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The effects of acid mine drainage at Britannia Beach, B.C., on fucus gardneri and associated intertidal… Marsden, Allan Dale 1999

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T H E EFFECTS OF ACID MINE DRAINAGE AT BRITANNIA B E A C H , B.C., ON FUCUS GARDNER! AND ASSOCIATED INTERTIDAL A L G A E by A L L A N D A L E M A R S D E N B.Sc, Simon Fraser University 1997 A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF T H E REQUIREMENTS FOR T H E DEGREE OF MASTER OF SCIENCE in T H E F A C U L T Y OF G R A D U A T E STUDIES (Department of Botany) We accept this thesis as conforming to the required standard T H E UNIVERSITY OF BRITISH COLUMBIA April 1999 © Allan Dale Marsden 1999 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 The University of British Columbia Vancouver, Canada Date A^]{ ZCJJJJ DE-6 (2/88) Abstract Copper ore was extracted from Britannia Mine, British Columbia, Canada, from 1902 until the mine ceased operations in 1974. Rain, snowmelt and groundwater now percolate through the mine tunnels, producing an acidic solution of dissolved metals known as Acid Mine Drainage (AMD). A portion of the A M D from the mine flows into Britannia Creek, which in turn flows into Howe Sound, 50 km north of Vancouver, B.C. This study examined the effects of this effluent on the distribution of intertidal macroalgae with a focus on Fucus gardneri Silva, a seaweed which thrives 2 km from the mouth of Britannia Creek but is absent from the shore near the Creek. F. gardneri provides habitat and food for benthic invertebrates, which are a major food source for chum salmon fry and chinook salmon fry and smolts. Algal communities were quantified at 7 intertidal stations to the north and south of the mouth of Britannia Creek and at 6 similar stations at nearby Furry Creek, a reference site. Algal cover was almost non-existent within 200 m of the mouth of Britannia Creek, and F. gardneri was completely absent on the 600 m of shoreline south of Britannia Creek and on 1000 m of shoreline north of the same Creek. There was, however, a heavy cover of filamentous green algae, mainly Enteromorpha compressa (L.) Link, at sites 300 m and 700 m south and north of Britannia Creek, respectively, suggesting that these algae can better tolerate A M D . Experimental work consisted of transplanting F. gardneri-covered cobbles from a control site to Britannia Beach and monitoring the plants' growth, survivorship and copper content. In five experiments conducted from June 1997 to November 1998, plants moved to within 100 m of Britannia Creek generally had lower survivorships and lower growth rates than plants at the control site, as well as higher tissue copper concentrations. Survivorship and growth rates of plants moved to areas farther from Britannia Creek (300-700 m) were not significantly different from control plants. ii Table of Contents Abstract ii Table of Contents iii List of Tables v List of Figures vii Acknowledgments . x C H A P T E R 1. G E N E R A L INTRODUCTION 1 1.1. Motivation for Research ...1 1.2. Background Information 1 1.2.1. Acid Mine Drainage 1 1.2.2. Local Oceanography 5 1.2.3. Copper Chemistry 7 1.2.4. Interactions between Copper and Algae 11 1.2.5. Fucus gardneri 15 1.2.6. Statistical Methods 18 1.3. Research Hypotheses 21 C H A P T E R 2. DISTRIBUTION AND R E P R O D U C T I V E F U N C T I O N OF A L G A E N E A R BRITANNIA C R E E K IN R E L A T I O N T O ACID MINE D R A I N A G E 22 2.1. Introduction 22 2.2. Methods 23 2.2.1. Dissolved Copper in Water 23 2.2.2. Quantification of Fucus gardneri length and algal cover 24 2.2.3. Oocyte Release 26 2.2.4. Copper body burden in Fucus gardneri 27 2.2.5. Statistical analyses 28 2.3. Results 29 2.3.1. Dissolved copper in water 29 2.3.2. Fucus gardneri cover 31 2.3.3. Filamentous green algal cover 31 2.3.4. Fucus gardneri plant length ; 33 2.3.5. Oocyte release 35 2.3.6. Copper body burden in Fucus gardneri 38 2.4. Discussion 39 2.4.1. Dissolved copper in water 39 2.4.2. Fucus gardneri cover... 40 2.4.3. Filamentous green algal cover 42 2.4.4. Length of Fucus gardneri 46 iii 2.4.5. Oocyte release 47 2.4.6. Copper body burden in Fucus gardneri 48 2.4.7. Statistical Methods I 50 2.5. Conclusions ...51 CHAPTER 3. TRANSPLANTATION OF FUCUS GARDNERI TO BRITANNIA BEACH 53 3.1. Introduction.... 53 3.2. Methods 53 3.2.1. Transplant of Adult Fucus gardneri 53 3.2.2. Measurements 57 3.2.3. Copper body burden 57 3.2.4. Data manipulation and statistical analysis 58 3.3. Results 58 3.3.1. Percent Cover 59 3.3.2. Survivorship 62 3.3.3. Growth 65 3.3.4. Copper body burden 68 3.3.5. General observations 69 3.4. Discussion 70 3.4.1. Severe Effects: November 1997 and February 1998 70 3.4.2. No Detectable Effects: June and August 1998... 73 3.4.3. General Discussion of Transplant Experiments.... 76 3.4.4. Power Analysis 78 3.5. Conclusions 80 CHAPTER 4. GENERAL CONCLUSIONS 81 References 86 Appendix I — Raw Data from Chapter 2 94 Appendix II — Raw Data from Chapter 3 143 iv List of Tables Table 2.1. Statistical analysis of Fucus gardneri percent cover from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Stations BB-C, BB-D, BB-E and BB-F were omitted in all months due to complete lack of F. gardneri. Station BB-B was omitted in December only. SNK tests were conducted if the A N O V A detected significant differences. All stations within a set of square brackets comprise a homogeneous group. See section 2.2.5 for explanation of SNK grouping 33 Table 2.2. Statistical analysis of filamentous green algal cover from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping. 35 Table 2.3. Statistical analysis of Fucus gardneri plant length from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping 37 Table 2.4. Statistical analysis of oocyte release, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping 38 Table 2.5. Statistical analysis of copper body burden in Fucus gardneri. a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping 38 Table 2.6. Correlation analysis of mean green algal percent cover and Fucus gardneri percent cover at Britannia Beach and Furry Creek stations. Stations BB-D and BB-E were excluded due to the presumed severe influence of A M D on filamentous green algae, n = 9 stations for June and July, n = 11 stations for August 43 Table 2.7. Concentrations of copper in Fucus spp. and Ascophyllum spp., both fucoids. All 1 2 studies rinsed algae in seawater before analysis. From: Forsberg et al. 1988, Soderlund et al. 1988,3Pedersen 1984, 4Barreiro etal. 1993,5Bryan 1983, 6Foster 1976,7Riget etal 1995,8Ho 1984, 9Morris & Bale 1975,10Dunn etal 1992, "Current study 49 Table 3.1. Statistical analyses of percent cover results from all transplants. All analyses use one-tailed, paired Student's t-tests to compare percent cover of Fucus gardneri on rocks transplanted to Britannia Beach with that on rocks transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a - 0.05 in all tests. Significant differences are in bold type 62 v Table 3.2. Statistical analyses of survivorship results from all transplants. All analyses use one-tailed, paired Student's t-tests to compare survivorship of Fucus gardneri on rocks transplanted to Britannia Beach with that of individuals transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant differences are in bold type 65 Table 3.3. Statistical analyses of growth rate results from all transplants. All analyses use one-tailed Student's t-tests to compare growth rates of Fucus gardneri on rocks transplanted to Britannia Beach with that of individuals transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant differences are in bold type. Tests marked with * were conducted using unpaired t-tests. All other tests were paired t-tests. 67 Table 3.4. Statistical analyses of copper body burden from the November 1997 transplant. All analyses use one-tailed Student's t-tests to compare copper body burden of Fucus gardneri transplanted to Britannia Beach with that in plants transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant. a = 0.05 in all tests. Significant results are in bold type 69 Table 3.5. Power of the statistical analyses of growth rate measurements during the June 1998 transplant. Power values are calculated using equation 3.1, with s = 0.40 mm d"1 and the 5, n and a values shown. All results of the current experiments were analyzed with n = 10 and a = 0.05. Power analyses using other parameters are shown for comparison 79 vi L i s t of Figures Figure 1.1. Relative location of Britannia and Furry Creeks, mine features and geographical features mentioned in the text. Adapted from Price et al. (1995) and Grout et al. (1998) 3 Figure 1.2. Combined daily loadings of Cu, Zn and Cd from Britannia Creek and the submerged outfall into Howe Sound. Data from R. McCandless (unpublished data). Graph compiled by J. Grout 4 Figure 1.3. Dissolved copper concentrations in Howe Sound near the mouth of Britannia Creek from March 1994 to January 1995. Values are mean of 8 samples taken along a transect covering the entire mixing zone of Britannia Creek, within approximately 400 m of the Creek mouth. Error bars are the maximum and minimum values recorded at each time point. From Chretien (1997) 4 Figure 1.4. General surface circulation patterns in upper Howe Sound. Longer arrows indicate faster flow rates. From Thomson (1981) ; 7 Figure 2.1. Locations of stations where dissolved copper was measured. From Grout and Levings (in preparation). Map from Canadian Hydrographic Service Chart #3526 23 Figure 2.2. Locations of transect stations at Britannia Beach (left) and the corresponding control stations at Furry Creek (right). 1 cm - 350 m. From Canadian Hydrographic Service Chart #3526 .24 Figure 2.3. Concentration of dissolved copper in Howe Sound surface from April-June 1998. Zero values indicate concentrations below the detection limit of 5 pg Cu L" 1. Stations 7 and 8 comprised 3 samples each; error terms are the standard errors for these measurements. Al l other stations comprised only one sample per time point. Values for station 7 are reported as numbers to allow clear presentation of values at other stations, which were much lower. From Grout and Levings (in preparation) 30 Figure 2.4. Percent cover of Fucus gardneri as measured from June to December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Sampling was not conducted at sites B until August. Error bars are ± 1 standard error; n = 9 for all stations 32 Figure 2.5. Percent cover of filamentous green algae as measured from June to December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Sampling was not conducted at sites B until August. Error bars are + 1 standard error, n = 9 for all stations 34 Figure 2.6. Mean length oi Fucus gardneri as measured from June to December 1998. A, June 24-25; B July 21-23; C, August 17-19; D October 8-10, E, December 1-3. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no plant lengths are given. Sampling was not conducted at sites B until August. Error bars are ± 1 standard error, n varied from 5 to 9, averaging 7.9, at sites withF. gardneri 36 vn Figure 2.7. Oocyte release from Fucus gardneri as measured in October and December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no oocyte counts are given. In October, n = 3 at all sites; in December, n = 3 at BB-G and FC-G, n = 4 at FC-C, n = 5 at all other stations. All values are normalized to the surface area of the source plant material. ..37 Figure 2.8. Copper body burden in dry weight of Fucus gardneri collected August 1998. BB = Britannia Beach. FC = Furry Creek. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no plants were collected. Error bars are ± 1 standard error, n = 3 at all sites 38 Figure 3.1. Transplant sites at Britannia Beach (left) and matching sites at Furry Creek (right). See Figure 1.1 for locations of creeks in Howe Sound. 1 and 2: November 1997; 3: February 1998; 4 and 5: June 1998; 6 and 7: August 1998. Sites BB-1 and FC-1 were -50 m from the mouths of Britannia and Furry Creeks, respectively. Sites BB-2, BB-3, FC-2 and FC-3 were all -100 m from their respective creek mouths. 1 cm = 350 m. From Canadian Hydrographic Service Chart #3526 56 Figure 3.2. Percent cover of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 7 rocks 60 Figure 3.3. Percent cover of Fucus gardneri on rocks transplanted February 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 6 rocks 60 Figure 3.4. Percent cover of Fucus gardneri on rocks transplanted June 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 8 rocks except day 75, when n = 7 rocks at site 4 61 Figure 3.5. Percent cover of Fucus gardneri on rocks transplanted August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 8 rocks until day 35, n = 7 thereafter at site 6. All rocks but one lost at FC-6 on day 100 61 Figure 3.6. Survivorship of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 10 rocks 63 Figure 3.7. Survivorship of Fucus gardneri on rocks transplanted February 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 6 rocks 63 Figure 3.8. Survivorship of Fucus gardneri on rocks transplanted June 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks except day 75, when n = 9 at site 4 64 Figure 3.9. Survivorship oi Fucus gardneri on rocks transplanted August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks 64 viii Figure 3.10. Growth rates of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks 66 Figure 3.11. Growth rates of Fucus gardneri on rocks transplanted June 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 10 rocks except day 75, when n = 9 at site 4. 66 Figure 3.12. Growth rates of Fucus gardneri on rocks transplanted August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks 67 Figure 3.13. Copper body burden of Fucus gardneri on rocks transplanted June 1997 to BB-1 and FC-1. BB = Britannia Beach. FC = Furry Creek. Data are mean ± 1 standard error, n = 3 plants, except day 39, when n = 2 68 ix Acknowledgments So many people, so many thanks and so little space. I should first thank my supervisory committee. Rob DeWreede has been a remarkable supervisor, teacher and friend for the duration of my thesis work. His rational thinking, friendly demeanor and sense of humour have made my work at U.B.C. incredibly rewarding, both personally and intellectually. Colin Levings provided much insight into all aspects of my work at Britannia Beach, was kind enough to take me on to the project despite having never met me, and, perhaps most importantly, was always diligent in making sure that I got paid. A l Lewis is an amazing critical thinker with an uncanny ability to make me think about things that I don't like to think about. His depth of knowledge in the chemistry and biological effects of copper has been invaluable in the planning and execution of my thesis work. Thanks also to George Haughn, who acted as my Graduate Advisor, to Iain Taylor, the chair of my thesis defense, and to Gary Bradfield, who agreed to be a part of my examining committee. The D.F.O. Science Branch, through Colin Levings, provided the bulk of the funding for the research. Additional funding was provided through Rob DeWreede's NSERC grant and through a teaching assistantship at U.B.C. There is a huge cast of characters who have been involved, in large and small ways, in producing this work. Dr. Colin Dunn of the Geological Survey of Canada kindly arranged for the analysis of the copper content of my transplanted seaweed. Carol-Ann Borden went above the call of duty in helping to identify certain mysterious algae. Christy Stesky and Annabel Forsythe did most of the grunt work on the February 1998 transplant as part of an undergraduate course at U.B.C. Christy has also been very helpful in discussing various ideas, both half-baked and otherwise, while she has been working on her honour's degree on A M D and algae. A long list of others have taken other roles. Some are graduate students and scientists at U.B.C. who have provided insight into my work: Sophie Boizard, Ligia Collado-Vides, Len Dyck, Dana Haggerty, Kristen Milligan, Sherri Rodgers, Anne Salomon and Andrea Sussmann. Others are current and former D.F.O. personnel who have been so helpful in conducting the experiments: Tracy Anderson, Andrea Coombs, Jeff Grout, Sharon Jeffery, Terry Maniwa, Nara Mehlenbacher, Troy Newton, Bruce Nidle, Beth Piercey, Steven Shorig, Doug Swanston and Christy Wright. I offer special thanks to my small army of undergraduate volunteers, most of whom came out to the field on rainy, cold nights to write down the data which I yelled at them: Drew Barker, Kelsey Cramer, Merran Hague, Lawrence Lee, Dorothy Liu, Joanne Roberts and Stephanie Seah. On a more personal note, I should thank my wonderful roommates, Sarah North and Andrea Tanaka who have helped with field work and put up with my mood swings, temper tantrums, rants and assorted other trials and tribulations that come with having a graduate student in your home. Finally, I thank my family, especially my Mom and Dad, for always pushing me to the limits of my abilities and for being honest in not even pretending to be interested when I talk about seaweed. 1. GENERAL INTRODUCTION 1.1. Motivation for Research The research in this thesis was part of a comprehensive examination by the Department of Fisheries and Oceans (D.F.O.) of the impact of acid mine drainage (AMD) on the marine ecosystem at Britannia Beach, British Columbia. D.F.O. is particularly interested in the effects of A M D on salmon and salmon habitat in the surrounding waters. The D.F.O. project involved measurement of environmental factors and metal concentrations in the area and the impacts of A M D on salmon food organisms. An important food source for the salmon is the community of benthic invertebrates that are closely associated with the intertidal brown alga Fucus gardneri Silva (Nassichuk 1975). Therefore, F. gardneri may be critical for the maintenance of the salmon productive capacity of the area. The shoreline near Britannia Beach is almost completely devoid of F. gardneri which does, however, inhabit beaches just a few kilometers from the mouth of the Creek. The research presented here aimed to determine the effects that A M D has on algae in the Britannia Beach area, with a particular focus on F. gardneri . These effects, if detrimental, may then directly or indirectly decrease the productive capacity of the Britannia Beach area for salmon by alteration of their food web. 1.2. Background Information 1.2.1. Acid Mine Drainage Acid Rock Drainage (ARD) is the naturally occurring oxidation of pyrite (FeS2), the metal-containing ore which was mined at Britannia Mine during its operation. The reaction 1 equation (Boulteral. 1994) is: 4 FeS 2 (s) + 15 0 2 + 8 H 2 0 => 2 F e 2 0 3 + 16 H + + 8 S0 4 2" This process has two primary consequences: (1) Production of large quantities of sulfuric acid, and (2) Release of heavy metal ions (e.g., Cu , Zn t , C d ' + ) that are bound up in the pyrite ore. While ARD is a naturally occurring process, the tunneling that necessarily takes place during mining activities can increase its severity by increasing the surface area of rock that is exposed to oxygen and water. Acid mine drainage (AMD) is the special case of ARD that is significantly aggravated by mining activities. Subsequent reference to this process will be to acid mine drainage, as the mine workings are the major source of drainage at Britannia Beach (see below). At Britannia Mine, rainwater, spring snow-melt and groundwater enter the mine workings mostly through open pits at the top (Figure 1.1; Chretien 1997). This water then flows through the underground workings of the mine, reacts with the ore, and exits the mine through two portals (Priceetal. 1995): • The 2200 portal, which emerges from the mine 700 m above sea level. The effluent from this location flows into Jane Creek, which then merges with Britannia Creek approximately 6 km from Howe Sound. • The 4100 portal, which exits the mine 67 m above sea level. This effluent is combined with local sewage and flows to a 30 m-deep outfall in Howe Sound. 2 Figure 1.1. Relative location of Britannia and Furry Creeks, mine features and geographical features mentioned in the text. Adapted from Price et al. (1995) and Grout et al. (1998). Studies by R. McCandless of Environment Canada (unpublished data) estimate the average loading of copper, zinc and cadmium in Britannia Creek as 0.328, 0.316 and 0.005 tonnes per day, respectively. R. McCandless (unpublished data; Figure 1.2) and Chretien (1997; Figure 1.3) found a distinct seasonal variation in metal loading in the Creek with peaks in the spring and fall seasons. 3 700 32 60 bO c 1 300 c N U a— Copper <3 Zinc A Cadmium 3.0 60 May 96 Jun 96 Jul 96 Aug 96 Sep 96 Oct 96 Nov 96 Dec 96 Figure 1.2. Combined daily loadings of Cu, Zn and Cd from Britannia Creek and the submerged outfall into Howe Sound. Data from R. McCandless (unpublished data). Graph compiled by J. Grout. ON ON k-ov ON k-ON Feb-Mar-Apr-May-Jun ON ON 1 ON k-ON ON k-ON Aug Sep- O O Nov-Dec-U") ON C in ON I Figure 1.3. Dissolved copper concentrations in Howe Sound near the mouth of Britannia Creek from March 1994 to January 1995. Values are mean of 8 samples taken along a transect covering the entire mixing zone of Britannia Creek, within approximately 400 m of the Creek mouth. Error bars are the maximum and minimum values recorded at each time point. From Chretien (1997). 4 Chretien (1997) explains the seasonality of metal loadings based on differential flow rates of water into the mine. During winter most precipitation falls as snow at higher elevations; the resulting low flow rates into the mine lead to low metal loadings out of the portals. However, some water stays in the mine, where it continually reacts with ore in tunnel walls and accumulates oxidation products. As spring snow-melt occurs at high elevations, the meltwater flowing through the mine flushes the accumulated A M D from the tunnels, resulting in a pronounced peak in metal loading. During late summer, little precipitation or meltwater enters the mine, so water again accumulates in the mine. Increased precipitation in the fall again flushes this accumulated A M D from the mine and into Britannia Creek, resulting in a second, albeit less pronounced, loading peak (Chretien 1997). There are three major metal components to the A M D at Britannia Beach. While it is beyond the scope of this study to causally link any effects observed in field experiments at Britannia Beach to a specific metal, this study will focus on copper. Cadmium has a very low loading in Britannia Creek relative to copper (Figure 1.2). Zinc has been generally found to be much less toxic to algae than copper (Hargreaves & Whitton 1976; Munda & Hudnik 1986; Lobban & Harrison 1994). It appears, then, that copper is the most likely component of A M D which might cause biological effects. Consequently, much of the data and discussion in this thesis will concern copper. It should be kept in mind, however, that results can only be attributed to A M D in general, and not to any specific component thereof. 1.2.2. Local Oceanography Britannia Beach lies on the eastern shore of Howe Sound approximately 10 km south of the mouth of the Squamish River (see Figure 1.1). The oceanography of the Sound is strongly 5 influenced by this river, which is the major source of freshwater to the Sound. Chretien (1997) examined the salinity and temperature of the waters near Britannia Beach. He found a strong thermocline from May to September, with temperatures of 15 °C at the surface and 7-12 °C at depths below 10 m. There was also a pronounced halocline, with salinities of <3 at the surface and -30 below 10 m. These patterns of stratification were virtually the same at sites near the shore and in the middle of Howe Sound. The halocline and thermocline indicate the presence of a freshwater lens, caused by the Squamish River during freshet (Chretien 1997). This stratification virtually disappears, however, from October to April; surface temperatures during this period were 5-10 °C, while those at depths greater than 10 m were ~8 °C. Salinities were ~22 at the surface and -30 at depths greater than 10 m. Buckley (1977) and Stronach etal. (1992) examined the surface flow pattern of the waters near Britannia Beach (Figure 1.4). There is a net down-inlet current due to the flow from the Squamish River. The main current in upper Howe Sound flows southward along the western side of the Sound. There is, however, a counterclockwise gyre, which breaks from the main southward flow near Porteau Cove and carries water northward along the shoreline at Britannia Beach. This gyre is observed at low tide when surface flow is dominated by river-induced currents. Different flow patterns may be observed, however, due to wind and tidal flows (Buckley 1977). Pond (1992) states that wind-induced currents totally dominate the surface flow pattern. These flow patterns may have important ramifications for the transport of A M D as it flows into the Sound, as the effluent from Britannia Creek and the submerged outfall may be predominantly carried in one direction or another upon entry to the estuary depending on the prevailing conditions at the time. 6 Figure 1.4. General surface circulation patterns in upper Howe Sound. Longer arrows indicate faster flow rates. From Thomson (1981). 1.2.3. Copper Chemistry Wherever pollutants are released into a marine system, chemical processes may affect the degree to which those pollutants act upon nearby communities. Heavy metals may exist in many different forms in aquatic environments. The partitioning of metals amongst these forms is dependent on the physical and chemical conditions of their environment. It is helpful to consider this partitioning of metals as it can significantly alter the ability of the metal to impact the biota of interest. Given the focus of this study, the following review deals only with copper. I.2.3.A. Copper Speciation in Aquatic Systems Copper species can be broadly categorized into three groups of decreasing size: 7 particulate, colloidal and dissolved forms. Particulate species of copper are those which are retained by a 0.45 pm filter (Spear & Pierce 1979). Copper may become part of the particulate phase by precipitating with inorganic ligands, forming insoluble organic complexes, or adsorbing on surfaces of other particles (e.g., clay minerals, hydrous metal oxides, organic matter). Some materials, such as humic acids and hydrous iron oxides, can form colloids in aqueous systems (Home 1969). Copper is known to bind to some of these colloids, and substantial portions of the total copper can be present in a colloidal form (Sholkovitz 1976; Sholkovitz 1978). These colloids, while not truly dissolved forms of the metal, are sometimes inadvertently included in measurements of the latter, as they can often pass through a 0.45 pm filter. Copper can be found in various dissolved forms in aqueous systems. The simplest aqueous form of copper is the hydrated ion, Cu(H 2 0) 6 2 + , also know as "free" copper. One or more of the six water molecules associated with the C u 2 + ion can then be replaced by a variety of ligands. These ligands may be organic (e.g., amino acids) or inorganic (e.g., carbonate, hydroxide). I.2.3.B. Estuarine Changes in Speciation Estuaries are areas of rapid transition; river water, with its low pH (typically 5.0-7.0, occasionally up to 8.0, in various unpolluted creeks near Britannia Beach; Price et al. 1995) and relatively low solute concentration, mixes with seawater which has a high pH (typically 7.5-8.5; Valiela 1995) and is a much more concentrated solution of salt ions (Leckie & Davis 1979). A significant alteration of the speciation of trace metals, including copper, might be expected during this mixing process. Copper has been observed in many situations to behave in a non-conservative manner during estuarine mixing (Windom 1975; Boyle 1976; Boyle 1979; Girvin et al. 1911; Sholkovitz 1978; Sholkovitz & Copland 1981; Hunt 1983), that is, its concentration is 8 changed by factors other than mixing and dilution. Non-conservative processes can result in addition or removal of copper from the dissolved pbol. Addition of copper to the dissolved pool would likely occur through the dissociation of copper from various complexes and particulates, while removal would entail the formation of complexes or adsorption to particulates and their subsequent removal from the water column by sedimentation. As metal-loaded freshwater from organic-poor rivers, such as those flowing into Howe Sound, mixes with relatively metal-free seawater in an estuary there may be a trend towards dissociation of copper from suspended matter (Girvin etal. 1977; Thomas & Grill 1977) and from organic substances (Nelson 1985; van den Berg etal. 1989). However, this apparent release of copper into the dissolved pool might actually be a result of the metal's association with iron colloids, which are themselves released from suspended particulates during estuarine mixing (Fletcher et al. 1983). The dissociation of copper from suspended and organic matter is a result of increased competition by seawater anions and cations for metals and adsorption sites, respectively. As might be expected, the decrease in prevalence of adsorbed and organically bound fractions is concurrent with an increase in inorganic complexation (Spear & Pierce 1979). The change in ionic strength upon mixing with seawater can result in the coagulation and sedimentation of colloidal particles, a process referred to as flocculation. Since copper is strongly bound by these colloids (Steeman Nielsen & Kamp-Nielsen 1970; Boyle etal. 1977; Huang etal. 1977), much of the total copper in river water can be removed through flocculation. Laboratory and field experiments have shown that 30-40% of copper can be removed through flocculation (Sholkovitz 1976; Boyle 1979; Sholkovitz & Copland 1981). The behaviour of copper as it enters an estuary is likely to be quite variable among different areas. While the ratios of major elements in the oceans are quite uniform, many other 9 components which affect copper speciation, such as dissolved organic matter, suspended particulate matter and other metals, can be quite variable. It is therefore extremely difficult to compare metal concentrations of any type in one area to those in another. For example, the percentage of dissolved copper which is biologically available at Britannia Beach may be much higher or lower than that at other polluted estuaries worldwide. This difficulty must be kept in mind when metal concentrations are measured and interpreted. I.2.3.C. Copper chemistry at Br i t ann ia Beach Chretien (1997) conducted detailed surveys of copper speciation in and near Britannia Creek. He found that 92% of the copper was in dissolved form 100 m upstream from the Creek mouth. The amount of copper transferred from the dissolved phase to the particulate phase during estuarine mixing varied over the year from 0-82%, with a mean and median of -30%. The amount of copper which precipitated in the mixing zone was strongly correlated with the concentration of copper in Britannia Creek such that higher concentrations of copper in the Creek resulted in more copper being removed from solution. This removal of copper from solution was attributed mainly to precipitation as hydroxides, hydroxy-carbonates, and hydroxy-sulphates. Dissolved copper concentrations in Howe Sound, 400 m from the mouth of Britannia Creek, were higher in surface waters (mean =13 pg/L) than in subsurface waters (mean = 1.9 pg/L at depths >10 m). Overall, Chretien (1997) found that the dispersion of metals from Britannia Creek occurs within the surface layer of Howe Sound rather than in the entire water column. This concentration of metals in the surface layer may have serious consequences for intertidal organisms as metal-contaminated Creek water will be diluted over a much larger area than if mixing was more homogenous with depth. 10 1.2.4. Interactions between Copper and Algae 1.2.4. A. Bioavailability of Copper Species A metal is considered to be in a biologically available state when it can be taken up by an organism and can react with its metabolic machinery (Campbell et al. 1988). The bioavailability of almost any substance is strongly organism-specific (Campbell etal. 1988). Perhaps the most general statement that can be made about bioavailability of metals is that it is critically dependent on speciation (Sunda & Guillard 1976; Stauber & Florence 1987; Phinney & Bruland 1994; Gledhill et al. 1997). It has been generally accepted that free copper (II) is the form of copper that is most available to almost all organisms (Sunda & Guillard 1976; Anderson & Morel 1978; Sunda & Lewis 1978; Brand etal. 1986; Nor 1987; Campbell etal. 1988). However, Sylva (1976) expresses some reservations regarding this hypothesis, which he asserts is unproven and possibly too exclusive (also, see below). I.2.4.B. Uptake Cellular uptake of copper can be by active, facilitated or passive transport (Campbell et al. 1988; Phinney & Bruland 1994). As with other organisms, the principal source of copper to macroalgae is generally recognized to be free cupric ion (Luoma 1983; Gledhill etal. 1997). However, there is some evidence that sediment-bound metal may be taken up or "scavenged" from sediments by Fucus spp. (Luoma et al. 1982). There is seasonal variation of uptake rate and concentration of several heavy metals mAscophyllum nodosum (Eide & Myklestad 1980) mdFucus vesiculosus (Rigetetal. 1995), bothfucoids (Fucales, Phaeophyceae). There is also interannual variation; body burdens of copper, zinc and cadmium varied over 3 years by factors of 2.5, 4 and 2, respectively (Riget et al. 1995). 11 I.2.4.C. Functions and Requirements Copper, despite its widely documented harmful effects on organisms when present in high concentrations, has been shown to be essential to life (Bowen 1966; Lewis & Cave 1982). It has several functions in the biochemistry of plants (Bidwell 1979). In plants, copper plays an exclusively catalytic role. It is a part of a number of important enzymes, including polyphenol oxidase and ascorbic acid oxidase. It is also present in plastocyanin in chloroplasts and may be involved in nitrate reduction (Bidwell 1979). Gledhill et al. (1997) state in their review that free copper concentrations of 6.3 x 10"6 to 6.3 x 10"4 pg L"1 are thought to be optimal for marine microalgae. Despite its undoubted importance to organisms, copper is probably never a limiting nutrient in natural waters (Lewis & Cave 1982). I.2.4.D. Detrimental Effects Sorentino (1979) outlined the stages of toxicity to phytoplankton which occur as the concentration of copper increases. Low concentrations affect the permeability of the cell membrane, causing K + loss and changes in cell volume. Higher concentrations of copper may be transported to the cytoplasm and then to chloroplasts, where copper inhibits photosynthesis by uncoupling electron transport to NADP + . If even higher concentrations are present, the copper binds chloroplast proteins and other cell proteins, causing degradation of chlorophyll and other pigments. At the highest concentrations irreversible damage to chloroplast lamellae occurs, preventing photosynthesis and eventually causing death. The effects of copper are variable amongst species of macroalgae; for example, the sensitivities of five fucoid seaweeds to copper varied by a factor of three (Stromgren 1980). This author found a 50% reduction in growth of all species at copper concentrations of 50-75 ug/L. 12 Copper and zinc have been observed to have reduced toxicity in the presence of algal exudates (Ragan etal. 1980; Schramm 1993). Macroalgae release metal-binding compounds, such as polyphenols (Gledhill etal. 1997), and some metal binding compounds also exist in the cell wall (Lobban & Harrison 1994). Trace metals may interact with each other in macroalgae, causing a variety of changes in uptake and toxicity of one or both metals. A series of metals were applied singly and in pairs to Fucus vesiculosus at a salinity of 31.5 in experiments by Munda & Hudnik (1986). Copper induced mortality within 20 days at concentrations of 2500 and 5000 pig L" 1, but manganese and cobalt reduced copper lethality at lower copper levels, allowing minimal growth. Copper uptake did not appear to be affected by the presence of other metals, including zinc and cadmium. Also, copper was accumulated to the same degree when applied with other metals as when applied on its own. Salinity can affect the toxicity of copper to some algae. An increase in salinity has been shown to reduce copper toxicity in Cladophora sp., a green alga (Betzer & Kott 1969). This may be due to a change in copper speciation such that less copper is in bioavailable forms. Alternatively, the additional cations in waters of higher salinity may compete with copper ions for binding sites on the surface of the alga. This effect may be important at Britannia Beach, as F. gardneri is exposed to different salinities at different times of the year (see section 1.2.2). Thus, toxicity may vary seasonally, with more severe copper toxicity during the spring freshet. One aspect of copper toxicity to macroalgae which must be considered is the differential sensitivities of various life history stages to copper. Microscopic stages such as gametophytes, gametes and zoospores appear to be especially sensitive. This may be due to their high surface-area-to-volume ratio relative to macroscopic stages. Zoospore release and development of 13 gametophytes were sensitive to copper in the brown algae Laminaria saccharina (Chung & Brinkhuis 1986) and Macrocystispyrifera (Anderson et al. 1990), while settlement and germination of spores were relatively tolerant in Laminaria saccharina (Chung & Brinkhuis 1986). Scanlan & Wilkinson (1987) found that spermatozoa and newly fertilized eggs of Fucus spp. were more sensitive to several biocides than 3-week-old germlings and adult plants. This trend may also hold for copper. Copper may inhibit algal reproduction by reducing the ability of the sperm to find the egg, perhaps by interfering with a pheromone which is thought to be involved in this process (Maier & Muller 1986; Lobban & Harrison 1994). Contradictory results have been obtained when examining the effect of pH on toxicity of copper to microalgae. In a review of the literature, Campbell & Stokes (1985) found that uptake and toxicity of metals in a variety of aquatic biota, including microalgae, decreased with decreasing pH. Toxicity of copper to the green alga Chlorella pyrenoidosa was lower at acidic pH in the experiments of Steeman Nielsen & Kamp-Nielsen (1970). Inhibitory effects of copper on Scenedesmus quadricauda, another green alga, increased 76-fold from pH 5.0 to 6.5 (Peterson et al. 1984). These findings are unexpected given that copper activity normally increases with decreasing pH. The authors of the above studies explained their observations by a mechanism involving competition between H + and C u 2 + for cellular binding sites. In contrast to the above studies, copper toxicity to Aphanizamnion gracile and Oscillatoria redekei, both cyanobacteria, in experiments by Liideritz & Nicklisch (1989) increased at acidic pH. Toxic effects of copper on Scenedesmus quadricauda were also enhanced at acidic pH (Starodub et al. 1987). This discrepancy points out the importance of site-and species-specific factors in determining copper toxicity. 14 1.2.4. E. Tolerance Macroalgae are likely more tolerant of copper than are microalgae (Nor 1987; Gledhill et al. 1997). Some of this tolerance may be conferred by exudates (Ragan et al. 1980; Xue & Sigg 1990; Schramm 1993; Gledhill etal. 1997). In some instances macrophytes can tolerate copper ion activities up to 10"5 M (Nor 1987). Ship-borne populations of Ectocarpus siliculosus (Phaeophyceae) which have been exposed to copper from anti-fouling paints are tolerant to copper concentrations ten times higher than populations from unpolluted areas (Russell & Morris 1970). Estuarine fucoid algae from 'high copper' areas were more tolerant of copper toxicity than the same species from 'low copper' estuarine situations (Bryan 1971; Gledhill 1997). Correa et al. (1996) examined the heritability of copper tolerance m. Enteromorpha compressa (Chlorophyta) in Chile. Plants from copper-polluted shores were able to withstand much higher concentrations of copper. However, their progeny showed no such tolerance. A possible mechanism of copper tolerance in Fucus vesiculosus and F. serratus is through internal detoxification. Smith et al. (1986) found much of the copper in these plants, collected from acid-mine-drainage-polluted shores, to be localized in physodes, which are membrane-bound organelles containing high concentrations of metal-binding polyphenols. Smith et al. (1986) suggest this may reduce the toxicity of the copper by sequestering it in an area where it can have relatively little effect on the metabolism of the plant. 1.2.5. Fucus gardneri Fucus gardneri Silva is a prominent feature of the littoral zone in most of Howe Sound. The plant grows on variably sized substrata, from bedrock to small pebbles which are sometimes 15 buried in sand. It is the dominant alga in the mid-intertidal in most of upper Howe Sound, from Porteau Cove to Watts Point on the east side of the Sound, and from Potlatch Creek to Woodfibre Creek on the west side (pers. obs.; see Figure 1.1). The alga is subject to quite variable salinities throughout the year. In Howe Sound, surface salinities were observed as low as 3 in areas with healthy growth of F. gardneri, while winter salinities reached highs of 20 (pers. obs.). Some fucoids live in estuaries where they are subject to both full strength seawater and fresh water in half a day (Chapman 1995). These algae are also quite tolerant of desiccation. Johnson etal. (1974) found thatF. distichus had photosynthetic rates up to 6 times higher in air than in water. Under the more extreme conditions of summer midday low tides, F. gardneri thalli in Howe Sound were observed to be apparently completely desiccated such that the fronds were black and extremely brittle and shriveled (pers. obs.). Yet, after repeated desiccation to this point, the same plants were observed in apparently healthy condition after the tide came up. The plant's remarkable tolerance of extremes in desiccation and salinity lead to its ability to survive in the mid-intertidal and in upper Howe Sound, respectively. I.2.5.A. Importance F. gardneri is of interest in the current DFO project because of its associated fauna and as a source of organic carbon to the estuarine environment. Nassichuk (1975) examined the relationships between F. gardneri and invertebrate fauna in Howe Sound. He found a wide variety of invertebrates associated with the seaweed, including gastropods (littorines and limpets), isopods, amphipods and chironomid larvae. Some members of the last two groups, as well as certain copepods, are important food sources for juvenile chinook (Oncorhynchus tshawytscha) and chum (Oncorhynchus keta) salmon (Levings & McDaniel 1976; Levings & 16 Riddell 1992), which feed in the intertidal zone after leaving freshwater. It is possible that, regardless of direct effects of A M D on salmon and their prey* F. gardneri biomass may be reduced, thereby reducing habitat for salmon food organisms and ultimately resulting in a lower productive capacity of the area for salmon. A study by Levings and McDaniel (1976) determined that the invertebrate community near Britannia Beach was extremely reduced when compared to unpolluted sites. They found a wide variety of organisms, including molluscs, copepods, amphipods, isopods and several insect groups, on either side of the Creek at distances of 1.5 km. In contrast, only a single insect group and one pelagic amphipod were found on the beach at Britannia Creek. Numbers of organisms were also extremely low; only one organism of one species was found in each of three quadrats. While this study was conducted when the mine was still in operation and would therefore be expected to show more severe detriments to the local biota due to other mine-related disturbances, it nevertheless raises the possibility of reductions in intertidal productivity. I.2.5.B. Life history and Biology F. gardneri is in the order Fucales, class Phaeophyceae. The plant is a perennial which lives up to five years (van den Hoek et al. 1995). It has a gametic life history with only a diploid vegetative stage. This monoecious gametophyte produces antheridia and oogonia in cavities (called conceptacles) on the thallus surface (van den Hoek et al. 1995). The gametes are large, non-motile eggs (typically 75 um in diameter) and small, biflagellate spermatozooids. Fertilization has been found to occur both within the conceptacle (McLachlan et al. 1971; Nassichuk 1975) and after the gametes have been released (Pollock 1969). Once fertilization has occurred the eggs must be dispersed. Burrows & Lodge (1951) 17 demonstrated that propagules of Fucus spp. can colonize at least 23 m from parent plants on a shore without seaweed. Chapman (1995) reviewed work which suggests that most settlement occurs within ~ 1 m of the parent plant. After fertilization, the zygote secretes a glue-like substance which helps it adhere to the substratum upon settlement (Nassichuk 1975). 1.2.6. Statistical Methods It is useful to briefly address the statistical methods which will be used to assess the significance of results obtained in the following experiments. Two types of errors can occur in a statistical test of a null hypothesis: a Type I error, in which the null hypothesis is rejected when it is true, or a Type II error, in which the null hypothesis is not rejected when it false. While the probability of a Type I error is usually determined through the selection of a significance level or a (typically 0.05), the probability of a Type II error, (3, is often not addressed (Underwood 1996). This (3 value determines the power of a statistical test, which is calculated as 1 - (3, and is best described as the probability of detecting a difference between two means when a difference actually exists. As an example of how various parameters of an experiment will determine the probabilities of various outcomes, the following is the equation provided by Zar (1996) to calculate the power of a one-tailed one-sample Student's t-test, which is identical to the paired t-tests in Chapter 3: tp(i),v = [SMs2/n)]-ta(1),v (eqn. 1.1) where tp(1) v is the t-value associated with a given (3 at v degrees of freedom, 6 is the minimum detectable effect size, s is the standard deviation of the attribute under examination and t a ( 1 ) v is the one-tailed t-value associated with a given a. The value of t|3(1) v obtained is translated into a value of (3 and the power of the test is calculated as 1 - (3. 18 There is a tradeoff in any statistical analysis in the selection of experimental parameters. One can assume that the variability of the population, s ,is fixed and that the sample size, n, will be set to the maximium possible given resource-related constraints. If there is sufficient previous information, the minimum detectable effect size can be set at a desirable level. If these steps are taken, there are only two parameters left undefined: a and (3. A decrease in a translates into an increase in (3, and vice versa. Therefore, for any given experiment, the researcher is left with a choice of: (1) setting one of these parameters to a pre-determined level, or (2) attempting to balance the two parameters. Most researchers choose the first option and set a - 0.05. The result of this choice is that the ability of the statistical test employed to detect a biologically significant difference (i.e., the power of the test) may be quite low. In the context of environmental monitoring, the consequences of the power of statistical tests, and of setting a and (3, are critical (Underwood 1996). For the purpose of illustration, consider a simple experiment which compares the growth rates of plants exposed to A M D with those at a control site. A typical experiment would set a at 0.05 with the result, given other constraints on experiments, that (3 might become 0.25. In this case, the chance of a Type II error (i.e., of not detecting a difference that actually exists between control and treatment plants) is 5 times that of a Type I error (i.e., detecting a difference that does not actually exist). The consequence of this discrepancy in error probabilities is that there is an inherent bias in the experiment toward making an error that would, in this example, erroneously fail to detect an effect of A M D on plant growth rates. The alternative procedure is to set a and (3 equal to one another. In the experiment described above, a and (3 would be set at 0.122. In this case, either type of error is equally likely. In an examination of environmental effects of A M D , this would seem to be a more logical 19 procedure, as there is no obvious justification for biasing the experiment towards a Type II error. If a and |3 are set to different values, there should be some justification for introducing the bias into an experiment. All parameters should be set according to the specific circumstances of the particular study. The values of both a and |3, as well as the justification for these values, should be reported. Unfortunately, many studies are limited in the amount of background information which is available. This information is necessary to determine what minimum detectable effect size and variance values are appropriate. If no previous examinations have been conducted on the system, it will be very difficult to determine the level at which these parameters should be set. In such studies, a priori power analyses, as described above, are very difficult to apply objectively. A different approach which may be taken is an a posteriori"power analysis. This approach consists of gleaning effect sizes and variances from the experiments, then estimating the power of the statistical tests after they have been conducted. While this approach does not allow a genuine adjustment of a and (3 values, it does provide useful information on the likelihood that the tests resulted in errors. The results can then be applied to the design of any subsequent experiments. Furthermore, they can provide insight into the usefulness of the experiments in inferring conclusions about the questions being examined. 20 1.3. Research Hypotheses Given all the information outlined above, it is possible to formulate a series of hypotheses which can be tested mensuratively and experimentally: 1. F. gardneri plants constitute a smaller proportion of cover near Britannia Beach than in reference areas. 2. Filamentous green algae constitute a smaller proportion of surface cover near Britannia Beach than in reference areas. 3. F. gardneri plants are shorter near Britannia Beach than in reference areas. 4. Extant F. gardneri plants near Britannia Beach produce fewer gametes than those in reference areas. 5. F. gardneri plants near Britannia Beach will have a higher body burden of copper than those at reference sites. 6. Growth rates of adult F. gardneri transplanted to Britannia Beach will be lower than those at control sites. 7. Survivorship of adult F. gardneri transplanted to Britannia Beach will be lower than that at control sites. 8. Copper body burden of F. gardneri transplanted to Britannia Beach will be higher than that at control sites. Hypotheses 1 through 5 will be addressed in Chapter 2; hypotheses 6 through 8 will be addressed in Chapter 3. 21 2. D I S T R I B U T I O N A N D R E P R O D U C T I V E F U N C T I O N O F A L G A E N E A R B R I T A N N I A C R E E K I N R E L A T I O N T O A C I D M I N E D R A I N A G E 2.1. Introduction At the outset of this investigation, there had been some quantification of the intertidal fauna near Britannia Beach (e.g., Levings and McDaniel 1976; Grout et al. 1998, 1999), but no such work had been done on algae. There was a need, then, to quantify the algal community to determine what, if any, differences exist between Britannia Beach and other, uncontaminated sites. Aside from a thin layer of green algae in the high intertidal, there is no apparent life within approximately 100 m of Britannia Creek. Farther from this creek, a dense cover of filamentous green algae and a few scattered barnacles appear on cobbles and boulders. Still farther from Britannia Creek, about 1000 m north and 600 m south, there is the first sign of Fucus gardneri Silva. These plants appear to be somewhat diminutive in both length and breadth. As one moves past this point, F. gardneri cover gradually increases to levels similar to those at other places in upper Howe Sound, (pers. obs.) A series of hypotheses (section 1.3) concerning F. gardneri plant length, cover and gamete production and filamentous green algal cover were formulated and tested. The sampling program was designed to examine both the areas where F. gardneri is absent and the fringe areas where it grows nearest to Britannia Creek. Sampling was also done in several seasons to determine if patterns may be visible at some times but not at others. 22 2.2. Methods 2.2.1. Dissolved Copper in Water Dissolved copper concentrations in water at a series of stations along the east and west shores of Howe Sound (Figure 2.1) were measured in an associated study of the effects of A M D on the blue mussel (Mytilus edulis; Grout et al. 1999). Collections were done at four times in 1998: April 29, May 15, May 28 and June 8. Samples were collected from surface water and immediately filtered through a 0.45 um polycarbonate filter into acid-washed polyethylene bottles. This filtration removed particulate copper, as dissolved copper is thought to better reflect the amount of metal available to organisms (see section 1.2.4.A). Samples were acidified with 1 mL of concentrated nitric acid and returned to the laboratory. Metal analysis was conducted by graphite furnace atomic absorption spectrometry (GF-AAS). Figure 2.1. Locations of stations where dissolved copper was measured. From Grout and Levings (in preparation). Map from Canadian Hydrographic Service Chart #3526. 23 2.2.2. Quantification of Fucus gardneri length and algal cover The locations of transect stations are indicated in Figure 2.2. All further references to individual transect stations are abbreviated as, for example, BB-D or FC-D, where BB and FC represent Britannia Beach and Furry Creek, respectively, and the last letter corresponds to the transect station itself. The labels are such that stations with the same label roughly match each other for distance from the appropriate creek mouth and substratum. For example, sites BB-B and FC-B are -800-1000 m north of their respective creeks and both consist of a relatively steep rocky intertidal zone. Figure 2.2. Locations of transect stations at Britannia Beach (left) and the corresponding control stations at Furry Creek (right). 1 cm = 350 m. From Canadian Hydrographic Service Chart #3526. Stations were chosen in pairs at each of the two sites, Britannia Beach and Furry Creek. A gradation of distances from Britannia Creek was desired based on the assumption that there is 24 a gradient of decreasing A M D concentration as one moves away from Britannia Creek. Stations at Britannia Beach were chosen to assess this gradient. A similar gradation was then sought at Furry Creek. The Furry Creek stations were intended to act as a control for the effects of the freshwater inflow from Britannia Creek. Where the influence of the Creek on salinity and turbidity was minimal due to the much greater influence of the Squamish River (i.e., during freshet), the sites could be considered as spatial replicates. Stations at Furry Creek were therefore chosen to match the distance from Britannia Creek as closely as possible. At the same time, substratum type and size can have important effects on distribution of F. gardneri and other algae. An attempt was thus made to select matching stations in each area with similar substratum types. Finally, an important consideration was safe access to the stations in all weather conditions and at all times of day. Access to the north of Britannia Beach was hindered by a train tunnel, while that north and south of Furry Creek was hindered by the extreme steepness of the intertidal and the backshore areas. Sampling was conducted over two or three days on the lowest tides of each month. Transect surveys were conducted in June, July, August, October and December 1998. The area of study at each transect station consisted of a 20 m horizontal reach of the intertidal zone. The vertical limits of the transect station were set by the limits of the F. gardneri zone. The upper and lower limits of the F. gardneri zone (typically 3.0 m and 1.5 m above chart datum, respectively) were measured at all stations where this alga grew and then extrapolated to stations from which it was absent. A horizontal transect line was laid along the top or bottom of the station. Three vertical transects were then randomly placed along the horizontal line and a 25 cm by 25 cm quadrat was randomly placed at three points along each vertical line, for a total of nine quadrats per transect station. The quadrat was pre-strung with nylon string in a grid 25 pattern with squares 1.5 cm x 1.5 cm, and 20 of the intersection points were randomly marked with pieces of coloured wire. Percent "canopy" cover estimates were obtained by recording the organism or substratum type under each of the marked points in the quadrat. Percent "surface" cover was then obtained by recording the object under all points after the canopy of F. gardneri had been moved aside. Categories assessed were: F. gardneri, filamentous green algae, rock (any size >2 mm), sand, mussel and barnacle. Due to the difficulty of identifying filamentous green algae in the field, all species were grouped into one functional group for purposes of cover data. Samples were taken at several stations during each collection period, returned to the laboratory and identified using the keys of Gabrielson et al. (1989). Other species of algae were occasionally encountered and were recorded as miscellaneous algae, then identified in the laboratory. F. gardneri plant lengths were then measured. Five plants were selected randomly in each quadrat as the individual that was closest to each of five randomly marked intersection points. Plant length was measured as the distance from the holdfast to the end of the longest frond. 2.2.3. Oocyte Release Sampling was conducted concurrently with the transect surveys. Collections took place in July, August, October and December 1998. Plants were collected haphazardly from the middle of the F. gardneri zone within 20 m of the transects. Fronds were sought which appeared to be reproductive, namely those that had swollen receptacles and conceptacles, were dark in colour and were covered in a large amount of exudate (Pollock 1969; DeWreede, pers. comm.). Three samples were collected at each station. Plants were pulled from the substratum by hand, 26 placed in open plastic bags and transported on ice to the laboratory. Oocyte release was induced by following the procedures of Pollock (1969) with slight modifications. Plants were left in the dark in their open collection bags for 48 hours at 4 °C. Non-reproductive portions of the fronds were then removed by cutting off areas where no swollen conceptacles were visible. The reproductive tips (called conceptacles) were then gently hand-scrubbed in running freshwater (dechlorinated West Vancouver city water, temperature 13-15 °C) for one minute, and were then repeatedly doused with this same running water in a 300 mL, 8 cm dish for one minute. The tissue was then immersed in still freshwater for 10 minutes. This water was removed and replaced with seawater from the West Vancouver Laboratory (W.V.L.) seawater system (salinity 27-33, temperature 7-10°C). After sitting in seawater for 10 minutes, receptacles were removed and rinsed into the sample dish. Oocytes were then counted under a dissecting microscope. For samples with large numbers of oocytes, a subsample was counted by mixing vigourously, then counting a delineated portion of the dish. Receptacle tissue was kept for surface area measurement. This was done by pressing tissue between two sheets of plexiglass, photocopying them, and scanning the photocopies using a computer image scanner. The area of each sample was calculated using SigmaScan Pro 5.0. Oocytes counts were then normalized to the surface area of the plant tissue which had produced them. 2.2.4. Copper body burden in Fucus gardneri F. gardneri were collected from the middle of their vertical distribution, approximately 2.0 m above chart datum, in August 1998 concurrently with the transect data collection. Three plants were collected at each station. These plants were returned to the laboratory, rinsed and 27 hand scrubbed three times with seawater from the W.V.L. seawater system and sent to A S L Laboratories (Vancouver, B.C.) for copper content Analysis. Fresh plants were air dried in a laminar flow fumehood at ambient temperature, ground and then ashed at 470°C for 24 hours in a muffle furnace. Ashed material was digested in a 1:1 mixture of concentrated nitric acid and concentrated hydrochloric acid at 95°C for 2 hours. Copper analysis was carried out using Flame Atomic Absorption Spectrophotometry (FAAS). 2.2.5. Statistical analyses All statistical comparisons were carried out in one of three computer programs: Microsoft Excel 5.0, Systat 7.0 or SPSS 8.0. F. gardneri percent cover, plant length, oocyte release, copper body burden and filamentous green algal cover were all compared across sites using a one-way analysis of variance (ANOVA) with a = 0.05. At some stations, certain measurements yielded values of zero in all quadrats; for example, percent cover of F. gardneri at stations BB-C through BB-F was consistently 0. These measurements could not be analyzed by A N O V A as they have no variance. These stations were omitted from the A N O V A , but the results can be interpreted based on their obvious contrast with stations with high values for the measurement in question. If significant differences were detected, a Student-Newman-Keuls (SNK) test was used. The SNK test compares all stations to each other and forms homogeneous groups of stations. These groups are constructed so that stations within a given group are not significantly different from each other at a = 0.05, but are significantly different from stations in all other groups. Occasionally, the A N O V A will detect differences, but the SNK test will be unable to form any homogeneous groups due to the latter test's lower power. Some violations of the assumptions of parametric statistics, namely the assumptions of 28 equal variances and of normal distributions, were detected using Levene's test and the Kolmogorov-Smirnov test, respectively. Transforming the data did not remove these violations. Most non-parametric procedures, while not requiring normality of the data, still require equal variances in all samples. While A N O V A and SNK tests do make the assumptions listed above, they are recognized to be very robust to departures from normality and equality of variances (Zar 1996, Underwood 1997). Therefore, parametric A N O V A and SNK tests are used despite violations of assumptions. 2.3. Results 2.3.1. Dissolved copper i n water Figure 2.1 indicates the locations of sampling for dissolved copper concentrations, and Figure 2.3 shows the concentration of copper found at these stations. Concentrations of dissolved copper are consistently highest at station 7, which is nearest the mouth of Britannia Creek. Copper concentrations are elevated to the north of the Creek mouth (stations 8 and 9) during all data collections but one (station 9 on April 29). No such elevation is evident to the south of Britannia Creek (station 6). A temporal trend, consistent with Chretien (1997) and R. McCandless' (unpublished data) findings, is evident with copper concentrations increasing to a peak in late May, then declining. 29 200 150 100 J 50 431 ± 29 ; , April 29 j i § • . „ 9 1 -+ i n | n , n pn I s i j M | ra | in > 200 2 3 4 5 6 7 8 9 10 11 Station 3 U T3 <D _> "o oo oo •3 00 55. 150 100 50 J L 797 ± 1 1 9 May 15 U 2 3 4 5 6 7 8 9 10 11 Station 200 3 U T3 cu "o oo 0O bO 3. 150 100 50 1009 ± 44 i , n,n r May 2H 3 4 5 6 7 8 9 10 11 Station 200 150 3 u I 100 oo oo •3 bO 3. 50 823 ± 52 n n n June 8 n 2 3 4 5 6 7 8 9 10 11 Station Figure 2.3. Concentration of dissolved copper in Howe Sound surface from April-June 1998. Zero values indicate concentrations below the detection limit of 5 ug Cu L" 1. Stations 7 and 8 comprised 3 samples each; error terms are the standard errors for these measurements. All other stations comprised only one sample per time point. Values for station 7 are reported as numbers to allow clear presentation of values at other stations, which were much lower. From Grout and Levings (in preparation). The variability in metal dispersal is apparent in several aspects of these data. First, the large error bars at station 8 on May 15 indicate variability of copper concentrations on a small (<10 m) spatial scale. Second, the variation in differences between copper levels at stations 8 30 and 9 indicates the complexity of circulation processes which, in one case (June 8), leads to slightly higher metal concentrations at the station farther north of Britannia Creek. This variability leads to great difficulty in trying to correlate any other observations with specific concentrations of copper. 2.3.2. Fucus gardneri cover Percent cover estimates for F. gardneri are presented in Figure 2.4. Results from statistical tests are shown in Table 2.1. Cover of F. gardneri was quite variable at all sites and at all time points. No F. gardneri was found at any time at stations BB-C, BB-D, BB-E or BB-F. Outside of this zone of F. gardneri absence, however, no significant differences were detected among estimates of F. gardneri cover at any stations, with one exception. This lack of significant differences occurs despite an apparent trend toward lower cover of F. gardneri at stations BB-G and BB-H than at FC stations. A general temporal trend is visible as a peak in F. gardneri cover in July, with steadily decreasing cover to a minimum in December. 2.3.3. Filamentous green algal cover Percent cover of filamentous green algae is presented in Figure 2.5. All numbers were obtained from surface cover data as opposed to canopy cover to avoid bias at sites with high cover of F. gardneri. Results of statistical analyses are listed in Table 2.2. Filamentous green algae generally comprised two groups: Enteromorpha intestinalis (L.) Link and Ulothrix spp. Further identification of the latter was not possible due to variability in specimens within the ranges listed in the keys. In addition to these two species, a thin layer of Chlorella spp., a single-celled green microalga, was found on the rocks within 10 m of the mouth of Britannia Creek. This alga is regarded as a freshwater alga (Stein 1975), which explains its 31 I o t-(D > o U 80 70 60 50 40 30 20 10 0 a B B D F C "June 24-25 -— 1 -H I L i -v.-I M-c O Ul CD > O U-80 70 60 50 40 30 20 10 0 a B B D F C July 21-23 —-i TJ - j I D E G H C D E F G H 80 " 70 I 60 I o Ul CD > O U 50 40 30 20 10 0 _ m BB August 17-19 • FC I _ i 1 1 --I— - H — H — -+ - -4 I O Ul CD > o U 80 70 60 50 40 30 20 10 0 a B B D F C - - : October 8-10 Ij I 1 1 1 • i - H — , ffil B C D E F G H B C D E F G H CD o Ui CD > O U 80 70 60 50 40 30 20 10 0 a B B ' • F C December 1-3 JL + B C D E F G H Figure 2.4. Percent cover of Fucus gardneri as measured from June to December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Sampling was not conducted at sites B until August. Error bars are ± 1 standard error; n 9 for all stations. 32 Table 2.1. Statistical analysis of Fucus gardneri percent cover from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Stations BB-C, BB-D, BB-E and BB-F were omitted in all months due to complete lack of F. gardneri. Station BB-B was omitted in December only. SNK tests were conducted if the A N O V A detected significant differences. All stations within a set of square brackets comprise a homogeneous group. See section 2.2.5 for explanation of SNK grouping. Sampling P-value from SNK grouping Period ANOVA ,. June <0.001 no homogeneous groups detected July 0.051 N/A N/A August 0.008 no homogeneous groups detected October <0.001 [FC-E] All other BB and FC December 0.307 N/A N/A limited distribution at Britannia Creek. Furthermore, it formed a layer only 2-3 cells thick on rocks in a very small portion of the study area and thus contributes very little to marine primary production in this area. Consequently, this alga is not discussed further here. Filamentous green algal cover tended to be higher at all stations near Britannia Creek than at stations near Furry Creek. Cover of filamentous green algae was significantly higher at stations BB-C and BB-F than at all other stations in June, July and August. An additional statistically significant difference was detected in December, although the relatively high cover of filamentous green algae at station BB-F in December appeared to be composed entirely of senescing plants. Cover of filamentous green algae was highest at all sites in the summer months, with a decline in October and an almost complete lack of filamentous green algae in December. 2.3.4. Fucus gardneri plant length Lengths of plants at each station are presented in Figure 2.6. Some sample sizes were <9 because some quadrats contained no F. gardneri and so were not included in the analysis. Most 33 90 ^ 80 .•Jp 70 < 60 CD 2 50 O o CD > O U 40 30 20 10 0 r n B B • FC July 21-23 D H C D E F G H CD cd c CD CD S-l CD >• O U B C D E F G H B C D F G H CD cd 00 C CD CD o M - i O S i CD > O U B C D E F G H Figure 2.5. Percent cover of filamentous green algae as measured from June to December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Sampling was not conducted at sites B until August. Error bars are + 1 standard error, n = 9 for all stations. 34 Table 2.2. Statistical analysis of filamentous green algal cover from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping. Sampling P-value from S N K grouping Period A N O V A June <0.001 [BB-C, BB-F] All other BB and FC July <0.001 [BB-C, BB-F] All other BB and FC August <0.001 [BB-C, BB-F] All other BB and FC October 0.199 N /A N /A December 0.002 [BB-F] [FC-F.FC-G] of the sample sizes, however, were >7. Results of statistical tests are presented in Table 2.3. Plant lengths, like the other measures taken, are quite variable at most times and locations. F. gardneri at Britannia Beach stations tended to be shorter than those at Furry Creek, but, while some differences were detected by the A N O V A , they were not elucidated by the SNK test. Plants appeared to be longest in June and July, and length then declined to a minimum in December. 2.3.5. Oocyte release July and August collections yielded extremely low numbers of oocytes. For example, of three F. gardneri samples from a given site, often only one would yield oocytes, with usually less than 5 egg cells released. In contrast, October and December samples yielded very high numbers of oocytes, frequently more than 600 in a given sample. Therefore, only October and December data are presented and analyzed (Figure 2.7). Results of statistical tests are presented in Table 2.4. Significantly more oocytes were released by F. gardneri from station BB-B than at all other stations in October, while there were no differences among other stations. No 35 12 10 i 8 bp 6 c CD d 4 c CO - D F C June 24-25 1 1 i i 12 10 J. s^ & 6CD s 4 to 2 0 a B B D F C July 21-23 i D G H i — i C D E F G H 12 10 3, 8 X ! & 6 CD ^ 4 ca H B B D F C December 1-3 B D E G H Figure 2.6. Mean length of Fucus gardneri as measured from June to December 1998. A, June 24-25; B July 21-23; C, August 17-19; D October .8-10, E , December 1-3. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no plant lengths are given. Sampling was not conducted at sites B until August. Error bars are + 1 standard error, n varied from 5 to 9, averaging 7.9, at sites with F. gardneri. 36 Table 2.3. Statistical analysis of Fucus gardneri plant length from transect study, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping. Sampling P-valuefrom SNK grouping Period ANOVA . . June <0.001 no homogeneous groups detected July <0.001 no homogeneous groups detected August 0.001 no homogeneous groups detected October 0.157 N/A N/A December 0.350 N/A N/A 1* o l-H CD O H CO CD •M >> o o O 16 14 12 10 8 6 4 2 0 a B B D F C October 8-10 fl 1 1 B C D E F G H 16 14 12 u 10 ' CD^  • co 8 CD -4-> o o o ° 4 a'BB D F C December 1-3 ri-,. -—f ._JL B C D E F G H Figure 2.7. Oocyte release from Fucus gardneri as measured in October and December 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no oocyte counts are given. In October, n = 3 at all sites; in December, n = 3 at BB-G and FC-G, n = 4 at FC-C, n = 5 at all other stations. All values are normalized to the surface area of the source plant material. significant differences were detected among oocyte releases from December samples. There is no general tendency toward higher or lower oocyte releases at Britannia Beach. 37 Table 2.4. Statistical analysis of oocyte release, a = 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping. Sampling P-value from SNK grouping Period ANOVA , : October 0.006 [BB-B] All other BB and FC December 0.105 N/A N/A 2.3.6. Copper body burden in Fucus gardneri Results of copper body burden measurements are shown in Figure 2.8. Results of statistical comparisons are shown in Table 2.5. Copper body burdens in F. gardneri from all Britannia Beach stations are significantly higher than those in algae from Furry Creek stations. 800 B C D E F G H Figure 2.8. Copper body burden in dry weight oi Fucus gardneri collected August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. Some stations had no F. gardneri (see Figure 2.4), so no plants were collected. Error bars are ± 1 standard error, n = 3 at all sites. Table 2.5. Statistical analysis of copper body burden in Fucus gardneri. a - 0.05 in all tests. BB = Britannia Beach. FC = Furry Creek. See Figure 2.2 for locations of sampling sites. SNK tests were conducted if the A N O V A detected significant differences. See section 2.2.5 for explanation of SNK grouping. Cu in plants Sampling P-value from SNK grouping Period ANOVA August <0.001 [BB-B] [BB-G] [BB-H] All FC 38 2.4. Discussion 2.4.1. Dissolved copper in water The measurements of copper concentrations in seawater are limited in several respects. First, the samples were taken over only a five week period. While this is very useful information for this particular interval, these data give limited insight into metal dispersal at other times of the year. The data from Chretien (1997; Figure 1.3) provide a better picture of dissolved copper levels over the entire year, but no spatial trend can be inferred from his data. A second limitation of the copper concentration data is that they measure only dissolved copper. Due to the highly variable speciation of copper in estuarine water, it is extremely difficult to use one measure of copper concentration to accurately assess and predict effects on biota. Measuring dissolved copper as opposed to total copper is one way of reducing error, as free copper is likely the form of the metal that is most available to organisms. However, dissolved copper includes not only free metal, but also some organically bound, inorganically complexed and colloidal copper, forms which can be less available for uptake by organisms. Additionally, Luoma etal. (1982) have found evidence that Fucus vesiculosus can accumulate particulate copper. If this occurs with F. gardneri at Britannia Beach, dissolved copper would not be the only source of the metal to the algae. Much useful information can, however, be gathered from these data. A spatial trend of copper is clearly visible (Figure 2.3). The most striking feature in all four graphs is the spike of copper at site 7, directly off the mouth of Britannia Creek. It is quite apparent that the high loading of copper in the Creek translates into elevated concentrations of dissolved copper in Howe Sound, at least in the immediate vicinity of Britannia Creek. 39 In addition, the dominant dispersal pattern of copper from Britannia Creek is visible in the elevated dissolved copper levels at stations 8 and 9 and the concurrent lack of consistently high copper levels at station 6. This pattern is consistent with the surface circulation pattern near Britannia Beach described by Buckley (1977; Figure 1.4), which causes the plume of Britannia Creek to flow northward as it enters Howe Sound. The dispersal of metals might also be responsible for the asymmetry in F. gardneri cover around the Creek, as is evident in Figure 2.4. 2.4.2. Fucus gardneri cover A conspicuous absence of F. gardneri from the shoreline near Britannia Creek was apparent at the outset of this study in the spring of 1997. Stations BB-C, BB-D, BB-E and BB-F were all completely devoid of this alga (see Figure 2.2). The area where F. gardneri was absent, measured in a straight line from the Creek mouth, extended over a total of approximately 1600 m. Stations BB-B and BB-G are the nearest locations to the north and south of Britannia Creek, respectively, where F. gardneri grows. BB-B is approximately 1000 m north of the Creek mouth, while BB-G is approximately 600 m to the south of the Creek. The cover of F. gardneri at these two stations, along with an additional station to the south of the Creek (BB-H) was monitored and compared to a similar series of stations near Furry Creek. An analysis of variance detected significant differences in the cover of F. gardneri among stations which supported the alga in three of five months: June, August and October 1998 (Table 2.1). The SNK test only formed homogeneous groups in October, however, with FC-E having a higher percent cover of F. gardneri than all other sites. This general lack of differences is due to the high variances that were present at all sites. F. gardneri is patchily distributed on the scale of the quadrats used (25 cm x 25 cm; pers. obs.), resulting in high variances and, therefore, low power 40 of any statistical tests. A trend is evident in the data, however. In all months except December, the percent cover of F. gardneri at all the Britannia Beach stations where these algae were present was lower than at all Furry Creek stations, although the differences were not significant (Figure 2.4; Table 2.1). This suggests that there may be a small effect of A M D on the cover of F. gardneri at these stations which was not detectable using the methods and sample sizes in this study. The biological importance, in terms of algal biomass and associated fauna, of so small a difference as that observed at these stations is debatable. Measurement of biomass might be a more appropriate way of determining this biological significance. The lack of any trend toward lower F. gardneri cover at Britannia Beach in December is the result of a general decrease in algal cover which occurs at most other sites. The absence of F. gardneri from the 1600 m surrounding Britannia Creek, which was not included in the statistical analysis, is obvious from the data presented in Figure 2.4. The asymmetry of the area from which F. gardneri is absent, with a greater area of absence north of Britannia Creek than south, is consistent with the dispersion of metals and circulation patterns in Howe Sound, as described above. Any relationship between the distribution of A M D and the absence of F. gardneri which might be inferred from these observations is exclusively correlative. However, A M D appears to be the most likely cause of the disrupted distribution of this alga near Britannia Creek. All other creek mouths observed in upper Howe Sound, including Furry, Potlatch and Woodfibre Creeks (see Figure 1.1 for locations), are dominated by a thick cover of F. gardneri (pers. obs.). An alternative explanation for the dearth of F. gardneri at Britannia Creek relative to Furry Creek might be the higher turbidity and lower salinity the plants would encounter at the former location due to its proximity to the Squamish River, which 41 dominates the salinity regime in upper Howe Sound during freshet. However, extremely dense populations of F. gardneri were found to the north of Britannia Creek at Watts Point (see Figure 1.1 for location; pers. obs.), which is much closer to the Squamish River. As well, the west side of Howe Sound (e.g., the mouth of Woodfibre Creek) encounters higher turbidity and lower salinity than Britannia Beach during freshet (Stockner et al. 1977, Grout et al. 1999). Despite these patterns of freshwater distribution, however, the mouth of Woodfibre Creek has F. gardneri percent cover in excess of 80% in some locations (pers. obs.). Salinity and turbidity do not appear to be viable explanations for the lack of F. gardneri at Britannia Beach. 2.4.3. Filamentous green algal cover In general, filamentous green algal assemblages in areas where F. gardneri was present comprised both Enteromorpha intestinalis and Ulothrix spp., while those at the stations near Britannia Creek from which F. gardneri was absent contained only E. intestinalis. These filamentous green algae have a seasonal pattern of abundance, taking advantage of high sunlight and warm temperatures during the summer months, then dying back in the fall and winter. This temporal trend is very evident in the measurements shown above (Figure 2.5), and is consistent with findings elsewhere (Prange 1976; Pomeroy 1977; Munda & Markham 1982) that filamentous green algae tend to have sharp seasonal peaks in biomass. These species appear to be well adapted to summer growth; several studies (Kim & Lee 1993; Einav etal. 1995; Kim & Lee 1996) have found that Enteromorpha spp. are very tolerant of high temperatures, desiccation and fluctuations in salinity. At stations BB-D and BB-E (see Figure 2.2 for locations of stations) on either side of the mouth of Britannia Creek, cover of all algae was generally very low (Figures 2.4 and 2.5). It 42 appears that A M D concentrations at these stations are high enough, at least periodically, to preclude any growth by marine algae. Farther from the Creek, at stations BB-C and BB-F, filamentous green algal cover is significantly higher than that at all other stations (Figure 2.5; Table 2.2). Still farther away from Britannia Creek, cover of green algae was again lower where F. gardneri occurred (Figures 2.4 and 2.5). Filamentous green algal andF. gardneri cover in June, July and August were negatively correlated when stations BB-D and BB-E were excluded (Table 2.6). Table 2.6. Correlation analysis of mean green algal percent cover and Fucus gardneri percent cover at Britannia Beach and Furry Creek stations. Stations BB-D and BB-E were excluded due to the presumed severe influence of A M D on filamentous green algae, n = 9 stations for June and July, n = 11 stations for August. Month Spearman correlation P-value for coefficient correlation June -0.966 " <0.001 July -0.929 <0.001 August -0.862 0.001 The most likely explanation for the heavy cover of Enteromorpha intestinalis near Britannia Creek is that this alga can tolerate higher concentrations of A M D than can F. gardneri. Several authors have found Enteromorpha spp. to be much more tolerant of copper than other algae. Correa et al. (1996) exposed Enteromorpha compressa to varying concentrations of copper in culture and found growth rates to be unaffected by <635 pg copper L" 1. Reed & Moffat (1983) exposed E. compressa from both ship-fouling (i.e., copper paint-exposed) and non-fouling populations to varying concentrations of copper. They found that the growth rate of plants which had previously been exposed to the metal were unaffected by <609 ug copper L" 1, while that of plants from non-fouling populations was-decreased by 114 pg copper L" 1. These 43 results suggest that tolerance to copper is built up through exposure. In comparison with Reed & Moffat's (1983) results, heavy E. intestinalis cover was found near Britannia Creek at sites which were exposed to dissolved copper levels <190 pg Cu L" 1. Direct comparison is difficult, however, as the speciation of the metal at Britannia Beach is likely different from that in Reed & Moffat's (1983) laboratory experiments. Furthermore, the temporal variability of dissolved metal levels in Howe Sound and the different species of algae used in the two studies makes any direct comparison overly ambitious. Castilla (1996) examined algal communities at Caleta Palito, Chile over a 19-year period. From 1975 to 1990, untreated mine tailings with 6000-7000 pg copper L"1 in the tailing water were discharged directly onto the intertidal near the Salado River. Castilla (1996) found that Enteromorpha compressa cover in the intertidal increased from <10% to >90% within 3 years of the initiation of tailing dumping. After 1990, tailings were treated to remove solids, after which the "clear tailing water" was released into the Salado River. Castilla (1996), in 1994, measured 2400 pg dissolved copper L"1 at the mouth of this river and 26.8-31.8 pg dissolved copper L"1 at a site 50 m south of the river mouth. He found that E. compressa remained the dominant alga in the area four years after this modification of dumping procedures. Finally, he observed that E. compressa dominated the intertidal up to 7 km north of the river mouth, comprising >80% of the cover. These data suggest that Enteromorpha spp. often persist—and sometimes flourish—in copper-polluted environments where other algae cannot survive. Several authors (Say et al. 1990; Castilla 1996) cite this alga as a "sentinel" species: one which can survive where other algae cannot in areas of pollution by copper and other metals. This also appears to be the case at Britannia Beach, where Enteromorpha intestinalis is the dominant organism in terms of cover and biomass in the intertidal areas where F. gardneri is excluded. 44 Filamentous green algae are generally very limited in their distribution, however, wherever F. gardneri is abundant, often living in the high intertidal at the fringe of F. gardneri''s vertical limits (pers. obs.). Previous studies have examined the interactions among Fucus spp., ephemeral (i.e., seasonal) algae, including Enteromorpha spp., and herbivores such as Littorina spp. (Lubchenco 1983). Lubchenco (1983) proposed that in the absence of littorine grazing, ephemeral algae may inhibit the recruitment of juvenile Fucus spp. The inhibition is only temporary, however, as the seasonal ephemeral algae die out in the fall, allowing recruitment of Fucus spp. Qualitative observations at all sites in Howe Sound suggest that littorines are absent from the areas without F. gardneri (pers. obs.). Whether this absence is a result of acid mine drainage or the lack of perennial algae (i.e., food items) is not clear. It might be suggested that A M D eliminates littorine grazers from the intertidal, resulting in inhibition of F. gardneri by filamentous green algae. However, the extreme seasonal variation in the abundance of the latter would likely rule out this possibility. A more likely scenario, suggested by Lubchenco's (1983) study, is that Fucus spp. and littorines are excluded from the area near Britannia Beach by A M D , allowing Enteromorpha spp. unhindered access.to intertidal habitat. At other locations, Littorina spp. preferentially feed on ephemeral species (Granado & Caballero 1991) and reduce the cover of filamentous green algae. While this model may apply at Britannia Beach, it is very speculative to extrapolate results obtained with different species of grazers and fucoids in New England to British Columbia. Chapman (1995) reviews several studies which suggest that Lubchenco's (1983) model may be site-specific, as other researchers have found results contrary to hers. Lubchenco's (1983) model is suggested here as one possible explanation for the patterns seen in Howe Sound; much experimental work would be necessary to confirm or refute its applicability to the findings in this study. 45 Another possible explanation for the heavy cover of filamentous green algae at stations BB-C and BB-F is that they are limited by copper in natural environments. Thus, the concentrations of copper at stations BB-D and BB-E may be toxic to these algae, while those at stations BB-B and BB-G were limiting. This hypothesis is not, however, supported by the work of Correa etal. (1996), who found that is. compressa'm Chile grew equally well in 0, 63.5 and 635 u.g copper L" 1. The concentrations of dissolved copper found at stations BB-B and B B - G (<5-25 p,g copper L"1) are well within this range. As well, Lewis & Cave (1982) state, based on an extensive review of the literature, that copper is probably never a limiting nutrient in natural waters. 2.4.4. Length of Fucus gardneri Qualitative examination of F. gardneri nearest Britannia Creek prior to the outset of this study suggested that they were shorter than those at Furry Creek. Analyses of variance detected significant differences in mean length of F. gardneri among stations in June, July and August, but these differences were not clarified by the SNK test (Figure 2.6, Table 2.3). This finding is similar to that for F. gardneri cover (section 2.4.2); there was a trend toward shorter plant lengths at some Britannia Beach stations, although this trend was less pronounced and less consistent than that for F. gardneri cover. These differences might be statistically significant if larger sample sizes had been used, but the biological significance of such differences could again be questioned. A temporal trend in plant lengths was evident in plants at Furry Creek, with a peak in plant length in July, then a steady decline into December. Thorn (1983) found a similar trend in F. gardneri in Puget Sound, Washington State, where plants were shortest in February and 46 longest inMay and June. Interestingly, Ang (1991) found thatF. gardneri (reported as F. distichus) in False Creek, British Columbia, were generally shorter than those at Furry Creek, and were longer on average in winter than in summer. The reason for this discrepancy over so short a geographical distance (-50 km) is not clear. 2.4.5. Oocyte release July and August samples yielded very low numbers of oocytes. While the specimens had the characteristics of reproductively mature plants, they may have been in the early stages of maturity. As a consequence, it is possible that not many oocytes were mature enough to be released. By October, however, the plants may have been mature enough to release oocytes. Plants at only one station released significantly more oocytes than plants at others: plants at BB-B released significantly more oocytes than plants at other stations in October (Figure 2.7; Table 2.4). This is the opposite result of that which was hypothesized. It is possible that the populations were simply undersampled, and that the result was obtained by chance. Alternatively, the population at BB-B may comprise a homogenous cohort of plants which all enter their reproductive peak simultaneously. It is difficult to support or refute either of these possibilities because no F. gardneri were found at BB-B during the December survey. Another critical consideration is the viability of the oocytes produced at stations BB-B, BB-G and BB-H. It is possible that these oocytes would not germinate in any area as a result of A M D toxicity during their pre-release development. Also, spermatozoa may be detrimentally affected by A M D , leading to a reduced reproductive capacity for the plants which is not detected by counting oocytes. Several studies reviewed in section 1.2.4.D (Chung and Brinkhuis 1986; Scanlan and Wilkinson 1987; Anderson etal. 1990) raise the possibility that microscopic algal 47 propagules may be very sensitive to copper. There are no conclusive differences among the stations which support the original hypothesis that plants near Britannia Beach would produce fewer oocytes. However, this does not rule out other possible reproductive effects. 2.4.6. Copper body burden in Fucus gardneri Plants at all three stations near Britannia Creek had significantly higher body burdens of copper than those near Furry Creek (Figure 2.8, Table 2.5). The term "body burden" is used here to describe the amount of copper which has been internalized by the plant, as well as metal which has been strongly bound outside the thallus. The plants Were rinsed in seawater, as opposed to more aggressive chemical treatments, to remove any metal which is loosely associated with the plant, but to leave metal which is more strongly bound. The weaker rinse was employed for two reasons: (1) to make results comparable with literature values, and (2) copper does not necessarily need to be internalized to exert effects on organisms. Sorentino (1979) cites studies which show that low concentrations of copper can bind cell proteins and affect the permeability of the cell membrane. This suggests that external copper should be considered along with internal copper. There have been numerous studies of metal content in Fucus spp. (Table 2.7), although most have examined habitats in Europe and fucoid algae other than F. gardneri. The levels of copper in this study agree well with those measured by Dunn etal. (1992) in Howe Sound. The slightly higher levels of copper Dunn et al. (1992) found at most sites may be partly attributed to the higher concentration of dissolved copper which would likely have been present at the time they collected, as A M D from the 4100 mine level was being discharged directly into Britannia Creek. Overall, the copper concentrations found in F. gardneri in Howe Sound appear to be 48 Table 2.7. Concentrations of copper in Fucus spp. mdAscophyllurn spp., both fucoids. All 1 2 studies rinsed algae in seawater before analysis. From: Forsberg etal. 1988, Sbderlunde? al. 1988, 3Pedersen 1984, 4Barreiro et al. 1993, 5Bryan 1983, 6Foster 1976, 7R\get etal 1995,8Ho 1984, 9Morris & Bale 1975,10Dunn et al 1992, nCurrent study. Organism ppm Cu in dry wt. Location Comments F. vesiculosus 4.0-6.5 1 Baltic Sea Unpolluted, exposed sites F. vesiculosus 5.7-17.3 1 Baltic Sea Sheltered, more polluted sites F. vesiculosus 1.6-9.8 2 Baltic Sea Expansion of sampling from 1 F. vesiculosus 5.0-9.8 3 Norway West coast fjord F. vesiculosus 2-8 4 Spain Lower estuary, mine water-polluted F. ceranoides 30-60 4 Spain Upper estuary, mine water-polluted F. vesiculosus 4-37 5 England Other estuaries F. vesiculosus 293 5 England Fal Estuary (mine water-polluted) F. vesiculosus 7.4-10 6 England Menai Straits A. nodosum 6-18 6 England Menai Straits F. vesiculosus 1.30-3.30 7 Greenland Unpolluted site F. vesiculosus (18) 8 England Tamar Estuary (unpolluted) A. nodosum (26) 8 England Tamar Estuary (unpolluted) F. vesiculosus (534) 8 England Fal Estuary (mine water-polluted) A. nodosum (476) 8 England Fal Estuary (mine water-polluted) F. vesiculosus 3.82-14.3 9 England Bristol Channel F. gardneri >300-96010 Howe Sound Near Britannia Beach F. gardneri 60-70 1 0 Howe Sound Lower Howe Sound F. gardneri 209-803 1 1 Howe Sound Near Britannia Beach F. gardneri 30.3-56.2 1 1 Howe Sound Near Furry Creek substantially higher than those found in similar seaweeds in Europe. This is likely due in part to higher levels of copper in Howe Sound water relative to other places. However, F. gardneri may not be directly comparable to the species examined in Europe, as it may accumulate more metal from similar concentrations in water. As well, differences in salinity, pH and other parameters affecting copper speciation and uptake may differ in the areas considered. Uptake of copper by this alga is further examined in Chapter 3. While bulk collection of algae without consideration of age of tissue may introduce some errors, several studies (Forsberg et al. 1988; Soderlund et al. 1988; Carvalho et al. 1997) have 49 found no consistent difference between copper levels in growing and older tissues. It seems that a sample incorporating tissues of a variety of ages is a valid method for obtaining an average body burden of copper. 2.4.7. Statistical Methods Despite some deviations from normality and equality of variances among samples, parametric statistical procedures were employed throughout this chapter. Analyses of variance and Student-Newman-Keuls tests are quite robust to violations of assumptions (Zar 1996, Underwood 1997). Underwood (1996) makes a strong argument for applying the precautionary principle in environmental monitoring. This principle, in the context of this study, is applied in trying to balance the probabilities of Type I and Type II errors (see section 1.2.6). Non-parametric procedures would reduce the probability of making a Type I error because they make fewer assumptions about the populations being examined. However, non-parametric procedures are inherently less powerful than parametric tests (Zar 1996). Additionally, the non-parametric equivalent of analysis of variance, the Kruskal-Wallis test, requires equal variances in all populations, an assumption which was sometimes violated in the above experiments. Parametric tests were employed in all analyses above to maximize the power of the analyses. In numerous tests, the A N O V A detected significant differences, but the post-hoc test failed to elucidate them. The only data in which consistent patterns were detectable using statistical tests was the higher cover of filamentous green algae at stations BB-C and BB-F than at all other sites. Given the distinct, visible nature of this phenomenon upon qualitative observation of the beach, it seems unlikely that the pattern detected is an artifact of the statistics employed. Indeed, if any problem may be suggested, it is that the power of the methods used 50 was insufficient to detect more subtle differences which may exist in all community parameters. This does not appear to be a problem in the statistical methods themselves, but rather in the survey methods employed. Due to the patchiness of the intertidal at the scale examined, larger and more quadrats would likely have reduced the standard errors of the measurements taken. 2.5. Conclusions Dissolved copper concentrations were extremely elevated in the immediate vicinity of Britannia Creek, but fell off quickly within 200-300 m. There were also elevated concentrations of copper further to the north of the Creek, up to 1000 m away. A series of hypotheses were presented in section 1.3, and the first five of these hypotheses are addressed in this chapter. F. gardneri is completely absent on 1000 m of coastline to the north of the Creek and 600 m south of the Creek. At the fringes of this zone, however, no differences were detected in mean length of F. gardneri or in percent cover between Britannia Beach stations and reference sites at Furry Creek. Thus the results support hypotheses 1 and 3 in this zone of F. gardneri absence, but not elsewhere. Filamentous green algae were sparse very close to Britannia Creek, and reached very high cover levels farther from the Creek where F. gardneri was absent. Cover of these green algae was much lower at sites where F. gardneri is abundant. Hypothesis 2, therefore, appears to apply to sites within 100 m of Britannia Creek, but the opposite situation—higher cover of green algae at Britannia Beach sites than at Furry Creek—exists 300-700 m from Britannia Creek. F. gardneri plants near Britannia Beach did not produce fewer oocytes than those at reference stations, but copper burden of F. gardneri tissues were elevated in relation to those at reference sites. Hypothesis 4 is therefore not supported by the above experiments, while Hypothesis 5 is strongly supported by the very 51 high levels of copper in algal tissue. No indisputable causative links can be drawn between any of these observations and A M D from Britannia Mine. However, all the evidence in this study suggests that A M D plays a substantial role in producing the above results. The causative role of A M D in F. gardneri distribution will be examined using results from transplant experiments in Chapter 3. With regard to the Department of Fisheries and Oceans' study of salmon productive capacity, the above findings have important implications. Assuming that there is a strong dependence of juvenile salmon onF. gardneri and its associated fauna, A M D emanating from the Britannia mine may be severely or totally eliminating this food-organism habitat. Effects of A M D on the algal communities farther from the Creek appear to be limited to enrichment of metal content. This does not rule out the potential for direct effects of A M D on the fish or on their food organisms. Another potential ramification of A M D could be vertical food chain transfer of metals from algae through their food organisms to salmon. While there is a possibility of biomagnification, Kay (1984) and Campbell et al. (1988) conclude from literature reviews that this phenomenon is quite rare for copper. In any case, there appears to be a strong probability that A M D reduces salmon productive capacity in the Britannia Beach area. 52 3 . T R A N S P L A N T A T I O N O F FUCUS GARDNERI T O B R I T A N N I A B E A C H 3.1. Introduction Chapter 2 examines the communities of algae at Britannia Beach and compares them to reference areas near Furry Creek. The most striking feature of the Britannia Beach area is the complete absence of Fucus gardneri Silva, the dominant species at nearby sites in upper Howe Sound. Combined with elevated levels of copper in plants <1000 m from the Creek and the dispersion pattern of Acid Mine Drainage (AMD) from the Creek, these observations suggest that A M D in the Creek may be responsible for the atypical algal community structure. This hypothesis, however, is based only on the correlation of community structure with copper concentrations in seawater. To better establish a causative link between A M D and F. gardneri distribution, experimental manipulation of the plants is necessary. While experimental manipulation can only be carried out to a very limited degree in the field, one way of examining the effects of A M D on F. gardneri is to move previously unaffected plants into the zone of A M D influence and observe changes in certain ecological parameters. This approach was taken in this study by transplanting adult and juvenile F. gardneri from near the mouth of Furry Creek to several sites on either side of the mouth of Britannia Creek and measuring their growth rates, survival, percent cover and copper body burden. 3.2. Methods 3.2.1. Transplant of Adult Fucus gardneri Five adult transplants were conducted over an 18 month period on the following dates: 53 June 23 1997, November 2 1997, February 15 1998, June 12 1998 and August 10 1998. The beach near Furry Creek has abundant growth of F. gardneri, typical of much of Howe Sound, and was selected as a source area for plants to move to Britannia Beach. It was also the recipient site for control plants, which were transplanted in the same manner as those moved to Britannia Beach, but were moved to other sites at Furry Creek. Much of the F. gardneri near Furry Creek, especially at stations FC-C, FC-D, FC-E and FC-F (see Figure 2.2), grew on rocks 5-35 cm in diameter. These rocks provided useful experimental units whereby an entire rock could be moved to the appropriate site with its associated plants. This largely avoided the possibility of damaging the plants during their removal and re-attachment, an alternative transplant method. It could also arguably make the results more generalizable to natural, unmanipulated populations of F. gardneri, as the cobble provides a micro-environment to which the plant is normally exposed. Rocks with F. gardneri growing on them were selected haphazardly in pairs at the source area. An attempt was made to select plants from over their entire'vertical range (typically 1.5-3.0 m above chart datum), as well as over a 100-200 m horizontal area to make results more generalizable to the entire population. A trade-off was necessary in the size of rocks to make them small enough to carry by hand, but large enough so as not to be swept away by wave action after transplant. Experience showed that, for these particular sites, rocks 15-25 cm in diameter usually fulfilled both requirements. Rocks were also selected based on percent cover of F. gardneri; a reasonably high (>80%) cover was deemed necessary. Finally, rocks were selected in pairs, where each member was similar in size (± 5 cm), percent cover and average plant length of F. gardneri (visual estimate), and within 1 m of each other in their original position on the gently sloped beach. The purpose of this pairing was to justify using paired 54 statistics in analyzing results, which is generally a more powerful method than unpaired statistics, and requires fewer assumptions about the data. The elevation from which each rock was taken was estimated using a leveling pole to measure vertical distance from the water line, then relating this measurement to predicted tides (Canadian Hydrographic Service 1997, 1998). Selected rocks were tagged with numbered flagging tape and grouped on the beach. Rocks were then randomly assigned to treatments (i.e., transplant sites) so that one member of each pair was moved to a site near Britannia Beach, while the other was moved to the appropriate control site near Furry Creek. Depending on the time of year, a boat or van was used to move the rocks to Britannia Beach. In June 1997, June 1998 and August 1998 rocks were moved in the bottom of a small, open boat. Due to the need to work at night in November 1997 and February 1998, rocks were placed in a van and driven 5 km to Britannia Beach. Rocks were moved from the recipient site to the control site at Furry Creek by carrying them along the beach by hand or on a wooden platform. The sites chosen for the transplant experiments are shown in Figure 3.1. The experimental sites to which the rocks were moved were selected based on a number of criteria. The first was the presumed gradient of A M D effects with distance from Britannia Creek. Once an appropriate site was chosen, the location on the beach was chosen to fall at the mean elevation of all rocks' original positions at Furry Creek. The site was also selected for its substratum composition, so that it would be possible to dig a hole in which to place the rocks. Control sites were then chosen at Furry Creek so that they and the Britannia Beach sites were similar distances from their respective creeks. Ice cream pails, 5.7 L and 22 cm in diameter, were used as a base in which to place the rocks. The pails could be dug into the substratum much deeper than the rocks 55 Figure 3.1. Transplant sites at Britannia Beach (left) and matching sites at Furry Creek (right). See Figure 1.1 for locations of creeks in Howe Sound. 1 and 2: November 1997; 3: February 1998; 4 and 5: June 1998; 6 and 7: August 1998. Sites BB-1 and FC-1 were -50 m from the mouths of Britannia and Furry Creeks, respectively. Sites BB-2, BB-3, FC-2 and FC-3 were all -100 m from their respective creek mouths. 1 cm = 350 m. From Canadian Hydrographic Service Chart #3526. alone, which helped to prevent movement and loss of the transplants. The pail also served to elevate the rock somewhat from the substratum to minimize the effect of sand scouring. A possible drawback of digging holes in which to place pails is that this may expose copper-containing sediments. Metals in the these deeper sediments may be highly bioavailable due to the anoxic state of the sediment. However, the pails likely shield the plants from most of this metal, and the deep sediments are re-covered by surface sediments after the first tidal inundation. Rocks and pails were placed in a group at the site, with the greatest horizontal distance between any two rocks being 5 m. 56 3.2.2. Measurements Plants were monitored for several parameters at varying intervals. A series of measurements were taken on each rock during each data collection. Percent cover of F. gardneri was measured using a 25 cm x 25 cm quadrat. The quadrat was pre-strung with nylon string in a grid pattern (with squares 1.5 cm x 1.5 cm), and 20 of the intersection points had been marked with pieces of wire. After haphazardly placing the quadrat on a rock, the object immediately under each point was recorded as F. gardneri or "other." Percent cover measurements were taken only on rocks from which no plants were taken for copper content analysis. Survivorship of F. gardneri was determined by monitoring tagged plants. Six plants were tagged on each rock during the first data collection, and their presence or absence was scored at each subsequent data collection. Growth rate of F. gardneri was estimated by measuring the length of the six tagged plants from the holdfast to the end of the longest frond. This provided an estimate of the average rate of growth over the entire interval between measurements. 3.2.3. Copper body burden Copper body burden of copper in F. gardneri thalli was measured in two or three plants from each site. The plants were plucked from the rock, placed into plastic bags and returned to the laboratory. They were rinsed three times with seawater from the West Vancouver Laboratory system and frozen at -20°C. When all samples had been collected they were dried in a fumehood at ~40°C for ~4 hours and sent to the Analytical Chemistry Laboratories of the Geological Survey of Canada (Ottawa, Ontario). They were digested in 20 mL concentrated nitric acid and 2 mL perchloric acid, reduced to near dryness and then additionally digested with 10 mL concentrated hydrochloric acid, 10 mL concentrated hydrofluoric acid and 2 mL concentrated 57 perchloric acid. Metal analysis was conducted by inductively-coupled plasma mass spectrometry (ICP-MS). 3.2.4. Data manipulation and statistical analysis Survivorship scores estimated the proportion of plants surviving on each rock. Growth rate was averaged over all plants on a given rock. Individual plants could not be counted as replicates for either growth or survivorship because they are not independent of each other; growth and survival of any given plant may be correlated with that of others due to biological interactions or genetic relatedness. Percent cover, proportion of tagged plants surviving and growth rates were then compared by paired or unpaired one-tailed Student's t-tests, as noted in the results. Copper body burden was compared using unpaired one-tailed Student's t-tests. Direction of one-tailed hypotheses was determined by the predictions made in section 1.3. The end point for the first four transplants was determined by mortality of F. gardneri plants; almost all plants at one or both of the Britannia Beach and Furry Creek sites had died and detached by the final time point shown in the results. The end of the August 1998 transplant occurred when most of the rocks at all sites could not be located, apparently due to a combination of wave action and shifting of the substratum. 3.3. Results Several difficulties were encountered during the June 1997 experiment. Plants were moved to the north and south sides of Britannia Creek, but within two weeks the transplants on the north side of the Creek were completely destroyed by human disturbance or some other physical disturbance. These samples therefore provided no useful data. Furthermore, control plants at Furry Creek were not transplanted in a manner similar to those at Britannia Beach, but 58 rather were picked up and replaced in their original location. This presents serious difficulties in separating transplant effects from those of A M D . The data from this transplant are therefore not presented here. It should be said, however, that the results observed were quite similar to those seen in the November 1997 transplant. 3.3.1. Percent Cover Results of percent cover measurements for transplanted F. gardneri are shown in Figures 3.2 through Figure 3.5. Results of statistical comparisons are shown in Table 3.1. The November 1997 and February 1998 experiments show similar trends of a rapid loss of F. gardneri cover on rocks transplanted to Britannia Beach. This decrease in percent cover becomes evident about 30 days after the transplant. In contrast, the June and August 1998 experiments show no significant differences between cover of F. gardneri on rocks transplanted to Britannia Beach and that on rocks moved to Furry Creek; percent cover decreased on rocks at both sites. At site FC-5, percent cover of F. gardneri appears to be lower than that at site BB-5. This difference is not detected by the statistical tests, however, as one-tailed t-tests were used. 59 0 20 40 60 0 20 40 60 Days post-transplant Days post-transplant Figure 3.2. Percent cover of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 7 rocks. 100 > O O $ 40 20 40 Days post-transplant 60 Figure 3.3. Percent cover of Fucus gardneri on rocks transplanted February 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 6 rocks. 60 100 CD > o O $ 40 100 20 40 60 Days post-transplant 80 20 40 60 Days post-transplant 80 Figure 3.4. Percent cover of Fucus gardneri on rocks transplanted June 1998. BB - Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 8 rocks except day 75, when n = 7 rocks at site 4. 100 CD > O U 40 50 100 Days post-transplant -1 £ 40 50 Days post-transplant 100 Figure 3.5. Percent cover of Fucus gardneri on rocks transplanted August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 8 rocks until day 35, n = 7 thereafter at site 6. All rocks but one lost at FC-6 on day 100. 61 Table 3.1. Statistical analyses of percent cover results from all transplants. All analyses use one-tailed, paired Student's t-tests to compare percent cover oi Fucus gardneri on rocks transplanted to Britannia Beach with that on rocks transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant differences are in bold type. November 1997 Feb 1998 June 1998 August 1998 Time Site 1 Site 2 Time Site 3 Time Site 4 SiteS Time Site 6 Site 7 4 >0.500 >0.500 0 0.388 2 >0.500 >0.500 1 >0.500 0.292 11 >0.500 >0.500 22 0.128 18 >0.500 >0.500 14 0.366 >0.500 18 >0.500 0.339 34 <0.001 29 0.270 >0.500 24 0.483 >0.500 25 >0.500 0.291 48 <0.001 39 0.313 >0.500 35 >0.500 >0.500 39 0.167 <0.001 46 0.500 >0.500 44 >0.500 >0.500 57 <0.001 <0.001 54 0.358 >0.500 54 0.482 >0.500 63 0.476 >0.500 61 0.384 >0.500 75 0.380 >0.500 74 100 >0.500 >0.500 >0.500 3.3.2. Survivorship Results of survivorship measurements for transplanted F. gardneri are shown in Figures 3.6 through 3.9. Results of statistical comparisons are shown in Table 3.2. During the August 1998 transplant, tagging strings were poorly attached to the base of the plants and were observed to be falling from the plants, resulting in excessively high mortality estimates. Plants were therefore re-tagged on day 24 and previous data were disregarded. The same general trends are seen in survivorship data as in the percent cover data. Plants at sites BB-1, BB-2 and BB-3 begin to detach after approximately 30 days during the November 1997 and February 1998 experiments, while plants at the control Sites maintain survivorship >75%. During the June and October 1998 experiments, plants at all sites sustain heavy mortality so that survivorship is <25% and <40% during June and October, respectively. 62 0 20 40 Days post-transplant 60 0 20 40 Days post-transplant 60 Figure 3.6. Survivorship of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error. n= 10 rocks. bO C > C o '•S o O H o (-1 PM 20 40 Days post-transplant 60 Figure 3.7. Survivorship of Fucus gardneri on rocks transplanted February 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 6 rocks. 63 0 20 40 60 80 0 20 40 60 80 Days post-transplant Days post-transplant Figure 3.8. Survivorship of Fucus gardneri on rocks transplanted June 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks except day 75, when n = 9 at site 4. 0 0 20 40 60 Days post-transplant 80 20 40 60 Days post-transplant 80 Figure 3.9. Survivorship oi Fucus gardneri on rocks transplanted August 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean ± 1 standard error, n = 10 rocks. 64 Table 3.2. Statistical analyses of survivorship results from all transplants. All analyses use one-tailed, paired Student's t-tests to compare survivorship of Fucus gardneri on rocks transplanted to Britannia Beach with that of individuals transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant differences are in bold type. November 1997 Feb 1998 June 1998 August 1998 Time Site 1 Site 2 Time Site 3 Time Site 4 Site 5 Time Site 6 Site 7 11 0.172 >.500 22 0.093 18 0.065 >0.500 35 >0.500 >0.500 18 0.041 >.500 34 0.005 29 >0.500 >0.500 44 0.486 >0.500 25 0.296 >.500 48 <0.001 39 0.453 >0.500 54 >0.500 0.380 39 0.024 <0.001 46 >0.500 >0.5()0 61 >0.500 0.071 57 <0.001 <0.001 54 >0.500 >0.500 74 0.463 0.167 63 >0.500 >0.500 75 >0.500 >0.500 3.3.3. Growth Results of growth rate measurements for transplanted F. gardneri are shown in Figures 3.10 through 3.12. Results of statistical comparisons are shown in Table 3.3. Time points are expressed as the mid-point of the interval over which the growth rate is calculated (see Appendix II for further explanation). Growth rates were not measured during the February 1998 transplant. Growth rates of F. gardneri transplanted to BB-1 and BB-2 during November 1997 were significantly lower than those of control plants moved to FC-1 and FC-2 during all measurement intervals except the last interval at site 2. Plants transplanted to BB-4 had lower growth rates than control plants during 4 of 7 measurement intervals, while plants moved to BB-5 (farther from Britannia Creek) grew slower during 2 of 6 measurement intervals. Only one difference was detected between growth rates at the two Creeks in the August 1998 transplant, with growth being faster at FC-6 than at BB-6 during the fifth interval. 65 1.5 1.0 m/d) 0.5 £ -»-» 0.0 cd Xi -0.5 o O -1.0 -1.5 10 20 30 40 Days post-transplant 10 20 30 40 Days post-transplant Figure 3.10. Growth rates of Fucus gardneri on rocks transplanted November 1997. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 10 rocks. Figure 3.11. Growth rates of Fucus gardneri on rocks transplanted June 1998. BB = Britannia Beach. FC = Furry Creek. See Figure 3.1 for locations of sampling sites. Data are mean + 1 standard error, n = 10 rocks except day 75, when n = 9 at site 4. 66 -0.4 0 20 40 60 80 100 Days post-transplant Figure 3.12. Growth rates of Fucus gardneri on Britannia Beach. FC = Furry Creek. See F are mean ± 1 standard error, n = 10 rocks. 0.8 0.6 -0.4 1 — I 0 20 40 60 80 100 Days post-transplant rocks transplanted August 1998. BB = igure 3.1 for locations of sampling sites. Data Table 3.3. Statistical analyses of growth rate results from all transplants. All analyses use one-tailed Student's t-tests to compare growth rates of Fucus gardneri on rocks transplanted to Britannia Beach with that of individuals transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant differences are in bold type. Tests marked with * were conducted using unpaired t-tests. All other tests were paired t-tests. November 1997 Time Site 1 Site 2 Time June 1998 Site 4 Site 5 Time August 1998 Site 6 Site 7 8 <0.001 0.039 10 0.001 >0.500 8 0.094 0.217 15 0.001 0.001 24 0.003 0.006 19 >0.500 0.269 22 0.009 <0.001 34 0.057 0.001 30 >0.500 >0.500 32 <0.001 0.141 43 <0.001* 0.398* 40 0.295 0.084 50 0.099* 0.434* 49 0.018 0.410 59 >0.500* 0.136* 58 >0.500 >0.500 69 0.019* 68 >0.500 0.239 87 >0.500 67 3.3.4. Copper body burden Copper body burden measurements were done only during the November 1997 transplant (Figure 3.13). The results of statistical analyses are shown in Table 3.4. Body burden of copper increased very rapidly during the first two weeks after being transplanted to the mouth of Britannia Creek. The rate of uptake began to decrease at this point, but copper was still being taken up 39 days after the transplant, after which no plants were available to be sampled. There was no apparent change in the copper body burden of plants at Furry Creek. 0 10 20 30 40 Days post-transplant Figure 3.13. Copper body burden of Fucus gardneri on rocks transplanted November 1997 to BB-1 and FC-1. BB = Britannia Beach. FC = Furry Creek. Data are mean ± 1 standard error, n = 3 plants, except day 39, when n = 2. 68 Table 3.4. Statistical analyses of copper body burden from the November 1997 transplant. All analyses use one-tailed Student's t-tests to compare copper body burden of Fucus gardneri transplanted to Britannia Beach with that in plants transplanted to Furry Creek. See Figure 3.1 for site locations. Time is days after transplant, a = 0.05 in all tests. Significant results are in bold type. Nov. 1997 Time Site! 11 0.001 18 <0.001 25 <0.001 39 0.001 57 0.014 3.3.5. General observations Some changes in the condition of the plants became apparent after they were transplanted. The most obvious change occurred in the colour and texture of plants which were moved to the Britannia Beach area. After 2-30 days of exposure to A M D , the entire thallus took on a reddish brown colour which contrasted remarkably with the olive green colour of plants at Furry Creek and other sites. The plants at Britannia Beach also produced large amounts of exudates, resulting in a layer of slime on the outside of the plant. Concurrently, the thalli of plants at Britannia Beach had a very soft texture relative to plants at Furry Creek. These changes in plant appearance and texture became apparent at different times: 2 days post-transplant for all plants in the June 1997 experiment, 14 days for all plants in the November 1997 experiments and 30 days for the plants at site 4 for the June 1998 transplant. Plants at site 5, 6 and 7 showed no such changes at any time. 69 3.4. Discussion The results of the transplant experiments showed two distinct patterns. The discussion is organized based on these patterns. 3.4.1. Severe Effects: November 1997 and February 1998 F. gardneri transplanted to the immediate vicinity of Britannia Creek in November 1997 showed no significant change in percent cover (Figure 3.2) or survivorship of tagged plants (Figure 3.6) for the first month after transplant. After this time, however, cover and survivorship drop off rapidly; by 57 days after the transplant almost all tagged plants had detached. The cover found on the last day was composed of holdfasts and stipes without fronds. These tissues are capable of neither reproduction nor growth. An identical pattern was seen in the cover (Figure 3.3) and survivorship (Figure 3.7) of the plants transplanted in February 1998. No differences between plants moved to Britannia Beach and Furry Creek are significant at a = 0.05 until more than 30 days post-transplant. Growth rates in the November 1997 transplant showed a more immediate onset of effects among the plants moved to Britannia Beach than is revealed in cover or survivorship. The first set of growth rate measurements, averaging growth from day 4 to day 11, showed that plants moved to Britannia Beach had significantly lower growth rates than those moved to Furry Creek (Figure 3.10; Table 3.3). While the growth rates are variable at different time points, the difference between growth at the two sites remains fairly consistent over the entire experiment. A notable feature of growth rates at both Britannia Beach sites in November 1997 is that they are negative or zero at all but one time point. This is not an artifact of plant breakage, as all detached plants were removed from the growth rate data set. Rather, the decrease in plant length appeared 70 to be the result of plant erosion. Concurrent with the change in colour and texture of the plants (section 3.3.5), the tips of the receptacles were clearly being worn away, presumably due to a combination of poor plant health and wave action. The implication of this phenomenon for growth rate measurements is evident: the length of the plant will become shorter at each successive sampling date. Since F. gardneri grows from an apical meristem, removal of tissue from the plant's tips will dislodge the meristem, precluding further growth. It appears that the growth rate of F. gardneri responds more rapidly to A M D than percent cover or survivorship. In particular, the use of tagged plants is a relatively conservative measure of their survivorship because a plant may die from toxicity quite quickly but remain attached to its substratum for a period of time before its holdfast or stipe gives way. Percent cover gives a very crude estimate of existing biomass, but like tagged plants, gives no information on the viability or health of this biomass. Periodic collection of plants for copper content analysis provided an opportunity to examine the speed with which plants would take up the metal when exposed to high concentrations and how the levels of metal at these sites compared with concentrations in unmanipulated plants farther from Britannia Beach. The results (Figure 3.13) clearly showed large increases in the copper body burden of individuals transplanted to Britannia Beach. Furthermore, the uptake was very rapid in the first 11 days, after which it appeared to slow considerably. The concentration was still increasing up to day 39, after which no samples could be collected due to the death of all plants at Britannia Beach. When compared to copper levels in existing F. gardneri at sites near Britannia Creek (Figure 2.8; 500-700 ppm in dry wt. in the plants nearest the Creek), the transplants had 2-3 times more copper after just 11 days. Given that the levels of copper found in the extant plants nearest to Britannia Creek likely represents an . 71 approximate maximum that the plants can withstand, severe degradation of the plants' health at >2000 ppm copper is expected. Some work has been done involving the transplantation of algae into areas with relatively high copper concentrations. Ho (1984) moved Fucus vesiculosus and Ascophyllum nodosum, two fucoid algae, from an unpolluted site in Cornwall, England, to Restronguet Creek, which was contaminated with zinc and copper from past mining activities. Ho (1984) observed copper concentrations of <20 ppm in control F. vesiculosus, while those moved to the polluted area contained 100, 150 and 230 ppm copper in dry weight after 15, 35 and 57 days, respectively. A. nodosum contained 60, 60 and 100 ppm copper at the same time intervals, while control plants contained <35 ppm copper. These amounts of copper are approximately one tenth of those seen in plants that were moved to Britannia Beach from Furry Creek, presumably due to lower concentrations of available copper (not reported) at Ho's (1984) site. However, interspecific differences are apparent within Ho's experiment; these differences may also be important when comparing Pacific F. gardneri with Atlantic F. vesiculosus. Forsberg et al. (1988) moved F. vesiculosus from Bjorkskar, an unpolluted site in Sweden, to Lordan, a relatively polluted site near Stockholm. After 12 months, there was no significant change in copper concentration in young tissues at Lordan relative to untransplanted controls, but there was significantly more copper in older tissues at the more polluted site (15.6 ± 1.0 ppm copper in dry weight at Lordan [mean ± standard deviation], 5.7 ± 4.1 ppm copper at Bjorkskar). Ambient concentrations of copper in the two areas were not reported. The change in colour and texture of the plants soon after transplant to the mouth of Britannia Creek seemed to be an indication of the quickly declining health of the plants. The change in colour is consistent with Sorentino's (1979) finding that very high concentrations of 72 copper cause degradation of chlorophyll and other pigments in some algae, which would lead to a change in colour. A change in plant texture may be a result of copper binding cell proteins such as those which maintain the plant's structure and which control the osmotic permeability of the cell (Sorentino 1979). The increased production of exudates may be one way the plant has of detoxifying the metals (Ragan et al. 1980; Schramm 1993). 3.4.2. No Detectable Effects: June and August 1998 Unlike the previous sets of transplant experiments, those conducted in June and August of 1998 to areas more distant (300-700 m) from Britannia Creek showed no consistent pattern of deleterious effects of A M D . In 33 comparisons using paired one-tailed t-tests, the null hypothesis of equal percent cover of Furry Creek and Britannia Beach transplants was never rejected (Table 3.1), while the null hypothesis of equal survivorship at Furry Creek and Britannia Beach was also never rejected (Table 3.2) in 24 comparisons. In fact, a quick glance at the graphical data suggests that percent cover and surviorship of F. gardneri may be lower at Furry Creek, at least at sites 4 through 7 for certain time points (Figures 3.4, 3.5, 3.8 and 3.9). This result is quite unexpected given the high cover and survivorship of plants at Furry Creek relative to those at Britannia Beach in all previous transplants. However, there are several possible explanations for the results observed. An important consideration in the June 1998 and, to a lesser extent, the August 1998 transplant is physical factors such as desiccation. Howe Sound has a semi-diurnal tide pattern, and during the summer months the lowest tides occur at or near midday. The summer of 1998 was hot and sunny in upper Howe Sound, resulting in severe desiccation of F. gardneri thalli, sometimes on a daily basis (pers. obs.). This repeated desiccation is: tolerated by most plants, but 73 appears to result in high mortality rates in some populations. As well, the desiccation may have been aggravated by moving the plants into a different microclimate on a beach with less F. gardneri cover. This would likely have resulted in lower moisture levels in the immediate area of the plants. Combined with brisk, warm southerly winds on summer afternoons (Jackson & Steyn 1992; pers. obs.), these factors may have combined to cause severe desiccation of F. gardneri, especially at the Furry Creek sites, which had a more southerly aspect than the Britannia Beach sites. This might have contributed to greater overall mortality at Furry Creek than at Britannia Beach. Alternatively, the desiccation-induced mortality might have been about equal at Furry Creek and Britannia Beach, but very severe at both sites. This high rate of "background" mortality could then overshadow AMD-induced mortality at the Britannia Beach sites with the result that the latter would not be detected. Another possible contributing factor is natural senescence. The population of F. gardneri at Furry Creek from which the June 1998 transplants were obtained sustained very high mortality in June-October 1998; the percent cover of F. gardneri declined from 70% to 15% over 4 months (Figure 2.4). If this mortality was caused by a natural senescence of this population, the mortality of the transplanted F. gardneri would be expected to be high regardless of any effect of A M D . Yet another possibility was the relative amounts of dissolved copper in Howe Sound at different times of year. Assuming that the temporal pattern of loading from Britannia Creek into Howe Sound (Figures 1.2 and 1.3, respectively) is reasonably consistent from year to year, there might be relatively little dissolved copper present at sites BB-4 through BB-7 in June and August. If the experiments had been done during the peak in metal loadings, very different results may have been observed. 74' One observation which suggests some degree of A M D effects at one of the sites was the onset of the characteristic colour and texture changes after 30 days in plants at site J3J3-4 (see section 3.3.5). These changes were not evident at any of the other sites during the June or August 1998 transplants, at Furry Creek or at Britannia Beach. As well, the receptacles of F. gardneri at site BB-4 were clearly eroding, as those at sites BB-1, BB-2 and BB-3 had in previous experiments, while plants at all other sites showed no such tissue degradation. This erosion was evident in the growth rates observed in the June 1998 transplant, which were significantly lower at BB-4 than FC-4 at 4 of 7 intervals (Figure 3.11 and Table 3.3). This difference is not consistent among intervals, which may reflect the lower severity of A M D effects at greater distances from Britannia Creek. The various explanations discussed above do not, however, address the possibility that A M D is having a minimal effect on adult F. gardneri at sites farther from the Creek. In fact none of the data from the June and August 1998 transplants, aside from qualitative observations of plant health, provide strong evidence that A M D affected adult F. gardneri growth or survivorship at the sites examined. This was not an entirely unexpected result, as copper concentrations at three of these sites (sites BB-5, BB-6 and BB-7 correspond to stations 6, 8 and 9 in Figures 2.2 and 2.3) are often <l-20% of those at the mouth of Britannia Creek (sites BB-1 through BB-3). However, the lack of transplant data from other seasons and the confounding factor of desiccation prevent any strong conclusions from being drawn from the results. While there is no evidence that A M D can adversely affect adult F. gardneri at sites, this does not eliminate the pollution as a possible causative agent for the lack of F. gardneri near Britannia Creek (see Figures 2.1 and 2.4). The findings of many authors (Chung & Brinkhuis 1986; Maier & Muller 1986; Scanlan & Wilkinson 1987; Anderson etal. 1990) suggest that the 75 microscopic reproductive stages of brown algae, including Fucus spp., may be much more susceptible than adult plants to toxins such as copper. Since F. gardneri zygotes must settle and germinate in an area to establish a viable population, any interference of A M D with this process may preclude the establishment of the algae in an area. Thus, A M D may be the major reason for the lack of F. gardneri in the entire area of absence near Britannia Creek despite the apparent lack of effects on adult plants at sites >300 m from the Creek mouth. 3.4.3. General Discussion of Transplant Experiments Due to the high temporal and spatial variability in dissolved concentrations of copper at Britannia Beach, as well as the intricacies of copper speciation in the field and in the laboratory, it is difficult to associate the transplant data with any specific exposure to copper. However, it is useful to compare the above results with previous examinations of copper effects on algae. While several authors (e.g., Ho 1984; Forsberg etal. 1988) have transplanted fucoids and measured copper content, there has apparently been no in situ work examining the effects on other biological parameters such as growth and survivorship. There have been numerous laboratory studies of copper effects on algae, however. Stromgren (1979) exposed Ascophyllum nodosum to a variety of concentrations of copper in a flowing seawater system and found that its growth rate was decreased to 80% of controls when exposed to 66 pg copper L"1 for 4 days, while growth rates of plants exposed to 340 pg copper L"1 were reduced to 20% of controls after 4 days. The same author also examined the effect of copper on other fucoid algae (Stromgren 1980). He saw significant reduction of growth rate of Pelvetia canaliculata and Fucus spiralis at 12 pg copper L" 1, of F. serratus at 25 pg copper L" 1, and of F. vesiculosus at 50 ug copper L" 1. Chung & Brinkhuis (1986) found that release of meiospores from Laminaria saccharina was 76 reduced by >50 ug copper L" 1. Settlement and germination of the meiospores were not affected by any of the concentrations tested (<500 u.g copper L"1). However, development of the microscopic gametophytes was delayed in >50 \ig copper L" 1, and growth of sporophytes was inhibited at copper concentrations >10 ug L" 1. Munda & Hudnik (1986) found 2500 ug copper L"1 at a salinity of 31.5 to be lethal to Fucus vesiculosus within 20 days. Anderson et al. (1990) observed decreases in sporophyte production in Macrocystis pyrifera at 10 pg copper L" 1, in sporophyte growth at 32 pig copper L" 1, in germ-tube growth at 18 pg copper L"1 and in percent of spores germinated at 100 ug copper L" 1. Andersson & Kautsky (1996) studied the effects of copper on Baltic Sea Fucus vesiculosus. At salinities of 6 and 20, 20 pg copper L"1 caused a 70-80% decline in germination. At a salinity of 14, which the authors recognized as the optimum for this alga, no reduction in germination was observed. When <60 ug copper L"1 was added 24 hours after fertilization, the zygotes were much more resistant to copper toxicity. Considering these previous studies, the results of transplant experiments at the mouth of Britannia Creek (sites BB-1, 2 and 3), where dissolved concentrations can locally exceed 1000 pig L" 1, are not surprising. The levels of copper at other sites (BB-4 through 7; <50-200 ug L"1) are more in line with those examined in laboratory work. However, the bioavailability of copper in these laboratory studies is likely different from that at Britannia Beach; direct comparisons of data are therefore difficult. Andersson and Kautsky's (1996) findings thatF. vesiculosus propagules are extremely susceptible to copper toxicity—especially at low salinities—lend support to the hypothesis that, while A M D at sites 300-500 m from Britannia Creek may not affect adult plants, it may completely preclude growth of juvenile plants. 77 3.4.4. Power Analysis The calculation and implications of the power of statistical tests are discussed in section 1.2.6. The consequence of low power is that there is an inherent risk of not detecting serious environmental degradation. This does not appear to be a problem in the data from the November 1997 and February 1998 transplants, as effects were quite obvious after one month, and were verified by statistical tests. In the June and October 1998 experiments, however, there appeared to be some degradation of some plants that was not consistently detected by statistical tests. To gain some insight into these results, an a posteriori power analysis is conducted below on the growth rate analysis at BB-4 and BB-5 in June 1998. An a priori power analysis was not used in this study for two reasons. First, there was little previous data which could be realistically used to estimate biologically significant effect sizes or variability in the population. Second, there was a practical limit on the number of samples which could be monitored to a sufficient degree to give meaningful results. Instead, an alternative approach is taken here, which is to measure in an a posteriori manner the power of tests already conducted. The analysis can be conducted with several different sample and effect sizes to look for weaknesses in the analyses. To remain consistent with most current ecological work, a is set at 0.05, but 0.10 will also be considered. Assuming that 0.05 is an "acceptable" rate of error, a value of (3 <0.05, and therefore a power >0.95, will also be deemed "acceptable." The standard deviation of growth rates in this experiment varied from 0.20 to 0.50 mm d"1; a relatively conservative value of 0.40 mm d"1 will be used in this analysis. Sample sizes were 10 at each site, but n = 20 will also be considered. Three effect sizes (i.e., mean differences in growth rate between F. gardneri 78 transplanted to Furry Creek and Britannia Beach), which are representative of those observed in the June 1998 experiments, will be used: 0.20, 0.35 and 0.50 mm d"1. Table 3.5. Power of the statistical analyses of growth rate measurements during the June 1998 transplant. Power values are calculated using equation 3.1, with s = 0.40 mm d"1 and the 6, n and a values shown. All results of the current experiments were analyzed with n = 10 and a = 0.05. Power analyses using other parameters are shown for comparison. n a 6 (mm/d) 0.20 0.35 0.50 10 0.05 0.10 <0.500 0.687 0.937 <0.500 0.813 0.968 20 0.05 0.10 0.556 0.958 >0.999 0.691 0.979 >0.999 It is clear that the statistical power of this particular set of tests was low. The effect size (the amount by which growth rates at Britannia Beach are less than those at Furry Creek) would have to be relatively large (>0.50 mm d"1) before it would be reliably (power > 0.95) detected using the methods described. Larger sample sizes would make smaller effect sizes (> 0.35 mm d"1) apparent. Increasing a to 0.10 appears to have relatively little effect on power in this test, and is not a desirable option as it increases the chance of a Type I error. The ability of these experiments and statistics to detect subtle differences (0.20 mmd"1) in growth rates of F. gardneri appears to be extremely low. The low statistical power of the methods used must be kept in mind when examining the results of these experiments; the lack of detectable differences in the June 1998 transplant may be more a function of the methods used than of an actual response of the algae. Furthermore, the lack of power may be even more important in the August 1998 experiment, where there is much more variability in the data (error bars in Figures 3.11 and 3.12). 79 3.5. Conclusions All of Hypotheses 6, 7 and 8 (see section 1.3) are supported by experiments very close to Britannia Creek, but experiments farther from this creek do not support any of the hypotheses. The results of these experiments provide strong evidence for the deleterious effects of A M D on F. gardneri growth rates, cover and survivorship at the high concentrations within 100 m of the mouth of Britannia Creek. No effects are apparent at greater distances (>300 m), but these latter observations are confounded by other effects which were unfortunately inherent in the timing and design of experiments. However, evidence from the literature suggests that A M D may affect reproductive stages of F. gardneri more severely than adult plants, resulting in exclusion of the plants from areas of high A M D concentrations. The results of the transplant experiments build on the correlative evidence of A M D effects on F. gardneri provided by the measurements described in chapter 2. The use of experimental evidence adds weight to the hypothesis that A M D is at least partially responsible for the lack of F. gardneri near Britannia Creek. In terms of salmon productive capacity, A M D appears to be capable of severely limiting F. gardneri cover, survivorship and growth. Based on the assumption that salmon productivity is critically dependent on F. gardneri for the alga's associated fauna and contribution to the organic carbon pool, A M D would therefore also reduce salmon productive capacity. 80 4. G E N E R A L C O N C L U S I O N S The overall objective of this thesis is to determine if acid mine drainage is responsible for the absence of Fucus gardneri Silva in the Britannia Beach area. The observations described in Chapter 2 showed a complete absence of F. gardneri from the shores 1000 m to the north of Britannia Creek and 600 m to the south of the Creek, despite the. apparent suitability of abiotic and biotic conditions. However, there was no apparent difference between populations of F. gardneri at Britannia Beach which grow near this zone of absence and those which grow elsewhere in terms of percent cover, plant length or reproductive capacity. The plants near Britannia Creek, however, did contain up to 20 times more copper than those at Furry Creek. To examine the causative role of A M D in excluding F. gardneri near Britannia Creek, plants were transplanted into various areas near the Creek mouth and their health was monitored. In general, the health of plants transplanted to the immediate area near the mouth of Britannia Creek declined rapidly, but, when placed farther away, the plants fared no worse than controls. The results at greater distances from the Creek may be confounded, however, by the harshness of abiotic conditions (e.g., temperature, humidity) at the time of the experiment. Additionally, the statistical power of the experiments was low, raising the possibility that effects were not detectable with the sampling protocol used. The above measurements and experiments provide strong correlative evidence for the suggestion that A M D is the major determinant of algal community structure within 600-1000 m of the Creek mouth. The absence of F. gardneri is not likely the result of unfavourable abiotic factors such as turbidity, salinity or temperature as the alga is found growing in equally or even more stressful areas in terms of all these factors (see Section 2.4.2). The transplant of F. gardneri into the area where it is absent examined one further possibility: that dispersal-81 related problems can exclude the alga. Dispersal ability is irrelevant at sites BB-1 through BB-3, as plants moved there died within two months; even if propagules settled at these sites they would not survive the extremely high concentrations of A M D . The possibility of dispersal ability limiting this alga's distribution is not, however, excluded farther from the Creek. While F. gardneri would not be expected to disperse rapidly given that most of its propagules settle within 1 m of the parent plant (Chapman 1995), other studies (Burrows & Lodge 1951; Paine et al. 1996) suggest that propagules can settle >1 km from the nearest population of adult F. gardneri. Given that the total shoreline length from which F. gardneri is absent covers only about 1600 m and the variety of circulation patterns which are observed, it seems very unlikely that propagules are not being carried to the AMD-affected shoreline by currents. Acid mine drainage is the most probable feature of the Britannia Beach ecosystem which could bring about the observed distribution of algae; the combined weight of all the evidence points to A M D . The work in this thesis did not establish a model by which A M D may exclude F. gardneri in terms of life history stages. Work that attempted to elucidate this issue was unsuccessful because algal propagules could riot be cultured. Given the lack of effects found in the June and August 1998 transplants, however, it seems that the peak in metal loadings in early spring (Figures 1.2 and 1.3; Chretien 1997; McCandless, unpublished data), possibly in combination with enhanced effects of A M D on juvenile F. gardneri, is necessary to explain the observed lack of the alga at Britannia Beach. Other seasonal characteristics of metal dispersion and toxicity which occur at Britannia Beach concurrently with the loading peak would likely aggravate the toxicity of the metal. The sharp density gradient in Howe Sound during the spring freshet of the Squamish River results in the majority of copper from Britannia Creek remaining in the surface layer of the Sound (Chretien 1997). This means that intertidal organisms such as 82 F. gardneri will be exposed to more copper than if the metal was dispersing into the entire water body. Additionally, the low salinities in this surface layer (often <5; Chretien 1997; this study, not presented) during the metal peak may increase the toxicity of copper to the algae (Betzer & Kott 1969). However, it is important to recognize two important considerations. The first is that the effects seen cannot be attributed solely to the metals coming from Britannia Creek. There is also a submerged outfall at 30 m depth, approximately 50 m from the Creek mouth, which releases A M D and municipal sewage. The metal loadings in this outfall are 30-100% of those in the Creek, depending on the season (R. McCandless, unpublished data). During the spring freshet, the density gradient appears to keep this effluent below intertidal depths (Chretien 1997). However, at other times of the year, this water may rise to the surface and interact with biota. While diverting all A M D flow from the mine to the 4100 level and into the submerged outfall might partially mitigate the effects on intertidal organisms, there is no evidence for this in the present study. A more fundamental issue in the above experiments involves the true separation of causation from correlation. This can be a problem in almost any ecological investigation, including the assessment of the effects of point-source pollution. The difficulty essentially rests in the fact that, while plants were moved to Britannia Beach and exposed to A M D , they were also subjected to all other biotic and abiotic elements which the Britannia Creek estuary ecosystem might impose on them. Thus, the absence of F. gardneri from the area near Britannia Creek might not be caused by A M D but by some other characteristic of the Creek's estuarine ecosystem. In this sense, all the evidence obtained from this study is simply correlative. However, the prospect of indisputably establishing causation is quite daunting. One 83 possibility would be to conduct laboratory experiments where various life stages of F. gardneri are subjected to different mixtures of A M D and seawater and the effects observed. Even if the various technical obstacles could be overcome, it is always difficult to generalize laboratory results to the field. Field experiments that eliminate the possibility of other Creek attributes confounding the results would be even more difficult. One could argue that the only way to remove the effects of Britannia Creek itself would be to add acid mine drainage from Britannia Creek to several other freshwater streams and investigate effects on biota in their estuaries. Even setting aside the obvious ethical prohibitions, such an experiment would be completely impracticable. It was decided at the outset of this study to conduct field experiments as opposed to laboratory experiments. The inherent difficulty of mimicking field conditions in the laboratory was exacerbated in this study because of the very subtle nature of copper chemistry. Field experiments, by definition, exposed the plants to the natural conditions which may play a role in mitigating or intensifying the effects of A M D . An important sacrifice is made in terms of the conclusions, however, in that the results cannot be generalized outside of the Britannia Beach area. Furthermore, there is still a dependence on correlative evidence, however strong this correlation may be. As a general summary in relation to the Department of Fisheries and Oceans' study, the results of this work provide strong evidence for the role of acid mine drainage from the Britannia Mine in causing the absence of F. gardneri from the shores of Howe Sound near Britannia Creek. While the experiments described here did not unequivocally establish A M D as the sole cause of F. gardneri's absence, there is strong correlative evidence for this hypothesis. More definitive results could be obtained if steps were taken to abate the A M D from Britannia Mine. 84 It would be quite informative to monitor any changes in algal communities which might result from such an operation. It would be even more ideal to periodically transplant F. gardneri into the area and compare their progress to the results in this study. Experiments of this sort could largely alleviate the shortcomings of the current results. 85 References Anderson, B.S., Hunt, J.W., Turpen, S.L., Coulon, A.R. & Martin, M. i990. Copper toxicity to microscopic stages of giant kelp Macrocystispyrifera: interpoptilation comparisons and temporal variability. Mar. Ecol. Prog. Ser. 68:147-156. Anderson, D.M. & Morel, F.M.M. 1978. Copper senstivity of Gonyaulax tamarensis. Limnol. Oceanogr. 23: 283-295. 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New Jersey. 93 A p p e n d i x I — • R a w Data f r o m Chapter 2 This appendix presents a portion of the raw data from the transect studies conducted from June to December 1998: (1) percent cover of all items, and (2) oocyte release from Fucus gardneri. Percent Cover Data The methods for the collection of these data are described in section 2.2.2 and the results for F. gardneri and green algal cover are summarized, statisically analyzed and discussed in sections 2.3.2, 2.3.3, 2.4.2 and 2.4.3. Locations of sampling sites are shown in Figure 2.2. Abbreviations used in the data tables: Q # = quadrat number. Mus. = Mussel. Barn. = Barnacle. Gr.Al. = Filamentous green algae. S.E. = Standard error of the mean. The values presented are the percent cover of the specific organism or substratum type in a single quadrat. Data were collected at all stations in August - December, but stations BB-B and FC-B were not monitored in June or July. Oocyte Release D a t a The methods for the collection of these data are described in section 2.2.3 and the results are summarized, statistically analyzed and discussed in sections 2.3.5 and 2.4.5. Locations of sampling sites are shown in Figure 2.2. Abbreviations used in the data tables: S # = sample number. S.E. = Standard error of the mean. Data were collected from July to December, but samples in July and August yielded very low quantities of oocytes. Therefore, only October and December data are presented. 94 C A N O P Y C O V E R — JUNE 1998 Station BB-C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 3 5 10 55 2 0 0 0 10 0 9 0 3 0 0 0 2 0 0 8 0 4 0 0 0 10 0 9 0 5 0 0 0 4 5 0 55 6 0 0 0 15 0 85 7 0 0 5 2 0 10 6 5 8 0 0 0 3 0 0 70 9 0 0 0 2 5 0 75 M E A N 0.0 0.0 0.6 2 3 . 3 2.2 7 3 . 9 S.E. 0.0 0.0 0.6 3.9 1.5 4 .5 Station Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 5 0 0 5 0 2 0 0 0 4 0 0 6 0 3 0 0 0 8 0 5 15 4 o 0 0 75 15 10 5 0 0 0 8 0 15 5 6 0 0 0 0 0 100 7 0 0 0 9 5 5 0 8 0 0 0 7 0 2 0 10 9 0 0 0 9 5 0 ; . 5 M E A N 0.0 0.0 0.0 6 5 . 0 6.7 2 8 . 3 S.E. 0.0 0.0 0.0 10 .2 2.6 11 .4 Station BB-E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 7 0 10 15 2 0 0 0 8 0 0 0 3 0 0 0 7 0 5 10 4 0 0 0 6 5 35 0 5 0 0 0 15 0 o 6 0 0 0 7 5 0 0 7 0 0 0 15 8 5 0 8 0 0 0 6 0 0 0 9 0 0 0 6 0 0 0 M E A N 0.0 0.0 0.0 5 6 . 7 15 .0 2.8 S.E. 0.0 0.0 0.0 8.2 9 .5 1.9 9 5 CANOPY COVER — JUNE 1998 Station BB-F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 20 0 80 2 0 0 0 45 0 55 3 0 0 0 45 0 55 4 0 0 0 35 0 65 5 o 0 0 5 0 95 6 0 0 0 0 0 100 7 0 0 0 20 0 80 8 0 0 0 95 0 0 9 0 0 0 85 0 0 M E A N 0.0 0.0 0.0 38.9 0.0 58.9 S.E. 0.0 0.0 0.0 11.0 0.0 12.3 Station BB-G Q# Fucus Mus. Barn. Rock Sand Gr.Al. l 0 0 40 40 0 20 2 15 0 45 0 0 40 3 0 10 80 10 0 0 4 40 0 60 0 0 0 5 10 0 45 10 5 30 6 30 5 5 35 0 25 7 45 0 5 45 0 • 5 8 60 0 5 35 0 0 9 5 0 5 70 20 0 M E A N 22.8 1.7 32.2 27.2 2.8 13.3 S.E. 7.3 1.2 9.4 7.9 2.2 5.2 Station JrJJri-H Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 25 10 45 20 0 2 10 5 35 35 0 15 3 5 0 0 55 5 35 4 35 5 15 40 5 0 5 5 0 10 45 20 20 6 5 0 30 30 0 35 7 15 0 25 45 15 o 8 0 0 10 5 0 85 9 0 0 0 95 0 5 M E A N 8.3 3.9 15.0 43.9 7.2 21.7 S.E. 3.7 2.7 4.2 7.9 2.9 9.2 96 C A N O P Y C O V E R — J U N E 1998 Station F C - C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 6 0 2 5 10 5 0 0 2 2 0 6 0 2 0 0 0 o 3 6 5 2 5 5 5 0 0 4 5 5 2 0 2 5 0 0 0 5 9 5 5 0 0 0 o 6 35 5 0 5 10 0 0 7 2 0 0 0 2 0 6 0 0 8 2 0 75 5 0 0 0 9 4 0 6 0 0 0 0 0 MEAN 45.6 35.6 7.8 4 . 4 6.7 0.0 S.E. 8.5 8.8 3.0 2 .3 6.7 0.0 Station FC-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 2 0 7 0 10 0 2 8 0 5 15 0 0 o 3 0 0 25 4 0 10 2 5 4 0 3 0 2 5 4 0 0 5 5 5 10 3 0 5 5 0 0 6 0 0 2 5 3 5 10 3 0 7 35 0 0 6 5 0 0 8 7 0 0 10 2 0 0 0 9 6 0 5 35 0 0 0 M E A N 2 7 . 8 5.6 2 0 . 6 36.1 3.3 6.7 S.E. 11 .3 3.3 3.6 8.5 1.7 4 .0 Station F C - E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 5 5 0 15 10 0 2 0 2 9 0 0 10 0 0 0 3 15 0 15 4 5 2 0 5 4 2 0 0 4 0 2 5 10 " 5 5 8 0 5 15 0 0 0 6 8 5 10 5 0 0 0 7 3 0 2 0 5 0 0 0 o 8 1 0 0 0 0 0 0 0 9 2 0 0 0 3 5 4 5 0 M E A N 55.0 3.9 16 .7 12 .8 8.3 3.3 S.E. 11.5 2.3 5.8 5 .9 5.1 2 .2 9 7 C A N O P Y C O V E R — JUNE 1998 Station F C - F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 5 25 5 65 0 0 2 45 20 10 25 0 0 3 80 0 0 20 0 0 4 90 0 5 5 0 0 5 60 0 0 5 35 0 6 95 0 5 0 0 o 7 90 0 5 5 0 0 8 95 0 5 0 0 0 9 55 0 25 20 0 0 M E A N 68.3 5.0 6.7 16.1 3.9 0.0 S.E. 10.1 3.3 2.5 6.9 3.9 0.0 Station F C - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 85 5 0 5 0 5 2 0 70 30 0 0 0 3 0 25 30 30 0 15 4 70 20 10 0 0 0 5 0 85 15 0 0 0 6 25 20 20 35 0 0 7 0 0 20 60 0 20 8 20 10 15 30 0 25 9 35 20 40 5 0 0 M E A N 26.1 28.3 20.0 18.3 0.0 7.2 S.E. 10.7 9.8 4.0 7.1 0.0 3.3 98 S U R F A C E C O V E R — J U N E 1998 Stat ion B B - C Q# Mus. Barn. Rock Sand Gr.Al. l 0 0 35 10 55 2 0 0 10 0 90 3 0 0 20 0 80 4 0 0 10 0 90 5 0 0 45 0 55 6 0 0 15 0 85 7 0 5 20 10 65 8 0 0 30 0 70 9 0 0 25 0 75 M E A N 0.0 0.6 23.3 2.2 73.9 S.E. 0.0 0.6 3.9 1.5 4.5 Stat ion B B - D Q# Mus. Barn. Rock Sand Gr.Al. l 0 0 50 0 50 2 0 0 40 0 60 3 0 0 80 5 15 4 0 0 75 15 10 5 0 0 80 15 5 6 0 0 0 0 100 7 0 0 95 5 0 8 0 0 70 20 10 9 0 0 95 0 5 M E A N 0.0 0.0 65.0 6.7 28.3 S.E. 0.0 0.0 10.2 2.6 11.4 Stat ion WJ-1.! Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 70 10 15 2 0 0 80 0 0 3 0 0 70 5 10 4 0 0 65 35 0 5 0 0 15 0 0 6 0 0 75 0 0 7 0 0 15 85 0 8 0 0 60 0 0 9 0 0 60 0 0 M E A N 0.0 0.0 56.7 15.0 2.8 S.E. 0.0 0.0 8.2 9.5 1.9 99 S U R F A C E C O V E R — JUNE 1998 Station BB-F Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 20 0 80 2 0 0 45 0 55 3 0 0 45 0 55 4 0 0 35 0 65 5 0 0 5 0 95 6 0 0 0 0 100 7 0 0 20 0 80 8 .0 0 95 0 0 9 0 0 85 0 0 M E A N 0.0 0.0 38.9 0.0 58.9 S.E. 0.0 0.0 11.0 0.0 12.3 Station B B - G Q# Mus. Barn. Rock Sand Gr.Al. l 0 40 40 0 20 2 0 50 10 0 40 3 10 80 10 0 0 4 0 90 10 0 0 5 0 50 15 5 30 6 5 15 55 0 25 7 0 35 60 0 5 8 0 50 50 0 0 9 0 5 75 20 0 M E A N 1.7 46.1 36.1 2.8 13.3 S.E. 1.2 9.1 8.4 2.2 5.2 Station Q# Mus. Barn. Rock Sand Gr.Al. 1 0 40 40 0 20 2 0 50 10 0 40 3 10 80 10 0 0 4 0 90 10 0 0 5 0 50 15 5 30 6 5 15 55 0 25 7 0 35 60 0 5 8 0 50 50 0 0 9 0 5 75 20 0 MEAN 1.7 46.1 36.1 2.8 13.3 S.E. 1.2 9.1 8.4 2.2 5.2 100 S U R F A C E C O V E R — J U N E 1998 Station F C - C Q# Mus. Barn. Rock Sand Gr.Al. 1 35 25 40 0 0 2 80 20 0 0 0 3 80 20 0 0 0 4 55 45 0 0 0 5 70 30 0 0 0 6 95 0 5 0 0 7 5 0 20 75 0 8 95 5 0 0 0 9 90 0 10 0 0 M E A N 67.2 16.1 8.3 8.3 0.0 S.E. 10.2 5.3 4.6 8.3 0.0 Station F C - D Q# Mus. Barn. Rock Sand Gr.Al. 1 0 20 70 10 0 2 30 50 20 0 0 3 0 25 40 10 25 4 30 25 40 0 5 5 15 30 55 0 0 6 0 25 35 10 30 7 0 5 95 0 0 8 10 35 55 0 0 9 30 55 15 0 0 M E A N 12.8 30.0 47.2 3.3 6.7 S.E. 4.6 5.1 8.3 1.7 4.0 Station F C - E Q# Mus. Barn. Rock Sand Gr.Al. l 5 20 60 15 0 2 15 20 60 5 0 3 0 25 50 20 5 4 0 45 30 20 5 5 40 60 0 0 0 6 75 0 25 0 0 7 40 60 0 0 0 8 30 15 20 35 0 9 0 0 35 65 0 M E A N 22.8 27.2 31.1 17.8 1.1 S.E. 8.6 7.6 7.6 7.1 0.7 101 S U R F A C E C O V E R — J U N E 1998 Station F C - F Q# Mus. Barn. Rock Sand Gr.Al. l 25 5 65 5 0 2 20 30 40 10 0 3 10 20 70 0 0 4 40 30 25 5 0 5 0 15 5 80 0 6 10 15 35 40 0 7 60 25 15 0 0 8 35 45 10 10 0 9 15 55 30 0 0 M E A N 23.9 26.7 32.8 16.7 0.0 S.E. 6.2 5.2 7.6 8.9 0.0 Station F C - G Q# Mus. Barn. Rock Sand Gr.Al. 1 65 20 10 0 5 2 70 30 0 0 0 3 25 30 30 0 15 4 75 25 0 0 0 5 85 15 0 0 0 6 20 20 60 0 0 7 0 20 60 0 20 8 10 30 35 0 25 9 70 30 0 0 0 M E A N 46.7 24.4 21.7 0.0 7.2 S.E. 10.8 1.9 8.5 0.0 3.3 102 C A N O P Y C O V E R — J U L Y 1998 Station BB-C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 : 4 0 0 6 0 2 0 0 0 5 0 9 5 3 0 0 0 0 0 1 0 0 4 0 0 0 5 5 0 4 5 5 0 0 0 0 0 100 6 0 0 0 2 0 0 8 0 7 0 0 0 5 0 95 8 0 0 0 5 0 95 9 0 0 0 15 0 8 5 M E A N 0.0 0.0 0.0 16.1 0.0 8 3 . 9 S.E. 0.0 0.0 0.0 6.4 0.0 6.4 Station BB-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 4 5 0 5 5 2 0 0 0 8 0 5 15 3 0 0 0 7 0 10 2 0 4 0 0 0 95 0 . 5 5 0 0 0 100 0 0 6 0 0 0 8 0 5 15 7 0 0 0 6 5 5 3 0 8 0 0 0 1 0 0 0 0 9 0 0 0 8 0 5 15 M E A N 0.0 0.0 0.0 7 9 . 4 3.3 17.2 S.E. 0.0 0.0 0.0 6 .0 1.2 5.7 Station BB-E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 0 2 0 0 0 1 0 0 0 o 3 0 0 0 1 0 0 0 0 4 0 0 0 1 0 0 0 0 5 0 0 0 1 0 0 0 0 6 0 0 0 1 0 0 0 0 7 0 0 0 9 5 5 0 8 0 0 0 1 0 0 0 0 9 o 0 0 100 0 o M E A N 0.0 0.0 0.0 9 9 . 4 0.6 0.0 S.E. 0.0 0.0 0.0 0.6 0.6 . 0 . 0 1 0 3 C A N O P Y C O V E R — J U L Y 1998 Station BB-F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 o 2 0 0 0 65 0 35 3 0 0 0 15 0 85 4 0 0 0 5 0 95 5 0 0 0 25 0 75 6 0 0 0 0 0 100 7 0 0 0 60 0 40 8 0 0 0 5 0 95 9 0 0 0 90 0 10 M E A N 0.0 0.0 0.0 40.6 0.0 59.4 S.E. 0.0 0.0 0.0 12.9 0.0 12.9 Station B B - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 5 0 20 35 0 40 2 25 25 40 10 0 0 3 55 5 10 0 0 30 4 40 0 20 25 0 15 5 30 0 30 20 0 20 6 5 10 30 55 0 0 7 20 0 5 50 0 25 8 25 0 50 5 0 20 9 0 30 65 5 0 0 M E A N 22.8 7.8 30.0 22.8 0.0 16.7 S.E. 6.0 3.9 6.4 6.7 0.0 4.8 Station B B - H Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 35 0 65 2 40 0 5 40 0 15 3 25 5 20 45 0 5 4 60 15 25 0 0 0 5 15 5 50 20 0 10 6 0 0 5 80 15 0 7 5 0 0 65 15 15 8 10 0 0 60 10 20 9 5 0 20 45 5 • : • 2 5 M E A N 17.8 2.8 13.9 43.3 5.0 17.2 S.E. 6.8 1.7 5.6 8.0 2.2 6.6 104 C A N O P Y C O V E R — J U L Y 1998 Station F C - C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 30 40 15 15 0 0 2 90 0 5 5 0 0 3 35 50 0 15 0 0 4 30 40 15 5 10 0 5 90 0 5 5 0 0 6 50 25 0 10 15 0 7 55 20 25 0 0 0 8 55 20 25 0 0 0 9 25 30 0 20 25 0 M E A N 51.1 25.0 10.0 8.3 5.6 0.0 S.E. 8.2 5.8 3.4 2.4 3.1 0.0 Station FC-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 25 5 45 25 0 0 2 0 15 40 40 0 •• 5 3 30 30 25 15 0 o 4 50 0 25 25 0 0 5 90 0 5 5 0 0 6 0 0 10 65 0 ,25 7 100 0 0 0 0 o 8 100 0 0 0 0 0 9 25 10 20 25 10 10 M E A N 46.7 6.7 18.9 22.2 1.1 4.4 S.E. 13.5 3.4 5.5 7.0 1.1 2.8 Station F C - E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 10 60 5 25 2 20 0 0 35 15 30 3 30 0 0 40 15 15 4 100 0 0 0 0 0 5 100 0 0 0 0 0 6 85 0 0 10 5 0 7 0 0 10 75 15 0 8 0 5 5 55 35 0 9 10 0 40 35 !5 o M E A N 38.3 0.6 7.2 34.4 11.7 7.8 S.E. 14.6 0.6 4.3 8.9 3.6 4.1 105 C A N O P Y C O V E R — J U L Y 1998 Station F C - F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 90 0 5 '5 0 0 2 80 0 10 10 0 • • • • 0 3 25 25 0 30 15 5 4 45 30 10 15 0 0 5 75 5 0 5 15 0 6 35 5 0 30 30 0 7 85 0 0 15 0 0 8 10 0 0 40 50 0 9 70 10 0 20 0 0 M E A N 57.2 8.3 2.8 18.9 12.2 0.6 S.E. 9.7 3.8 1.5 4.1 5.9 0.6 Station F C - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 25 0 30 35 0 10 2 35 15 20 30 0 0 3 15 0 30 55 0 0 4 55 0 25 20 0 0 5 40 15 0 45 0 0 6 60 5 15 20 0 0 7 0 0 0 100 0 o 8 25 45 20 10 0 0 9 15 0 5 80 0 0 M E A N 30.0 8.9 16.1 43.9 0.0 1.1 S.E. 6.5 5.0 4.0 10.0 0.0 1.1 106 S U R F A C E C O V E R — J U L Y 1998 Station BB-C Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 40 0 60 2 0 0 5 0 95 3 0 0 0 0 100 4 0 0 55 0 45 5 0 0 0 0 100 6 0 0 20 0 80 7 0 0 5 0 95 8 0 0 5 0 95 9 0 0 15 0 85 M E A N 0.0 0.0 16.1 0.0 83.9 S.E. 0.0 0.0 6.4 0.0 6.4 Station BB-D Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 45 0 55 2 0 0 80 5 15 3 0 0 70 10 20 4 0 0 95 0 5 5 0 0 100 0 0 6 0 0 80 5 15 7 0 0 65 5 30 8 0 0 100 0 0 9 0 0 80 5 15 M E A N 0.0 0.0 79.4 3.3 17.2 S.E. 0.0 0.0 6.0 1.2 5.7 Station BB-E Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 100 0 0 2 0 0 100 0 0 3 0 0 100 0 0 4 0 0 100 0 0 5 0 0 100 0 0 6 0 0 100 0 0 7 0 0 95 5 0 8 0 0 100 0 0 9 0 0 100 0 0 M E A N 0.0 0.0 99.4 0.6 0.0 S.E. 0.0 0.0 0.6 0.6 0.0 107 S U R F A C E C O V E R — J U L Y 1998 Station BB-F Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 100 0 0 2 0 0 65 0 35 3 0 0 15 0 - 85 4 0 0 5 0 95 5 0 0 25 0 75 6 0 0 0 0 100 7 0 0 60 0 40 8 0 0 5 0 95 9 0 0 90 0 10 M E A N 0.0 0.0 40.6 0.0 59.4 S.E. 0.0 0.0 12.9 0.0 12.9 Station B B - G Q# Mus. Barn. Rock Sand Gr.Al. 1 0 20 35 0 45 2 45 45 10 0 0 3 20 50 10 0 20 4 0 30 45 0 25 5 0 30 40 0 30 6 10 35 55 0 0 7 0 10 60 0 30 8 0 70 10 0 20 9 30 65 5 0 0 M E A N 11.7 39.4 30.0 0.0 18.9 S.E. 5.5 6.6 7.2 0.0 5.3 Station B B - H Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 35 0 65 2 0 5 85 0 10 3 5 20 70 0 5 4 65 30 0 0 5 5 5 65 20 0 10 6 0 5 80 15 0 7 0 0 70 15 15 8 0 0 65 15 20 9 0 20 50 5 25 M E A N 8.3 16.1 52.8 5.6 17.2 S.E. 7.1 7.1 9.7 2.4 6.5 108 S U R F A C E C O V E R — J U L Y 1998 Station F C - C Q# Mus. Barn. Rock Sand Gr.Al. 1 70 10 20 0 0 2 35 30 15 20 0 3 75 10 15 0 0 4 70 15 5 10 0 5 35 35 20 10 0 6 25 20 25 30 0 7 30 70 0 0 0 8 60 40 0 0 0 9 30 0 25 45 0 M E A N 47.8 25.6 13.9 12.8 0.0 S.E. 6.8 7.0 3.3 5.3 0.0 Station FC-D Q# Mus. Barn. Rock Sand Gr.Al. 1 5 60 35 0 0 2 15 40 40 0 5 3 55 25 20 0 0 4 25 35 40 0 0 5 15 75 10 0 0 6 0 10 65 0 25 7 30 30 40 0 0 8 10 50 40 0 0 9 10 30 35 15 10 M E A N 18.3 39.4 36.1 1.7 4.4 S.E. 5.5 6.5 5.1 1.7 2.8 Station F C - E Q# Mus. Barn. Rock Sand Gr.Al. 1 0 10 60 5 25 2 0 0 45 20 35 3 0 0 30 15 55 4 20 35 15 30 0 5 0 10 25 65 0 6 20 15 55 10 0 7 0 10 75 15 0 8 5 5 55 35 0 9 0 45 40 15 0 M E A N 5.0 14.4 44.4 23.3 12.8 S.E. 2.9 5.2 6.3 6.1 6.9 109 S U R F A C E C O V E R — J U L Y 1998 Station F C - F Q# Mus. Barn. Rock Sand Gr.Al. 1 40 20 35 5 0 2 30 50 20 0 0 3 15 0 55 30 0 4 45 25 25 5 0 5 10 30 30 30 0 6 0 5 50 45 0 7 10 10 70 10 0 8 0 0 45 55 0 9 5 10 60 25 0 M E A N 17.2 16.7 43.3 22.8 0.0 S.E. 5.7 5.5 5.7 6.4 0.0 Station F C - G Q# Mus. Barn. Rock Sand Gr.Al. 1 0 45 50 0 5 2 15 55 30 0 0 3 0 40 60 0 0 4 0 75 25 0 0 5 25 25 50 0 0 6 5 25 70 0 0 7 0 0 100 0 0 8 40 55 5 0 0 9 0 20 80 0 0 M E A N 9.4 37.8 52.2 0.0 0.6 S.E. 4.8 7.6 9.8 0.0 0.6 110 C A N O P Y C O V E R — A U G U S T 1998 Station BB-B Q# Fucus Mus. Barn. Rock Sand GrlAI. l 0 0 20 55 0 25 2 25 5 35 25 0 10 3 0 0 25 0 0 75 4 15 5 45 10 0 ; 25 5 25 5 35 25 0 10 6 20 0 55 25 0 0 7 10 0 40 50 0 0 8 0 0 25 25 0 50 9 5 0 15 10 0 70 M E A N 11.1 1.7 32.8 25.0 0.0 29.4 S.E. 3.5 0.8 4.3 6.0 0.0 9.6 Station BB-C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 10 35 5 50 2 0 0 0 30 5 65 3 0 0 0 45 30 25 4 0 0 0 30 0 70 5 0 0 5 15 0 80 6 0 0 0 35 55 10 7 0 0 0 5 0 95 8 0 0 0 5 10 . 85 9 0 0 0 40 15 45 M E A N 0.0 0.0 1.7 26.7 13.3 58.3 S.E. 0.0 0.0 1.2 4.9 6.1 9.4 Station BB-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 0 2 0 0 0 75 5 20 3 0 0 0 60 0 40 4 0 0 0 80 20 0 . 5 0 0 0 85 0 15 6 0 0 0 65 0 35 7 0 0 0 85 0 15 8 0 0 0 70 30 0 9 0 0 0 85 10 • 5 M E A N 0.0 0.0 0.0 78.3 7.2 14.4 S.E. 0.0 0.0 0.0 4.1 3.6 5.0 111 C A N O P Y C O V E R — AUGUST 1998 Station BB-E Q# Fucus Mus. Barn. Rock Sand Gr.Al, 1 0 0 0 55 45 0 2 0 0 0 100 0 o 3 0 0 0 100 0 o 4 0 0 0 55 45 0 5 0 0 0 90 10 0 6 0 0 0 55 45 o 7 0 0 0 100 0 0 8 0 0 0 90 10 o 9 0 0 0 90 10 0 M E A N 0.0 0.0 0.0 81.7 18.3 0.0 S.E. 0.0 0.0 0.0 6.8 6.8 0.0 Station Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 10 0 90 2 0 0 0 0 0 100 3 0 0 0 90 0 10 4 0 0 0 55 0 45 5 0 0 0 10 0 90 6 0 0 0 15 0 85 7 0 0 0 10 0 90 8 0 0 0 25 0 75 9 0 0 0 100 0 0 M E A N 0.0 0.0 0.0 35.0 0.0 65.0 S.E. 0.0 0.0 0.0 12.5 0.0 12.5 Station B B - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 25 60 10 0 ' 5 2 0 0 5 15 0 80 3 0 0 5 10 0 85 4 0 0 0 0 0 100 5 20 20 40 10 0 10 6 70 0 5 20 0 • 5 7 0 . 0 10 90 0 o 8 55 0 10 10 25 0 9 15 5 55 25 0 0 M E A N 17.8 5.6 21.1 21.1 2.8 31.7 S.E. 8.9 3.3 7.9 8.9 2.8 14.3 112 C A N O P Y C O V E R — A U G U S T 1998 Station BB-H Q# Fucus Mus. Barn. Rock , Sand Gr.Al. l 0 0 0 45 15 40 2 10 0 5 55 15 15 3 5 0 0 90 0 5 4 10 10 50 10 15 5 5 0 10 15 55 10 10 6 0 0 0 80 5 15 7 0 0 10 90 0 0 8 25 0 10 65 0 o 9 30 0 0 70 0 o M E A N 8.9 2.2 10.0 62.2 6.7 10.0 S.E. 3.8 1.5 5.3 8.4 2.4 4.2 113 C A N O P Y C O V E R — A U G U S T 1998 Station F C - B Q# Fucus Mus. Barn. Rock * Sand Gr.Al. l 40 0 10 50 0 o 2 50 15 10 25 0 0 3 10 0 15 75 0 0 4 5 0 10 85 0 0 5 25 20 10 40 5 0 6 85 0 5 10 0 0 7 0 20 35 45 0 0 8 45 0 15 40 0 0 9 5 5 60 30 0 0 M E A N 29.4 6.7 18.9 44.4 0.6 0.0 S.E. 9.4 3.0 5.9 7.8 0.6 0.0 Station F C - C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 45 30 15 5 5 0 2 75 25 0 0 0 0 3 40 40 5 0 15 o 4 30 60 10 0 0 0 5 40 20 5 10 25 0 6 30 35 30 5 0 o 7 75 20 5 0 0 0 8 30 55 0 0 15 0 9 85 0 15 0 0 0 M E A N 50.0 31.7 9.4 2.2 6.7 0.0 S.E. 7.4 6.2 3.2 1.2 3.1 0.0 Station FC-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 65 5 5 15 0 10 2 85 5 0 10 0 0 3 40 5 35 20 0 o 4 0 5 5 80 10 0 5 0 15 30 50 5 0 6 60 35 5 0 0 0 7 60 15 5 20 0 o 8 0 15 10 70 5 0 9 70 5 15 10 0 0 M E A N 42.2 11.7 12.2 30.6 2.2 1.1 S.E. 11.2 3.3 4.1 9.6 1.2 1.1 114 C A N O P Y C O V E R — A U G U S T 1998 Station F C - E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 35 15 10 20 20 0 2 20 20 0 45 15 0 3 5 15 20 50 10 o 4 45 0 0 20 35 o 5 30 5 0 50 15 0 6 0 0 5 45 45 • . 5 7 0 10 0 55 35 0 8 60 10 0 25 5 o 9 95 0 5 0 0 0 M E A N 32.2 8.3 4.4 34.4 20.0 0.6 S.E. 10.4 2.5 2.3 6.3 5.1 0.6 Station F C - F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 40 0 5 55 0 0 2 70 0 0 30 0 0 3 50 15 5 25 5 0 4 70 0 10 20 0 o 5 60 15 25 0 0 0 6 40 0 5 40 15 0 7 75 5 15 5 0 0 8 0 20 10 30 40 0 9 0 20 20 45 15 0 M E A N 45.0 8.3 10.6 27.8 8.3 0.0 S.E. 9.5 3.0 2.7 6.0 4.5 0.0 Station F C - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 20 80 0 o 2 10 25 10 55 0 0 3 70 25 0 5 0 0 4 45 25 30 0 0 0 5 45 5 35 15 0 0 6 0 0 25 75 0 o 7 80 15 5 0 0 0 8 10 10 55 20 0 5 9 0 65 35 0 0 0 M E A N 28.9 18.9 23.9 27.8 0.0 0.6 S.E. 10.6 6.7 5.8 11.0 0.0 0.6 115 S U R F A C E C O V E R — A U G U S T 1998 Station BB-B Q# Mus. Barn. Rock Sand Gr.Al. l 0 20 55 0 25 2 5 55 30 0 10 3 0 25 0 0 75 4 5 55 10 0 30 5 5 60 30 0 5 6 0 65 35 0 0 7 0 45 55 0 0 8 0 25 25 0 50 , 9 0 15 15 0 70 M E A N 1.7 40.6 28.3 0.0 29.4 S.E. 0.8 6.4 6.2 0.0 9.8 Station BB-C Q# Mus. Barn. Rock Sand Gr.Al. 1 0 10 35 5 50 2 0 0 30 5 65 3 0 0 45 30 25 4 0 0 30 0 70 5 0 5 15 0 80 6 0 0 35 55 10 7 0 0 5 0 95 8 0 0 5 10 85 9 0 0 40 15 45 M E A N 0.0 1.7 26.7 13.3 58.3 S.E. . 0.0 1.2 4.9 6.1 9.4 Station BB-D Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 100 0 0 2 0 0 75 5 20 3 0 0 60 0 40 4 0 0 80 20 0 5 0 0 85 0 15 6 0 0 65 0 35 7 0 0 85 0 15 8 0 0 70 30 0 9 0 0 85 10 5 M E A N 0.0 0.0 78.3 7.2 14.4 S.E. 0.0 0.0 4.1 3.6 5.0 116 S U R F A C E C O V E R — A U G U S T 1998 Station BB-E Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 55 45 0 2 0 0 100 0 0 3 0 0 100 0 0 4 0 0 55 45 0 5 0 0 90 10 0 6 0 0 55 45 0 7 0. 0 100 0 0 8 0 0 90 10 0 9 0 0 90 10 0 M E A N 0.0 0.0 81.7 18.3 0.0 S.E. 0.0 0.0 6.8 6.8 0.0 Station BB-F Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 10 0 90 2 0 0 0 0 100 3 0 0 90 0 10 4 0 0 55 0 45 5 0 0 10 0 90 6 0 0 15 0 85 7 0 0 10 0 90 8 0 0 25 0 75 9 0 0 100 0 0 M E A N 0.0 0.0 35.0 0.0 65.0 S.E. 0.0 0.0 12.5 0.0 12.5 Station B B - G Q# Mus. Barn. Rock Sand Gr.Al. 1 25 60 10 0 5 2 0 5 15 0 80 3 0 5 10 0 85 4 0 0 0 0 100 5 30 50 10 0 10 6 0 25 70 0 5 7 5 20 65 0 10 8 5 20 65 0 10 9 5 70 25 0 0 M E A N 7.8 28.3 30.0 0.0 33.9 S.E. 3.8 8.5 9.4 0.0 13.8 117 S U R F A C E C O V E R — A U G U S T 1998 Station B B - H Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 45 15 40 2 0 10 55 20 15 3 0 0 95 0 5 4 10 60 10 15 5 5 10 15 55 10 10 6 0 0 80 5 15 7 0 10 90 0 0 8 25 20 55 0 0 9 0 0 100 0 0 M E A N 5.0 12.8 65.0 7.2 10.0 S.E. 2.9 6.4 9.6 2.6 4.2 118 S U R F A C E C O V E R — A U G U S T 1998 Station F C - B Q# Mus. Barn. Rock Sand Gr.Al. 1 0 25 75 0 0 2 35 10 55 0 0 3 0 15 85 0 0 4 0 10 90 0 0 5 " 40 10 45 5 0 6 25 50 25 0 0 7 20 35 45 0 0 8 0 45 55 0 0 9 5 65 30 0 0 M E A N 13.9 29.4 56.1 0.6 0.0 S.E. 5.5 6.8 7.7 0.6 0.0 Station F C - C Q# Mus. Barn. Rock Sand Gr.Al. 1 40 35 25 0 0 2 75 15 10 0 0 3 45 30 5 20 0 4 75 25 0 0 0 5 40 15 20 25 0 6 40 55 5 0 0 7 60 35 5 0 0 8 85 10 0 5 0 9 25 35 30 10 0 M E A N 53.9 28.3 11.1 6.7 0.0 S.E. 6.9 4.6 3.7 3.2 0.0 Station FC-D Q# Mus. Barn. Rock Sand Gr.Al. 1 10 25 50 0 15 2 15 45 40 0 0 3 10 60 30 0 0 4 5 5 80 10 0 5 15 30 50 5 0 6 85 15 0 0 0 7 25 25 50 0 0 8 15 10 70 5 0 9 50 10 40 0 0 M E A N 25.6 25.0 45.6 2.2 1.7 S.E. 8.6 6.0 7.7 1.2 1.7 119 SURFACE COVER — AUGUST 1998 Station FC-E Q# Mus. Barn. Rock Sand Gr.Al. 1 10 0 60 30 0 2 20 5 50 25 0 3 15 20 55 10 0 4 0 0 40 60 0 5 5 0 60 35 0 6 10 0 55 35 0 7 10 0 55 35 0 8 25 0 60 15 0 9 25 5 35 35 0 M E A N 13.3 3.3 52.2 31.1 0.0 S.E. 2.9 2.2 3.0 4.8 0.0 Station FC-F Q# Mus. Barn. Rock Sand Gr.Al. 1 0 5 70 25 0 2 0 15 65 20 0 3 30 0 45 25 0 4 20 20 60 0 0 5 25 70 5 0 0 6 5 10 60 25 0 7 15 55 25 5 0 8 20 10 30 40 0 9 20 20 45 15 0 M E A N 15.0 22.8 45.0 17.2 0.0 S.E. 3.6 7.9 7.2 4.5 0.0 Station FC-G Q# Mus. Barn. Rock Sand Gr.Al. 1 0 20 80 0 0 2 30 10 60 0 0 3 70 25 5 0 0 4 45 50 5 0 0 5 15 75 10 0 0 6 0 25 75 0 0 7 70 20 10 0 0 8 10 55 30 0 5 9 65 35 0 0 0 M E A N 33.9 35.0 30.6 0.0 0.6 S.E. 9.8 7.0 10.8 0.0 0.6 120 CANOPY COVER — OCTOBER 1998 Station BB-B Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 0 2 25 0 55 20 0 0 3 0 0 35 65 0 0 4 5 0 40 55. 0 o 5 10 0 45 45 0 0 6 15 0 45 40 0 0 7 0 0 10 90 0 0 8 25 5 55 15 0 o 9 0 0 15 85 0 0 M E A N 8.9 0.6 33.3 57.2 0.0 0.0 S.E. 3.5 0.6 6.7 10.1 0.0 0.0 Station BB-C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 10 45 0 45 2 0 0 0 65 35 0 3 0 0 0 45 55 0 4 0 0 0 75 25 0 5 0 0 30 70 0 0 6 0 0 0 90 10 0 7 0 0 5 95 0 • o 8 0 0 0 35 0 65 9 0 0 5 50 0 45 M E A N 0.0 0.0 5.6 63.3 13.9 17.2 S.E. 0.0 0.0 3.3 7.0 6.7 8.8 Station BB-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 . o 2 0 0 0 5 95 0 3 0 0 0 45 0 55 4 0 0 0 80 20 0 5 0 0 0 40 50 10 6 0 0 0 85 0 15 7 0 0 0 85 0 15 8 0 0 0 90 10 0 9 0 0 0 65 5 30 M E A N 0.0 0.0 0.0 66.1 20.0 13.9 S.E. 0.0 0.0 0.0 10.2 10.8 6.2 CANOPY COVER — OCTOBER 1998 Station BB-E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 90 0 10 2 0 0 0 100 0 0 3 0 0 0 95 0 5 4 0 0 0 30 0 70 5 0 0 0 100 0 0 6 0 0 0 80 0 20 7 0 0 0 100 0 0 8 0 0 0 0 20 80 9 0 0 0 80 20 0 M E A N 0.0 0.0 0.0 75.0 4.4 20.6 S.E. 0.0 0.0 0.0 11.9 2.9 10.6 Station BB-F Q# Fucus Mus. Barn. Rock Sand Gr.Al. l 0 0 0 100 0 0 2 0 0 0 20 0 80 3 0 0 0 70 0 30 4 0 0 0 100 0 0 5 0 0 0 0 0 100 6 0 0 0 0 0 100 7 0 0 0 100 0 0 8 0 0 0 100 0 . 0 9 0 0 0 100 0 0 M E A N 0.0 0.0 0.0 65.6 0.0 34.4 S.E. 0.0 0.0 0.0 15.2 0.0 15.2 Station B B - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 45 0 15 40 0 0 2 5 10 85 0 0 0 3 50 5 5 40 0 0 4 5 25 70 0 0 0 5 5 0 50 45 0 0 6 30 0 25 40 0 5 7 0 0 25 75 0 0 8 0 0 0 100 0 . ;.: 0 9 25 0 25 50 0 0 M E A N 18.3 4.4 33.3 43.3 0.0 0.6 S.E. 6.6 2.8 9.7 10.6 0.0 0.6 122 C A N O P Y C O V E R — O C T O B E R 1998 Station B B - H Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 70 0 0 30 0 o 2 5 0 0 95 0 0 3 0 0 5 95 0 0 4 80 0 0 20 0 0 5 40 0 25 20 15 0 6 0 0 5 95 0 0 7 60 10 15 15 0 0 8 10 o 10 80 0 0 9 0 0 0 100 0 0 M E A N 29.4 1.1 6.7 61.1 1.7 0.0 S.E. 11.1 1.1 2.9 12.8 1.7 0.0 123 C A N O P Y C O V E R — O C T O B E R 1998 Station FC-B Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 0 0 0 100 0 0 2 5 5 0 90 0 o 3 0 0 10 90 0 0 4 0 0 10 90 0 0 5 0 0 50 50 0 0 6 0 5 45 50 0 o 7 5 0 60 35 0 o 8 50 0 25 25 0 0 9 45 15 5 35 0 o M E A N 11.7 2.8 22.8 62.8 0.0 0.0 S.E. 6.8 1.7 7.7 9.8 0.0 0.0 Station F C - C Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 15 20 35 25 5 0 2 20 70 10 0 0 0 3 0 60 10 30 0 o 4 50 10 20 20 0 0 5 70 25 5 0 0 0 6 0 15 20 60 5 o 7 20 65 15 0 0 o 8 15 50 10 20 5 o 9 0 65 15 20 0 0 M E A N 21.1 42.2 15.6 19.4 1.7 0.0 S.E. 8.0 8.1 2.9 6.4 0.8 0.0 Station FC-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 15 70 10 5 0 0 2 0 5 5 90 0 o 3 0 15 10 75 0 0 4 90 5 0 5 0 o 5 0 40 0 50 10 0 6 35 0 25 40 0 o 7 50 10 25 15 0 0 8 50 5 20 25 0 0 9 65 5 10 20 0 :. 0 M E A N 33.9 17.2 11.7 36.1 1.1 0.0 S.E. 10.8 7.7 3.2 10.1 1.1 0.0 124 CANOPY COVER — OCTOBER 1998 Station FC-E Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 15 10 10 65 0 0 2 70 0 0 15 15 0 3 65 0 0 35 0 . o 4 25 20 20 35 0 0 5 95 5 0 0 0 0 6 80 0 20 0 0 0 7 60 20 0 20 0 o 8 75 0 10 15 0 0 9 75 0 5 20 0 0 M E A N 62.2 6.1 7.2 22.8 1.7 0.0 S.E. 8.7 2.9 2.8 6.7 1.7 0.0 Station FC-F Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 5 10 5 65 15 0 2 0 30 40 30 0 0 3 5 25 25 30 15 0 4 5 30 10 40 15 0 5 0 10 45 45 0 ..; o 6 0 15 25 55 5 0 7 5 10 45 40 0 0 8 5 5 0 75 15 0 9 30 15 10 40 5 0 M E A N 6.1 16.7 22.8 46.7 7.8 0.0 S.E. 3.1 3.1 5.8 5.1 2.4 0.0 Station FC-G Q# Fucus Mus. Barn. Rock Sand Gr.Al. 1 100 0 0 0 0 0 2 65 0 10 20 0 5 3 20 0 65 10 0 \.5 4 55 0 15 0 0 30 5 10 15 20 55 0 0 6 40 0 20 20 0 20 7 15 0 25 45 0 15 8 10 0 85 5 0 0 9 0 15 20 65 0 0 M E A N 35.0 3.3 28.9 24.4 0.0 8.3 S.E. 11.0 2.2 9.2 8.2 0.0 3.6 125 SURFACE COVER — OCTOBER 1998 Station BB-B Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 100 0 0 2 0 60 40 0 0 3 0 35 65 0 0 4 o 45 55 0 0 5 0 55 45 0 0 6 0 55 45 0 0 7 0 10 90 0 0 8 5 50 45 0 0 9 0 15 85 0 0 M E A N 0.6 36.1 63.3 0.0 0.0 S.E. 0.6 7.4 7.6 0.0 0.0 Station BB-C Q# Mus. Barn. Rock Sand Gr.Al. 1 0 10 45 0 45 2 0 0 65 35 0 3 0 0 45 55 0 4 0 0 75 25 0 5 0 30 70 0 0 6 0 0 90 10 0 7 0 5 95 0 0 8 0 0 35 0 65 9 0 5 50 0 45 M E A N 0.0 5.6 63.3 13.9 17.2 S.E. 0.0 3.3 7.0 6.7 8.8 Station BB-D Q# Mus. Barn. Rock Sand Gr.Al. l 0 0 100 0 0 2 0 0 5 95 0 3 0 0 45 0 55 4 0 0 80 20 0 5 0 0 40 50 10 6 0 0 85 0 15 7 0 0 85 0 15 8 0 0 90 10 0 9 0 0 65 5 30 M E A N 0.0 0.0 66.1 20.0 13.9 S.E. 0.0 0.0 10.2 10.8 6.2 126 SURFACE COVER — OCTOBER 1998 Station BB-E Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 90 0 10 2 0 0 100 0 0 3 0 0 95 0 5 4 0 0 30 0 70 5 0 0 100 0 0 6 0 0 80 0 20 7 0 0 100 0 0 8 0 0 0 20 80 9 0 0 80 20 0 MEAN" 0.0 0.0 75.0 4.4 20.6 S.E. 0.0 0.0 11.9 2.9 10.6 Station BB-F Q# Mus. Barn. Rock Sand Gr.Al. 1 0 0 100 0 0 2 0 0 20 0 80 3 0 0 70 0 30 4 0 0 100 0 0 5 0 0 0 0 100 6 0 0 0 0 100 7 0 0 100 0 0 8 0 0 100 0 0 9 0 0 100 0 0 M E A N 0.0 0.0 65.6 0.0 34.4 S.E. 0.0 0.0 15.2 0.0 15.2 Station B B - G Q# Mus. Barn. Rock Sand Gr.Al. 1 0 30 70 0 0 2 10 90 0 0 0 3 5 15 80 0 0 4 25 75 0 0 0 5 0 50 50 0 0 6 0 40 55 0 5 7 0 25 75 0 0 8 0 0 100 0 0 9 0 35 65 0 0 M E A N 4.4 40.0 55.0 0.0 0.6 S.E. 2.8 9.4 11.5 0.0 0.6 127 S U R F A C E C O V E R — O C T O B E R 1998 Station BB-H Q# Mus. Barn. Rock Sand Gr.Al. 1 0 5 95 0 0 2 0. 0 100 0 0 3 0 5 95 0 0 4 0 40 60 0 0 5 0 30 45 25 0 6 0 5 95 0 0 7 20 30 50 0 0 8 0 10 90 0 0 9 0 0 100 0 0 M E A N 2.2 13.9 81.1 2.8 0.0 S.E. 2.2 5.1 7.5 2.8 0.0 128 SURFACE COVER — OCTOBER 1998 Station FC-B Q# Mus. Barn. Rock Sand Gr.Al. l 0 0 100 0 0 2 5 0 95 0 0 3 0 10 90 0 0 4 0 10 90 0 0 5 0 50 50 0 0 6 5 45 50 0 o 7 0 65 35 0 0 8 0 70 30 0 0 9 35 35 30 0 0 M E A N 5.0 31.7 63.3 0.0 0.0 S.E. 3.8 9.2 10.0 0.0 0.0 Station FC-C Q# Mus. Barn. Rock Sand Gr.Al. 1 30 40 25 5 0 2 85 15 0 0 0 3 60 10 30 0 0 4 20 60 20 0 0 5 55 35 10 0 0 6 15 20 60 5 0 7 70 25 5 0 .0 8 50 20 25 5 0 9 65 15 20 0 0 M E A N 50.0 26.7 21.7 1.7 0.0 S.E. 7.9 5.3 5.8 0.8 0.0 Station FC-D Q# Mus. Barn. Rock Sand Gr.Al. l 80 10 10 0 0 2 5 5 90 0 0 3 15 10 75 0 0 4 25 45 30 0 0 5 40 0 50 10 0 6 5 45 50 0 0 7 25 30 45 0 0 8 15 50 35 0 0 9 25 20 55 0 0 M E A N 26.1 23.9 48.9 1.1 0.0 S.E. 7.7 6.4 7.9 1.1 0.0 129 SURFACE COVER — OCTOBER 1998 Station FC-E Q# Mus. Barn. Rock Sand Gr.Al. l 10 10 80 0 0 2 5 5 80 10 0 3 5 30 60 5 0 4 40 20 40 0 0 5 40 40 20 0 0 6 10 80 10 0 0 7 30 15 55 0 0 8 15 20 65 0 0 9 15 20 45 20 0 M E A N 18.9 26.7 50.6 3.9 0.0 S.E. 4.7 7.5 8.1 2.3 0.0 Station FC-F Q# Mus. Barn. Rock Sand Gr.Al. 1 10 5 70 15 0 2 30 40 30 0 0 3 25 25 35 15 0 4 30 10 45 15 0 5 10 45 45 0 0 6 15 25 55 5 0 7 10 50 40 0 0 8 5 5 75 15 0 9 20 30 45 5 0 M E A N 17.2 26.1 48.9 7.8 0.0 S.E. 3.1 5.6 5.1 2.4 0.0 Station F C - G Q# Mus. Barn. Rumpus Sand Gr.Al. 1 15 40 45 0 0 2 0 45 45 0 10 3 0 75 20 0 5 4 0 50 0 0 50 5 15 20 65 0 0 6 0 30 45 0 25 7 0 40 45 0 15 8 0 95 5 0 0 9 15 20 65 0 0 M E A N 5.0 46.1 37.2 0.0 11.7 S.E. 2.5 8.3 7.9 0.0 5.6 130 C A N O P Y C O V E R — D E C E M B E R 1998 Station BB-B Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 0 100 0 0 0 2 0 0 0 100 0 o 0 3 0 0 0 100 0 o 0 4 0 0 0 100 0 0 0 5 0 0 0 100 0 0 0 6 0 0 0 100 0 0 0 7 0 0 0 100 0 0 0 8 0 0 0 100 0 0 0 9 0 0 0 100 0 . 0 0 M E A N 0.0 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Station BB-C Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 20 80 0 o 0 2 0 0 0 100 0 o 0 3 0 0 0 100 0 0 0 4 0 0 0 90 10 0 0 5 0 0 0 95 5 0 0 6 0 0 0 95 5 o 0 7 0 0 10 90 0 0 0 8 0 0 0 100 0 0 0 9 0 0 0 100 0 0 0 M E A N 0.0 0.0 3.3 94.4 2.2 0.0 0.0 S.E. 0.0 0.0 2.4 2.3 1.2 0.0 0.0 Station BB-D Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 0 100 0 0 0 2 0 0 0 100 0 0 0 3 0 0 0 100 0 0 0 4 0 0 0 100 0 o 0 5 0 0 0 100 0 0 0 6 0 0 0 100 0 0 0 7 0 0 0 100 0 • o 0 8 0 0 0 100 0 0 0 9 0 0 0 100 0 o 0 M E A N 0.0 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 * Other = Porphyra spp. C A N O P Y C O V E R — D E C E M B E R 1998 Station BB-E Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* l 0 0 0 100 0 0 0 2 0 0 0 100 0 0 0 3 0 0 0 100 0 0 0 4 0 0 0 100 0 0 0 5 0 0 0 100 0 o 0 6 0 0 0 100 0 0 0 7 0 0 0 100 0 o 0 8 0 0 0 100 0 0 0 9 0 0 0 100 0 0 0 M E A N 0.0 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 0.0 0.0 Station BB-F Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* l 0 0 0 100 0 0 0 2 0 0 0 70 0 30 0 3 0 0. 0 65 0 35 0 4 0 0 0 100 0 0 0 5 0 0 0 55 0 45 0 6 0 0 0 100 0 0 0 7 0 0 0 85 0 15 0 8 0 0 0 95 0 5 0 9 0 0 0 70 0 30 0 M E A N 0.0 0.0 0.0 82.2 0.0 17.8 0.0 S.E. 0.0 0.0 0.0 5.8 0.0 5.8 0.0 Station B B - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 10 0 5 85 0 0 0 2 20 0 0 80 0 0 0 3 0 0 70 30 0 0 0 4 20 0 35 45 0 0 0 5 0 0 40 60 0 0 0 6 5 0 0 95 0 0 0 7 20 0 50 30 0 0 0 8 15 0 15 70 0 o 0 9 20 10 35 35 0 0 0 M E A N 12.2 1.1 27.8 58.9 0.0 0.0 0.0 S.E. 2.9 1.1 8.1 8.3 0.0 0.0 0.0 * Other = Porphyra spp. 132 C A N O P Y C O V E R — D E C E M B E R 1998 Station B B - H Q# Fucus Mus. Barn. Rock *' Sand Gr.Al. Other* 1 0 0 0 100 0 0 0 2 5 0 5 90 0 0 0 3 0 0 0 0 100 0 0 4 5 5 20 70 0 0 0 5 0 0 5 95 0 0 0 6 5 0 0 80 15 0 0 7 60 0 0 40 0 0 0 8 15 0 0 75 10 0 0 9 0 0 0 15 85 0 0 M E A N 10.0 0.6 3.3 62.8 23.3 0.0 0.0 S.E. 6.5 0.6 2.2 12.0 13.3 0.0 0.0 * Other = Porphyra spp. 133 C A N O P Y C O V E R — D E C E M B E R 1998 Station FC-B Q# Fucus Mus. Barn. Rock ' Sand Gr.Al. Other* 1 60 0 0 40 0 0 0 2 0 0 0 100 0 o 0 3 0 0 50 35 0 0 15 4 0 0 10 90 0 0 0 5 0 0 0 100 0 o 0 6 0 0 0 95 0 0 5 7 0 0 50 45 0 o 5 8 15 0 0 85 0 0 0 9 0 0 55 30 0 0 15 M E A N 8.3 0.0 18.3 68.9 0.0 0.0 4.4 S.E. 6.7 0.0 8.4 10.1 0.0 0,0 2.1 Station F C - C Q# Fucus Mus. Barn. Rock Sand Gr.AL Other* 1 35 30 30 5 0 0 0 2 15 65 10 10 0 0 0 3 50 10 10 25 5 o 0 4 20 55 20 5 0 0 0 5 25 50 15 10 0 0 0 6 0 55 20 25 0 0 0 7 0 90 10 0 0 0 0 8 20 45 25 10 0 0 0 9 0 35 5 60 0 0 0 M E A N 18.3 48.3 16.1 16.7 0.6 0.0 0.0 S.E. 5.7 7.5 2.7 6.1 0.6 0.0 0.0 Station F C - D Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 5 95 0 o 0 2 0 10 15 75 0 0 0 3 35 15 5 45 0 0 0 4 0 10 5 85 0 0 0 5 0 20 5 70 5 o 0 6 0 5 0 95 0 o 0 7 0 5 0 45 50 0 0 8 0 5 15 80 0 o 0 9 5 5 5 65 20 0 0 M E A N 4.4 8.3 6.1 72.8 8.3 0.0 0.0 S.E. 3.9 2.0 1.8 6.2 5.7 0.0 0.0 * Other = Porphyra spp. 134 C A N O P Y C O V E R — D E C E M B E R 1998 Station F C - E Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 70 0 5 25 0 V ' " o 0 2 10 0 0 25 65 o 0 3 40 25 10 5 20 0 0 4 15 5 0 75 5 0 0 5 0 0 0 100 0 0 0 6 10 30 0 45 15 0 0 7 5 5 5 80 5 o 0 8 65 0 10 0 25 0 0 9 20 0 5 55 20 o 0 M E A N 26.1 7.2 3.9 45.6 17.2 o.o 0.0 S.E. 8.7 3.9 1.4 11.6 6.7 0,0 0.0 Station F C - F Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 0 20 0 45 35 0 0 2 50 0 10 40 0 0 0 3 30 0 0 40 25 ;•' 5 0 4 55 0 0 35 10 0 0 5 0 0 0 70 30 0 0 6 0 10 10 45 35 o 0 7 5 5 5 85 0 0 0 8 0 0 5 95 0 0 0 9 0 0 15 85 0 0 0 M E A N 15.6 3.9 5.0 60.0 15.0 0.6 0.0 S.E. 7.7 2.3 1.9 7.9 5.3 0.6 0.0 Station F C - G Q# Fucus Mus. Barn. Rock Sand Gr.Al. Other* 1 20 0 10 70 0 0 0 2 0 0 0 100 0 0 0 3 50 0 0 50 0 0 0 4 10 0 30 55 0 0 5 5 0 0 10 85 0 0 5 6 0 0 0 95 0 . -5 0 7 0 15 20 65 0 o 0 8 0 0 55 45 0 0 0 9 0 0 25 75 0 0 0 M E A N 8.9 1.7 16.7 71.1 0.0 0.6 1.1 S.E. 5.6 1.7 6.1 6.5 0.0 0.6 0.7 * Other = Porphyra spp. 135 S U R F A C E C O V E R — D E C E M B E R 1998 Station BB-B Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 100 0 0 0 2 0 0 100 0 0 0 3 0 0 100 0 0 0 4 0 0 100 0 0 0 5 0 0 100 0 0 0 6 0 0 100 0 0 0 7 0 0 100 0 0 0 8 0 0 100 0 0 0 9 0 0 100 0 0 0 M E A N 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 . 0.0 Station BB-C Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 0 20 80 0 0 0 2 0 0 100 0 0 o 3 0 0 100 0 0 0 4 0 0 90 10 0 0 5 0 0 95 5 0 0 6 0 0 95 5 0 o 7 0 10 90 0 0 0 8 0 0 100 0 0 0 9 0 0 100 0 0 0 M E A N 0.0 3.3 94.4 2.2 0.0 0.0 S.E. 0.0 2.4 2.3 1.2 0.0 0.0 Station BB-D Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 0 0 100 0 0 0 2 0 0 100 0 0 o 3 0 0 100 0 0 0 4 0 0 100 0 0 o 5 0 0 100 0 0 ; 0 6 0 0 100 0 0 0 7 0 0 100 0 0 0 8 0 0 100 0 0 .. o 9 0 0 100 0 0 0 M E A N 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 0.0 * Other = Porphyra spp. 136 S U R F A C E C O V E R — D E C E M B E R 1998 Station BB-E Q# Mus. Barn. Rock Sand Gr.Al. Other* l 0 0 100 0 0 0 2 .0 0 100 0 0 0 3 0 0 100 0 0 o 4 0 0 100 0 0 o 5 0 0 100 0 0 o 6 0 0 100 0 0 0 7 0 0 100 0 0 0 8 0 0 100 0 0 o 9 0 0 100 0 0 0 M E A N 0.0 0.0 100.0 0.0 0.0 0.0 S.E. 0.0 0.0 0.0 0.0 0.0 0.0 Station BB-F Q# Mus. ' Barn. Rock Sand Gr.Al. Other* 1 0 0 100 0 0 o 2 0 0 70 0 30 o 3 0 0 65 0 35 0 4 0 0 100 0 0 0 5 0 0 55 0 45 0 6 0 0 100 0 0 o 7 0 0 85 0 15 0 8 0 0 95 0 5 0 9 0 0 70 0 30 0 M E A N 0.0 0.0 82.2 0.0 17.8 0.0 S.E. 0.0 0.0 5.8 0.0 5.8 0.0 Station B B - G Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 0 5 95 0 0 0 2 0 0 100 0 0 0 3 0 70 30 0 0 0 4 0 50 50 0 0 0 5 0 40 60 0 0 0 6 0 0 100 0 0 0 7 0 60 40 0 0 0 8 0 15 85 0 0 0 9 10 55 35 0 0 0 M E A N 1.1 32.8 66.1 0.0 0.0 0.0 S.E. 1.1 9.3 9.7 0.0 0.0 0.0 * Other = Porphyra spp. 137 S U R F A C E C O V E R — D E C E M B E R 1998 Station B B - H Q# Mus. Barn. Rock Sand , Gr.Al. Other* 1 0 0 100 0 0 0 2 0 5 95 0 0 0 3 0 0 0 100 0 0 4 5 20 75 0 0 o 5 0 5 95 0 0 o 6 0 0 85 15 0 0 7 0 0 100 0 0 0 8 0 0 85 15 0 0 9 0 0 15 85 0 0 M E A N 0.6 3.3 72.2 23.9 0.0 0.0 S.E. 0.6 2.2 12.6 13.2 0.0 0.0 * Other = Porphyra spp. 138 S U R F A C E C O V E R — D E C E M B E R 1998 Station F C - B Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 0 10 90 0 0 0 2 0 0 100 0 0 0 3 0 50 35 0 0 15 4 0 10 90 0 0 0 5 0 0 100 0 0 0 6 0 0 95 0 0 •\ 5 7 0 50 45 0 0 5 8 0 0 100 0 0 0 9 0 55 30 0 0 15 M E A N 0.0 19.4 76.1 0.0 0.0 4.4 S.E. 0.0 8.2 10.0 0.0 0.0 2.1 Station F C - C Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 50 40 10 0 0 0 2 80 10 10 0 0 0 3 35 20 45 0 0 0 4 60 35 5 0 0 : 0 5 60 25 15 0 0 o 6 55 20 25 0 0 0 7 90 10 0 0 0 0 8 55 35 10 0 0 0 9 35 5 60 0 0 0 M E A N 57.8 22.2 20.0 0.0 0.0 0.0 S.E. 6.1 4.2 6.7 0.0 0.0 0.0 Station FC-D Q# Mus. Barn. Rock Sand Gr.Al. Other* l 0 5 95 0 0 0 2 10 15 75 0 0 0 3 35 5 60 0 0 0 4 10 5 85 0 0 0 5 20 5 70 5 0 0 6 5 0 95 0 0 0 7 5 0 45 50 0 o 8 5 15 80 0 0 0 9 5 5 70 20 0 0 M E A N 10.6 6.1 75.0 8.3 0.0 0.0 S.E. 3.6 1.8 5.4 5.7 0.0 0.0 * Other = Porphyra spp. 139 S U R F A C E C O V E R — D E C E M B E R 1998 Station F C - E Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 15 20 65 0 0 0 2 5 0 30 65 0 0 3 30 10 40 20 0 0 4 5 5 85 5 0 0 5 0 0 100 0 0 0 6 30 5 45 20 0 0 7 10 5 80 5 0 0 8 0 25 30 45 0 o 9 0 10 70 20 0 0 M E A N 10.6 8.9 60.6 20.0 0.0 0.0 S.E. 4.0 2.9 8.5 7.4 0.0 0.0 Station F C - F Q# Mus. Barn. Rock Sand Gr.Al. Other* 1 20 0 45 35 0 0 2 0 10 55 35 0 0 3 0 0 55 40 5 o 4 0 0 85 15 0 0 5 0 0 70 30 0 o 6 10 10 45 35 0 0 7 5 5 90 0 0 0 8 0 5 95 0 0 0 9 0 15 85 0 0 0 M E A N 3.9 5.0 69.4 21.1 0.6 0.0 S.E. 2.3 1.9 6.6 5.8 0.6 0.0 Station F C - G Q# Mus. Barn. Rock Sand Gr.Al. Other* l 0 20 80 0 0 o 2 0 0 100 0 0 0 3 0 15 85 0 0 0 4 0 40 55 0 0 5 5 0 10 85 0 0 5 6 0 0 95 0 5 0 7 15 20 65 0 0 0 8 0 55 45 0 0 0 9 0 25 75 0 0 0 M E A N 1.7 20.6 76.1 0.0 0.6 1.1 S.E. 1.7 6.0 6.1 0.0 0.6 0.7 * Other = Porphyra spp. 140 FUCUS GARDNERI O O C Y T E R E L E A S E Station BB-B Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 660 88.9 7.42 2 864 62.4 13.85 3 1356 87.0 15.59 M E A N 960 79.4 12.29 S.E. 207 8.5 2.48 Station B B - H Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 252 155.8 1.62 2 324 176.2 1.84 3 60 158.7 0.38 M E A N 212 163.6 1.28 S.E. 79 6.4 0.45 Station F C - C Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 996 215.5 4.62 2 180 187.2 0.96 3 1212 217.4 5.57 M E A N 796 206.7 3.72 S.E. 314 9.8 1.41 Station F C - E Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 1200 222.6 5.39 2 372 186.5 1.99 3 72 203.8 0.35 M E A N 548 204.3 2.58 S.E. 337 10.4 1.48 Station F C - G Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 720 156.0 4.62 2 720 133.9 5.38 3 2040 189.8 10.75 M E A N 1160 159.9 6.91 S.E. 440 16.3 1.93 O C T O B E R 1998 Station B B - G Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 732 110.4 6.63 2 336 174.4 1.93 3 386 130.2 2.96 M E A N 485 138.3 3.84 S.E. 125 18.9 1.43 Station F C - B Plant surface . S# Oocytes area (cm2) Oocytes cm"2 1 60 203.3 0.30 2 60 174.1 0.34 3 900 167.9 5.36 M E A N 340 181.8 2.00 S.E. 280 10.9 1.68 Station F C - D Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 1896 213.6 8.88 2 792 215.1 3.68 3 804 184.9 4.35 M E A N 1164 204.5 5.64 S.E. 366 9.8 1.63 Station F C - F Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 204 171.2 1.19 2 1164 217.0 5.36 3 1380 238.1 5.80 M E A N 916 208.8 4.12 S.E. 361 19.7 1.47 141 FUCUS GARDNERI OOCYTE RELEASE — DECEMBER 1998 Station BB-G Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 400 49.2 8.13 2 240 38.4 6.25 3 4 5 112 66.3 1.69 M E A N 251 51.3 5.36 S.E. 83 8.1 1.91 Station FC-B Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 136 98.3 1.38 2 16 48.2 0.33 3 128 55.9 2.29 4 136 80.0 1.70 5 32 72.8 0.44 M E A N 90 71.0 1.23 S.E. 35 11.5 0.48 Station FC-D Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 128 72.5 1.77 .2 136 80.6 1.69 3 32 58.3 0.55 4 656 93.4 7.02 5 16 59.0 0.27 M E A N 194 72.8 2.26 S.E. 153 8.6 1.59 Station FC-F Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 152 108.7 1.40 2 208 80.7 2.58 3 288 130.7 2.20 4 552 95.1 5.80 5 104 58.3 1.78 M E A N 261 94.7 2.75 S.E. 102 15.8 1.02 Station BB-H Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 64 30.0 2.13 2 16 70.6 0.23 " 3 24 91.9 0.26 4 64 131.4 0.49 5 72 81.0 0.89 M E A N 48 81.0 0.80 S.E. 15 21.2 0.46 Station FC-C Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 152 68.4 2.22 2 -3 544 76.6 7.10 J 4 160 57.0 2.81 5 48 104.0 0.46 M E A N 226 76.5 3.15 S.E. 126 11.6 1.63 Station FC-E Plant surf ace S# Oocytes area (cm2) Oocytes cm"2 1 496 181.5 2.73 2 56 79.7 0.70 3 392 89.0 4.40 4 208 89.0 2.34 5 168 87.3 1.92 M E A N 264 105.3 2.42 S.E. 102 24.7 0.78 Station FC-G Plant surface S# Oocytes area (cm2) Oocytes cm"2 1 448 116.4 3.85 2 112 40.9 2.74 3 4 5 208 71.4 2.91 M E A N 256 76.2 3.17 S.E. 100 21.9 0.34 142 Appendix II — Raw Data from Chapter 3 This appendix presents a portion of the raw data from the transplant experiments conducted from November 1997 to November 1998: (1) survivorship of Fucus gardneri, and (2) growth rates of F. gardneri. Survivorship Data The methods for the collection of these data are described in section 3.2.2 and the results are summarized, statisically analyzed and discussed in sections 3.3.2, 3.4.1, 3.4.2 and 3.4.3. Locations of sampling sites are shown in Figure 3.1. Abbreviations used in the data tables: R # = rock number. S.E. = Standard error of the mean. Dates are abbreviated as D D M M Y Y . e.g., 131197 = 13 November 1997. The values presented are the proportion of plants on a given rock which survived to that data collection. Growth Rate Data The methods for the collection of these data are described in section 3.2.2 and the results are summarized, statistically analyzed and discussed in sections 3.3.3, 3.4.1, 3.4.2 and 3.4.3. Locations of sampling sites are shown in Figure 3.1. Abbreviations used in the data tables: R # = rock number. S.E. = Standard error of the mean. Dates are abbreviated as D D M M Y Y . e.g., 131197 = 13 November 1997. The values presented are the mean growth rate of all plants on a given rock. These rates are calculated as: GR = (L t - L t . x)/T where GR is the growth rate in mm d"1, L t is the length of the plant at the time of data collection, L t_! is the length of the plant at the last data collection, and T is the number of days between the two data collections. 143 FUCUS GARDNERI SURVIVORSHIP — N O V E M B E R 1997 TRANSPLANT B B - 1 Date F C - 1 Date R# 131197 201197 271197 111297 080198 R# 131197 201197 271197 111297 080198 1 1.00 1.00 1.00 0.83 0.00 1 1.00 1.00 1.00 1.00 1.00 4 1.00 1.00 1.00 0.50 0.00 4 1.00 1.00 1.00 1.00 0.83 9 1.00 1.00 1.00 1.00 0.00 9 1.00 1,00 1.00 1.00 1.00 10 1.00 1.00 1.00 1.00 0.17 10 1.00 1.00 0.83 0.83 0.83 13 0.83 0.83 0.83 0.50 0.00 13 1.00 1.00 1.00 1.00 1.00 15 1.00 0.83 0.83 0.33 0.00 15 1.00 1.00 1.00 1.00 1.00 16 1.00 1.00 1.00 0.83 0.00 16 1.00 1.00 1.00 1.00 1.00 18 1.00 0.83 0.83 0.33 0.00 18 1.00 1.00 0.83 0.83 0.83 19 1.00 1.00 1.00 0.83 0.00 19 1.00 1.00 1.00 1.00 0.83 20 1.00 1.00 1.00 1.00 0.00 20 1.00 1.00 1.00 0.83 0.83 Mean 0.983 0.950 0.950 0.717 0.017 Mean 1.000 1.000 0.967 0.950 0.917 S.E. 0.017 0.025 0.025 0.086 0.017 S.E. 0.000 0.000 0.022 0.025 0.028 B B - 2 Date F C - 2 Date R# 131197 201197 271197 111297 080198 R# 131197 201197 271197 111297 080198 2 0.83 0.83 0.83 0.00 0.00 2 1.00 1.00 1.00 1.00 1.00 3 1.00 1.00 1.00 0.00 0.00 3 0.83 0.83 0.50 0.50 0.50 5 1.00 1.00 0.17 0.00 0.00 5 0.83 0.83- 0.83 0.83 0.83 6 1.00 1.00 0.83 0.00 0.00 6 1.00 1.00 1.00 1.00 1.00 7 1.00 1.00 1.00 0.00 0.00 7 1.00 1.00 0.83 0.83 0.50 8 1.00 1.00 1.00 0.00 0.00 8 0.83 0.83 0.67 0.67 0.67 11 1.00 1.00 0.83 0.00 0.00 11 0.83 0.83 0.83 0.83 0.83 12 1.00 0.83 0.67 0.00 0.00 12 1.00 1.00 1.00 1.00 1.00 14 1.00 1.00 1.00 0.17 0.00 14 1.00 1.00 1.00 1.00 0.50 17 1.00 1.00 1.00 1.00 0.00 17 0.83 0.83 0.83 0.83 0.83 Mean 0.983 0.967 0.833 0.117 0.000 Mean 0.917 0.917 0.850 0.850 0.767 S.E. 0.017 0.022 0.082 0.100 0.000 S.E. 0.028 0.028 0.052 0.052 0.067 FUCUS GARDNERI SURVIVORSHIP — F E B R U A R Y 1998 TRANSPLANT B B - 3 Date F C - 3 Date R# 090398 210398 040498 R# 090398 210398 040498 1 1.00 0.75 0.50 1 1.00 1.00 1.00 2 1.00 1.00 0.50 2 1.00 1.00 1.00 3 0.50 0.25 0.25 3 1.00 1.00 1.00 4 1.00 0.50 0.25 4 1.00 1.00 1.00 5 0.75 0.75 0.50 5 1.00 1.00 1.00 6 1.00 0.75 0.50 6 1.00 1.00 1.00 Mean 0.875 0.667 0.417 Mean 1.000 1.000 1.000 S.E. 0.085 0.105 0.053 S.E. 0.000 0.000 0.000 FUCUS GARDNERI SURVIVORSHIP — JUNE 1998 TRANSPLANT BB-4 Date R# 290698 100798 200798 270798 040898 130898 250898 2 0.83 0.33 0.33 0.33 0.17 0.17 0.00 6 0.67 0.67 0.67 0.67 0.50 0.33 * 8 1.00 0.83 0.67 0.67 0.50 0.33 0.17 9 1.00 0.33 0.33 0.17 0.17 6.17 0.00 10 0.67 0.33 0.33 0.33 0.17 0.17 0.00 12 1.00 0.83 0.83 0.83 0.83 0.67 0.50 14 0.83 0.83 0.83 0.67 0.50 0.50 0.50 15 0.83 0.83 0.83 0.83 0.67 0.50 0.33 16 0.83 0.50 0.50 0.50 0.33 0.33 * 17 0.67 0.67 0.67 0.67 0.50 0.50 0.33 Mean 0.833 0.617 0.600 0.567 0.433 0.367 0.229 S.E. 0.043 0.070 0.067 0.071 0.071 0.054 0.069 F C - 4 Date R# 290698 100798 200798 270798 040898 130898 250898 2 1.00 1.00 1.00 0.83 0.50 0.17 0.00 6 0.83 0.33 0.17 0.00 0.00 0.00 0.00 8 1.00 0.40 0.40 0.20 0.00 0.00 0.00 9 0.67 0.40 0.40 0.00 0.00 0.00 0.00 10 0.67 0.33 0.33 0.00 0.00 0.00 0.00 12 1.00 0.83 0.80 0.80 0.40 0.20 0.00 14 1.00 0.67 0.50 0.25 0.25 0.25 0.25 15 1.00 1.00 1.00 0.80 0.60 0.20 0.00 16 1.00 0.80 0.60 0.40 0.40 0.00 0.00 17 1.00 1.00 0.50 0.33 0.33 0.33 0.17 Mean 0.917 0.677 0.570 0.362 0.248 0.115 0.042 S.E. 0.045 0.091 0.089 0.107 0.074 0.041 0.028 * data were not collected — there was a large log on top of these two rocks 145 FUCUS GARDNERI SURVIVORSHIP — JUNE 1998 TRANSPLANT BB-5 Date R# 290698 100798 200798 270798 040898 130898 250898 1 0.83 0.83 0.83 0.83 0.67 0.67 0.33 3 1.00 0.83 0.67 0.60 0.00 0.00 0.00 4 1.00 1.00 1.00 1.00 1.00 0.75 0.75 5 1.00 0.80 0.60 0.60 0.20 0':20 0.20 7 1.00 0.83 0.83 0.60 0.50 0.00 0.00 11 1.00 1.00 0.33 0.20 0.17 0.00 0.00 13 0.83 0.83 0.83 0.83 0.83 0.33 0.17 18 1.00 1.00 1.00 0.80 0.50 0.33 0.17 19 1.00 1.00 0.80 0.60 0.40 0.20 0.20 20 1.00 1.00 1.00 1.00 0.60 0.60 0.60 Mean 0.967 0.913 0.790 0.707 0.487 0.308 0.242 S.E. 0.022 0.029 0.066 0.076 0.098 0.089 0.081 F C - 5 Date R# 290698 100798 200798 270798 040898 130898 250898 1 1.00 0.83 0.50 0.17 0.17 0.17 0.00 3 1.00 0.83 0.80 0.17 0.00 0.00 0.00 4 0.83 0.50 0.50 0.17 0.00 0.00 0.00 5 1.00 0.83 0.60 0.00 0.00 0.00 0.00 7 0.83 0.17 0.00 0.00 0.00 0.00 0.00 11 1.00 0.50 0.33 0.00 0.00 0.00 0.00 13 0.83 0.50 0.50 0.17 0.20 0.20 0.00 18 1.00 1.00 0.50 0.00 0.00 0.00 0.00 19 1.00 0.80 0.80 0.40 0.20 0.20 0.00 20 1.00 0.67 0.50 0.00 0.00 0.00 0.00 Mean 0.950 0.663 0.503 0.107 0.057 0.057 0.000 S.E. 0.025 0.078 0.072 0.042 0.029 0.029 0.000 146 FUCUS GARDNERI SURVIVORSHIP — AUGUST 1998 TRANSPLANT BB-6 Date R# 140998 230998 031098 101098 231098 2 0.83 0.83 0.83 0.83 0.67 3 0.67 0.33 0.33 0.33 0.00 4 0.67 0.33 0.33 0.17 0.17 7. 0.83 0.83 0.50 — — 8 0.50 0.50 0.50 0.50 0.33 9 0.83 0.67 0.67 0.00 0.33 13 0.83 0.83 0.40 0.40 0.40 15 0.67 0.50 0.50 0.50 0.33 16 1.00 0.83 0.83 0.83 0.67 17 0.67 0.80 0.40 0.20 0.00 Mean 0.750 0.647 0.530 0.419 0.322 S.E. 0.045 0.067 0.059 0.090 0.077 FC-6 Date R# 140998 230998 031098 101098 231098 2 0.83 0.83 0.50 0.50 0.50 3 0.67 0.67 0.33 0.00 0.00 4 0.83 0.83 0.83 0.67 0.50 7 o 1.00 0.83 0.83 0.33 0.33 o 9 0.83 0.67 0.50 0.50 0.33 13 1.00 0.83 0.67 0.67 0.67 15 0.50 0.50 0.17 0.17 0.17 16 0.50 0.33 0.00 0.00 0.00 17 0.67 0.50 0.50 0.50 0.50 Mean 0.759 0.667 0.481 0.370 0.333 S.E. 0.060 0.059 0.089 0.082 0.075 — : no data due to complete mortality of plants on these rocks 147 FUCUS GARDNERI SURVIVORSHIP — AUGUST 1998 TRANSPLANT B B - 7 Date R# 140998 230998 031098 101098 231098 1 0.67 0.67 0.67 0.50 0.50 5 1.00 0.83 0.33 0.33 0.17 6 0.83 0.67 0.33 0.17 0.17 10 1.00 0.67 0.50 0.33 0.33 11 0.67 0.50 0.33 0.17 0.17 12 0.83 0.67 0.67 0.67 0.67 14 0.83 0.83 0.67 0.20 0.20 18 1.00 0.67 0.50 0.50 0.50 19 0.83 0.83 0.67 0.67 0.50 20 0.83 0.67 0.67 0.50 0.17 Mean 0.850 0.700 0.533 0.403 0.337 S.E. 0.039 0.033 0.048 0.061 0.060 F C - 7 Date R# 140998 230998 031098 101098 231098 1 0.83 0.83 0.50 0.33 0.33 5 0.33 0.33 0.33 0.17 0.17 6 0.50 0.33 0.33 0.33 0.33 10 0.83 0.83 0.83 0.83 0.67 11 0.83 0.67 0.67 0.60 0.60 12 0.83 0.67 0.67 0.80 0.60 14 0.67 0.33 0.40 0.40 0.00 18 0.83 0.83 0.83 0.83 0.67 19 0.50 0.50 0.50 0.50 0.50 20 0.67 0.50 0.50 0.50 0.17 Mean 0.683 0.583 0.557 0.530 0.403 S.E. 0.058 0.067 0.059 0.074 0.075 148 FUCUS GARDNERI G R O W T H R A T E — N O V E M B E R 1997 TRANSPLANT B B - 1 Date F C - 1 Date R# 131197 201197 271197 111297 R# 131197 201197 .271197 111297 1 0.12 -0.36 0.12 -0.30 1 1.10 0.81 1.21 0.29 4 -0.98 0.26 -0.17 -0.46 4 -0.07 0.40 0.81 0.31 9 -0.43 0.00 -0.43 0.07 9 -0.29 1.31 0.71 0.48 10 0.02 -0.57 0.95 -0.39 10 0.86 0.33 0.54 0.31 13 0.11 0.26 -0.26 -0.54 13 0.71 -0.05 -0.17 0.38 15 -0.26 -0.49 0.11 -0.64 15 -0.02 0.69 0.00 0.35 16 0.07 -0.60 0.31 -0.79 16 0.45 0.12 1.36 0.27 18 0.02 -0.26 0.43 -1.21 18 0.69 0,57 0.43 0.43 19 -0.19 0.19 0.10 -0.96 19 0.43 0.69 0.64 0.25 20 -0.60 -0.12 -0.24 -0.86 20 0.45 0.23 0.74 0.17 Mean -0.210 -0.168 0.093 -0.609 Mean 0.431 0.511 0.628 0.324 S.E. 0.115 0.106 0.128 0.116 S.E. 0.139 0.124 0.149 0.028 B B - 2 Date F C - 2 Date R# 131197 201197 271197 111297 R# 131197 201197 271197 111297 2 0.17 -0.09 -0.76 — 2 1.07 0.81 0.86 0.43 3 0.02 0.14 -0.76 — 3 0.00 1.20 0.86 0.24 5 -0.24 0.05 -0.52 — 5 0.71 0.77 0.46 0.17 6 0.00 -0.14 -0.37 — 6 -0.52 1.83 . : 0.74 0.13 7 -0.43 -0.19 -0.86 — 7 -0.79 1.68 0.49 0.33 8 -0.29 0.40 -0.74 — 8 0.40 0.51 0.57 0.11 11 -0.40 0.37 -0.36 — 11 0.21 1.57 0.71 0.39 12 -0.45 0.52 -0.80 — 12 -0.02 0.90 0.83 0.30 14 -0.43 -0.20 -0.23 -0.64 14 0.23 0.26 0.26 0.01 17 -0.02 -1.38 -1.00 -1.57 17 -0.06 1.29 0.09 0.27 Mean -0.207 -0.052 -0.640 -1.107 Mean 0.124 1.083 0.586 0.237 S.E. 0.073 0.169 0.080 0.464 S.E. 0.172 0.163 0.084 0.042 — : no data due to complete mortality of plants on these rocks 149 FUCUS GARDNERI G R O W T H R A T E — JUNE 1998 TRANSPLANT B B - 4 Date R# 290698 100798 200798 270798 040898 130898 250898 2 0.25 0.14 -0.60 -0.36 -0.38 0.22 — 6 0.18 0.50 -0.23 -0.64 -0.29 0.72 — 8 0.14 -0.04 -0.88 -0.54 0.00 -0.28 -0.58 9 -0.04 -0.14 0.05 -1.71 -0.38 0.33 — 10 0.26 -0.18 -0.80 -0.43 -0.38 -0.44 — 12 0.03 -0.16 -0.56 -0.34 -0.50 -0.31 -0.86 14 0.38 -0.42 -0.88 -0.29 -0.46 0.15 -0.14 15 -0.25 0.25 -0.80 -0.29 0.22 -0.07 -0.33 16 0.55 0.06 -0.80 -2.33 1.88 -0.17 — 17 -0.07 0.91 -0.58 -0.96 0.25 -0.30 -0.29 Mean 0.142 0.092 -0.607 -0.789 -0.003 -0.014 -0.442 S.E. 0.074 0.122 0.097 0.221 0.226 0.115 0.127 F C - 4 Date R# 290698 100798 200798 270798 040898 130898 250898 2 0.50 0.77 0.25 0.43 0.50 0.78 — 6 0.78 0.32 -3.10 — — — — 8 0.73 1.82 0.50 2.14 -0.63 — — 9 0.68 0.55 0.65 — — — — 10 0.16 1.00 0.65 — — • — — 12 0.42 0.62 0.72 0.25 0.50 -0.89 — 14 0.48 0.36 0.97 0.57 0.63 0.00 — 15 0.32 0.45 0.22 0.62 0.79 -1.22 — 16 0.65 0.58 0.50 0.36 0.19 — — 17 0.43 0.94 -0.03 0.21 1.00 -0.89 0.50 Mean 0.514 0.741 0.133 0.655 0.426 -0.444 0.500 S.E. 0.061 0.139 0.371 0.254 0.200 0.367 n/a — : no data due to complete mortality of plants on these rocks 150 FUCUS GARDNERI G R O W T H R A T E — JUNE 1998 TRANSPLANT BB-5 Date R# 290698 100798 200798 270798 040898 130898 250898 1 0.17 0.47 -0.08 0.40 -0.03 0.47 0.33 3 0.07 0.25 0.12 0.29 — — — 4 0.52 0.24 0.22 -1.32 0.44 -0.89 0.50 5 0.57 0.18 0.43 0.05 -0.25 o;67 0.08 7 0.20 0.44 0.22 -0.11 0.29 — — 11 -0.54 0.80 -0.60 0.57 2.75 — — 13 0.07 0.65 0.00 -0.21 -0.10 -1.94 -3.67 18 0.30 0.24 0.45 0.09 0.29 0.00 0.17 19 0.23 0.50 -0.20 0.67 0.25 -2.11 -1.92 20 0.29 0.55 -0.18 0.29 -0.19 0.00 -0.44 Mean 0.189 0.432 0.039 0.070 0.384 -0.544 -0.706 S.E. 0.096 0.064 0.102 0.178 0.307 0.426 0.582 F C - 5 Date R# 290698 100798 200798 270798 040898 130898 250898 1 0.11 1.16 0.53 0.43 0.50 -0.11 — 3 0.07 0.71 0.33 -0.14 — — — 4 -0.20 1.15 0.73 0.43 — — — 5 0.10 0.49 0.73 — — — — 7 0.17 0.82 — — — — 11 0.30 0.82 -0.05 — — — — 13 0.03 0.21 0.70 -0.29 0.50 1.11 — 18 -0.46 1.43 0.80 — — — — 19 0.01 0.95 0.13 0.29 — — — 20 -0.18 1.84 0.53 — — — — Mean -0.006 0.958 0.493 0.143 0.500 0.500 n/a S.E. 0.069 0.148 0.100 0.150 0.000 0.611 n/a — : no data due to complete mortality of plants on these rocks 151 FUCUS GARDNERI G R O W T H R A T E — AUGUST 1998 TRANSPLANT B B - 6 Date R# 240898 030998 140998 230998 031098 101098 231098 181198 2 0.18 0.23 0.36 0.16 0.48 0.29 0.42 0.26 3 0.29 0.50 0.50 -0.50 -0.15 0.07 —- — 4 0.11 0.35 0.48 -1.22 0.25 0.43 0.69 0.19 7 0.03 0.50 -0.16 0.16 -0.37 — —- — 8 0.50 0.40 0.48 -0.41 1.00 0.05 0.31 -0.23 9 -0.01 0.25 0.44 -0.08 0.45 — 0.05 -0.15 13 0.58 0.65 -0.27 0.11 -0.25 0.79 -0.46 0.19 15 0.50 0.58 -0.20 -0.52 -0.30 1.05 0.15 -0.42 16 -0.72 0.10 0.42 0.11 0.75 0.30 0.29 0.25 17 0.50 0.00 0.25 0.18 -0.20 -0.43 ' — — Mean 0.196 0.356 0.230 -0.202 0.166 0.317 0.208 0.012 S.E. 0.123 0.067 0.100 0.145 0.154 0.161 0.136 0.105 F C - 6 Date R# 240898 030998 140998 230998 031098 101098 231098 181198 2 0.31 0.10 0.18 -0.04 0.33 -1.00 -0.21 — 3 0.27 -0.80 -0.14 -0.11 0.20 — — — 4 0.70 0.83 0.15 0.49 0.60 -0.43 0.41 — 7 0.81 0.70 0.23 0.13 0.24 0.14 0.62 — 8 0.35 0.45 9 0.13 0.37 0.33 0.53 0.30 0.67 0.08 0.54 13 1.25 — 0.67 0.09 0.33 0.36 0.40 -4.02 15 -0.25 0.55 0.12 0.30 0.00 0.57 0.15 — 16 0.18 0.07 -0.24 0.28 • — — — 17 — 0.40 -0.05 -1.67 0.00 0.29 — — Mean 0.417 0.297 0.138 -0.001 0.250 0.085 0.243 -1.740 S.E. 0.146 0.160 0.090 0.220 0.069 0.225 0.120 2.279 — : no data due to complete mortality of plants on these rocks. 152 FUCUS GARDNERI G R O W T H R A T E — AUGUST 1998 TRANSPLANT B B - 7 Date R# 240898 030998 140998 230998 031098 101098 231098 181198 1 -0.14 1.25 0.09 0.17 0.07 0.00 0.10 0.23 5 0.79 -0.50 0.23 0.02 0.33 0.33 . — — 6 0.44 0.00 0.49 0.28 -0.10 0.00 0.15 10 0.21 -0.05 0.23 0.19 0.63 -0.86 -0.04 -0.75 11 0.19 -0.43 0.23 0.17 0.65 0.29. 0.85 0.42 12 0.00 — 0.36 0.39 0.68 0.07 1,06 0.18 14 -0.04 0.50 0.29 -0.29 -0.05 0.00 -0.15 0.15 18 -0.08 0.53 0.15 -0.47 0.20 0.43 0.08 — 19 0.21 0.20 0.40 -0.33 -0.35 -0.38 0.23 -0.33 20 0.89 0.30 0.67 0.53 0.05 0.24 0.08 0.12 Mean 0.247 0.199 0.314 0.066 0.212 0.012 0.261 0.003 S.E. 0.113 0.179 0.055 0.104 0.112 0.121 0.137 0.152 F C - 7 Date R# 240898 030998 140998 230998 031098 101098 231098 181198 1 0.38 -0.30 0.02 0.58 -0.20 0.14 0.50 — 5 0.08 1.3.0 0.41 -0.61 0.20 -1.43 1.00 -0.19 6 0.36 -0.10 0.18 1.17 -0.15 -0.71 1.00 — 10 0.54 0.95 0.44 0.29 0.34 -0.17 0.38 — 11 0.80 0.27 0.51 0.42 0.63 0.33 0.97 0.44 12 0.27 0.75 0.22 0.14 0.27 -0.22 -0.03 -0.21 14 0.30 0.30 0.18 -0.39 0.45 -0.14 — — 18 0.46 — 0.20 0.76 0.42 0.23 0.63 0.42 19 0.28 0.77 0.33 0.11 -0.03 0.00 -0.23 — 20 0.23 0.47 0.20 0.74 0.43 0.05 0.62 0.46 Mean 0.370 0.489 0.269 0.320 0.235 -0.193 0.539 0.185 S.E. 0.062 0.170 0.047 0.170 0.088 0.165 0.147 0.158 — : no data due to complete mortality of plants on these rocks. 153 

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