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Mechanisms Underlying the Effects of Marine Herbivores Implications for a Low Intertidal Kelp Community

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Mechanisms Underlying the Effects of Marine Herbivores Implications for a Low Intertidal Kelp Community MECHANISMS UNDERLYING THE EFFECTS OF MARINE HERBIVORES IMPLICATIONS FOR A LOW INTERTIDAL KELP COMMUNITY by RUSSELL W. MARKEL B.Sc, The University of British Columbia, 1992 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of B otany) We accept this thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA October 1996 © Russell W. Markel, 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 t5T~-rcVlO>v/ The University of British Columbia Vancouver, Canada Date }Q)bQ>- I I'H & DE-6 (2/88) MECHANISMS UNDERLYING THE EFFECTS OF MARINE HERBIVORES: IMPLICATIONS FOR A LOW INTERTIDAL KELP COMMUNITY ABSTRACT Rocky low intertidal communities structured by the canopy forming kelp Hedophyllum sessile and the generalist herbivore Katharina tunicata are typical of semi- exposed coasts of British Columbia. Katharina tunicata is capable of removing the community supporting canopy formed by H. sessile . Mechanisms which determine and mediate the interaction between this alga and herbivore are investigated. Densities of Katharina tunicata were manipulated within the range of densities found on the shore, without the use of artificial barriers, and the percent cover, density, population structure and demography of Hedophyllum sessile were monitored to determine the effects of these manipulations. Differential susceptibility of H. sessile holdfast size classes are shown to account for the rapid decline of H. sessile abundance in areas of high chiton density. These data are used to quantify the mechanism referred to as 'an escape in size.' Several lines of evidence suggest that the mechanism of K. tunicata interaction with H. sessile adults is an indirect effect whereby holdfast integrity is degraded, making individuals of H. sessile more susceptible to wave-induced mortality. The implications of this mechanism for observed geographic variation in K. tunicata interaction strength are discussed. The potential role of polyphenolics (anti-herbivore secondary metabolites) in the interaction between Hedophyllum sessile and Katharina tunicata is examined. Three experiments determined that: (1) H. sessile is a phenolic-rich species (5-7% H. sessile dry weight), (2) induction of phenolic production does not occur within three days of simulated wounding, (3) individuals of H. sessile from areas of naturally high and low herbivore densities do not differ in phenolic content, and (4) juvenile H. sessile tissue contains significantly more phenolics than adult vegetative tissue but adult reproductive and ii vegetative tissues do not differ. These results are discussed in terms of both present-day and evolutionary ecological importance. By monitoring the Hedophyllum sessile understory community during the chiton density manipulation experiment, and performing descriptive investigations during the same time period, several direct and indirect interactions were revealed which are potentially important determinants of the structure of this community. These interactions are used to construct three interaction webs which distinguish between the interactions experienced by juvenile and adult H. sessile. A model of the annual successional trajectory of this low intertidal kelp community is used to summarize the findings of this thesis. iii TABLE OF CONTENTS Page ABSTRACT ii LIST OF FIGURES v LIST OF TABLES vii ACKNOWLEDGEMENTS ix CHAPTER 1: General introduction to algal/herbivore interactions , 1 CHAPTER 2: Effects of herbivore density on the demography and population structure of an intertidal kelp, Hedophyllum sessile (C. Agardh) Setchell 9 CHAPTER 3: Polyphenols content of the intertidal kelp Hedophyllum sessile: present-day implications and evolutionary frameworks • 49 CHAPTER 4: Direct and indirect interactions regulating a low intertidal kelp community: implications for juveniles vs. adult kelps 67 CHAPTER 5: General conclusions and summary: A successional model of the mechanisms and processes regulating a low intertidal kelp community 103 iv LIST OF FIGURES Page Figure 2.1 Katharina tunicata density (#/m2) within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date. Sampling date abbreviations are: JL95 (July 1995), S95 (September 1995), 095(October 1995), D95 (December 1995), F96 (February 1996), A96 (April 1996), JU96 (June 1996), JL96 (July 1996) 30 Figure 2.2 Hedophyllum sessUe percent cover within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 31 Figure 2.3 Hedophyllum sessile density within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 32 Figure 2.4 Hedophyllum sessile blade number / individualwithin low, control and high chiton density manipulation areas. Mean blade number / individual was averaged over each 25 x 25 cm quadrat. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 33 Figure 2.5 Hedophyllum sessile blade length / individualwithin low, control and high chiton density manipulation areas. Mean blade length / individual was averaged over each 25 x 25 cm quadrat. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 34 Figure 2.6 Blade length (a) and blade number (b) of Hedophyllum sessile individuals of the 1-4 cm maximum holdfast diameter size class within low, control and high chiton density manipulation area. Mean blade number and length / individual were averaged over each 25 x 25 cm quadrat. Data are means ± SE OF 1-4 cm size-class individuals pooled over each chiton density manipulation on each sampling date 35 Figure 2.7 Hedophyllum sessile recruitment densities (#/m") within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 36 Figure 2.8 Comparison of survivorship curves of the July 1995 cohort of Hedophyllum sessile (a) <1 cm (b) 1-4 cm and (c) >4 cm holdfast size classes of as a function of low, control and high chiton density 37 Figure 2.9 Results of the Kaplan-Meier survival analysis of the Hedophyllum sessile <1 cm, 1-4 cm and >4 cm maximum holdfast diameter size classes as a function of low, control and high chiton densities. Data are mean survival time (weeks) ±95% confidence intervals 38 Figure 2.10 Comparison of survivorship curves of the July 1995 cohort of Hedophyllum sessile within (a) low, (b) control and (c) high Katharina tunicata density treatment areas as a function of holdfast size 39 Figure 3.1 Phenolic content of Hedophyllum sessile wounded and control blades 1, 2, and 3 days after wounding. Data are mean + SE of 14 replicates for days 1 and 3 and of 6 replicates for day 2 65 v Figure 4.1 Articulated coralline algae percent understory cover within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 86 Figure 4.2 Crustose coralline algae percent understory cover within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 87 Figure 4.3 Bare rock percent understory cover within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 88 Figure 4.4 Fugitive algae percent understory cover within low, control and high chiton density manipulation areas. Data are the mean ± SE of 6 replicates for density manipulations at each sampling date 89 Figure 4.5 February 1996 comparison of Hedophyllum sessile percent cover, density, holdfast diameter and blade length between the high and mid H. sessile zones. Letters represent the transition from the high H. sessile zone (A) to the mid H. sessile zone (D) 90 Figure 4.6 Comparison of interaction webs involving juvenile (A) and adult (B) Hedophyllum sessile 91 Figure 4.7 Types of indirections occuring between the chiton Katharina tunicata, adult and juvenile Hedophyllum sessile, and articulated (AC) and encrusting (EC) coralline algae 92 Figure 4.8 Direct and indirect interactions occuring between juvenile and adult Hedophyllum sessile, articulated (AC) and encrusting (EC) coralline algae, Katharina tunicata (Chiton) and wave force 93 Figure 5.1 Summary model of annual successional trajectory of the temperate low intertidal community dominated by the kelp Hedophyllum sessile and the chiton Katharina tunicata 112 vi LIST OF TABLES Page Table 2.1 Results of ANOVA comparing Katharina tunicata densities between chiton density treatment areas at each sampling date 40 Table 2.2 Results of ANOVA on the treatment effect of Katharina tunicata density on Hedophyllum sessile percent cover 41 Table 2.3 Results of ANOVA on the treatment effect of Katharina tunicata density on Hedophyllum sessile density
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