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Download The MICROHABITAT SELECTION AND REGIONAL COEXISTENCE IN WATER-STRIDERS (HETEROPTERA: GERRIDAE) by JOHN RICHARD SPENCE B.A., Washington and Jefferson College, 1970 M.S., University of Vermont, 1974 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE FACULTY OF GRADUATE STUDIES (Department of Zoology) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA January, 1979 (c) John Richard Spence 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 Zool°gy The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date January 30, 1979 ii ABSTRACT This study considers the natural history and ecology of water-strider species occurring on the Fraser Plateau of south- central British Columbia. The overall aim was to assess the effects of spatial heterogeneity on factors controlling the distribution and relative abundance of gerrid species. The relationships among temperature, population dynamics and habitat use were investigated. From a regional perspective, spatial heterogeneity allows species population dynamics to converge in time while keeping them separate in space. , Laboratory rearing studies were used to calculate physiological time-scales for developmental processes. Patterns of mating behaviour, fecundity and fertility are described for Serris comatus and G. pinqreensis in the laboratory and G- incoqnitus in the field. Egg laying and juvenile growth are shown to be strongly temperature dependent in all species studied. Temperature thresholds for development differ, both among species, and often among stages of a particular species. Low thresholds recorded for G. pinqreensis can lead to significant growth advantages for this species during early spring. Instar differences seem to be adapted to seasonal temperature regimes experienced by gerrids. Gerris species and instars showed distinct optimum temperatures for survival. These optima vary with developmental thresholds. It is suggested that species may be best adapted for growth under different temperature regimes. A method was developed for estimating absolute densities in the field from relative abundance measures using linear regression techniques. Gerrid size and presence or absence of vegetation markedly affect capture rates. No effect of species or type of emergent cover was demonstrated. Availability for capture varies with leg^length in G. buenoi and G. pinqreensis. This relationship is used to estimate availability constants for other water-strider species. Field surveys between 1975 and 1977 established that G. buenoi, G. comatus and G. pinqreensis were the most abundant water-strider species in- the study area. Each of these was strongly associated with a single type of vegetation in the field; G. buenoi with grass/sedge habitats, G. comatus with floating vegetation and G. pinqreensis with bulrush habitat. Limnoporus dissortis and L. notabilis were commonly encountered on small, temporary ponds, G. incognitus was first taken during 1976 in the study area and small populations are confined to brushy, well-shaded habitats. G. buenoi, G. comatus and G. pinqreensis are all potentially bivoltine in the study area; Limnoporus spp. are univoltine. Generation timing varies tremendously among lakes and periods of maximum abundance for each species are not separated in time. Strong between-lake habitat associations in the field result proximately from habitat fidelity at the time of spring colonization. The tendency of gerrids to overwinter near the iv mother pond and trial and error habitat selection during spring dispersal enforce habitat fidelity during colonization. Species distributions within lakes are affected by habitat availability. Habitat preference experiments demonstrate that G. pinqreensis and G. comatus have active preferences for emergent cover and open habitats respectively. G. buenoi is a habitat generalist but its distribution can be affected by a tendency to avoid other species. Smaller stages of each species are found close to shore and often in areas of dense emergent vegetation. Enclosure experiments demonstrated that G. pinqreensis can exclude G. buenoi and G. comatus from bulrush habitats, which are most favorable for the growth and development of all species. Fiftgainsh wheinstan confiner G. dbueno in ithei anr dcharacteristi G. comatus cshowe habitatsd greates. tHabitat weigh-t specific differences in foraging efficiency among late instars may help produce the habitat associations observed for these two species. Fifth instar G. pinqreensis showed poor survival when enclosed in freshwater habitats, suggesting the hypothesis that its distribution is restricted by the presence of surface-feeding predators other than water-striders. It is suggested that competition for space, predation, density-independent mortality and colonization dynamics all interact on the template of spatial heterogeneity to produce regional patterns of distribution and abundance. V TABLE OF CONTENTS ABSTRACT . ii TABLE OF CONTENTS ............. V LIST OF TABLES xi LIST OF FIGURES xiv ACKNOWLEDGEMENTS X.vii CHAPTER I. INTRODUCTION 1 A. Coexistence of species ... A plot 1 B. Water-striders . The cast 4 1. Natural history background 4 2. Geographical distributions 9 3. The problem and the approach 11 C. The study area . The stage ,. 13 1. Sites . 13 i 2. Weather 20 3. Lakes 21 CHAPTER II. THE EFFECTS OF TEMPERATURE ON WATER-STRIDER GROWTH AND DEVELOPMENT 31 INTRODUCTION • —• 31 METHODS AND MATERIALS ...... 33 A. Egg production 33 1. Effects of temperature 33 2. Mating behaviour and fecundity in the laboratory 35 3. Fecundity in the field ....................... 37 B. Growth and development 38 1. Development and temperature 38 vi a. Eggs .............. ... 38 b. Larvae 38 2. Growth thresholds ... 39 RESULTS ....... 4 0 A. Egg production 40 1. Effects of temperature 40 2. Mating behaviour and fecundity in the laboratory . 48 3. Fecundity in the field 55 B. Growth and development 59 1. Development and temperature .................. 59 2. Growth thresholds ............................ 64 DISCUSSION 69 A. Mating behaviour • • 69 B. Fecundity 70 C. Growth and Development 74 CHAPTER III. DENSITY ESTIMATES FOR GERRIDS 77 INTRODUCTION . .......... 77 METHODS 79 A. Seasons, species, and habitats 79 B. Relative abundance estimates 80 C. Absolute abundance estimates 84 D. Test estimates 86 RESULTS 87 A. Differences between seasons and size classes .... 87 B. Differences between habitats • 96 C. Differences between species 96 D. Test estimates 99 vii DISCUSSION . 100 CHAPTER IV. COMPARATIVE ECOLOGY OF WATER-STRIDERS ON THE FRASER PLATEAU OF BRITISH COLUMBIA 107 INTRODUCTION 107 MATERIALS AND METHODS 109 A. Lakes and species studied - - 109 1. Field temperatures 109 2. Egg production and alary morphism 113 3. Population dynamics 113 4. Habitats ..... .......... 115 RESULTS . 117 A. Field temperatures .117 B. Egg production and alary morphism ............... 120 C. Population dynamics ................ 125 D. Habitats 140 1. Vegetation .. ...... .. .. ... 140 2. Surface conductivity .....143 3. Lake permanence 145 DISCUSSION .... 148 A. Life cycles and population dynamics 148 B. Comparative ecology ....152 1. Habitats and timing .......................... 152 2. Habitat permanence and adaptive strategies ... 153 3. Habitat and regional coexistence ............. 155 a. Gerrids in British Columbia 156 b. Gerrids in eastern and western North America 158 4. Conclusions .........160 viii CHAPTER V. EXPERIHENTAL ANALYSIS OF MICROHABITAT SELECTION IN WATER-STRIDERS 161 INTRODUCTION 161 MATERIALS AND METHODS ...........163 A. Dispersal ... Habitat selection among lakes ... 163 B. Habitat selection within lakes 167 1. Field distributions ......... — 167 a. Habitat differences among species ........... 167 b. Habitat differences within specie's.. 167 2. Laboratory experiments . Responses to artificial habitat structure ...... 168 a. Laboratory conditions and apparatus ......... 168 b. Species tendencies to enter complex habitats 169 c. Selection of artificial habitat mimics ...... 181 d. Habitat structure and foraging success ...... 181 RESULTS ..' , ' 187 A. Dispersal . Habitat selection between lakes . 187 1. Immigration 187 2. Emigration ........... 189 B. Habitat selection within lakes 195 1. Field distributions . 195 a. Habitat differences among species ........... 195 b. Habitat differences within species .......... 197 2. Laboratory experiments . .Responses to artificial structure 202 a. Species tendencies to enter complex habitats 202 b. Selection of habitat mimics ................. 210 c. Habitat structure and foraging success ...... 216 ix DISCUSSION ............. ............. 219 A. Dispersal 219 B. Habitat selection within lakes 222 1. Species differences 222 2. Instar differences 226 3. Species morphology and habitat structure 227 CHAPTER VI. PERSISTENCE, POPULATION PERFORMANCE AND INTERSPECIFIC COMPETITION ... 229 INTRODUCTION •• --• •-- 229 MATERIALS AND METHODS 231 A. Colonization, persistence and population success
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