentrations, possibly due to the time lag between periods of high chlorophyll biomass and zooplankton response. At- ii tempts were made to estimate the instantaneous mortality of L. helicina, and the seasonal changes experienced from spring to summer. Our estimates were complicated by a combination of 1.) inherent patchiness of L. helicina, 2.) ad- vection, and 3.) merged recruits. Generally, there were no cases of significant mortality throughout the seasons however, short term mortality was observed after peak spawning. It is plausible that the smallest size-groups of L. helicina experiences the highest mortality after peak spawning. Our findings show that in Rivers Inlet, L. helicina has a life cycle spanning 1{1.5 years with spring and summer peak spawning activities. The spring cohort is likely spawned by the summer cohort from the previous year. It utilizes the spring phytoplankton bloom to reach sexual maturity and spawn the summer cohort. iii Preface This thesis is ultimately a product of the collaborative effort of many in- vestigators from the University of British Columbia, Point Grey campus, of the inter-disciplinary RIES - Rivers Inlet Ecosystem Study conducted from 2008 to 2010 (riversinlet.eos.ubc.ca/People.html). The aims of this thesis are separate and not directly connected with the main objectives of the RIES. Additional winter sampling from 2010{2011 was performed by Wayne Jacobson with direc- tion from Brian Hunt and the Hakai Institute. Results from other studies in the RIES were used throughout Chapters 2 and 3 to help rationalize our findings. Formalin preserved zooplankton samples for the bi-weekly and monthly sam- pling in Chapter 2, as well as the daily zooplankton samples used in Chapter 3, were processed by myself. Prior to being considered for this thesis, the daily zooplankton samples in Chapter 3 was used by an undergraduate student as the basis of a directed studies project, guided by Evgeny Pakhomov. The cohort analysis in Chapters 2 and 3, performed using the mixdist sta- tistical package (for the R statistical programming language), was conducted by myself with guidance from Evgeny Pakhomov and Brian Hunt. Complicated de- tails concerning the use of mixdist were clarified through email communication with the author, Peter MacDonald. iv Table of Contents Abstract .................................... ii Preface ..................................... iv Table of Contents .............................. v List of Tables ................................. ix List of Figures ................................ xi Acknowledgements ............................. xiii Dedication ................................... xv 1 An Introductory Review ........................ 1 1.1 What Are Pteropods . 1 1.2 Taxonomy . 2 1.3 Thecosome Morphology . 3 1.4 Swimming and Buoyancy Regulation . 4 1.5 Food and Diet . 5 1.6 Reproductive Biology . 6 1.7 Ecological Role, Ecological Threats, and Knowledge Gaps . 7 1.7.1 Ocean Acidification . 7 1.7.2 Life Cycle of L. helicina { Knowledge Gaps . 8 v 2 Life-Cycle Dynamics of Limacina helicina in Rivers Inlet B.C. 10 2.1 Introduction . 10 2.1.1 Research Aims . 10 2.1.2 Limacina helicina: Past Life Cycle Investigations . 11 2.1.3 Rivers Inlet: Historical Context . 13 2.1.4 Goals and Aims . 14 2.2 Methods . 15 2.2.1 Study Area { Rivers Inlet . 15 2.2.2 Seawater Temperature, Fluorescence, Salinity . 15 2.2.3 Sample Collection and Selection . 16 2.2.4 Sample Preparation and Enumeration . 17 2.2.5 Size Frequency Histograms and Identification of Cohorts . 18 2.2.6 Estimation of Growth and Life-Cycle Longevity . 19 2.3 Results . 25 2.3.1 Environmental Parameters - Temperature, Salinity, Fluo- rescence . 25 2.3.2 L. helicina Abundance: Seasonal and Inter-annual Variation 25 2.3.3 L. helicina Size Structure: Seasonal Development . 26 2.3.4 Spawning Activity . 27 2.3.5 Estimate of Life-Cycle Longevity . 28 2.3.6 Rivers Inlet Spatial Analysis . 31 2.4 Discussion: Life-cycle re-evaluation . 37 2.4.1 Seasonal Spawning and Recruitment . 37 2.4.2 Seasonal Growth and Environmental Correlations . 38 2.4.3 Rivers Inlet Spatial Analysis . 40 2.4.4 Potential Sampling Errors . 40 3 Seasonal Growth and Mortality { Spring vs. Summer ..... 42 vi 3.1 Introduction . 42 3.2 Goals . 44 3.3 Methods . 46 3.3.1 Study Area & Sample Collection . 46 3.3.2 Daily Fluorescence . 46 3.3.3 Size-Frequency Histograms & Identification of Cohorts . 47 3.3.4 Spawning Events . 48 3.3.5 Shell Size Growth and Mortality . 48 3.4 Results . 51 3.4.1 Daily Chlorophyll . 51 3.4.2 Daily Population Abundance . 51 3.4.3 Daily Population Size-Structure . 52 3.4.4 Spawning Events . 52 3.4.5 Cohorts Identified and Tracked . 53 3.4.6 Seasonal Growth . 54 3.4.7 Daily Mortality . 54 3.5 Discussion . 59 3.5.1 Spawning, Cohorts, and Size-Structure Development: Com- parison to Chapter 2 and Relevant Literature . 59 3.5.2 Caveats to Estimating Daily Growth . 62 3.5.3 Estimating Daily Mortality and Problems Encountered . 63 3.5.4 Potential Sampling Errors . 65 4 Life Cycle of L. helicina: A Conceptual Model and General Conclusions ................................ 69 4.1 A Conceptual Model . 71 4.2 General Conclusions . 74 vii Bibliography ................................. 77 Appendices .................................. 84 A Chapter 2: Supplementary Data ................... 85 A.1 2010{2011 Winter-Transition . 85 A.2 Seasonal Correlations - Physical Parameters & Population Abun- dance . 87 B Chapter 3: Supplementary Data ................... 90 B.1 Daily Data Sampling Dates . 90 B.2 Size-Frequency Histograms - March, April, May, June, July . 93 B.3 Finite Mixture Distributions { Daily Data . 98 B.4 What Are Finite Mixture Distributions . 98 B.5 Fitting Finite Mixtures . 99 B.6 Finite Mixture Distributions { Statistical Output . 109 B.7 Life Tables for Cohorts Tracked . 115 B.8 Seaonal Growth Rate . 135 B.9 Environmental Connection . 137 B.10 Seasonal Mortality . 139 viii List of Tables 2.1 Limacina shell size summary statistics (max, min, mean) for L. helicina individuals enumerated in the fort-nightly and monthly samples at station DFO 2 . 21 2.2 Survey sampling dates for the samples processed, for spatial anal- ysis of stations DFO 1, DFO 2, DFO 3, DFO 4, and DFO 5 in Rivers Inlet . 23 2.3 A life cycle table for the cohorts C1 and C2, tracked from March 2008 to July 2010. 29 A.1 Linear regressions testing the relation between 30 m depth aver- aged temperature and salinity, and 30 m depth averaged fluores- cence { 2008 season . 87 A.2 Linear regressions testing the relation between 30 m depth aver- aged temperature and salinity, and 30 m depth averaged fluores- cence { 2009 season . 88 A.3 Linear regressions testing the relation between 30 m depth aver- aged temperature and salinity, and 30 m depth averaged fluores- cence { 2010 season . 89 A.4 Linear regressions testing the relation between 30 m depth aver- aged temperature and salinity, and 30 m depth averaged fluores- cence { 28 February to 2 June, 2009 . 89 ix B.1 Daily dates of sample collection . 90 B.3 Life Tables of Population components tracked . 116 B.4 Regression table for observed periods of increased shell growth in the population . 136 B.5 Regression table for observed periods of increased shell growth in the population . 137 B.6 Statistical results of linear regressions testing the relation be- tween chlorophyll and the daily variation in population abun- dance, for each month . 138 B.7 Statistical results of linear regressions testing for periods of in- creased mortality, for cohorts identified and followed in the daily time series . 139 x List of Figures 2.1 L. helicina with measure of shell diameter . 17 2.2 Map of west coast of British Columbia and of Rivers Inlet . 20 2.3 2-panel figure showing A. the seasonal distribution of 30 m depth- averaged salinity and 30 m depth averaged temperature for 2008, 2009, and 2010, and B. the seasonal distriburtion of L. helicina abundance throughout 2008, 2009, 2010, and the 2010{2011 win- ter. Also shown is the seasonal variation of 30 m depth integrated fluorescence . 32 2.4 Finite-mixture distributions showing the seasonal development of the L.
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