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Calochortus)From Southern British Columbia Demographic perspectives on the rarity and persistence of two mariposa lilies (Calochortus)from southern British Columbia M<chael Miller B.A., Queen's uhversity at Kingston, 1989 A Dissertation Submitted in Partial Fulfilment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY in the Department of Biology O Michael Miller, 2004 University of Victoria All right reserved. This dissertation may not be reproduced in whole or in part, by photocopying or other means, without the permission of the author. Abstract Co-supervisors: Dr. Geraldine Allen, Dr. Joe Antos The dynamics of peripheral populations provide insight into range limits and rarity. I studied sympatric populations of two species of mariposa lilies with contrasting distributions: Calochortus lyallii, a regional endemic of central Washington and southern British Columbia, and C. macrocarpus, a more widespread taxon. Marked plants were monitored for five years in the Okanagan highlands, at the northern range limit for C. Zyallii and near the elevational limit for C. macrocarpus. Life table data were used to generate stage-classified matrix population models for three populations of each species and, for C. lyallii, different microsites and plant densities. My objectives were to evaluate the demographic variation of C. lyallii among populations, microsite types and density classes, and to identify demographic differences between C. lyallii and C. macrocarpus that might contribute to their differing occurrence patterns. Annual population growth rates (lambdas) for C. lyallii ranged from 0.89 to 1.07 among populations, 0.87 to 1.29 among microsite types, and 0.86 to 1.07 among density classes. Life table response experiment analyses showed that inter-population and inter-microsite variances in lambda resulted mostly from variance in large adult fecundity, whereas the variance in lambda among density classes was mostly due to variance in vegetative stasis. Although differences were not significant, lambda tended to be highest in high density plots, arguing against a density-dependent equilibrium. Stochastic projections yielded long-term growth rates of near 1 for C. lyallii, whereas two of three C. macrocarpus populations were projected to decline rapidly in size. In both species, prolonged bulb dormancy was common (lasting up to 4 yr) and helped buffer population fluctuations. The relatively higher lambda of C. lyallii resulted primarily fiom higher flowering fi-equency. On average, C. macrocarpus had higher fruit set and more seeds per capsule than C. lyallii, but experienced higher fruit predation and had lower seedling establishment. Seedlings of C. lyallii were predicted to live longer, flower sooner, flower more frequently, and leave more offspring than C. macrocarpus seedlings. I conclude that differences in the local distribution and abundance between the two species can largely be explained by subtle differences in life history and demography. Table of Contents Title Page ....................................................................................................................... .i. Abstract ........................................................................................................................... ii ........................................................................................................... Table of Contents iv.. List of Tables ................................................................................................................ vii List of Figures .............................................................................................................. ix.. Acknowledgements ....................................................................................................... xi1 Dedication .................................................................................................................... xiv Chapter 1: General Introduction .................................................................................. 1 Chapter 2: Dormancy and flowering in two mariposa lilies (Calochorlus)with contrasting distribution patterns: implications for monitoring ................................. 10 Introduction ............................................................................................................... 10 Methods .................................................................................................................... 12 STUDY SPECIES AND HABITAT .................................................................................. 12 FELD METHODS ...................................................................................................... 15 DORMANCY DETERMINATIONS ................................................................................. 17 SELF COMPATIBILITY ............................................................................................... 17 VARIATION IN FLOWERING AND FRUITING ................................................................ 18 REPRODUCTION IN RELATION TO CLIMATIC FACTORS ................................................ 18 Results ................................................................................................................. 19 DORMANCY ............................................................................................................ 19 SELF COMPATIBILITY ............................................................................................... 21 VARIATION IN FLOWERTNG AND FRUIT SET ............................................................... 24 FLOWERING AND FRUITING IN RELATIONSHIP TO CLIMATE ........................................ 27 FATES OF BUDS, FLOWERS, AND FRUITS ................................................................... 31 Discussion ............................................................................................................... 31 DORMANCY ............................................................................................................ 31 REPRODUCTION....................................................................................................... 35 CONCLUSION .......................................................................................................... 37 Chapter 3: Effects of environmental heterogeneity on the demography of Calochortus lyallii (Liliaceae). a bulbous perennial with prolonged dormancy ........ 38 Introduction ............................................................................................................... 38 Methods ................................................................................................................... 41 THE SPECIES AND STUDY AREA ................................................................................ 41 FIELD SAMPLING ................................................................................................... 42 DELINEATION OF PLOT GROUPS ................................................................................ 44 STAGE CLASSIFICATION........................................................................................... 46 MA- CONSTRUCTION AND ANALYSIS................................................................... 50 SIGNEICANCE TESTING AND CONFIDENCE INTERVALS .............................................. 53 LIFE TABLE RESPONSE EXPERIMENT (LTRE) ANALYSIS .............................................. 54 Results ....................................................................................................................... 56 HABITAT-ABUNDANCE RELATIONSE-IIPS .................................................................... 56 GERMWATION......................................................................................................... 56 TRANSITION PROBABILITIES..................................................................................... 56 POPULATION PROJECTIONS ................................................................................... 68 POPULATION STRUCTURE AND STABLE STAGE DISTRIBUTIONS................................... 70 REPRODUCTIVE VALUES ................................................................................... 75 LTRE ANALYSIS ....................................................................................................... 76 ELASTICITY ANALYSIS ............................................................................................. 80 Discussion ................................................................................................................. 82 VARIATION IN TRANSITION MATRICES AND IN h ........................................................ 82 VARIATION IN STABLE STAGE DISTRIBUTIONS........................................................... 86 PROLONGED BULB DORMANCY ................................................................................ 88 HABITAT RELATIONSHIPS ......................................................................................... 89 ELASTICITY ANALYSIS ........................................................................................... 91 RELEVANCE TO SPATIAL MODELS OF DYNAMICS ....................................................... 92 Chapter 4: Comparative demography of two sympatric species of Calochortus (mariposa lily) with contrasting distribution patterns ............................................... 95 Introduction ............................................................................................................... 95 Methods .................................................................................................................
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