Diversity of Litter Spiders (Araneae) in a Successional Douglas-Fir Forest in British Columbia
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DIVERSITY OF LITTER SPIDERS (ARANEAE) IN A SUCCESSIONAL DOUGLAS-FIR FOREST IN BRITISH COLUMBIA by LISA J. BRUMWELL B.A., The University of British Columbia, 1994. A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (DEPARTMENT OF ZOOLOGY) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA October 1996 ©Lisa J. Brumwell, 1996 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 or her representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department of .Zooi-p The University of British Columbia Vancouver, Canada Date DE-6 (2/88) ABSTRACT Litter spiders were collected at two locations by pitfall trapping: Victoria Watershed South, and Koksilah, located on south-eastern Vancouver Island, British Columbia. Sites were in Douglas-fir forest (in the Coastal Western Hemlock region) and contained four forest successional stages: Regeneration (3-8 years), Immature (25-45 years), Mature (65-85 years), and Old-Growth (>200 years). Thirty-two species were collected in families selected to represent litter spiders. Several of the collected species are listed as potentially rare and endangered in British Columbia. The collection of Zora hespera is the first record of the family Zoridae in Canada. Intraspecific comparisons revealed three distinct distribution patterns: generalists, regeneration specialists, and intermediate forest specialists (spiders prefering the Immature and Mature successional stages). These patterns can be related to biological information regarding individual species. Species richness measures and diversity indices indicated that the regeneration sites have the greatest species richness. There was no significant difference between the remaining three successional stages. Factors affecting the distribution of spider species in the forest litter are discussed. TABLE OF CONTENTS Page ABSTRACT ii TABLE OF CONTENTS Hi LIST OF TABLES v LIST OF FIGURES vi ACKNOWLEDGMENTS x INTRODUCTION 1 MATERIALS AND METHODS Study Area 8 Traps and Trap Placement 9 S orting and Identification 12 Methods of Data Analysis Species Abundance Plotting 14 Intraspecific Comparisons 15 Diversity Indices 16 Similarity Coefficients and Cluster Analysis 16 RESULTS Number of Species 17 Species Abundance Models 17 Intraspecific Comparisons 17 Generalists 22 Regeneration Specialists 22 Intermediate Forest Specialists 30 Unresolved 30 Forest Successional Stage Comparisons Diversity Indices 36 Similarity Coefficients and Cluster Analysis 38 TABLE OF CONTENTS: Continued DISCUSSION Pitfall Trapping Methodology 41 Species Abundance Models 42 Intraspecific Comparisons 43 Generalists 43 Regeneration Specialists 45 Intermediate Forest Specialists 50 Unresolved Species 51 Uncommon Species 51 Forest Successional Stage Comparisons Diversity Indices 52 Similarity Coefficients and Cluster Analysis 54 Rare and Endangered Species 54 GENERAL DISCUSSION Factors Affecting Litter Spider Distribution 56 Dispersal Ability 56 Habitat Conditions 57 Structural Components 57 Climatic Components 59 Nutritional Components 60 SUMMARY 63 LITERATURE CITED 65 APPENDIX A 72 APPENDIX B 73 LIST OF TABLES Table 1. Pitfall trap collection dates. Table 2. List of spider families selected for specimen identification to species. Table 3. Total species collected - arranged according to distribution patterns. Table 4. Number of species, corrected number of species E(S)i67, with SD, and the Shannon- Wiener index (FT) for Victoria Watershed and Roksilah at all successional stages. Table 5. Number of species, and the Shannon-Wiener index (FT) comparing Victoria Watershed and Koksilah. Table 6. Matrix of the Simplified Morisita-Horn Index of Similarity Coefficients for Victoria Watershed and Koksilah at all successional stages. LIST OF FIGURES Figure 1. Locations of the forest chronosequences (referred to as Victoria Watershed South and Koksilah) where the ecological studies took place. Figure 2. Example of the plot layout and subplot assignments provided by the Canadian Forest Service. Circled areas numbered 1-8 were used for spider collection. Figure 3a. Species rank/abundance graph of the regeneration stage of Victoria Watershed and Koksilah site. Abundance of each species is plotted on a logarithmic scale against the species rank, ordered from most abundant to least abundant species. Figure 3b. Species rank/abundance graph of the immature stage of Victoria Watershed and Koksilah site. Abundance of each species is plotted on a logarithmic scale against the species rank, ordered from most abundant to least abundant species. Figure 3c. Species rank/abundance graph of the mature stage of Victoria Watershed and Koksilah site. Abundance of each species is plotted on a logarithmic scale against the species rank, ordered from most abundant to least abundant species. Figure 3d. Species rank/abundance graph of the old-growth stage of Victoria Watershed and Koksilah site. Abundance of each species is plotted on a logarithmic scale against the species' rank, ordered from most abundant to least abundant species. Figure 4a. Mean number of individuals (with standard error) of Antrodiaetus pacificus (Simon) trapped in each successional stage at the Victoria Watershed site. The Kruskal- Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 4b. Mean number of individuals (with standard error) of Antrodiaetus pacificus (Simon) trapped in each successional stage at the Koksilah site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 5a. Mean number of individuals (with standard error) of Cicurina simplex Simon trapped in each successional stage at the Victoria Watershed site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. vi Figure 5b. Mean number of individuals (with standard error) of Cicurina simplex Simon trapped in each successional stage at the Koksilah site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non- parametric multiple comparison test. Figure 6a. Mean number of individuals (with standard error) of Alopecosa kochii (Keyserling) trapped in each successional stage at the Victoria Watershed site. The Kruskal- Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 6b. Mean number of individuals (with standard error) of Alopecosa kochii (Keyserling) trapped in each successional stage at the Koksilah site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 7a. Mean number of individuals (with standard error) of Pardosa wyuta Gertsch trapped in each successional stage at the Victoria Watershed site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 7b. Mean number of individuals (with standard error) of Pardosa wyuta Gertsch trapped in each successional stage at the Koksilah site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non- parametric multiple comparison test. Figure 8. Mean number of individuals (with standard error) of Pardosa vancouveri Emerton trapped in each successional stage at the Victoria Watershed site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line, as determined by a non-parametric multiple comparison test. Figure 9a. Mean number of individuals (with standard error) of Castianeira longipalpa (Hentz) trapped in each successional stage at the Victoria Watershed site. The Kruskal-Wallis test statistic, H, and the statistical probability, P, are the result of intersuccessional stage comparisons. Stages which did not differ from one another are joined with a line,